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Hey tat I. A CaTaLoGuE OF THE SPECIES OF MOSSES FOUND UP TO THE PRESENT ¥
TIME ON THE NorTHWEST COAST OF THE UNITED STATES, AND Bere
ESPECIALLY IN CALIFORNIA...........- Sete Oe By Leo Lesquereux. Fe
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(Published January, 1868.)
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ERRATA.

Page 6, Ist line, for “‘ Wusiacese” read ‘* Weissiacew.’”
«6, 9th line, from top for ‘ Weisia” read ‘“‘ Weissia.”
Be east wp « « «© © Breyerianum” read ‘«Brewerianum.”
«15; 16th «« “© heterostychum ” read ‘‘ heterostichum.”
“ 96, 9th — “4 @thyphylla” read “ ithyphylla.”’
** 32, 10th, 18th, 19th - «© « «Breyerianum” read ‘‘ Brewerianum.” —
ot 3a bottom for ‘* Eurynchum ” read “«Burhynehum.”’
mee so, ; top ‘“ ‘‘commulatum” read “ commutatum.
‘© 37, 10th ; “ ‘<“Rhitidium” read “ Rhytidiam.” —

MEMOIRS

PRESENTED TO THE CALIFORNIA ACADEMY OF SCIENCES.

VOLUME I.

1. A Catalogue of the Species of Mosses found, up to the present time, on the North-
West Coast of the United States, and especially in California.

By Leo LeEsQuerEux.
[Presented Dec. 2d, 1867. ]

RELIMINARY NOTICE. This catalogue comprises all the species of Mosses

found, up to the present time, so far as I know, on the western coast of North Amer-
ica, from the Mexican boundary to British Columbia. Most of the species are from
California, gathered there by Dr. Bigelow, and Messrs. W. H. Brewer and H. N.
Bolander of the State Geological Survey. It is to the last-named gentleman that the
Bryology of the State of California is chiefly indebted. To render this list more com-
plete, I have admitted into it the species collected by Dr. Lyall, in Oregon, on the
Northwest Boundary Survey, along the 49th parallel, from the western base of the
Rocky Mountains to Vancouver Island included.

I have not attempted, in this enumeration, to suggest any peculiar system of
nomenclature or classification ; but have merely followed what I considered the most
natural grouping, copied from the ‘ Bryologia Europea,” and from Schimper’s Synop-
sis of European Mosses. New systems of classification are now originated in bryologic
science, as it appears to me, for the mere satisfaction of their authors, and without the
slightest advantage to science. Indeed, these new systems have had a most baneful
influence on the study of this most interesting department of botanical science. A
discussion of their merits—or rather, their demerits—will, especially in a catalogue
like this, be useless.

Some species of mosses, considered as new and distinct by authors of high repu-
tation, are included in this catalogue as mere varieties. The flora of California, in all
its departments, is liable to great local varieties, according to the peculiar atmospheric
and chemical conditions to which it is subjected. The more the phenerogamic flora of

MEM. CAL. ACAD. SCI. VOL. I. 1 Jan. 1868.

2 LESQUEREUX — CATALOGUE

that region is studied, the more the number of species is diminished. Having had access
to a very large number of specimens—thanks to Mr. Bolander’s zeal in collecting—I
have been able to compare forms from different stations, and to see characters con-
sidered by others as indicating difference of species, blended together, sometimes
even on the same specimen.

To prevent repetition of references, I will merely say that the species mentioned
from Mr. Mitten, have been published in his Bryologia of the 49th Parallel of Latitude,
in the Proceedings of the Linn. Soc. of London, Bot., Vol. VIII, —; the species from
Hampe in a pamphlet, Musei Californici Novi, 1860 ; those of Muller, not included in
his synopsis, in the Bot. Zeitung, No. 40; those of Sullivant in the Reports of Lieut. A. W.
Whipple's and Chs. Wilkes’ U.S. Expeditions ; and those of myself in the Transactions of
the Amer Philos. Soe. of Philadelphia, Vol.15. A few species given with short diagnoses
in the Musei Exsiecati Americant, Ed. 2d., by Sullivant and Lesquereux, are here copied
from this work, which has been published in a limited number of copies, and which is
not therefore accessible to every bryologist.

As this enumeration is intended as a representation of our actual knowledge of
the Bryology of the west coast of the United States, I have carefully mentioned the
habitat of the species, and the names of the botanists who have gathered them.

Cotumeus, O., 30th March, 1867.

(2)

OF PACIFIC COAST MOSSES. s)

COL TAS k1© Gow, Ei:

MUSCI.

SPHAGNES.

SPHAGNUM, Dill.
]. §S. acutifolium, Ehrh.
Hab. Sierra Nevada ; Upper Tuolumne Canon ; foot of Mt. Dana, 8,000 to
9,000 feet, Bol.
2. §. fimbriatum, Wills.
Hab. by a brook, at about 11,000 feet on Mt. Brewer, Brew.

S. rigidum, Schp.
Hab. Yosemite Valley, hillsides, in the spray of the Vernal Falls, Bol.

(3%)

Var. B. compactum, Schp.
Hab. rivulets, foot of Mt. Dana, Bol.
4. §. squarrulosum, Lsqx Ms.
S. squarrosum, var. B. Schp.
Hab. head of William Lake, near Lassen’s Peak, about 5,000 ft., Brew.

The first Sphagnum found in California. On this species Prof. Brewer re-
marks: ‘‘It forms a large bog, and is the only Sphagnum I have yet seen in the State
‘‘in more than 13,000 miles of peregrinations.”

The true value of this species is still uncertain. In his remarkable Memoire
pour servir a Uhistoire des Sphaignes, (1857) W. P. Schimper mentions it as possibly a
small marked form of S. sguarrosum, Pers., and in his Synopsis Muscorum Europeorum,
(1860) he joins it to this last species as var. B. The far different facies, long, slender
branches, similar to those of some forms of Sphagnum cuspidatum, Ehrh.; the longer,
narrower leaves, also indicate a specific difference. By details of analysis, S. squarru-
losum shows indeed in the cross-sections of the leaves the same kind of reticulation as S.
squarrosum, viz: large, round, primary cells with intermediate, elongated, narrow oval
ones. But thisis the only affinity; and even the cells are narrower in S. sguarrulosum,
and actordingly the leaves are proportionally thinner. The nearest relative to this
still uncertain species is S. rigidum, Schp. One of its varieties, from the Racoon Mts.
of Alabama, with branch leaves contracted above the middle and the point reflexed, is

>

(3)

4 LESQUEREUX — CATALOGUE

much like it, our species differing merely by more acute branch leaves and a broader
reticulation of the stem leaves—differences of little imporfance. Therefore if 8.
squarrulosum should be united to any species asa variety, it would rather go with S.
rigidum. But it is better to let the species stand as it is till its fruit is known, and it
can be thus thoroughly studied. It is right to remark also that it was originally found
in the bogs of the Jura Mountains, of Switzerland, where S. sguarrosum has never
been seen. For this species more generally inhabits the bogs and streams of primitive
rocks, and not the limestone.

5. §. subsecundum, Nees & Hrnsch.
Hab. wet meadows, Mariposa Big Trees, God.
Var. B. longifolium, ramis attenuatis longioribus, foliis angustis lanccolatis,
elongatis.
Hab. Mendocino City, swamps, Bol.
6. §. auriculatum, Schp.
Hab. in swamps, 8,000 to 9,000 ft., between forks of King’s River, Brew. In
pools, near Mendocino, Bol.

The form from Mendocino has a peculiar facies, which is not seen in any European
specimens. The leaves are longer and narrower, the branches more slender and at-
tenuated, similar to those of the floating large varieties of S. cuspidatum. But it has
the lanceolate-pointed shape of the leaves, especially the loosely reticulated branch
leaves, and the stem leaves with fibrillose stems filled with very numerous round small
pores, a character ascribed by Schimper to this species only. The differences may be
considered as mere local variations. In the California moss the auricles of the stem
leaves are not more developed than they are generally in S. subsecundum.

7. S. eymbifolium, Ehrh.
Hab. swamps near Mendocino City, Gol.

PHASCACE AS.

EPHEMERUM, Hampe.

8. E. serratum, Hainpe.
Hab. on the ground in fields and meadows, Mission Dolores, Bol.

SPH.®RANGIUM, Schp.
9. S. muticum, Schp.
Hab. with the former, Pol.

This species has not yet been found, to my knowledge, in the Atlantic States,
where it is represented by an intermediate form equally referable to both 8. muticum
and S. triquetrum, Schp. It is S. triquetrum, var. No. 31, of the Musci Exsice. Amer.,
Ed. 2d.

(4)

OF PACIFIC COAST MOSSES. 3)

PHASCUM, Schreb.

10. P. cuspidatum, Schreb.
Hab. with the former, Bol.; with var. piliferum, Fort Colville, Lyall.
11. P. bryoides, Dicks.
Var. B. piliferum, Schp. Synop.
Hab. south side of hills of Oakland, ol.
A few specimens only were sent, mixed with Anacalypta Starkcana and Weista
virtdula.

PLEURIDIUM, Brid.

12. P. subulatum, Brch. & Schp.
Hab. ditches and dry hills, San Rafael, Bol.
By the form of the capsule and of the leaves, the California moss is referable to
P. alternifolium, Brch. and Schp. Even the two years’ old plants bear slender inno-
yatine branches with short distant leaves, similar to the flagelliform branches of P.
alternifolium. Yet, the inflorescence is that of P. subulatum, the anthers being free in
the axils of the upper leaves.

BRUCHIA, Schweer.

13. B. Bolanderi, spec. nov. Monoica, dense gregaria, caespitulosa, pallide virescens.
Caulis brevis, vix linearis, basi tantum radiculosus, simplex. Folia caulina
remota, minuta, lanceolata ; comantia erecto-aperta lanceolata, brevi subu-
lata, costa plana sub apice obscure et obtuse serrulato desinens, reti basilari
quadrato-elongato, apiciali angustiore distincto ; folia perichzetialia late ovata,
amplectentia tubulosa, externa breviter acuminata, interna lanceolato-subu-
lato erecta, caulinis duplo longiora, reti laxiore. Capsula in pedicello robusto,
semipollicari subflexuoso erecta vel subobliqua, e collo elongato pallide viridi
oblonga angusta, virescens, in rostrum rectum pallidumque producta ; calyp-
tra tertiam partem capsule obtegens, basi laciniata. Inflorescentia vere
monoica sed primo intuitu dioica videtur, caule primario repente, radiculoso
masculas, femineasque gemmas radiculosas gestante. Flores masculi cras-
siores, foliis perigonialibus internis brunneis, ovato lanceolatis acutis, obsolete
nervosis.

Hab. near Big Tree Grove, Mariposa ; Westfall’s Meadow, 8,000 ft. Bol.

It is not easy to separate this species from Vogesiaca, Schw. ‘The inflorescence
is the same ; at least, male and female buds bear in their position the same relation.
The long pedicel, the form and color of the capsule, are also similar. The difference
is essentially in the shorter stems, shorter leaves, the shorter beak, and especially the
longer collum of the capsule. The pedicel is thicker than in the European species,
slightly flexuous when dry, straight when moistened. The color of the plants is of a
pleasant pale green,

LESQUEREUX — CATALOGUE

WUSIACE 4.
GY MNOSTOMUM, Brch. and Schp.

G. caleareum, Nees and Hrnsch.
Var. perpusillum, Sulliv. Whipple’s Exped. p. 189.
Hab. on clayey soil, near San Francisco, Big.
G. curvirostrum, Hedw.
Hab. on wet rocks, Leroux Springs, base of San Francisco Mts.;. Mogollon
Ridge, Bry.

WEISA, Hedw.
W. viridula, Brid.
Hab. on the ground, at and around San Francisco, Gol., Big.
Var. capsulis longioribus, ovato-cilindricis, regulariter plicato-striatis, viridibus.
Hab. on the ground, San Francisco, Bol.
W. cirrhata, Hedw.
Hab. region of the Redwoods, common, especially on burnt and decayed wood ;
Big Trees, etc., Bol., Big. ; Vancouver Island, Lyall; N. W. America,
Douglas.

W. crispula, Hedvw.
Hab. Galton Mts., near Fort Colville, Lyall.

DICRANUM, Auct.

21. CYNODONTIUM, Schp. Syn.

D. polycarpum, Ehrh.
Hab. Cascade Mts., Lyall.
D. virens, Hedw.
Var. G. serratum. Schp. syn.
Hab. Tuolumne Cation and Mono Pass, 9,000 ft., borders of snow-water
streams, Bol.; Galton Mts., Lyall, alt. 6,000 to 7,000 ft.

D. crispum, Hedw.
Hab. Galton Mts., Lyall.

22. DICRANELLA, Schp.
D. varium, Hedw.
Hab. on perpendicular sandstone rocks, watered by springs, near the bay of
San Francisco, Gol.

OF PACIFIC COAST MOSSES.

25. D. subulatum, Hedw.
Hab. foot of Mt. Dana, 8,000 to 10,000 ft., borders of ditches and stream-
lets, Bol. ; Galton Mts., Lyall.

24. D. heteromallum, Hedw.
Hab. slopes of a ditch cut through a bog, edge of the redwoods near Big

River City, Bol. ; Cascade Mts., Lyall.

23. DICRANUM, Schp.
D. strictum, Schl.
Hab. coniferous trees, Devil’s Canon, Forest Hill, Lol.; Fort Colville, Lyall.
26. D. albicans, Brch. and Schp.
Hab. N. W. America, Douglas.
27. D. fuscescens, Turn.
Hab. Redwoods, Bol.; Spokane River, Oregon, Wilkes’ U. S. Exp.; Galton
Mts., Lyall.

28. D.scoparium, Lin.
Hab. Oregon, Wilkes’ Hxp.; and a var. foliis vix denticulatis, in California? Bol.

bo
or

29. D. palustre, Lapyl.
Hab. Pend’ Oreille, Zyall ; Vancouver Island, Wood. Mr. Bolander has sent
it from Eureka, Humboldt County.

Var. foliis angustioribus, levibus, falcato-secundis, areolatione longiori. D.

Breverianum, Lsqx. in litter.
Hab. Deep Canon, Klamath River, Brew.

This variety is remarkable enough to be considered a proper species. Besides
the characters indicated above, it differs from D. palustre by its shorter stems, its
narrow, not at all, or scarcely undulate leaves, its more incurved, and more slender,
and longer capsule, the great quantity of male plants mtermixed in the cespites.
The male plants are slightly more slender than the fertile ones ; the flowers large, ter-
minal, or lateral by innovations ; the perigonial leaves broadly ovate, narrowed into a
pretty long point and nerved.

30. D. majus, Schweer.
Hab. Fort Discovery, Oregon, Wilkes’ U.S. Exped.

FISSIDENS, Hedw.

31. F.limbatus, Sulliv.

Hab. on shaded ground, near Oakland; common around San Francisco,
Bol., Big.

F. ventricosus, spec. nov. Monoicus, robustus, laxe lateque caespitosus, caespi-
tibus nigricantibus, innovationibus junioribus apice tantum atro-viridibus.
Caulis ultra pollicaris, e basi divisus, divisionibus simplicibus vel parce ramo-
sis. Folia multijuga, conferta, erecto-aperta, cultriformia, duplicatura di-

(7)

Oo
bo

8 LESQUEREUX — CATALOGUE

latata ad mediam laminam verticalem procedente, margine crasso, gibboso,
plerumque cum costa crassa in apiculum muticum confluente, raro sub apice
cum costa evanido. <Areolatio ovato-quadrata vel irregulariter polygona,
minuta. Flores utriusque sexus terminales ; folia perigonalia late ovata, sub
lamina verticali apiculiformi irregulariter dissecta ; folia pericheetialia con-
formia. Capsula in pedicello brevissimo, vix exserta, erecta, obovata, in
pedicello attenuata, obscure viridis. Peristomii dentes erecti latiores ; spor
ovales magnee.
Hab. on submerged rocks, Mendocino City, Bol.
By the ventricose or enlarged duplicature of its leaves, this species resembles
F. rufulus, Brch. & Schp. In our species the leaves are more obtuse, and generally
entirely surrounded by a thick margin like those of #. revularis of the same authors.
It is only in the upper leaves of new branches that the leaves appear similar to those
of F. rufulus. The division of the stems is generally from near the base and dichoto-
mous, sometimes the primary divisions being divided again near the top in short
branches. The axils of all the leaves bear tufts of radicles. The specimens sent to me
had but one capsule, already deoperculate.
33. F. adiantoides, Hedw.
Hab. Fort Colville, Lyall.
33a. F. grandifrons, Brid.
Hab. sea-shore, Mattole District, Humboldt Co.; rocks over which spring
water flows constantly, Gol., June, 1867.

POTTTACH A.

POTTIA, Ehrh.

34. P. subsessilis, Brch. & Schp.
Hab. Los Angeles, Lig.

35. BP. cavifolia, Ehrh.
Hab. Fort Colville, Lyall.
36. P.minutula, Breh. & Sehp.
Hab. Los Angeles, Big.
P. Heimii, Hedw.
Hab. ditches near the Soda Springs, alt. 9,000 ft., Bod.

ANACALYPTA, Roehl.

9

38. A. Starkeana, Nees & Hrnsch.
Hab. Mission Dolores, Oakland, ete., on clayey ground, Bol.

(8)

OF PACIFIC COAST MOSSES. 9

DIDYMODON, Iedw.

39. D. rubellus, Breh. & Schp.
Hab. Tuolumne Canon, near Soda Springs, 9,000 ft. ; foot of Mt. Dana, Bol.

DISTICHIUM, Brch. & Schp.

40. D. capillaceum, Breh. & Schp.
Hab. on shelving rocks, between Vernal and Nevada Falis, Yosemite Valley,
Bol., rare ; Cascade Mts., Lyall.

41. D. inelinatum, Hedw.
Ilab. Soda Springs, ol.

CERATODON, Brid.

42. C. purpureus, Brid.
Hab. California, common, Lol., Brew., Big. ; spray of Nevada Falls on rocks,
Bol. ; Vancouver Island, Lyall.
Var. B. xanthopus, Sulliv. & Lsqx.
Hab. on Sequoia sempervirens, extremely common; also, Big Trees, Bol.,
Big.

LEPTOTRICHUM, Hampe.

43. L. Schimperi, Spec. nov. Monoicum, subcespitosum, subsimplex, humile lu-
tescente-viride. Folia patentia, flexuosa, vel falcato-secunda, e basi ovata,
lanceolata, subulata, valde elongata, angusta, apice tantum subdenticulata, costa
tenui vix totam subulam occupante percursa.. Perichetii elongati folia late
amplectentia, interna tubulosa, in subulam longiorem attenuata. Capsula in
pedicello vix ultra pollicari lutescente, robusta, erecta, ovato-cylindrica
brunnea, operculo elongato conico recto vel subobliquo. Peristomi dentes
in membrana basilari lata fugaci pulcherrime aurantiaca vel rubella, gracil-
limi, pallide lutei filiformes, leves, versus basim unocrure diviso tripartiti
vel trabeculati. Annulus latior vix mediam partem membrane attinens.
Spore maxime.

Hab. Coast Range, Mendocino, Bol.

This species much resembles L. pallidum, Hmp. But, besides the difference in
the peristome, it is easily distinguished by its green color—the shorter, broader capsule
borne on a shorter, thicker pedicel, having a longer, dark-red operculum. The leaves,
also, of our species are much longer; their basilar reticulation larger, ete. The male
flowers are gemmiform, axillary, and their leaves large, ovate, short, lanceolate,
obtuse.

44. LL, flexicaule, Schweer.

Hab. Cascade Mts., Lyall.
B (9)

10 LESQUEREUX — CATALOGUE

TRICHOSTOMUM, Hedw.

45. T. rigidulum, Smith.
Hab. on rocks subject to overflow on Russian River, opposite Ukiah, Bol. ;
Vancouver Island, Lyall.
46. TL tophaceum, Brid.
Hab. limestone rocks, constantly watered by a spring, Ukiah City; Fort
Point, ete., Bol. ; Cajon Pass, By.
47. T. anomalum, Schp.
T. corniculatum, Wahl.
Hab. Oakland Cajion, Bol.; a few sterile stems mixed with Hypnwm leuco-
neurum ; Vancouver Island, Lyall; California, Coulter.

48. TT. flexipes, Brch. & Schp.

T. crassinerve, Hmp. Muse. Calif.
Hab. in shaded grounds and decayed trunks, Bol., Big., Bauer ; near San
Rafael, and also in the mountains of the Coast Range, Mendocino, Bol.

DESMATODON, Brid.

49. D. Californicus, Spec. nov.
D. nervosus, var. B. edentulus, Sulliv. & Lsqx., Muse. Exsice. Amer., 2d Ed.,
No. 121.
D. nervoso proximus, differt: caule breviore, foliis confertis, arctius imbri-
catis, brevi-acutis, vel nervo sub apice evanido obtusis, nervo medio inflato
latiori, peristomii dentibus albidis, capsulo annulato.
Hab. on decayed ground and old walls of clay, (adobe) San Francisco, Bol.

Though the differences between both species are slight, the presence of an
annulus, the broader, more inflated costa of the shorter generally obtuse leaves, appear
the specific characters of the California moss.

r

50. OD. latifolius, Hedw.
Hab. slopes of Tuolumne Cajon, Bol.
Var. B. mutieus, Breh. & Schp.

Hab. Mt. Dana, 11,000 to 11,500 feet ; Tuolumne Cafion, Bol. ; Silver Val-
ley, on exposed soil, 8,000 to 9,000 feet, Brew.; common in the mountains
along water courses, Cascade Mountains, Lyall.

51. D. systilius, Breh. & Schp.
Hab. foot of Mt. Dana, Bol.
52. D. Guepini, Brch. & Schp.
Hab. on the ground, bank of a run, Oakland, Pol.

OF PACIFIC COAST MOSSES. 1]

BARBULA, Hedw.

B. membranifolia, Hook.
Hab. dry ravines on William’s Fork of the Great Colorado, Lig.

on
Os

54. 8B. chloronotus, Schultz.
Hab. same as the last, Big.

55. 3B. fallax, Hedw.

Hab. Cajon Pass, Sierra Nevada ; also, near San Francisco, Lig.
56. B. subfallax, Muller, Bot. Zeit. No. 40, p. 338.

Hab. San José Valley, Lauer.

57. B. semitorta, Sulliv., Whipple’s Rep.
Hab. California, Bag.

58. B. brachyphylla, Sulliv., ibid.
Hab. California, big.

59. B. artocarpa, Lesqx.
Hab. Redwood Hills, Coast Range, Ukiah, ete., Bod.

60. B. vinealis, Brid.
Hab. near San Francisco, on the ground and stones, Lol., Bay.

61. B.rubiginosa, Mitt.
Hab. N. W. America, Douglas.

62. B. insulana? De Not.
Hab. California, Deechy.

A remarkable form, perhaps referable to this species, has been collected by
Mr. Bolander, on gravelly soil, near the Big Trees. It is subcespitose, with a robust
stem branching by innovations generally in two, opposite, from one-half to one-inch
long. Its color is brownish yellow ; the leaves when dry are appressed or slightly tor-
tuous, open-erect by humidity, loosely imbricated, ovate lanceolate, very entire, revo-
lute on the borders, with a strong excurrent nerve-forming point ; the basilar reticula-
tion is equilateral, elongated, pellucid ; in the upper part of the leaves, the areol
are opaque, minute and minutely papillose. The capsule borne on a short, straight,
reddish pedicel of half an inch long, is large, oblong, of a reddish color, with an erect
pointed operculum half the length of the capsule. The teeth of the peristome are
red, twice twisted on a short membrane; the annulus is double, pellucid or white ;
the male flowers terminal and thick. This form is more robust than any other of the
section of the Vineales, and is easily distinguished by its appressed, scarcely twisted
leaves, its thick, brown, reddish capsule on a proportionally short pedicel. The form
and areolation of the leaves are like those of B. vinealis ; but the peristome is twisted
twice and dark red. The forms of this section of Barbula found in California are so
numerous, that it will be necessary to reduce them to a few species, or to make a
monograph of them, embracing more than one dozen species. I have only mentioned

in this catalogue the most marked forms.
(11)

(=p)
(or)

72.

73.

B.

B.

LESQUEREUX — CATALOGUE

flexifolia, Hmp.
Hab. on sandy ground and boulders near the coast, San Francisco, Bol., Bauer.

virescens, Lesqx.
Hab. same as the former, Pol.

convoluta, Hedw.

Hab. gardens in San Francisco ; burnt logs, Ukiah, Bol. ; Oakland, Big.
cuneifolia, Brid.

Hab. clay soil, Oakland, God.
Wabhliana, Schultz.

Hab. Cajon Pass, Sierra Nevada, Big. ; Mt. Diablo, Gol.

Bolanderi, Lesqx.

Hab. on rocks, near San Francisco, Pol.

amplexa, Lesqx.

Hab. on sandstone around springs, in water, Lol.

marginata, Bryol. Eur.

Hab. on rocks near San Francisco, mixed with LB. flexifola, Gol.; on dry
rocks, Big. :

. B. brevipes, Spec. nov. Dioica, gregaria vel cespitulosa. Caulis brevissimus,

simplex. Folia in comam conferta, lingulata, vel oblongo-cuneiformia obtusa
margine revoluta, superne concavo-carinata, costa in pilum subleve exeunte
instructa, reticulatione basilari laxa, pellucida, quadrato-elongata, apicem
versus compacta obscura irregulariter rotundata, chlorophillosa, papillosa.
Capsula in pedicello longiori elongata, cylindrica, gracilis, subincurva, operculo
longe conico obtusiusculo. Peristomii dentes arcte convoluti, sanguinei,
in membrana longa quadrato-tessellata suffulti, Flos masculus in planta
graciliori terminalis foliis perigonialibus brevibus-latioribusque.

Hab. on mud walls, Mission Delores, in mats one inch in diameter or more ;

divide of the Russian River, ol.

This species is similar in appearance to Larbula marginata, but widely different
in its non-marginate leaves, and its capsule and peristome, which are syntrichial.

B.

B.

B.

B.

subulata, Lin.
Hab. Big Tree grove, Bol. ; dry ravines on Big William’s Fork of the Rio
Colorado, Big.
inermis, Brch.
Hab. base of mountains west of the Rio Colorado, Big. Sent also by
Bolander without label.
levipila, Brid.
Hab. railroad levees, Sacramento, Bol. ; Vancouver Island, Lyadl.
latifolia, Brid.
Hab. on Alnus viridis, borders of a creek near San Rafael, Bol.

OF PACIFIC COAST MOSSES. 13

76. B. ruralis, Hedw.
Hab. on rocks and trunks of trees from the plains to the mountains, very
common, Lol., Big., Brew., Lyall.
Var. gyantea, caule elongato, foliis e basi squarroso reflexis ; areolatione papil-
losa, papillis elongatis, furcatis ; costa dorso spinuloso-rugosa.
Hab. on moist rocks, Yosemite Valley, Bol.
. B. Muelleri, Brch.
Hab. same as the former, especially common in the plain, Biy., Bol., Brew., Lyall.

1
~I

GRIMMIACE/E.
GRIMMIA, Ebrh.

78. G. Scouleri, C. Mull.
‘coulerta aquatica, Hook.
Hab. on granite rocks in Merced River, at Clark’s, near the Big Trees, sterile,
Bol. ; in Columbia River, near Fort Colville, Lyall.
79. G. conferta, Funk.

Hab. on metamorphic rocks, Mt. Diablo, Bel.; rocks in caiions near Green
Valley, Brew.

80. G. apocarpa, Hedw.
Hab. on rocks, Yosemite Valley, Bol. ; crossing of the Colorado, on rocks,
Big.; N. W. America, Lyall, Douglas.
Var. rivularis, Breh. & Schp.
Hab. on wet rocks, Yosemite Valley, Bol.
Sl. G. pulvinata, Smith.
Hab. on metamorphic limestone-boulders, near Russian River, Ukiah ;
around Clear Lake, Bol.; rocks on Bill William’s Fork, Big.; Fort
Colville, Lyall.
82. G. contorta, Wahl.
G. uncinnata, Kaulf.
Hab. Big Tree grove, Bol.

I have received one specimen only with deoperculate capsule. The form of
the leaves, their reticulation, the black color of the plants, and the small, slightly
inclined capsule, mark the identity of this species.

83. G. Muhlenbeckii, Schp.
Hab. on a rock, specimen from Mt. Dana, 11,500 feet, Pol.
84. G. trichophylla, Grey.
Hab. on rocks, Redwood hills, common in California, Bol., Big.; Van-
couver Island, Lyall.
85. G. ancistrodes, Dur. & Mont.
(13)

14

86.

87.

88.

89.

90.

(14)

LESQUEREUX — CATALOGUE

Hab. on rocks, Dardanelles Caiion, Bol.

It agrees in every point with Muller’s description of this species.

G.

Californica, Sulliv.
Hab. on rocks, San Rafael, and around San Francisco, common, Bol., Big.

Var. foliis caulinis obtusiusculis vel acutis, epiliferis, capsula subglobosa, operculo

G.

longiore.
Hab. same as the normal form, Bol.

hamulosa, Spec. nov. Dioica, irregulariter laxe pulvinata, nigricans. Caulis
dichotome et innovante ramosus, subfasciculatim foliosus. Folia sicca homo-
mallo-faleata, madefacta erecta hamuloso-incurvata, subhomomalla, laxe
irregulariter imbricata, anguste lanceolato-subulata, apice plano obtusiuscula,
carinato-concava, margine plana vel vix revoluta, nervo valido sub apice
evanido intructa, reti basilari quadrato elongato, superiori irregulariter
quadrato. Folia perichetialia longiora, longius acuminato-subulata. Theca
in pedicello laterali medio curvulo, emersa, ovalis, levis, pachydermis, brun-
nea, operculo conico brevi; peristomii dentes breviores, lacerati vel perforato
divisi, annulo nullo.

Hab. gravelly soil, Mt. Dana, 10,000 ft., Bol.

It resembles @. contorta, Wahl., but is very distinct indeed by the hamulose and
homomallous leaves, with long, subulate, opaque points; by the large emerged capsule,
the curved pedicel, the absence of an annulus, ete.

G.

G.

G.

leucophea, Grev.
Hab. metamorphic rocks, Dardanelles Cation, Bol. ; Fort Colville, Lyall. .

montana, Brch. & Schp.
Hab. on rocks from the plain to the mountains; boulders near San Fran-
cisco, Mt. Diablo, Mono Pass, 7,000 to 8,000 ft., Bod.

alpestris, Schl.
Hab. Fort Colville, and Pend’ Oreille, Lyall.

RACOMITRIUM, Brid.

patens, Dicks.
Hab. between Fort Colville and the Rocky Mts., Lyall.

aciculare, Brid.
Hab. on granite rocks above the Yosemite Valley, Brew.

depressum, Spec. nov. Longe, lateque cespitosum, depressum, fusco
lutescens. Caulis elongatus, 4—5 pollicaris, parce dichotome ramosus, laxe folio-
sus. Folia siccitate appresssa, humiditate erecta, aperta, homomalla, e bast
late ovata, dilatato-semiauriculata decurrente, lanceolata, obtusa nervo lato
planiusculo sub apice evanido instructa, concavo-carinata, margine medio
revoluta, superne reflexa, apice denticulis remotis irregularibus instructa,
quandoque integra, retis alaribus quadratis vel equilateralibus latis, plus

OF PACIFIC COAST MOSSES. 15

minus ve granulosis, fuscis, basilaribus subtuberculoso linearibus continuis,
superioribus ovato rotundatis. Capsula in pedicello brevi innovationibus
duplo longioribus immersa, e basi brevicolla subcylindrica; peristomii dentes
irregulariter tripartiti, raro bifidi, cruribus inaequalibus vel liberis, vel
coherentibus pertusis, laevibus.

Hab. falls of the Yosemite Valley, Bod.

This species, remarkable for its large size, is intermediate between A. aciculare
and R. protensum, Brid. Its color is the same as that of var. eataractarum of the last,
but the leaves turned to one side, even in the dry state, are larger, broader, and more
obtuse, mostly denticulate, like those of FR. aciculare, and nearly with the same areo-
lation. The decurrent base of the leaves is slightly enlarged in a narrow auricle whose
reticulation is particularly broad, resembling that of a Dieranum. The capsule, open
at the mouth, is nearly exactly cylindrical, and sometimes slightly curved. The teeth
of the peristome, generally tripartite near the base, are irregular, like those of &. pro-
tensum, but narrow, smooth, with well marked articulations.

94. R. heterostychum, Brid.
Hab. Fort Colville, Lyall.

95. R. lanuginosum, Brid.

Hab. Vancouver's Island, Wood; Mt. Rainier, Oregon, U. S. Exp. Wilkes.
96. R. canescens, Brid.

Hab. Vancouver’s Island, Lyall.

Var. foliis pellucidis acuminatis, angustius reticulatis, vix papillosis.
Hab. on shaded rocks near the Paper Mill, Marin Co., Bol.

HEDWIGIA, Ehrh.

No)
-~I

H. ciliata, Ehrh.
Hab. on stumps of Sequoia sempervirens, Bol.; on rocky cliffs of Bill Williams’
Fork, Big. ; British Columbia, Lyall.

BRAUNIA, Brch. & Schp.

98. B.Californica, Lesqx.
Hedwigia pilifera, Mitt.
Hab. on metamorphic rocks on low mountains ; Mt. Diablo to 3,000 ft., very
common and variable, Bol. ; Vancouver’s Island, Lyall.

In my description of this species, I have omitted to mention a few characters
which, apparently important when one specimen only is considered, are, on the whole,
local peculiarities or varieties only. This omission has led Mr. Mitten to suppose that
his Hedwigia pilifera of Vancouver Island might be a different species, its capsule being
plicate and the branches not inflated at the top. The capsule is indeed sometimes
plicate, but generally smooth, at least when moistened. On rocks exposed to the sun,
the branches of this species are longer and thickened at the top, and the perichetial

(15)

16 LESQUEREUX — CATALOGUE

leaves are much shorter. When growing in the shade, the appearance of the plants
is different, the leaves being longer and not appressed closely to the stem, ete. But
the true specific characters of this species, as they are given in the description, are
recognizable in every form.

PTYCHOMITRIUM, Brch. & Schp.

99. BP. Gardneri, Spee. nov. Ramificatione, habitu, formaque foliorum, P. poly-
phyllo, Breh. & Schp., simillimum, differt: foliis brevioribus, latioribus,
siccitate tortilibus vel cirrhatis, reflexis humiditate, margine argutius
serratis, retis basilaribus longioribus, superioribus cellulis quadratis con-
fertioribus ; capsula in pedicello breviori, pallide ferruginea longiori,
operculo longiori rubello persistente, margine minus crenulato; annulo
angustior! revolubili; peristomi dentibus crassioribus, atro-sanguineis, ad
basim plerumque trifidis ; calyptra levi.

Hab. on rocks, Dardanelles Canon, Forest Hill, Bol.

The large size of this moss separates it at first sight from the European species.
The male flowers are rarely axillary, generally two to four at the base of the vaginule
within the perichetium. Its color is dark green.

ORTHOTRICHEZ.

ZYGODON, Brch. & Schp.

100. Z. Lapponicus, Breh. & Schp.

Hab. spray of the Nevada Falls, Yosemite Valley, Bol. ; Cascade Mts., Lyall.
101. Z. Californicus, Mull.

Hab. on shaded rocks, Marin Co. ; Dardanelles Caiion, ete., Bol.

Z. cuspitosus, Mitt. A new species formed on sterile specimens collected on
Vancouver Island by Lyall, appears, from the description, to belong to this
species.

ULOTA, Mohr.

102. U. phyllantha, Brid.

Hab. Vancouver Island, Wood.

This division of the Orthotrichee has not yet any representative from Cali-
fornia.
ORTHOTRICHUM, Hedw.

103. ©. cupulatum, Hoffn.

Hab. on metamorphic limestone rocks, near the Russian River, Ukiah, red-

woods, deep gulches, Lod.
(16)

OF PACIFIC COAST MOSSES. 17

104. O. Sturmii, Hoppe & Hrnsch.
Hab. on shaded rocks, Yosemite Valley, Lol.

105. O. Texanum, Sulliv.
Hab. on rocks and trees, common in California, and extremely variable, Lol.

Var. B. globosum, Lesqx. Capsula breviori, subglobosa, pedicello longiori emersa,
calyptra colore nitente brunnea, caule breviori, foliis humiditate erectis.
This form has a peculiar appearance, and seems, at first sight, a distinct species.
But I do not find any good character to separate it. The peristome has either fuga-
cious slender cilioli, or short incipient large ones, or none. A peculiar character of this
species, and its varieties, which at once separates it from O. anomalum, is the black color
of the spores.
106. O. speciosum, Nees.
Hab. on trees, Bol. ; Fort Colville, Lyall.

Among the numerous varieties of this species, two forms, collected by Mr. Bo-
lander, merit to be noted.
1st var. polyanthum, Lesqx. Leete viride ; foliis caulinis laxe imbricatis, humiditate
appressis, comalibus longioribus, floribus masculis pernumerosis.
It has long slender branches, and a male bud in the axils of nearly each leaf.
Hab. on rocks, Big Trees.

2d var. brevicaule, Lesqx. Pallide vel luteo viride; cespitibus compactis, sicut
pulvinatis ; caule brevi, theca terminali, pedicello longiori exserta.
Hab. same as the former.
M. Mitten quotes still 0. elegans, Schweer., (from British Columbia, Lyall) a
variety less distinct from the typical form than both the above described ones.
107. O. rupestre, Schl.
Hab. on rocks, Big Trees, Lol.

108. ©. rivulare, Turn.
Hab. roots of elm trees and posts in water, Anderson Valley, ete., quite
common, Bol.

109. O. cylindrocarpum, Lesqx. Pusillum, monoicum subpulvinatum. Caulis sim-
plex vel parce ramosus, obscure viridis, inferne nigrescens. Folia in sicco
appressa, madefacta erecto appressa, e basi latiori elongato-lanceolata acuta
vel subobtusa nervo valido cum apice evanido carinata, margine e basi fere
usque ad apicem revoluta, areolatione ovato-quadrata minuta papillosa
versus basim polygono-elongata. Folia pericheetialia vix longiora. Flos
masculus in ramo proprio sub perichtio innovante, raro axillaris. Capsula
in pedicello brevi, exserta, cylindrica, vel cylindrica-ovalis, pallide viridis vel
luteola. Peristomii dentes 16 per paria approximati, albidi ciliis 8 robustis,
basi dilatatis «quilongis, duplici serie cellularum formatis, separati. Oper-
culum conicum, margine rubellum vaginula calyptraque anguste conica
plicata. valde pilose.

Cc (17)

18 LESQUEREUX — CATALOGUE

Hab. on trees, Oakland, common, Bol.

This diagnosis completes the short comparative description given of this species
in the Trans. Amer. Phil. Soe., vol. 13, p. 6. It is very small, not larger than the
small forms of O. strangulatum, Beauy., with which it has some affinity by the peris-
tome and form of the leaves.

110. O. Kingianum, Spec. nov. Monoicum, cxspitoso pulvinatum. Caulis
erectus, pollicaris parce dichotome ramosus, inferne brunneus, superne
obscure viridis. Folia humida erecto-patentia, breviora, ovato-lanceolata
acutiuscula nervo carinata, e basi usque sub apice revoluta, reticulatione
inferiori quadrato-elongata vel rectangulari, superiore quadrata, obscure
granulosa et minute papillosa. Folia perichzetialia conformia vix longiora.
Capsula in pedicello 2-3 lineas longa, exserta, ovalis, brunnea levis, collo
brevi in pedicello attenuata, ore constricta, peristomii 16 dentes pallidi,
breves, obtusi vel erosi germinati ciliis 8 interpositis e duplo serie cellularum
coinpositis brevioribus. Calyptra elongata capsulam ad basim usque inte-
gens, subleevis.

Hab. rocks near the falls of the Yosemite Valley, Bol.

Distinguishable from O. anomalum, Hedw., which this species most resembles,
by its peristome, and from O. Hutchinsie, Hook. and Tayl., by the areolation of its
leaves and other indicated characters. Appears also nearly related to Q. consimile,
Mitt. But the author says of his species: theca siceitate piicata, ciliis simplicibus @quilon-
gis, calyptra ramentosa, characters at variance with those of our species.

Ill. O. consimile, Mitt.
Hab. Vaneccuver Island, on trees, Lyall.
112. O.Columbicum, Mitt.
Hab. same as the former, Lyall.
113. ©. Coulteri, Mitt.
Hab. California, Coulter.
114. O.pulechellum, Smith.
Hab. Clear Lake, Bol. Also communicated from Eureka and Vancouver
Island by the same.
115. O. Lyellii, Hook.
Hab. on trees, very common in California, Menzies, Bol., Big., Bauer, Lyall,
Wilkes’ U. 8. Exp., ete.
Var. foliis eorumque papillis longioribus, Sulliv. & Lesqx.
Musc. Exsice. Amer.; Ed. 2d, No. 185.
O. Menziesii, W. Hooker, Ms.
O. papillosum, Hampe.
Hab. same as the former.
All the authors who have examined this moss remark, that by its general ap-
pearance, it is indeed far different from the European species ; but the analysis fails to
(18)

OF PACIFIC COAST MOSSES. 19

show a specifie difference. Among the great number of specimens collected by Mr.
Bolander, all the transitional varieties may be followed from the European type to the
largest forms with very long stems, leaves, capsules and papilla. One of the most
marked varieties has short stems, longer, nearly cylindrical exserted capsules borne on
a longer pedicel, with the calyptra covering the whole capsule and part of the pedicel,
and spiniform or branching papilla of the leaves.

TETRAPHIS, Hedw.

116. TT. pellucida, Hedw.
Hab. redwoods, on logs near Big River City, very rare, Gol. ; Fort Colville,
Lyall.

ENCALYPTA, Schreb.

117. E. vulgaris, Hedw.
Hab. Monte del Diablo ; Oakland hills, Gol. ; Fort Colville, Lyall.

118. E. rhabdocarpa, Schw.
Hab. Cascade Mts., Lyall.

119. E. ciliata, Hedw.
Hab. spray of Nevada Falls, Yosemite Valley, in very large and fine speci-

mens, Gol. ; also banks of Russian River, near Ukiah.

?

SPLACHNACEZ.

TAYLORIA, Hook.

120. 'T. serrata, Hedw.
Hab. Fort Colville, Lyall.

SPLACHNUM, Linn.

121. §S. melanocaulon, Schw.
Hab. California, as indicated by Mitten.

FUNARIACE4®.

PHYSCOMITRIUM, Brid.
122. BP. pyriforme, Brid.
Hab. wet banks, Dardanelles, and swamps near San Rafael, Bol.
ENTHOSTODON, Schwegr.

123. E. Bolanderi, Lesqx.
(19)

20 LESQUEREUX — CATALOGUE

Hab. on clayey ground, near San Francisco, Bol. .

124. E. Templetoni, Schw.
Hab. swamp near Mendocino, Coast Range, very rare, Bol.

FUNARIA, Schreb.

125. F. Californica, Sulliv. & Lesqx., Musci Exsicc. Amer., Ed. 2d, No. 258. Pusilla ;
folia coronalia in gemmula congesta, vel erecta, oblonga, breviter acuta,
integra, fere continuo-costata ; capsula in pedicello brevi-recto, sicco sini-
strorsum torto, sub pyriformi-oblonga, erecta, equalis, vix asymetrica ;
operculum convexo-conicum, calyptra et peristomium generis, annulus
nullus.

Hab. on clayey soil, Auburn, Ukiah, ete., Bol.

Is much like a small Enthostodon, the capsule being generally straight and sym-
metrical. The peristome is double, the ternal being glued to the outer one to above
the middle of the teeth. The reticulation of the leaves is remarkably close for a
species of this genus.

126. ¥F. calearea, Wahl.

A var. B. patula, Schp.

Hab. on the ground, Mission Dolores; Mt. Diablo, Bol. Funaria Muhlen-
bergu, Schwegr, mentioned by Sullivant, as found by Dr. Bigelow near
the crossing of the Colorado, is referable to this species.

127. F. hibernica, Hook. :

Hab. Cajon Pass, Sierra Nevada, Big.

128. F. hygrometrica, Hedw.

Hab. California, Bol., Big., Wilkes’ U. S. Exp. ; Fort Colville and Vancouver,
Island, Lyall.

Var. C. calveseens, Schp.

Hab. rocks in canons of the American River, Brew.

129. F. convoluta, Hampe.

Hab. on the Sierra Nevada Mts., alt. 3,000 to 5,000 feet, Bauer.

130. F. microstoma, Breh. & Schp.

Hab. Soda Springs, Upper Tuolumne, 9,700 ft., Bol.

Appears to be abundant there ; the specimens are very fine.

BRYACE A.
BRYUM, Dill.
§ 1. LEPTOBRYUM, Schp

131. B. pyriforme, Hedw.
(20)

OF PACIFIC COAST MOSSES. 2]

Hab. around Clear Lake and Mt. Dana, common from 8,000 to 11,000 ft.,
Bol. ; moist banks of Sonora Pass, Brew.; banks of streams, Cajon Pass,

Big.; Cascade Mts., Lyall.

§ 2. WEBERA, Hedw.

132. B. polymorphum, Hoppe & Hrnsch.
Hab. Mt. Dana, mixed with B. nudicaule, Lesqx., rare, Lol.

133. B. longicollum, Swartz.
Hab. Cascade Mts., Lyall.

134. B. nutans, Schreb.
Hab. foot of Mt. Dana, Bol. ; Cascade and Galton Mts., Lyall.
Var. @. bicolor, Brch. & Schp.
Hab. Mt. Dana, Bol. ; banks of King’s River, 4,000 ft., Brew.

135. B.nudicaule, Spec. nov. Dioicum, cespitosum ; ceespites compacti, inferne
brunnei, superne virentes. Caulis basi tantum radiculosus, subpollicaris,
gracilis simplex vel raro ex apice innovans; plants antherigere graciliores
fructifereque e basi usque versus summitatem subnude, folia minima
squameformia, erecto-apressa, pellucida gerentes. Folia comantia dense
conferta, erecta, ovato lanceolata brevia, nervo valido sub apice evanido
carinato-concava, subintegra vel apice obsolete denticulata; floralia
intima breviora denticulata ; plante sterilis basi interrupte foliose folia
breviora. Theca in pedicello vix semi-pollicari, valde flexuoso horizontalis
vel pendula, ovata, subventricosa, ore haud vel vix constricta collo brevi
fusca, pachydermis. Peristomii interni dentes liberi angusti, elongati,
seepius laciniis irregularibus summo conati, ciliis nullis interpositis; opercu-
lum conicum vel plano convexum annulatum ; annulus pallidus compositus,
revolubilis.

Hab. Mt. Dana, 11,000 ft., Bol.

The male plants of this species are mixed in an abundance in the caspites,
and their terminal buds are thick with numerous anthers without or with a few short
paraphyses. The perigonial leaves are broad, oval, short-pointed or obtuse, brown
colored, and obscurely nerved. Some free anthers are also found, though rarely, in
the axils of the pericheetial leaves. The capsule is rarely symmetrical, but generally
more or less inflated on the lower side. This moss agrees so well with the excellent
description given by Muller (Bot. Zeit., N. 40, p. 328) of his B. Drummond: from the
Rocky Mts., that I should have considered it identical, if it was not that Muller’s moss
has cilia Weber duplicia brevia, while ours has no trace of cilioli, and that also the
former is exannulata, while the California moss has a pretty large white revolving annu-
lus. Muller also does not mention the peculiar stem leaves of the flowers bearing
plants, and the presence of anthers in the axils of the pericheetial leaves. The areola-
tion of the leaves is of a Webera, but short and broad.

(21)

22 LESQUEREUX — CATALOGUE

136. Bryum Bolanderi, Spec. nov. Dioicum, laxe cxespitosum, depressum, pallide
viride nitens. Caulis simplex, foliosus. Folia inferiora laxe imbricata,
erecta, lanceolata; superiora comantia conferta, longiora, anguste lanceo-
lata, nervo sub apice denticulato evanido. Planta mascula vix gracilior ;
folia perigonalia e basi late ovata concava in acumen longum flexuosum
angustatum producta subintegra. Folia pericheetialia conformia, vix lon-
giora. Capsula in pedicello longiori rubello inclinata vel horizontalis, brevi-
ovata, in collum sat longum attenuata; peristomii dentes externi late
brevesque, interni longiores ciliis vel nullis vel binis rudimentariis interjectis.
Operculum conicum apiculatum; annulus compositus.

Hab. foot of Mt. Dana, Bol.

Distinct from the former by its longer capsule of a thinner texture, narrowed
at the mouth, and borne on a longer pedicel; by its shining color, its long, narrow,
strongly denticulate leaves, the form also of the perigonial leaves, ete. The peri-
cheetial leaves do not have any anthers in their axils. A fine species, rather related to
B. crudum, Schreb.

157. B. cucullatum, Schweer.
Hab. Mt. Dana; rare, Gol.

138. B.crudum, Schreb.
Hab. around Clear Lake and near the Big Trees, on shady rocks, Bol. ; Fort
Colville, Lyall.
159. B. Ludwigii, Schweer.
Hab. foot of Mt. Dana, 9,000 to 10,000 ft., Bol. ; Cascade Mts., Lyall.
The form collected by Mr. Bolander is robust, has thick branches and open
leaves.

140. Bryum Tozeri, Grev.
Hab. on the ground, borders of ditches and roads around San Francisco,
Oakland, ete., Lol., Big.
141. B. albicans, Wahl.
Hab. wet banks, Dardanelles Cation, Bol. ; Galton Mts., Lyall.
Var. gracihor, capsulis brevioribus, ete.
Hab. on rocks watered by springs near San Francisco, Bol.

§ 3. BRYUM, Dill. emend

142. B areticum, Brch. & Schp.
Hab. foot of Mt. Dana, Bol.
It slightly differs by the shorter capsule. Specimens few and deoperculate.
143. B. cernuum, Breh. & Schp.
Hab. foot of Mt. Dana, Bol.
144. B. inclinatum, Brch. & Schp.
Hab. on rocks at Clark’s, Yosemite Valley, Bol.

OF PACIFIC COAST MOSSES. 29

145. B. Warneum, Brid.
Hab. foot of Mt. Dana, 9,000 to 10,000 ft., borders of ditches ;
abundant, Bol.
There is no difference whatever between this and the European moss. The
flagelliform, twisted, nearly naked branches are extremely numerous.

146. B. intermedium, Brid.
Hab. on wet ground, Big Trees; on the ground near Crescent City ; Mt.
Dana, ol. ; Sonora Pass, mixed with L. pyriforme, Brew. ; also found
by Bigelow—no locality marked.
147. B. cirrhatum, Hoppe & Hrsch.
Hab. in a meadow near the Big Trees, 8,000 ft; Mono Pass, Lol.
148. B. bimum, Schreb.
Hab. Fort Colviile and Galton Mountains, Zyadl.
149. B. torquescens, Brch. & Schp.
Hab. Oakland, opposite San Francisco, Lig.
150. B. pallescens, Schl.
Hab. Cascade Mountains, Lyall.

151. B. subrotundum, Brid.
Hab. on rocks, Yosemite Valley, Lol.
It differs from the European form only by the slightly longer capsule, rather
horizontal than pendent.

152. B. cespiticium, Linn.
Hab. on the ground, Yosemite Valley, Bol.; Silver Mts., and through the
Sierra Nevada, from 7,000 to 8,000 ft.; Brew.; Cascade and Galton
Mts., Lyall.

155. B. argenteum, Linn.
Hab. on the ground, San Francisco, Pol. ; dry ravines, 50 miles west of the
Colorado, Big. ; rare.
154. B. Californicum, Sulliv.
Hab. near Benicia, Big. ; common around San Francisco, on the ground, in
grassy places, Bol.
155. B. Bigelowii, Sulliv.
Hab. trunks of trees above Sonora; base of the Sierra Nevada, Pig.
156. B. miniatum, Spec. nov. Dioicum compacte czspitosum. Caulis parce radi-

culosus simplex vel pluries sub capitulis inflatis innovante ramosus, gracilis,
innovationibus quandoque gracillimis filiformibus. Folia sicca appressa,
caulina laxe imbricata superiora in capitulum conferta concava, ovato-obtusa
vel ovato-lanceolata obtusa, haud vel vix marginata, margine vix reflexa,
nervo valido sub apice evanido instructa, e cellulis polygonis parietibus crassis
reticulata. Folia pericheetialia comalibus vix longiora, angustiora. Capsula

(23)

24 LESQUEREUX — CATALOGUE

in pedicello longo rubello, inclinata, elongato-obconica, sub ore sub con-
stricta, rufescens, operculo mamillari rubello, annulo latiori, Peristomii
interni dentes angusti, ciliis crassis articulatis interpositis.

Hab. on moist rocks, Yosemite Valley.

This fine species is somewhat related to B. pseudo-triquetrum, with which it
grows, but easily separated by the enumerated characters. The ramification is the
same as in B. Bigelowit, Sulliv., from which it differs by the long capsule and the
form of the leaves. The top of the branches is purplish, abruptly crimsoned, as if
it had been plunged in carmine.

157. 3B. occidentale, Sulliv.

Hab. on ground, rocks, old logs, very common in California, Gol., Dig., Brew.

This species is most intimately related to L. capillare, Linn., presenting most of

the forms which are marked as varieties in the European species. I owe to the kind-
ness of Prof. Hooker, specimens of a Brywm named BL. eapillare, by Mr. Mitten, col-
lected in Vancouver Island, by Lyall, and which is evidently a large form of L. ocei-
dentale, and at the same time undistinguishable from B. eapillare. It would be too long
to enumerate here all the varieties of ths California moss. The most marked one is a
small form with slender innovations emerging from under more distinct and thicker
capsules having with generally longer pointed leaves, narrow, cylindrical proportionally
longer, broad red capsules and a proportionally broader annulus. But even the largest
forms, whose capsules are generally pale brown and the leaves short, pointed, bear in
the same tufts capsules of various color, from pale buff to dark red, with leaves also
elongated or rather contracted into a piliform point. Though, from its larger annulus
—especially its shorter pointed leaves and peculiar ramification—this species may be
separated with some right from B. capillare, it does not appear possible to subdivide
its varieties into species. One of these divisions is B. Baveri, Hampe, which, accord-
ing to the author, differs from B. capillare; statura minori, folias brevisribus immarginatis,
nervo excedente flexipilr.
158. B. Oreganum, Sulliv. Wilkes’ U. 8. Expedition, p 10, tab. vii.

Hab. Oregon, Wilkes’ U. S. Exp.
159. B. obconicum, Hrnsch.

Hab. flanks of Santa Cruz Mts., 2,000 feet, Brew.
160. B. Canariense, Brid.

B. Billarderti, Schweegr.

Hab. canons in Monte del Diablo, Bol.
161. B. pseudotriquetrum, Schweer.

Cum var B. gracilescens, Schp.

Hab. wet rocks, Big Trees, Bol. ; Fort Colville, Lyall.
162. B. Duvallii, Voit.

Hab. Hureka, Bol., in good though sterile specimens ; Fort Colville, Lyall.
163. B. turbinatum, Hedw.
(24)

164,

165,

166.

167.

170.

174.

OF PACIFIC COAST MOSSES.

Hab. Galton Mts., Lyall.
Var. @. latifolium, Schp.
B. Schleicheri, Schwiegr.
Hab. wet meadows, Big Trees, Lol.

>

MNIUM, Lion. emend.

M. affine, Bland.
Hab. near Clear Lake, sterile, Bol.

M. insigne, Mitt.

Hab. borders of creeks, Devil’s Caiion, Bol.; Vancouver Island, Lyall, Wood.

M. venustum, Mitt.

Hab. shaded rocks, Ukiah, etc., Bol.; Vancouver Island, Lyall; Oregon,

Wilkes’ U. 8. Eup.

M. medium, Breh. & Schp.
Hab. Fort Colville and Cascade Mts., Lyall.

M. spinulosum, Brch. & Schp.
Hab. Fort Colville, Lyadl.

M. punctatum, Linn.

Hab. Eureka. Sent by Mr. Bolander in good fruiting specimens.

M. Menziesii, Mull.

Hab. shaded rocks, banks of creeks, Bol.; Coast Range, By. ; Oregon,
Port Discovery, Wilkes’ U.S. Exp. First discovered in N. W. America

by Menzies, and gathered also by Seouler.

MEBESIA, Hedw.

M. uliginosa, Hedw.
Hab. found again in swamps, 9,000 ft., Bol.; swamp,
Mooyie River, Lyall.

M. tristicha, Breh. & Schp.
Hab. Tack River, Lyall.

M. longiseta, Hedw.
Hab. Oregon, Wilkes’ U. S. Exp.

AULACOMNIUM, Schweeer.

A. androgynum, Schweegr.

8,500 ft., Brew. ;

Hab. especially on burnt stumps of Sequota sempervirens, from the plains to

the mountains, very common, Sol., Lrew., Big. ; Vancouver Island and

Fort Colville, Lyall.

D

(25)

26 LESQUEREUX — CATALOGUE

175. A. palustre, Schweer.
Hab. swamps near Mendocino, Bol. ; Cascade Mts., Lyall.

BARTRAMIA, Hedw.

176. B. stricta, Brid.
Hab. on rocks and ground near San Francisco, Gol.

It differs from the European form only by its pedicel, which is round and not
square, obtuse under the capsule ; agreeing, therefore, with Bridel’s description, which
does not mention this peculiarity.

177. B. thyphylla, Brid.
Hab. foot of Mt. Dana, ditches, abundant, Bo/. It is the Alpine form, in
compact tufts, with short stems and pedicels ; also Cascade Mts., Lyall.

178. B. fontana, Brid.
Hab. Yosemite Valley, Big Trees; Clear Lake, on wet rocks, Bol., Brew.
Appears common in the Sierra Nevada Mts.; Fort Colville, Lyall.
179. B. Menziesii, Turn.
Hab. on shaded rocks ; very common in California, and gathered by all the
botanists who have explored that country, Menzies, Big., Brew., Bol.,
Bauer, ete.
Var. foliis latioribus, brevius acuminatis, capsula longiore, ovato-elongata, ore
Ja iori nudo, vel membrana integra circumdato.
Hab. on rocks, Bol.

With the normal form, whose capsule is oval, sometimes nearly round, Dr.
Hampe has made a new species, under the name of Glyphocarpa Baueri. But the eap-
sules of this normal form have generally a well developed peristome, though, indeed,
this peristome is sometimes replaced by a pellucid membrane, lacerated in the form
of irregular teeth, or even is totally wanting. The moss, which I consider as a variety
as described above, has generally a naked mouth, though sometimes it shows traces of
even a well formed peristome.

TIMMIA, Hedw.

180. T. megapolitana, Hedw.
Hab. Vancouver Island, Lyall.

POLYTRICHACE AS.
ATRICHUM, Tal. Beauv.

181. A. undulatum, Tal. Beauv.
Hab. on rocks, Mission Dolores, Bol. ; Oregon, Wilkes’ U. S. Exp. ; Fort
Colville, Lyall.
(26)

OF PACIFIC COAST MOSSES. ai

182. A. angustatum, Bryol. Hur.
Hab. banks, Santa Cruz Mts., alt. 2,200 ft., Drew.

POGONATUM, Tal. Beauv.

183. P. dentatum, Meuz.
Hab. pine woods of N. W. America, Menzies; clay banks in dense shade of
the redwoods near Crescent City, Brew.

Prof. C. Muller says of this species: calyptra leevis ; in my specimens it is
rugose, like that of P. urnigerum, Brid., and, accordingly, the only difference which
separates the Californian from the Huropean species is the glaucous color of the more
stronly dentate leaves.

184. P. alpinum, Roehl.
Var. D. brevifoluum, Schp.
Hab. foot of Mt. Dana, borders of ditches, Bol.
185. P. contortum, Menz.
Hab. on shores of N. W. America, Menzies.

POLYTRICHADELPHUS, GC. Mull.

186. P. Lyallii, Mitt. ]
Hab. foot of Mt. Dana, and arounfd Clear Lake, Bo/.; swamp, east side of
the Cascade Mts., Lyall.

POLYTRICHUM, Dill.
187. P. piliferum, Schreb.
Hab. rocky places above Sonora, big.; Vancouver Island, Big., Lyall.
Var. pilo brevi, serrato, foliis quandoque epiliferis.
Hab. Lassen’s Peak, alt. 8,000 ft., Brew.
Var. levipilum, P. levipilum, Hampe.
Hab. Mt. Diablo and Coast range, Bol., Bauer.

188. P.juniperinum, Hedw.
Hab. California, from the plains around Mendocino, to the Yosemite Valley,
common, /ol.; near Cajon Pass, big.; abundant in woods, from 7,000 to
9,000 ft., Mt. Brewer, Srew.; Vancouver Island, Lyadl,; Oregon, U. 8.
Wilkes’ Exped.

BUXBAUMIA, Haller.

189. B.aphylla, Haller.
Hab. Cascade Mts., Lyadl.

28 LESQUEREUX — CATALOGUE

FONTINALACEZS.

FONTINALIS, Dill.

190. F. antipyretica, Linn. j
Hab. Swamps, in stagnant water and Coast Ranges in rivulets, bol. big.,
Alpine Lake, near Silver Mts., Brew.

191. F.Mercediana, spec. nov. F. antipyretica habitu similis, differt : caule graciliori,
distycho, raro trigono folioso ; foliis acutius carinatis, plicatis, haud vel vix
decurrentibus ; areolis foliorum superne angustioribus, vacuis ; alaribus bre-
vioribus inflatis, paucis ; peristomil interni, reti clathrato imperfecto ; externi
dentibus brevioribus, articulis distantibus, 18 to 20; operculo duplo breviori,
capsulee colore pallida.

Hab. on rocks, Merced river, Bol.
This species has the facies, color and peristome of F’. dalecarliea, Brch. and

Schp., differing essentially from it by the entire (not lacunose) teeth, the longer capsules,

the flattened branches, ete.

DICHELYMA, Myr.

192. D. uncinatum, Mitt.
Hab. Fort Colville, Lyadl.

193. D. Swarzii, Lndb.

Hab. upland swamps between King’s and Kaweah rivers, Brew.; swamps near
Mendocino city, and pools at the foot of Mt. Dana, Bol.; no fruit found.
This moss agrees in every point with Schimper’s description of the species, and
g, with
long, scarcely branching stems, and appressed, nearly straight leaves. It grows in
prairie swamps, which are dry in summer. The other, black or brown colored, with

stems much divided in horizontal short branches and faleate-uncinate leaves.

presents two forms far different in appearance. One, pale green colored, shinin

NECKERACEZE.
NECKERA. Hedw.

194. WN. Menziesii, Hook.
Hab. on rocks, in deep shaded gulehes, Bol.; base of trees, near Crescent
City, Brew ; Fort Colville, Lyall.
From the Russian valley, Mr. Bolander has sent remarkable specimens, with
leaves and branches covered with capillary filaments resembling tufts of conferve, or
of Hypnum confervoides. These filaments are generally simple, emerging from the

(28)

OF PACIFIC COAST MOSSES. 29

axils of the branch leaves, and bear very small oval-pointed, entire or slightly dentic-
ulate leaves, without costa and with a loose reticulation. Other specimens also gath-
ered by Mr. Bolander, and marked ‘“‘shaded rocks, Yosemite Valley,” are of a pale
green color, with short stems, and without any trace of stolons.
195. Neckera Douglasii, Hook.
Hab. on trees, Marin Co., Mendocino City, ete., Bol.; Oregon, Wilkes’ U.S.
Lexp.; Vancouver Island, Wood.

ALSIA, Sulliv.
196. <A. Californica, Sulliv.
Neckera Californica, Hook. and Arn.
Hab. on trees and rocks ; very common along the coast, Bol., Big.; Oregon,
Withes’ U. S. Exp.

197. A. longipes, Sulliv. and Lesqx., Musci Exsice. Amer., Ed. 2d, No. 399 ; Dioica,
pinnato ramulosa, subfrondiformis, compressiuscula foliosa. Folia oblongo-
lanceolata, breviter acuta, superne grosse serrata, minute ovali-areolata,
costa ultramedia apice dorsali denticulato ; pericheetialia abrupte filiformi
attenuata ; capsula cylindracea, in pedicellum subuncialem perichxtio quad-
ruplo longiorem defluente ; peristomio 1-2 ciliolato, operculo calyptraque
A. abietine.

Hab. on rocks in deep canons, Oakland, Bol.
198. A. abietina, Sulliv.
Neckera abietina, Hook.
Hab. on trees, common in California, and gathered by all the botanists who
have explored the country.

ANTITRICHIA, Brid.
199. A. curtipendula, Brid.
Hab. Mt. Diablo, sterile, Bol. It is a peculiar form, with closely appressed
leaves and julaceous branches. Oregon, Wilkes’ U. S. Exp.; Vancouver
Island, Wood.

Var. C. gigantea, Sulliv. & Lesqx., Musci Exsicc. Amer., Ed. 2d, No. 356. Valde
robusta, atro-viridis, foliis latioribus, homomallofaleatis, capsula cylindrica
longiori.

Hab. on rocks and trunks of living trees, Redwoods, Bol.
200. A. Californica, Sulliv.
Hab. on shaded rocks, and sandstone boulders, Oakland, Bol.

HOOKERIACE2.

HOOKERIA, Tayl.

201. H. acutifolia, Hook.
(29)

30 LESQUEREUX — CATALOGUE

Hab. in a deep canon, Pol.
This moss is the same in every point as that described under that name from
the Atlantic States. No fruit has been found as yet.
202. H. anomala, Mull.
Hab. N. W. America, Menzes.

LESKEACE4®.

ANOMODON, Hook & Tayl.

203. A. Californicum, spec. nov. Czespites laxi prostrati vel dependentes, fusco-
lutei, inferne brunnei. Caulis parce ramosus, gracilis, sicco foliis appressis
angulatus. Folia quadrifariam imbricata, humida aperta, basi semi amplec-
tente auriculato-decurrentia, late ovata, acuta, margine replicata, nervo valido
sub pallido cum apice evanido carinata, cellulis alaribus oblongis elongatis,
inde superne ovato-quadratis utraque pagina papillosis, papillis ad auriculas
longioribus, spinosis.

Hab. on rocks, Mt. Diablo, sterile, Bol.
A fine species, without.near relation to any other described. The borders of
the leaves are abruptly plicate backward to above the middle of the leaves, and only
reflexed near the point.

FABRONIACEZE.

FABRONIA, Raddi.

204. F. pusilla, Raddi.
Hab. bark of trees, Oakland, Gol.

HYPNACEH2ZE.

PTERIGYNANDRUM, Hedw. emend.
205. BP. filiforme, Hedw.
Hab. on shaded boulders and trees, Bol., Big.
Var. nervo validiore elongato, foliis apice cristato-serratis. Leptohymenium cris-
tatum, Hampe.
Hab. on rocks, Sierra Nevada, Bauer.

PTEROGONIUM, Swartz.

206. BP. gracile, Swartz.
Hab. on rocks near the bay of San Francisco, common, Gol., Big. Cali-
fornia, Wilkes’ U. S. Eup.
Var. statura graciliori, ramis magis filiformibus, foliis duplicato-serratis.

(30)

OF PACIFIC COAST MOSSES. 51

Leptohymenium duplicato-serratum, Hampe.
Hab. on trees in California, Lauer.
This species is still more variable than the former.

HYPNUM, Dill.
21. THUIDIUM, Schp. Bryol, Hur.

207. H. Blandowii, Web. & Mohr.
Hab. Fort Colville, Lyall.

208. H. ecrispifolium, Hook.
Hab. N. W. America, Menz.; on shaded ground and rocks, Oakland, ete.,
Bol. ; Vancouver Island, Wood.

209. H. (Leskea) laxifolium, Hook.

Hab. coast of N. W. America, Venz.

Nothing like this species of Hooker has been found in California by recent col-
lectors. It may be a variety of the former species, or appears, at least, to belong to
this section.

210. H. remotifolium, Grev.

Hab. N. W. America.

An obscure species.

211. H. Whippleanum, Sulliv.

Hab. California, Big.

212. H. leuconeurum, Sulliy. et Lesqx., Musci Exsicc. Amer., Ed. 2d. Hypno
Whippleano peraffine, czespite densiore, operculo breviore, pedicello levi
etc, distinguendum.

Hab. on the trunks of Quercus agrifolia, Oakland ; in woods, but more com-
mon on moist soil of shaded hillsides, Oakland.

213. H. calyptratum, Sulliv.
Hab. near Los Angeles, on the ground, Big.

2 2. ISOTHECIUM, Brid.

214. H. myosuroides, Linn.
Hab. near San Francisco, dry woods, Big.
Perhaps a variety of the next.
215. H. stoloniferum, Hook.
Hab. on trees, in the Redwoods ; very common in California, Menzies, Bol.,
Big. Oregon, as var. B. of H. myosuroides, in Sulliv., Wilkes’ U. 8. Kup. ;
Vancouver Island, Wood.
This species is particularly polymorphous. In order to elucidate the remarka-

ble disposition of this moss to modify its form under peculiar circumstances, especially

(31)

32 LESQUEREUX — CATALOGUE

under the influence of wind and fog, Mr. Bolander sent me a large specimen, whose
branches on one side are elongated in slender filiform stolons, from six inches to one
foot long, while on the other side the stem and branches are thick, short, with large
leaves. The filamentous part, according to Mr. Bolander’s remarks, was hanging from
a branch exposed to wind and fog, while the other part, presenting a normal develop-
ment, was, by its extension to the other side of the trunk, preserved against this ac-
tion. Specimens of H. cireinale and H. Nuttallii are subjected to the same peculiar de-
velopment. Ican but therefore consider the multiplication of species from such poly-
morphous mosses as a hazardous task. I have admitted as a species the more distantly
‘related form of H. Breverianum, which, growing in dense tufts on dry rocks, has a
black color, short stems, short obtuse leaves, and in appearance is totally different
from H. stoloniferum. But even on dry rocks, the part of the cespites which is not di-
rectly exposed to the sun’s influence passes to a yellowish green color, and bears some
more elongated attenuate branches, even stolons, showing more and more an approach
to H. stoloniferum or H. myosuroides, for Professor Muller does not separate these spe-
cies. Mr. Mitten has described a number of forms which are more or less intimately
related to H. stoloniferum, and which, if the opinion of Muller is right, should be re-
united as well as H. Breverianum as varieties of H. myosuroides.

216. H. Breverianum, Lesqx.

Hab. on metamorphic sandstone around San Francisco, ol.

217. H.aggregatum, Mitt.
H. Breverianum var., Sulliv. and Lesqx. Musci Exsice. Amer., Hd. 2d, No. 427,
Hab. in deep canons; Oakland, on trunks, ol.
218. H. aplocladon, Mitt.
Hab. N. W. coast of America, Douglas.
219. H.lentum, Mitt.
Hab. same as the former, Douglas.

23. CAMPTOTHECIUM, Bryol. Eur.

220. H. lutescens, Hnds.
Hab. Vancouver Island, Lyall ; California, Coulter; N. W. Coast, Douglas.

Received from Victoria, Oregon, through Mr. Bolander.

221. H. Nuttallii, Wills.
Hab. on trees, California, common, Bol., Big. ; Vancouver Island, Lyadl,
Douglas.
Var. stoloniferum, ramis elongatis, aggregatis filiformibus lutescentibus.

Hab. on trees. Sent by Mr. Bolander. A very remarkable variety. The
stems are creeping, as in the normal form, and bear shorter and more slender capsules,
and the crowded branches are elongated filiform, depending from the branches of trees.
222. H. arenarium, Lesqx.

Hab. on sand, around bushes near the shores, San Francisco, Gol.

OF PACIFIC COAST MOSSES. 33

223. H. pinnatifidum, Sulliv. and Lesqx., Musci Exsice. Amer., Hd. 2d, No. 513. Ab
Hypno aureo affinissimo distinguitur: capsula cernua annulo majori, foliis
brevius filiformi acuminatis, reti multo laxiore denique inflorescentia revera
dioica.

224. H. Nevadense, Spec. nov. Dioicum, laxe cxspitosum, robustum, lutescente
viride. Caulis primarius repens pinnatim ramosus, ramis brevibus, horizon-
talibus vel longioribus, irregulariter divisis, arcuato curvatis. lolia lanceo-
lata, sensim brevi-acuminata, homomallo-curvata, apice minute serrata, costa
carinata, profunde biplicata, margine revoluta vel reflexa, cellulis alaribus
perpaucis, irregulariter ovalibus. Capsula in pedicello rubello erecta vel
sub cernua, ovato-cylindrica, longe rostrata, peristomii interni cilia vel rudi-
mentaria, vel nulla; annulus compositus.

Hab. on rocks in the spray of Nevada and Bridal Veil Falls, Lol.

Is distinguished from HZ. lutescens, Huds., by its green color, larger thick stems
and branches ; the evidently pennate ramification, less marked, however, than in //.
aureum ; the alar reticulation scarcely marked by a few oval cells; by the large capsule,
the peristome without cilia, ete. The leaves are more regularly and deeply plicate
than in any other Camplothecium.

24. BRACHYTHECIUM, Bryol. Eur.

225. H. letum, Brid.
Hab. the Yosemite Valley. The plants are sterile, and the species not un-
doubtedly ascertained.

226. H.salebrosum, Hoffm.
Hab. Fort Colville and Pend’ Oreille, Lyall.

227. H. collinum, Schl.
Hab. Cascade Mts., Lyall; Mr. Bolander sent it sterile from the Yosemite
Valley.

228. H. Hillebrandi, Spec. nov. Monoicum, tenellum, cespitibus intricatis conden-
satis sericeo-lutescentibus. Caulis erectus, plicatus, irregulariter subfascicu-
latim ramosus cum ramis undique divergentibus radiculosis. Folia laxe im-
bricata subsecunda, e basi ovata, sensim lanceolato-acuminata, concava,
basim tantum plano reflexa, toto margine serrulata, costa medio vel paulo
ultra evanida, areolatione angusta elongato-polygona, angulis dilatata, quad-
rato chlorophyllosa. Pericheetialia pellucida, laxe areolata, plerumque
enervia, e basi ovata, sensim in acumine longo, gracili erecto vel flexuoso sub-
integro attenuata. Capsula in pedicello breviusculo inferne rubello subru-
goso, superne pallido levi, brevis, turgide ovata, equalis, erecto raro in-
clinata, sicca ore dilitata, operculo conico brevi obtuso truncato. Peristomii
externi dentes longiores inferne pallide lutei, superne albidi, processus
tenerrimi pellucidi dentibus perforatis ciliis duobus articulatis interpositis ;
annulus ?

E (33)

34 LESQUEREUX — CATALOGUE

Hab. on rocks, Merced River, Bol.

A fine species related to HZ. collinum and H. saheinum. The leaves have about
the same form as in this last, but are broader, though not as broad as in LZ. collinum,
with a shorter costa. The essential difference is in the small, generally erect capsule,
with a pedicel rough from the middle downward. The operculum, though found in
plenty among the tufts, is detached from all the capsules. No trace of an annulus
is left.

229. H. declivum, Mitten.

Hab. Pend’ Oreille River, Lyall.

230. H. vallium, Sulliv. and Lesqx., Musci Exsicc. Amer., Ed. 2d, No. 506.
Hypno leto inflorescentia, statura, foliatione et habitu accedens ; distinctis-
simum autem caule rigidiore, pedicello seabro, foliorum cellulis quadratis ad
basis angulos late decurrentes paucioribus, haud granulosis, ete.

Hab. on shaded metamorphic rocks, in deep canons, Bol.

This species is apparently nearly related to the former, but the leaves are not
denticulate from the base, and the costa is also not denticulate on the back.

251. H. populeum, Hedw.

Hab. Sierra Nevada, Dr. Hillebrand.

The specimens were communicated by Mr. Bolander; all the capsules are
deoperculate.

232. H. Bolanderi, Lesqx.
Hab. on the ground, shaded by Oreodaphne Californica, rare, Oakland, Bol.

25. SCLEROPODIUM, Bryol. Eur.

253. H. illecebrum, Schweer.

Hab. rocks and ground, from the plain to the mountains, Gol. Extremely
variable. In compact dark brown tufts, with densely imbricated, ovate-
obtuse, nearly entire leaves and julaceous branches. Hab. in the water
of the Yosemite Valley. In subdendroidal divisions, with acute and
nearly entire leaves. Hab. deep cations, etc. It is remarkable that no
species of the genus Seleropedium, so widely distributed in California, has
been found, except by Mr. Bolander.

234. H. cespitosum, Wills.
Hab. on shaded rocks, and on the ground in the redwoods, Oakland, ete.,
Bol.
255. H. Californicum, Lesqx.
Hab. on shady, sandy ground, and sand around San Francisco, Gol.

26. EURYNCHUM, Bryol. Eur.
236. H. strigosum, Hoff.
Hab. wet ground, Big Trees, Bol. ; Fort Colville and Galton Mts., Lyall.

ew
Or

OF PACIFIC COAST MOSSES.

237. H. Stokesii, Turn.
Hab. on shaded ground and rocks, very common, Lol., Diy. ; Vancouver

Island, Lyall, Wood.

238. H. Oreganum, Sulliv.
Hab. in woods, on decaying logs or on the ground, Bol., Drew., Big.; Puget
Sound, Oregon, U. 8. Wikes’ Exp.; Vancouver Island, Wood.

27. RYNCHOSTEGIUM, Bryol. Kur.

239. H. rusciforme, Weis.
Hab. in a rivulet near Mt. Diablo, sterile, Dr. Hillebrand, Bol.

38. THAMNIUM, Bryol. Eur.

240. H. Bigelowii, Sulliv.
Hab. on shaded rocks, in canons, Big., Bol., Brew.

241. H. Neckeroides, Hook.
Hab. Vancouver Island, Wood.

According to Mitten, we have, in the United States, in this division of the Hyp-
nacee, H. Neckeroides, Hook., found by Drummond in St. Louis; H. alopecurum, sent
from Boston, and H. Alleghaniense, Mull. Though we have examined, with Mr. Sulli-
vant, a great number of specimens, either collected by ourselves or sent from various
points of the United States, we have never been able to find any species different from
that of H. Alleghaniense, Mull., which differs from H. alopecurum by the inflorescence
only. This 7. Alleghaniense is the same moss as No. 119 of Drummond, H. Neckeroides,
quoted by Mitten. And as the male flowers are rarely found on fruiting plants, in-
complete specimens are often taken for 7. alopecurum. I believe, therefore, that these
three species, quoted by Mitten, are, for American specimens, referable to the same
H. Alleghaniense, Mull. H. alopecurum from Europe, and H. Neckeroides, Hook., from
New Zealand, are indeed different, but not yet found in our country.

29. PLAGIOTHECIUM, Bryol. Eur.
242. H. pulchellum, Dicks.
Hab. Fort Colville, Lyall.
243. H. turfaceum, Lndb.
Hab. same as the former, Lyall.

244. H. elegans, Hook.
Hab. near Nootka, Vancouver Island, Menzies.

bo
TA
Or

H. denticulatum, Lin.
Hab. Coast Range, Mendocino City; on a fork of redwood tree, Jackson
Valley, Bol.; rare, Fort Colville, Lyall.

246. H. undulatum, Lin.

36 LESQUEREUX — CATALOGUE

Hab. swamps near the coast, Mendocino, Bol.; Oregon, U. S. Wilkes’ Exp. ;
Fort Colville, Zyail; indicated by Muller, as found at Cape Disappoint-
ment.

210. AMBLYSTEGIUM, Bryol. Eur.

247. H. compactum, Mull.
H. serpens var. compactum, Hook.
Hab. on the ground around springs, Big Tree grove, Bol.; near Fort Col-
ville, Lyall.
248. H. serpens, Lin.
Hab. on the roots of bushes, in swamps, Bo/.; on the ground, near Crescent
City, Brew. ; common in California, big.
249. H. radicale, Beauv.
Hab. Galton and Cascade Mts., Lyall.

According to Mr. Mitten, who mentions this species, which he calls a much mis-
understood species, it differs from H. serpens in its narrower and longer leaf cells. If
we admit the descriptions of Schimper and other authors, and rely for the nomencla-
ture on our American specimens as types, the contrary should have been asserted.
For, indeed, H. radieale, as it is generally understood, has the leaf cells shorter and
proportionally broader, and a longer and stronger costa than H. serpens, Lin. About
the understanding of this troublesome species, I readily admit what Mr. Sullivant says
in his Icones, p. 200: ‘‘ A. orthocladon, H. contextum, H. tenax, H. inordinatum, H.
“varium, attributed to this country only, are (together with a few other supposed
‘species, included H. radicale, found both here and in Europe) so variable in their
‘‘ characters, and pass so gradually into each other, that it appears quite impracticable
“to define their specific limits, or to separate them from large forms of H. serpens, to
‘which they have been referred by Muller and others.”

250. H. orthocladum, Beauv.
HI, varium, Hook. and Wills., in Drummond.
Hab. borders of springs, and among willows, Little Lake Valley, Bol. ; Pend’
Oreille River, Lyall.
H. riparium, Lin.
Hab. Mill Falls, Oakland, and Big Tree grove ; around springs, Bol. ; near
Crescent City, Brew.; wet places, Big.

t
NX
—

¢ 11. CAMPYLIUM, Sulliy.
252. H. hispidulum, Brid.
Hab. Mooyie River, Lyall.

412. HARPIDIUM, Sulliv.

253. H. aduncum, Hedw.
Hab. a swamp in San Francisco, Bol.

eo
~]

OF PACIFIC COAST MOSSES.

254. H. uncinatum, Hedw.
Hab. Bridal Veil Falls, Mono Pass, near Merced River, Big Tree grove,
and foot of Mt. Dana, /ol.; wet banks near King’s River, Brew. ;
Cascade Mts., Lyall.

255. H. commulatum, Hedw.
Hab. east side of Mono Pass, sterile, Lol.
256. HH. filicinum, Linn.
Var. Vallis-cluse.
Hab. Fort Colville, Lyall.

213. RHITIDIUM, Sulliv.

257. H. robustum, Hook.
Hab. Western Coast of North America, Menzies ; Fort Colville, Lyall.

214. DREPANIUM, Schp.
258. H. fertile, Sendt.
Hab. Big Trees, on humected rocks, Gol.
A small specimen only was received. It has the essential characters, the form
of the leaves, the areolation, capsule, ete. But no male flower could be found on the
plants. The species is therefore somewhat uncertain.

259. H. subimponens, Lesqx.
H. plumifer, (2?) Mitten.
Hab. on shaded rocks in woods, from the plain, Oakland, to the mountains,
Big Trees, Long Valley, Bol. ; Vancouver Island, Lyall.
It is still doubtful if this species is the same as that described by Mr. Mitten as
HT, plumifer (among other characters) with opereulo brevi subulato rostrato, and the gen-
eral appearance of A. Crista-Castrensis, Linn. In our moss the lid is conical, some-
times obtuse, sometimes more elongated, pointed, but never rostrate; and its likeness to
H. imponens is so remarkable that it is difficult to separate both species.
260. H. circinale, Hook.
Hab. base of trees, Siskiyou Mts., 4,000 to 5,000 ft., Grew. ; on living red-
woods exposed to winds and fogs, 2,000 to 2,500 ft., Lol. ; Oregon,
Wilkes’ U. S. Exp.; Vancouver Island, Lyall.

215. LIMNOBIUM, Bryol. Eur.

261. H. arcticum, Sommerf.
Hab. foot of Mt. Dana, entirely submerged, in streamlets, apparently very
rare, and seen but once, Bol.

#16. HYPNUM, Schp. Synop.

bo
for)
bo

H. giganteum, Schp.
Hab. Fort Colville, Lyall.
(37)

9

38 LESQUEREUX — CATALOGUE.

263. H. cuspidatum, Linn.
Hab. Pack River, Lyall.

217. HYLOCOMIUM, Bryol. Eur.
264. H. loreum, Dill.

Hab. Oregon, Wilkes’ U. 8. Exp.; Vancouver Island, Lyall, Wood.

265. H. splendens, Dill.

Hab. Oregon, Wilkes’ U. 8. Exp.; Vancouver Island, Lyall, Wood.

(38)

MEMOIRS

PRESENTED TO THE CALIFORNIA ACADEMY OF SCIENCES.

VOLUME I.

Il. Prineiples of the Natural System of Volcanie Rocks.
By F. Baron RiIcHTHOFEN, Dr. Pur.
[Presented, May 6th, 1867.]

| AERO ae Among the features peculiar to modern Geology may be noticed
a revival of that speculative tendency which prevailed among the cultivators of this
science at the close of the last century. But while in those early times imagination
exerted a dominant influence in the framing of hypotheses, and discussions between
the adherents of different doctrines were conducted with all the bitterness peculiar to
such struggles, when neither party has a firm basis upon which to found its arguments,
‘the constant ascendency of the spirit of the inductive method has imparted to those
theories more recently propounded a more logical and scientific form, while, at the
same time, the increasing amount of positive knowledge has given to the different
doctrines a more varied and more definite character, and enlarged the scope of dis-
senting views.

This renewed tendency to systematize and theorize, which is especially con-
spicuous in the records of the last twenty years, must be ascribed, partly, to the vast
amount of well-established facts gathered during the previous decades, and which
have since been multiplied and intensified in a constantly increasing ratio, as regards
depth and distinctness of observation as well as the geographical area over which they
extend ; partly, and in no less degree, to the rapid progress made by those sciences on
which geology has to draw for the general laws which are alone capable of affording a
philosophical guide to speculation on the basis of facts gained by observing and com-
paring. The advance of the chemical and physical sciences, especially, has had a

MEM. CAL. ACAD, SCI. VOL, 1. [2r] Jan. 1868, (89)

2 INTRODUCTORY.

powerful influence, by allowing an immediate application to geological problems of
such general laws as are established beyond any doubt. Besides, the improvement of
the practical methods applied in chemical and physical laboratories has given rise to
the execution of numerous experiments, which were made with a view of imitating the
processes applied in the great natural laboratories. They suggested frequently the
causes of facts disclosed by the examination of a certain region, or obtained by com-
paring the observations made in different countries on one certain subject.

It appears to be mainly due to these causes that a number of theories have
been proposed, in rapid succession, relating to the origin of rocks, to the mode and causes
of metamorphism, to the agencies of vulcanism, to the structure and mode of formation
of mountain ranges, to the structure of the entire globe, and other cognate subjects.
It will be admitted, even by those who are most strongly opposed to theorizing, that
geological science has in this way been promoted and enriched in various respects ;
since there is scarcely a theory which, even if insufficient to explain what it under-
takes, has not some truth in it, or is applicable to some extent in certain cases, or has
at least, even when proving to be erroneous itself, led to discussions on subjects of high
interest, which is indeed a result of no little value. In reviewing the theories pro-
posed on any particular subject, we find them, it is true, often in apparent contradic-
tion with each other; yet almost every one is based upon arguments drawn from
observed facts, and there are probably very few which will ultimately be entirely
abandoned. The immense range of varied processes as applied by nature allows the
applicability, in a limited way, of many a theory in certain instances, while in others
it may be refuted on no less valid grounds ; and the struggle between the defenders of
different doctrines is often founded only in the difference of their standing-points.
What appears to be true in one instance is frequently not applicable in others ; it is
the bold generalizations which render theories so often untenable in that form in which
they are usually first expressed. An instructive example is presented by the different
theories which have been proposed for explaining the mode of formation of mineral
veins. Almost every one of them was based upon a limited range of observations, and
was, from its first application to a few instances, extended to the generality of veins.
Numerous exceptions to it were then found, leading to the rejection of the first, and
the establishment of a different, theory, which, in its turn, shared a similar fate. Ob-"
scure as this subject still is, we are able to state this as certain, with our present state
of knowledge, that every mineral vein is the product, not of one simple but of complex
processes. Nearly every one of the theories proposed will, therefore, have its limited
range of applicability, inasmuch as the agent it suggests may have been especially
active in the formation of the veins of a certain order, while the same agent may have
played a subordinate part in regard to the origin of other veins which were chiefly
due to processes of another kind.

This instance points clearly towards the one principal cause of the divergence
of opinions in regard to some of the most important geological questions. This cause
is the want of latitude of the basis upon which arguments are founded. Conclusions
which are obtained by reasoning on geological subjects solely on the strength of chem-
ical analysis, are, when generalized, often found to be utterly in discordance with the
(40)

INTRODUCTORY. 5

facts revealed by geological observation ; and how unsatisfactory general theories may
be when based upon the latter alone, is sufficiently exemplified by the fantastical
attempts made in all ages of geological science to interpret the geological structure of
the world from that of a limited region. The basis for argumentation can therefore
never be broad enough, and its enlargement should be, as it indeed is, one of the
chief objects of geological science. But it is not sufficient to content ourselves with
an accumulation of primary observations, which are in fact being infinitely increased
by the conjoint labors of geologists in all countries: it should be a higher object of the
student of geology, to compare the established results of observation, and to investi-
gate their mutual relations. The study of the structure of one mountain range, or of
several ranges comprised within a limited district, may lead to the establishment of an
elaborate theory of the mode of their formation, which may apparently answer per-
fectly well in that one case, but may be found inadmissible when generalized, even in
those cases where, by imperfect observation, one would expect to detect a great simi-
larity to the structure first observed. But in determining those features which are
common to a number of mountain ranges, or to certain orders of them which we may
discern among their generality, we may aspire to form conclusions which are more
generally applicable. It is particularly the auxiliary branches of geology to which
these remarks apply. The value of observations made in limited regions, or from a
limited point of view, on subjects such as the outlines of the morphological feat-
ures of the continents, the occurrence of mineral springs, the structure of mineral
veins, the age of those among them which carry a certain metal, the generality of vol-
canic phenomena, the mode of action of earthquakes, the nature of certain kinds of
rocks, and their part in the structure of the surface of the globe—cannot be fully real-
ized unless the comparative method is applied in as wide a scope as we may be able to
do, and the mutual relations among the different modes of manifestation of force, or
among the properties of the kinds of matter upon which it acts, or the bearing of all
these relations to each other and to the evolution of the globe, are investigated from
as many points of view as we may detect, and in as many combinations as possible.
We may then be able to gain a foundation for argumentation on more involved prob-
lems, consisting not of imperfect premises, nor of a confused accumulation of facts,
but of established truths of a higher order.

The mode of origin of the non-foliated crystalline rocks, made up of silicates,*
is among those subjects which have at all times, but at no time more than of late,
commanded a great deal of attention, and given rise to the establishment of numerous
theories, each of which was applied in a general way, if not by its author, then by his
followers. It is well known how conflicting they apparently are, and what weighty ar-
guments have been brought in favor of as well as against each of them. The only
method, promising success, of weighing the merits of these different theories, or of
modifying them in accordance with the general advance of science, appears to be, to

* [have for these applied the name “eruptive rocks” in the following pages, considering that, wherever we have
occasion to observe them, they are not at their original seat, but ejected from it towards the surface. The reasons supporting
this position will be more fully mentioned in the chapter ou the origin of voleanic rocks.

(41)

{ INTRODUCTORY.

ascend from the examination of the nature of these rocks to that of their mutual
relations, to investigate these from as many points of view as we can discover, in re-
gard to physical and chemical properties, mode of occurrence and age, as well as in
regard to geographical distribution: that is, to try to establish the natural system of
eruptive rocks. The results so obtained may then, in accordance with what we just
remarked in a general way, be applicable to reasoning on remoter questions, of
which we can only attempt to find the most probable solution. They regard chiefly
the causes of those relations, the mode of origin of the eruptive rocks, and the
processes connected with their ejection. The intricate nature of the subject, and
the fact that the present changes on and below the face of the globe, as well as
the events of the past, are often but dimly and imperfectly perceptible to our ob-
servation, demand that we should concentrate our endeavors in exploring first the
laws of that which is definite and constant within the infinite range of phenomena, and
await further experience to arrive at an explanation of those isolated facts which form
apparent exceptions to the order of things.

It is with these views that the following pages were written. They extend
chiefly over the comparatively limited, and yet very extensive class of ‘‘ volcanic rocks,”
and are offered as a mere elementary attempt, which is necessarily very imperfect.
The application to exact reasoning of the numerous observations which have been made
on the subject of volcanic rocks in different countries, is nearly prevented by the ex-
traordinary discrepancy existing in regard to the mode in which the names of rocks
are used by different authors. The first condition of a uniform and harmonious mode
of observation on voleanie rocks, and the phenomena connected with them, is the ap-
plication of a uniform system of nomenclature.

In concluding these preliminary remarks, I dare express the hope that some in-
dulgence may be had with the imperfections of this essay, if it is taken into considera-
tion that it was written on the Pacific coast, where chemical laboratories are unknown,
libraries scarce, and little opportunity is afforded of becoming acquainted with the
current geological literature. I fulfill a deep-felt duty if I tender at this place my
sincere thanks to Professor J. D. Whitney, not only for allowing me the use of his
library and revising the manuscript of this essay, but also for the interest which he has
constantly taken in my pursuits, and for what I owe to his personal intercourse, especially
in a country where scientific communication is so extremely limited. The influence
of this intercourse will, long after this, be kept in grateful memory by all those who are
taking personally a part in the development of the California Academy, the members
of which kindly allowed this paper to be published in their Memoirs.

(42)

THE NATURAL SYSTEM OF VOLCANIC ROCKS.

In reviewing the various attempts which have been made towards a classifica-
that is, those crystalline rocks made up of silicates, which,

tion of eruptive rocks
without showing themselves any traces either of stratified deposition or foliation, enter
into the structure of the surface of the globe in such a way as to be unconformable
with the stratification of the neighboring sedimentary rocks, and as a rule to abut
we are struck by the observation that, if

against them without any gradual passage
they are based on any principles at all, these are usually artificial, while none but un-
satisfactory results have been obtained when the application of natural principles has
been tried. This want of success is the more striking if we consider that it is peculiar
to petrology, and that the efforts made in the same direction with other branches of
descriptive natural sciences have been attended by extraordinary results. In zodlogy
and botany, the natural system has long since been considered as the ultimate object
of scientific research ; and since the time when its first outlines were discovered, the
progress of these sciences has been admirable. Since then only have the developments
of their different branches codperated harmoniously towards one common end: the
profoundest investigations into the anatomy of animals and plants, the study of their
geographical distribution in modern time, and of their gradual development in past
ages, have in their final results but been subservient to the establishment of a founda-
tion of the natural system, and the ingenious deductions made by Mr. Darwin on the
origin of species are but its philosophical interpretation. As regards mineralogy, clas-
sification was for a long time a simple enumeration of minerals, governed by cer-
tain artificial principles. A new era was inaugurated for this science by the progress
of chemistry, and its application to mineralogy, by Berzelius. It led to a more correct
estimate of those principles which had been formerly applied, and to the discovery of
the existence of an intimate connection between crystallographical form and chemical
composition. The combination of these two principles gave rise to the natural system
of minerals, which since their adoption has been constantly gaining in completeness.
These are results which surpass in a surprising degree those obtained in regard
to the natural classification of eruptive rocks. Even the most recent and elaborate sys-
tematical arrangements, as those proposed by C. F. Naumann, F. Senfft, B. v. Cotta,
and J. Roth, though marking a conspicuous progress, are based on almost purely arti-
ficial principles. In no other branch of the descriptive natural sciences, it is true, do
difficulties arise so great as those which present themselves in petrology. Prominent

MEM. CAL, ACAD, SCI. VOL. I. F Jan. 1868. (43)

6 RICHTHOFEN — THE NATUR SYSTEM

among these is the entire absence of what we could call “ genus,” or ‘‘ species,” not to
mention ‘ individual.” If we should succeed in discovering some natural group to
which we might apply the term ‘‘ family,” (though even this can never be used in
petrology in as definite a sense as it is in the organic kingdoms) we should find
it made up of an infinite number of varieties; and if we should be able to
establish several groups, the main types of which are conspicuously distinct in
nature, we should find them linked together by gradual passage in chemical and min-
eral composition. It appears indeed utterly impossible to draw distinct boundaries
between cognate groups. Certain names, such as those of granite, syenite, quartzose
porphyry, trachyte, basalt and others, have been applied to designate distinct types of
crystalline rocks, which can easily be recognized wherever met with. But, practically,
they have to be used for larger groups of rocks, in which those distinct types appear
like luminous centers, surrounded by clouds of varieties blending with each other in
such a way as often to render it arbitrary whether to bring a certain rock within one
or the other denomination. But there are other groups of rocks, of larger dimensions
than the former, the nomenclature of which is far more indistinct, and which, in regard
to classification, may indeed be said to be still in an entirely nebulous state. The
vague and arbitrary mode in which different names are used for them shows plainly
that difficulties in regard to them are greater than with other rocks. As will be seen
in the sequel, this indistinctness of external character, as wellas of designation, applies
particularly (with the exception of basalt) to those rocks in the composition of which
silica takes a less prominent part, while those which are richer in silica offer much
more distinct characters. Most conspicuous among the names applied for the former
are those of “trap,” ‘‘ greenstone,” and ‘ porphyry.” The latter two may be con-
veniently used to designate groups of rocks having certain external characters in com-
mon, but as generic terms they should all be completely abolished. They never con-
vey a definite conception, as each of them is used for a great variety of rocks, and
they have only too often been made to serve as a convenient cloak to cover ignorance.

The perception of these difficulties has caused the idea to be almost universally
accepted, that only an artificial system of eruptive rocks can be established ; that is,
that classification should be made dependent on one certain principle previously as-

1This applies chiefly to the term “Trap,” (or trapp) which had originally a definite meaning, but has gradually been
extended with wonderful elasticity. It was first introduced by Torbern Bergmann for a very ancient, dark colored, augitic
rock of Uddevalla, in Sweden, which is arranged in superposed layers abutting against the slope of the hill in the shape of a
stairway (trappar in Swedish). This rock would be called “diabase” in modern nomenclature. The name “ trapp’” was
then applied to other, and gradually to all, dark colored eruptive rocks, particularly to such as were found occurring in
dykes ; afterwards the “ greenstones” (which name, too, has shown itself capable of wonderful distension) were included in
that denomination; and finally it has become still more comprehensive, though not quite to the same extent with every au-
thor. Its present meaning may best be seen from the following passage by Lyell, (Elements of Geology, 6th Hd., 1866, p.
601 of Am. Ed.): “This term (lava) belongs more properly to that (melted matter) which has flowed either in the open air
or on the bed of a lake or sea. If the same fluid has not reached the surface, but has been merely injected into fissures
below ground, it is called trap.” Thus, the same name which was originally applied to a distinct rock of ancient origin has
been generalized so as to express now a mode of occurrence, and, what is more remarkable, a mode of occurrence the very
reverse of that exhibited by the original type, which must be supposed to have been flowing on the bed of the sea, and the
position of which in relation to neighboring rocks can never be explained by assuming it to have been “ injected into fissures
below ground.” Would it not be better to drop such a name altogether ?

(44)

OF VOLCANIC ROCKS. 7

sumed as the point of issue. Taking crystalline texture, lack of stratification as well
as of foliation, and the fact of their being made up of silicates to be the characteristic
features of eruptive rocks in general, the most obvious external differences among
them are caused by the variations of texture and color. During the early stages of
petrographical science, rocks were therefore classified on these principles. The terms
trap, porphyry, pearlstone, obsidian, lava, amygdaloid, wacke, as well as green-
stone, black porphyry, green porphyry, and others, are the remnants, in our present
nomenclature, of that epoch when the more minute differences of rocks arising from
their mineral composition were but imperfectly investigated. This principle was
necessarily next in order for serving as point of issue for classification, since, as far as
regards external characters, it is only second to the former in value. The emer-
gence of petrology from a chaotic state, by the scientific application of this prin-
ciple, dates from the investigation of Gustav Rose on the feldspathic minerals entering
into the composition of rocks, and it has since been more generally applied for
establishing subdivisions than any other. The presence or absence of quartz, the
predominance among the feldspathic minerals of orthoclase, oligoclase, or labrador,
the presence of augite or hornblende, are the usual points of issue, even in the most
recent attempts at classification. The high value of mineralogy as a basis of classi-
fication cannot be denied. But its exclusive application has caused the combi-
nation into certain groups, of such rocks as from a geological point of view are
widely separated, while it has given rise to distinctions in cases where the results of
geological observation would demand close connection, as we shall have occasion to
illustrate in the following pages, with reference to those volcanic rocks which are com-
posed of hornblende and oligoclase. Gradually, those differences based on chemical
composition, not capable of being detected by the eye, and the knowledge of which
could only be obtained after chemistry had made the necessary advauces, have become
an object of scientific research. But this principle has not yet been used to any great
extent for classification. It can easily be demonstrated that, when exclusively ap-
plied, it leads to a systematical arrangement of rocks which is in even greater contra-
diction with the natural mode of occurrence than when the same is based upon
mineral composition alone, notwithstanding the fact that its great value has been con-
clusively demonstrated, especially by the important results which Bunsen obtained
from the chemical analysis of rocks, and which mark an era in petrology. To com-
bine granite, quartzose porphyry, and rhyolite into one class, because they resemble
each other in their chemical composition, and to place them at the head of the list
because containing the highest amount of silica observed among eruptive rocks, would
be to take no regard whatever of geological facts. Rhyolite is, mineralogically and
geologically, far nearer related to trachyte than to either granite or quartzose por-
phyry ; and these two are quite distinct from each other, while granite is closely allied,
by gradual passage, to syenite, and quartzose porphyry to porphyrite.

It is by slow degrees only that we can hope to reach a more scientific, that is,
a more natural system in this, the most intricate branch of descriptive natural sciences.
The natural differs from the artificial system in this, that it starts from the application

(45)

8 RICHTHOFEN — THE NATURAL SYSTEM

not of one only but of various principles, compares and weighs the results obtained by
each of them, and accepts them as final only when perfectly harmonizing among each
other. Itis then that it tries to determine what principles are most available for
establishing the higher orders, and which for the subdivisions. The singular complica-
tion which is peculiar to the classification of rocks, is, besides the reasons already
mentioned, due in a great measure to the fact that geology combines the double func-
tions of a historical and an inductive science, while in petrology we have besides the
requirements of a descriptive natural science. The natural system of rocks should
therefore be based, not only upon the entire range of their petrographical characters,
such as mineral composition, chemical composition, texture, and specific gravity, but
also upon their mode of origin and geological occurrence. Classification of objects
and classification of relations are, with them, closely connected, and should be made
to assist each other.

The question may be raised, whether a natural system of rocks based upon
such principles can be established at all, and if it can, whether it would be of any use
for the advancement of science. To the first question, the answer must be in the
negative, as far as sedimentary rocks are concerned. They have been formed by a
complexity of circumstances, and just so complex and infinite in variety are they, in
respect to chemical and mineral composition and all external characters. To analyze
in detail their mode of origin, and the sources from which their material has been
derived, transcends the faculty of human intellect, and it would be a hopeless task to
attempt to discover any laws regulating the boundless differences of their composition.
They are thus debarred from natural classification, though its principles may be applied
imperfectly to the establishment of some general groups. We arrive at similar con-
clusions in regard to those rocks, the sedimentary origin and subsequent metamorphism
of which can be proved. Accidental and local circumstances have played as conspic-
uous a part in their first deposition as was the case in regard to those sedimentary
rocks to which the term metamorphic has not been usually applied. But as meta-
morphie processes of a certain nature have ordinarily affected extensive tracts of
these rocks, and similarly pervaded great thicknesses of them, the local differences of
their action having been apparently more in degree than in mode, they have occa-
sioned a certain similarity of effect which partly conceals the original differences in
the composition of the rocks affected ; and it appears that the differences in the kind
and intensity of metamorphic action, though recognizable only in their final results,
will, when better known, afford a convenient principle for a classification which may
have some similarity with, but not the full requirements of, the natural system. It is
different with those rocks which on the surface of the globe appear as intrusive or
eruptive masses. Notwithstanding their infinite variety in character and composition,
they are connected by definite relations which bring their elementary composition
even within range of mathematical calculation. Their recurrence in the most widely
separated countries, with similar external character, identical chemical composition,
and in analogous relative order of succession, is another distinguishing feature of
eruptive rocks. For these reasons, as well as in virtue of other peculiar characters

(46)

OF VOLCANIC ROCKS. 9

which will be more fully mentioned in other pages, they appear to owe their present
positions to the action of general planetary processes, and to reveal by their own
nature that of the mineral matter participating in the original composition of the
globe, and by their order of succession, the mode in which the same is arranged
beneath the theater of those changes which since a remote period have been taking
place on its surface. Only presumptive evidence can be adduced in favor of this
common and yet much disputed theory. The probability of its approximating the
truth could hardly be better established by any other evidence than by the proof that
all eruptive rocks of the globe, taking their historical part into account, are capable
of being brought into a natural system; or, to express it more correctly, that they
form among themselves a natural system, the laws of which we may be capable of dis-
covering. Considering this in its widest bearing, as embracing all the definite correla-
tions of eruptive rocks, and being indeed their philosophical expression, we may
expect that it will make us acquainted with the history of one great feature in the
development of the globe. Bearing in their own character and system the imprint of
their origin, the eruptive rocks will, by their nature itself, allow well-founded con-
jectures as to the interior structure and composition of the earth. This, then,
together with a more perfect understanding of everything connected with the agencies
working below the surface of the globe, would be the philosophical use of the natural
system of eruptive rocks.

Only initiatory steps can be taken at the present time towards the establish-
ment of this system. I have confined myself in this essay to an attempt at classifying,
in a way as natural as experience will allow, the ‘ volcanic rocks,” that is, the eruptive
rocks of Tertiary and Post-Tertiary ages. The term ‘‘ volcanic rocks,” has been chosen,
because the rocky matter ejected by active voleanoes belongs altogether to this class,
and because almost every kind of rock, generated by eruptive activity during the
period indicated, has partially forced its way through volcanic vents. We must keep
the two-fold mode of occurrence of volcanic rocks clearly separated in our minds. We
see them at the present day flowing from craters in the shape of lava, or being thrown
out as scoria and rapilli; and there is abundant evidence that their mode of origin
was very frequently the same in past ages. But in other places, it is perfectly clear that
volumes of matter of the same kind have been forced to the surface through extensive
fissures and accumulated above them in elongated ranges, when the origin of the out-
breaks cannot be ascribed to voleanic activity. These eruptions are evidently similar
in nature to those by which the greater part of the granite, syenite, or quartzose por-
phyry ascended to the surface in ancient times. We shall distinguish the two modes
of eruptions, for the use in this present paper, as ‘‘ volcanie eruptions” and “ massive
eruptions,” and shall dwell in the sequel more fully on the difference between both
manifestations and their probable causes. Then, too, the reasons will be mentioned
which justify the uniting of all recent eruptive rocks into one separate class.

No other class of eruptive rocks offers greater difficulties for a systematical
arrangement, as none presents throughout so great a number of varieties, and so many
accidental modifications of texture and mineral composition. On the other hand, their

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10 RICHTHOFEN — THE NATURAL SYSTEM

classification is of especial importance, as it furnishes the key for deciphering the
natural system of the ancient eruptive rocks. Volcanic rocks issue from volcanic
vents under our very eyes, and the record of the history of those which have origi-
nated in past ages, as preserved in their geological relations, is far more distinct than
it is in regard to their ancient predecessors. No doubt exists in the mind of any ob-
server in respect to the eruptive origin of all basalt ; while in regard to granite, very
different views are entertained by distinguished geologists, and supported by weighty
arguments. In the former case, we have conclusive evidence, while in the second
speculation has a wider scope. The knowledge of volcanic rocks will, for this reason,
facilitate the correct interpretation of the nature of rocks generated in remote ages.

Volcanic rocks are widely spread over the face of the globe. It would be an
object of great interest to lay down their geographical distribution on maps, and to
explore the laws by which this distribution has been governed. This has been tried
in regard to active volcanoes, and the importance and interest attaching to the results
obtained are such as to have given rise at once to speculations as to the connection
between volcanoes and other phenomena. The value of those results would be in-
creased, if to the active craters were added the vastly greater number of those extinct
volcanoes, the mode of preservation of which still allows us to recognize their former
nature. Even then, however, the maps would convey but a remote idea of the gen-
eral distribution of voleanic rocks; the areas comprised by which should be marked
out with proper distinction of their main subdivisions. Besides an immediate bearing
on more special geological questions, the knowledge of these subjects promises to be
of high value for that entire department of geological science, by which the latter is
most closely connected in scope with the science of physical geography. Many weighty
problems, such as the causes of the present direction and extent of mountain ranges,
of the outlines of continents, of the position and shape of groups of islands, of the
secular oscillations of the surface of the globe, and many other questions, appear to be
intimately connected with the geographical distribution of voleanic rocks and their
mutual geological relations. The deduction from the latter of definite laws appears
to be the initiatory step towards understanding the laws of eruptive activity of
remote times, and, thereby, towards establishing a chapter in the history of the globe,
which is among the obscurest and least understood.

The following classification, in which existing names are retained, as nearly as
could be done with convenience, is chiefly founded on observations made in the Carpa-
thians and in the States of California and Nevada. In respect to the variety and the
distinctness of the mutual relations of the volcanic rocks, these two countries are hardly
surpassed by any in which this subject has, up to this time, been scientifically investi-
gated. Until recently, all voleanic rocks, at least those of more frequent occurrence,
were comprehended in the terms: trachyte, phonolite, trachydolerite, dolerite, basalt ;
while, besides, separate names were used to distinguish modifications of texture, such
as pumice-stone, obsidian, pearlite ; or varieties somewhat more distinct in point of min-
eral composition, such as leucitophyre. The classification as given by Al. von Humboldt,
in the fourth volume of the ‘‘ Cosmos,” may be considered as having represented the

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OF VOLCANIC ROCKS. 1]

most advanced stage of the science a few years ago, because it was guided by a definite
principle ; though when compared with contemporaneous works on petrology, it gives,
on the other hand, a remarkable illustration of the great difference in the way in which
the same names have been applied. Since then, the name ‘rhyolite ” (Richthofen,
Studien aus den ungarisch-siebenbiirgischen Trachytgebirgen, in Jahrbuch der K. K.
geologischen Reichsanstalt in Wien, Vol. XI [1860], pp. 153-277) has been introduced
for a very distinct class of volcanic rocks. Adding this to the previous list, there
results a number of names which, in geological treatises, are either grouped com-
pletely at random,? or in an arbitrary order, or arranged by artificial principles, when
the whole classification ordinarily comprises volcanic and ancient eruptive rocks pro-
miscuously. In order to establish a more natural system, we have, not to make groups,
but to find them. Dropping all of those @ prior? principles which may be conceived
having an artificial basis, we must endeavor to discover whether any great divisions
are established by nature herself, and if so, of what character they are. We may
then apply, as second in the order of their importance, those results which are obtained
in the laboratory or geological cabinet, for defining and subdividing those groups.
Most of the natural divisions which may be derived from geological observation, coin-
cide essentially with those based on artificial principles, but are more naturally
limited as regards each other. Hach of them has its own more or less independent
part in the architecture of mountain ranges, and a distinct geological age in reference
to the other groups. Hach of them comprehends a series of rocks, which, besides, are
closely connected by the relations of their petrographical characters, chemical compo-
sition, texture, specific gravity, and other properties. The test of the natural founda-
tion and general validity of these groups will be their recurrence, with mutual rela-
tions unchanged, in different parts of the globe, of which test we are never to lose
sight.

The following is the classification, the approach of which to a natural system
of volcanic rocks, I will endeavor to set forth in the course of this paper :

Orver First: Rhyolite.
Family 1. Nevadite, or granitic rhyolite.
‘2. Liparite, or porphyritic rhyolite.
3. Rhyolite proper, or lithoidic and hyaline rhyolite.

ce

Orper Seconp: Trachyte.
Family 1. Sanidin-trachyte.
‘2. Oligoclase-trachyte.

2 That this is even done in books of the highest standard, may be seen by reference to one so prominent as Lyell’s
Elements of Geology. The following is the order of names of which, under the head of “volcanic rocks,” definitions are
given (p. 592, pp. of 6th Am. ed., 1866) : Basalt, augite rock, trachyte, trachytie porphyry (in connection with which the
name “andesite” is mentioned), clinkstone, greenstone, porphyry, amygdaloid, lava, scoris or pumice, yolcanic tuff or trap-
tuff, agglomerate, laterite.

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ee RICHTHOFEN — THE NATURAL SYSTEM

Orper Tuirp: Propylite. _
Family 1. Quartzose propylite.
“2. Hornblendie propylite.
3. Augitic propylite.

ce

OrpvEerR FourtH: Andesite.
Family 1. Hornblendic andesite.
“2. Augitie andesite.

Orpver Firtu: Basalt.
Family 1. Dolerite.
ee Basalt
“3. Leucttophyre.

Orper First—RuyYOLITE.

‘

The name “rhyolite” was proposed, early in 1860, for certain rocks fre-
quently occurring on the southern-slope of the Carpathians, and distinguished, in min-
eral character, from trachyte, which they otherwise resemble, by the presence of quartz
as an essential ingredient, and an almost infinite variety of texture. Beudant* had,
long before, described certain varieties of these rocks as porphyre trachytique, pumice-
stone, pearlite, ete. In 1861, the name “‘liparite” was proposed by J. Roth® for rocks
of similar nature occurring on the Lipari Islands. The term ‘‘ rhyolite,” however,
being of prior date, has since been almost generally adopted, among others by F. v-
Hochstetter, for rocks from New Zealand, by C. Peters, G. Stache, and others for those
of Hungary and Transylvania, by Ferd. Zirkel for those of Iceland, by B. v. Cotta as
a general term in his ‘‘ Gesteinslehre.” The word ‘‘rhyolite” is designed to express
one of the prominent features of these rocks. It is this: that their chief varieties
have the appearance, as it were, of natural glasses, and bear evidence, more than any
other rocks do, to the unpracticed eye, of having been flowing in a viscous state.

Mode of Geological Occurrence —Rhy olite has had its distinct epoch of eruption in
relation to other volcanic rocks. Wherever it occurs it may be easily proved to have
been of more recent origin than either propylite, andesite, or trachyte, but to have
preceded basalt in age. As to its geographical distribution, it is confined to the im-
mediate neighborhood of one or all of those antecedent rocks, and oceurs ordinarily
within their very limits. Until lately, not much attention had been paid to it. But
since the establishment of the name, rhyolite has been found to be widely distributed,
though always occupying a subordinate position. In Hungary it usually skirts the
lower part of the flanks of andesitic ranges, forming hillocks and ridges of little
elevation, fillmg depressions, and issuing in currents from fractures, and, in general,
giving evidence of its entire dependency on the places of previous eruptions.
The greater portion of the rhyolite has, in that country, been evidently ejected
by volcanic activity. It appears that the same may be said in regard to the

3 Loe. Cit. 4 Voyage en Hongrie. Paris, 1820. 5 J. Roth, die Gesteins-analysen. Berlin, 1861.
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OF VOLCANIC ROCKS. 13

rhyolitic rocks of New Zealand, St. Paul, and Iceland. But it has not been
so generally the case on the western coast of America. The voleano of Lassen’s
Peak, and the environs of Mount Helena, in California, present graud instances
of a volcanic origin of rhyolite. But in the adjoining State of Nevada it ap-
pears to have been extensively brought to the surface by massive eruptions. It is of
unusually frequent occurrence along the eastern slope of the Sierra Nevada, and far-
ther east in the Great Basin. Ranges of hills are there completely built up of rhyo-
litic rocks, not always in as close proximity to such as are antecedent to them in age,
as is the case in the Carpathians.

It is one of the characteristic features of rhyolite, that it presents, more than
any other rock does, signs of having been in a state of what Daubrée has called
“aqueous fusion,” or the fusion of its mass by solution, under great pressure, in super-
heated water. Another peculiarity is the circumstance that the eruptions of rhyolite,
whether massive or volcanic, bear evidence of having been generally accompanied by
extremely violent solfataric action, which probably surpassed, on an average, that con-
nected with the ejection of other volcanic rocks. This action appears to have been
one of the chief agents in the formation of the rich silver-bearing veins of Hungary,
as well as of some in Mexico, and to have also been peculiarly characterized by the
occurrence of an unusually large amount of fluorine and chlorine among the escaping
gases.

Mineral Composition.—Rhyolite may be concisely defined as trachyte with an
addition of silica, not chemically combined, and which is either segregated into crystals
of quartz, or dissolved in the rock, and then no longer recognizable to the eye. It is,
owing to the high proportion of silica entering into its composition, the representative
of granite among voleanic rocks. Chemically, it is its complete counterpart, as far as
ascertained by analysis, and even in outward appearance certain varieties of rhyolite
offer at first sight a striking similarity with granite, though closer observation will at
once reveal well marked differences between both. Rhyolite outrivals any other rock
in respect to the truly astonishing number of its varieties, which are chiefly occasioned
by modifications of texture. It consists in general of a paste, with or without minerals
enclosed.

The paste, chiefly, is liable to variation. Its colors are: white, gray, yellow,
green, red, brown, which occur in all manner of shades; light ones prevail, while
perfect black has not been met with. The texture is as varied as the color. First
are to be noticed a number of hyaline varieties, which are represented by obsidian,
pumice-stone, and pearlite, and the frequent occurrence of which is a peculiar feature
of rhyolite. Though associated with volcanic rocks of every composition, these natu-
ral glasses, as they may be called, decrease in relative quantity and variety with the
decrease of silica.6 Obsidian when having the composition of rhyolite, offers little

g There is no better example of the artificial principles on which the classification of rocks has usually been
based than the fact, that accidental modifications of texture, which appear to result chiefly from the difference of the condi-
tions attending either the fusion or the cooling of the mass, have been considered as of equal value with other differences of
the greatest importance ; and pumice-stone, trachyte, basalt and pearlite have been considered as codrdinate subdivisions,
even long after classification had been made dependent chiefly on mineralogical principles.

. (51)

14 RICHTHOFEN—NATURAL SYSTEM

difference in character from that which is a modification of other voleanic rocks. In
regard to pumice-stone, however, Abich has proved that when formed of the material
of trachyte or andesite it has rounded pores, and ordinarily a green tint ; while those
' varieties which have the composition of rhyolite, excel by the elongated and irregular
shape of their cavities, which are enclosed in a fine tissue of fibres of silken appearance
and white color. Between both kinds of pumice-stone there are gradations apparently
dependent as to their character on the amount of silica entering into their composition.
Pearlitic texture is peculiar to rhyolitic rocks. From these more or less perfectly glassy
varieties there are gradations to the texture of enamel and porcelain, and to a certain
cryptocrystalline texture very frequent among volcanic rocks in general, and for which
we may apply the obsolete term ‘‘lithoid.” This passes into the microcrystalline, and
always more or less vesicular, texture of trachyte.

The paste constitutes occasionally alone the substance of the rock. But these
instances are rare. More frequently it contains enclosed mineral substances differing
from it in nature, and in a few instances these accumulate to such a degree as almost
to exclude the paste. Quartz is of the most general occurrence among those which
are crystallized. Sanidin is its almost unfailing companion. Oligoclase, usually of a
vitreous variety, and black mica, are, too, among the usual ingredients, while horn-
blende is generally less conspicuous. Besides these minerals, there are two substances
entering accidentally into the composition of rhyolite, which are, however, among the
characteristic features of the rocks of this order. One of them consists in small ,glob-
ular grains, from the size of a pin-head to that of a rifle bullet, called ‘‘ sphaerolites ”
by Beudant. They have a radial structure, and contain ordinarily a small crystal of
feldspar in the center. Certain hyaline, and, in a greater measure, lithoid varieties
of rhyolite contain them in large quantity. They occur also, though less frequently,
in other natural glasses not of rhyolitic composition, and may be produced artificially,
by allowing molten glass to cool very slowly to what is known by the term “ Réau-
mur’s porcelain.” The second formation frequently met with in rhyolitic rocks are
the ‘“‘lithophyse,” consisting in larger and smaller cavities filled by a substance
strangely inflated by some gaseous evolution which apparently originated in the mat-
ter itself (Richthofen, 1. c.). They constitute sometimes nearly the entire mass of
the rock.

The endless varieties of rhyolite appear to be due to the susceptibility of the fluid
mass to be influenced by accidental circumstances to which it may have been exposed,
partly before being ejected, and partly during the process of solidifying. A consid-
erable influence, which, however, has not yet been investigated, is probably exercised
by the difference in the amount of water which entered into the composition of the
molten mass, and partly expanded to steam in the instant of ejection. The vesicular
inflation proper to trachytic texture, the spongy inflation of pumice-stone, and the
concentric separation of infinitely fine lamin, as is often shown in perfect pearlitic
texture, are probably three different modes of manifestation of one slightly varied
eause, which may most likely be found in the conversion of water into steam, which
participated in the composition of the molten mass.

(52)

OF VOLCANIC ROCKS. 15

Difference of Rhyolite from other Rocks nearly related to t.—Several varieties of
rhyolite bear so close a resemblance to other rocks, that some mention must be made
of their distinguishing features. Rhyolite may be easily distinguished from granite. The
varieties by which it mostly approaches the same are those which contain crystals of
quartz, feldspar and mica in unusually large proportion and size. But in the case of rhy-
olite, the paste in which they are imbedded is never wanting. Moreover, the orthoclase
and oligoclase are of the vitreous varieties, and quartz is present either in crystals or in

‘rounded erystalline grains, while in granite it usually permeates the interstices between

the other component mine rals. Much closer is the affinity which certain other varie-
ties of rhyolite bear to quartzose porphyry, especially those which have a paste of
homogeneous appearance containing no crystals but those of quartz inclosed. Geo-
logical observation will never fail, in such instances, to establish the nature of the
questionable rock, as it will show its association either with true rhyolite or with
true porphyry. The same test has to be applied occasionally with respect to some
other varieties which contain the silica equally diffused through the paste, and bear a
close resemblance to trachyte. In this instance, however, even geological observations
will sometimes fail to determine the exact position. There is a gradual passage in
character between every two nearly related rocks, such as rhyolite and trachyte, or
granite and syenite, and it frequently happens that either name may be used with
equal right.

Subdivisions —In establishing the subdivisions of most orders of eruptive rocks,
mineral composition affords a principle, not only the most convenient for practical
application, but one that answers well the requirements of the natural system, when
made subordinate in value to those higher principles which determine the limits of
classes and orders. In the case of rhyolitic rocks, however, it is not as applicable as
in that of other orders. They should, from this point of view, be subdivided into
those which contain quartz and those which are devoid of it, or into such as carry
sanidin and such as contain both sanidin and oligoclase. But, since rhyolite of any
certain chemical composition may contain its surplus of silica, either visibly segregated in
crystals of quartz, or dissolved in the mass of the rock, and as the case may be similar
in regard to the occurrence of either species of feldspar, the application of this princi-
ple would lead us to combine into one group quartzose rotks differing considerably
among themselves as to the proportion of silica they contain, while another group
might comprehend rocks of virtually the same nature as those of the first, and differing
from them only accidentally in external character. More natural subdivisions of rhy-
olitic rocks are obtained by taking as a basis of classification their difference in texture,
which either approaches, to a certain degree, that of granite or is porphyritic or hya-
line. It is a singular fact, and one difficult of explanation, that rhyolite, at every place
where it has been hitherto observed, presents, either solely or chiefly, one of those
three modes of texture. lLassen’s Peak, for instance, presents the granitic variety
almost exclusively. Sonoma, in California, and the Tokay Mountains in Hungary,
only the hyaline, and other places exclusively the porphyritic varieties. This circum-
stance, which is peculiar to none but rhyolite among eruptive rocks, indicates the

es

16 RICHTHOFEN—NATURAL SYSTEM

dependency of the mode of texture upon deep-seated influences which acted at the very

source of the eruptive matter, and produced a certain molecular condition of the latter,
varying at each locality. Some light on this subject may be expected from minute

geological observation, accompanied by exact chemical and microscopical researches.

The following are the subdivisions which may be distinguished in regard to the
texture ;

Pani: 1st. Nevadite or Granitic Rhyolite—The name ‘‘ Nevadite ” is derived from
that of the State of Nevada, where these rocks have ,been first met with in larger
accumulations. The local derivation may answer in this instance, as granitic rhyolite
is little known from other countries, excepting the neighboring California. In the
Carpathians, it occurs isolated in Transylvania, but by no means as characteristic as in
Nevada. The name ‘“ granitic rhyolite” is designed to indicate the general resem-
blance of these rocks to granite, which is conspicuous in boulders, or on large exposed
faces, but disappears on closer examination. It is chiefly produced by the similarity
in color, which is of light shades of gray and red in Nevadite, and by some affinity
in mineral composition. Nevadite contains crystals of quartz in large proportion ; the
corners are usually rounded, and the quartz itself cracked, like glass when rapidly
cooled. Sanidin occurs in erystals of larger size than oligoclase, sometimes of an inch
in diameter. The crystals of both are often cracked throughout their mass, and
rounded at the corners. Black mica and hornblende are ingredients of frequent occur-
rence. These minerals are, in more or less quantity, enclosed in a paste which is
probably a partially Siocon elltag. and partially amorphic aggregation of the same
ingredients, and has a highly vesicular texture, rendering it rough to the touch, even
more so than is the case with trachyte. Geologically, Nevadite appears to have been
produced as frequently by voleanic activity as by massive eruptions. An interesting
occurrence is that at Lassen’s Peak, in California, where it was discovered by Prof. W.
H. Brewer and Mr. Clarence King.

Fam. 2d. Liparite, or Porphyritic Rhyolite—The name “ Liparite,” which was pro-
posed by J. Roth for this whole class, may be conveniently retained for those varieties
of rhyolite which approach quartzose porphyry in character, as they appear to occur
on the Liparic Islands, either solely or at least in larger proportion than other varieties.
They consist of a paste which has a similar texture to that of quartzose porphyry, and
incloses crystals either of quartz only, or of quartz and sanidin, or of quartz, sanidin,
oligoclase, and black mica, or of one or both kinds of feldspar, without quartz being
present. The crystals have sharp corners and are seldom cracked ; oligoclase is rarely
of the vitreous variety. Typical varieties of the rocks of this family occur largely in
the hills of Bereghszasz in Hungary.

Fam. 3d. Rhyolite proper, or Hyaline Rhyolite—The extensive range of varieties
afforded by all manner of modifications of hyaline texture are a peculiar feature of
rhyolite, distinguishing it from any other eruptive rock. In outward appearance they
remind one of artificial glasses cooled under the most varied conditions. Obsidian,
pumice-stone, and pearlite, constitute but a small portion of the varieties occurring ;

(54)

OF VOLCANIC ROCKS. Li

others resemble enamel and porcelain, or present appearances which are difficult to
describe. The occurrence of sphierolites and lithophyse add to the variety of their
aspect. Another feature peculiar to the rocks of this family consists in a foliated
structure, the folia being often thinner than paper, and presenting an endless variety
of color and modifications of texture. Pearlite alone does not participate in the pecu-
liar form of foliated structure.

All these varieties again contain, enclosed, all the different minerals before men-
tioned, or only a few of them, or they enclose no foreign substances at all. The rocks
of this family are chiefly of ptirely volcanic origin, but in some instances currents of
them appear to have been ejected through crevices in the older volcanic rocks.

OrpER Seconp—TRACHYTE.

The name ‘‘trachyte” was first used by Hany, in his academic lectures, to
designate the well-known volcanic rocks composing the Drachenfels on the Rhine.
But it did not come into general use, until Beudant, the pupil of the former, intro-
duced the name into geological literature, by his justly celebrated work ‘Travels
through Hungary,” a book which abounds in sagacious observations on the subject of
voleanic rocks. Beudant extended, however, the application of the name over a much
wider range than his teacher had done. Several years later, in 1835, L. v. Buch
introduced into literature the name ‘‘ andesite,” designating by it certain dark-
colored rocks, which were then known, especially through the collections of Al. v.
Humboldt and Boussingault, to enter largely into the composition of the voleanic por-
tions of the South American Andes. These rocks, which form obviously a part of
those which Beudant had comprised under the name ‘‘ trachyte,” were supposed to
be particularly distinguished by containing a peculiar species of feldspar which Abich
(in 1840) called ‘‘andesine.” When, however, a few years later, this mineral was no
longer considered to be a separate mineralogical species, and its name was dropped,
that of ‘‘ andesite ” became obsolete with it, notwithstanding its prior origin. Thence-
forth, the range of the varieties of volcanic rocks comprised in the term “ trachyte ”
has been even more enlarged than Beudant had proposed in his dissertation. Hruptive
rocks, widely differing in nature—in fact nearly all those of tertiary and post-tertiary
‘age, with the exception of basalt—have been united in it. But by no author, proba-
bly, was the application of the name as much extended as by Al. von Humboldt.’
On the other side, however, he was the first, by the establishing of numerous sub-
divisions, to draw attention upon the necessity of using separate names for more lim-
ited ranges of varieties. Recently, B. von Cotta, J. Roth, and others. have tried to_
demonstrate, how little reason there had been for dropping the term ‘‘ andesite.”
They re-introduced it into petrology; but, in drawing the limits between andesite and
cognate groups by principles of artificial classification, they used the name in a sense
differing to some extent from that in which it has been applied in the following pages.

7 Cosmos, vol. iv. ‘The first four orders of trachyte were proposed by G. Rose, the other two were added by
Humboldt.
(55)

18 RICHTHOFEN—NATURAL SYSTEM

There is plainly indicated, from a geological point of view, the existence of two large
groups within the limits of Beudant’s “ trachyte.” The two existing names, “ trachyte”
and ‘‘andesite,” may conveniently be used for their designation, since those rocks,
for which either of the two names was first introduced, are indeed the types of the
two natural groups.

Mode of Geological Occurrence—The trachytic rocks have had their independ-
ent epoch of eruption in every volcanic country. They preceded rhyolite and basalt
in age, and were posterior to the ejection of propylite and andesite. The trachytic
epoch was usually of long duration. In many localities, its later part blended with
the earlier of the rhyolitic epoch, which is manifest by the alternate emission of tra-
chytic and rhyolitic matter, during that time which was intermediate between the epochs
of the ejection ofthe principal bulk of either of them. As regards geographical dis-
tribution, trachyte is as much dependent upon, and as closely allied to, the preéxisting
masses of propylite and andesite, as is the case with rhyolite. It towers up in peaks,
cones, and ridges, which are distinguished by their rugged outlines, and rest, in the ma-
jority of cases, upon the summits or the flanks of the ranges composed of those older
volcanic rocks. But there are also numerous instances when these may be seen to be
accompanied, at some distance, by trachytic outbursts, when a cursory examina-
tion might make the latter appear to occupy an independent position. Trachyte does
probably not compose, by itself, any extensive mountain ranges, and it remains, in
general, greatly inferior in bulk to andesite. In Europe, its outbreaks were scattered
and isolated, and, though they have been quite numerous, the aggregate quantity of
trachytic rocks is not considerable. They occur in Hungary and Transylvania, on the
Lower Rhine, in Central France, in the Grecian Archipelago, and in other parts of
that continent, as well as in the adjacent portions of Asia. Specimens of trachyte,
owing to their beauty and varied aspect, are usually much more numerous in geological
cabinets than those of andesite—a fact which has frequently occasioned some miscon-
ception regarding the relative proportion and importance of trachyte and andesite
among voleanic rocks.

In the structure of the North American Andes, trachyte takes a more import-
ant part than in Europe. A continuous range of it, at least ten miles in extent, and
forming rugged crests, encircl:s the Washoe Mountains to the east, in the shape of a
crescent. Trachyte rests there on propylite, and its ejection has probably had an
intimate connection with the formation of the Comstock Lode. Other accumulations
of similar extent may be noticed at Esmeralda, on the eastern slope of the Sierra
Nevada, around Red Rock Cajion, south of Walker’s Pass, in the surroundings of Lake
* Tahoe and Sierra Valley, and at other places east and west of the Sierra Nevada. In
the northern provinces of Mexico, trachyte is known to occur quite extensively. At
all the places mentioned, the greater part of its bulk bears evidence of having been
brought into its present position by massive eruptions, while traces of extinct trachytic
voleanoes are scarcely wanting at any of them. Trachyte is still being ejected by a
number of active voleanoes. Among them may chiefly be mentioned those of Central
America, the greater part of the modern lava of which has been proved to have the
chemical and mineral composition of trachyte.

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OF VOLCANIC ROCKS. 19

Mineral Composition.—Trachyte is only inferior to rhyolite in the number of its
varieties. Yet its chemical composition is as simple as that of the latter, and ranges
within as definite limits. The rock appears to contain, on an average, from 60 to 65
per cent. of silica, and is in this respect, as in others, next allied to rhyolite. The
difference between the rocks belonging to both orders, having its fundamental cause
in the chemical composition, manifests itself externally in certain differences of charac-
ter, among which may be mentioned: the absence of quartz among the essential
ingredients of trachytic rocks, which probably contain no free silica at all ; the usual
predominance in them of oligoclase over sanidin; the larger proportion in which horn-
blende participates in their mineral composition, and the fact of their specific gravity
exceeding that of rhyolite. Like almost all volcanic rocks, trachyte consists of a paste
in which are imbedded various crystallized minerals. This paste is of various colors,
and has usually a more or less vesicular texture, which, by its property of imparting
to the rock a certain roughness of touch, gave origin to the name. Chief in order
among the enclosed minerals are sanidin, oligoclase, mica, and hornblende. They vary
considerably as regards their relative proportion, and therefrom arises a number of
varieties which have been partly distinguished by separate names. More numerous
varieties, however, are occasioned by the differences of texture. Pumice-stone is of
frequent occurrence among the latter, but it never exhibits that perfect long-fibrous
and silken nature peculiar to it when being a variety of rhyolite. Obsidian, in differ-
ent grades of perfection, is no unusual modification of the trachytic masses, and it is
often filled with spheerolites, while no pearlite has yet been found having the chemical
composition of trachyte. Foliated structure may frequently be met with; but the
folia are not. of that exquisite fineness which is peculiar to those of rhyolite. The
modes of texture which alternate in the folia, are chiefly obsidian, pumice-stone, and
microcrystalline varieties.

Subdivisions.—It appears that two natural groups of trachytic rocks may be
distinguished, which differ at the same time, from a mineralogical and chemical point
of view, and have therefore been arrived at similarly by the application of artificial
principles. We distinguish with B. v. Cotta:

8 I may here remark that I have endeavored to retain existing names, as much as possible, for designating the orders
and families distinguished in this present classification. What I have tried to establish as natural groups, may therefore, on
account of the similarity of nomenclature, coincide apparently, in many cases, with the groups established by artificial
principles, and named in accordance with them. ‘The former do indeed coincide in a few instances with the latter; but in
the majority of cases, the limits of the application of the names differ widely when established from the two points of view
mentioned. It would require too much space to go into detail on this point in regard to every name. I will, therefore,
confine myself to an illustration regarding the order of trachytie rocks. B. vy. Cotta, (with several other authors) unites
under the name of oligoclase-trachyte all voleanic rocks consisting chiefly of hornblende and oligoclase, with the exclusion
of certain dark-colored rocks, to which he applies the name andesite. ‘The former name, if used in the meaning of that
author, comprises, therefore, our order of propylite, together with all those rocks for which the term oligoclase-trachyte has
been here applied. It is evident, from the different geological positions occupied by propylite and trachyte, as well as from
the distinct petrographical character which either of them exhibits when occurring in large accumulations, how much their
union would be opposed to a correct representation of natural relations. It is obvious, besides, that the union is unprac-
tical. It would actually prevent the possibility of a clear and simple geological description of those countries in which
propylite and trachyte occur together with other voleanie rocks. The distinction between both is so great that it has
scarcely ever failed to be noticed by unbiased observers in those localities where both rocks occur. ‘To unite them because

. (57)

20 RICHTHOFEN—-NATURAL SYSTEM

Fam. lst. Sanidin-trachyte—The color of the paste varies, but it usually pre-
sents light shades of gray, reddish, and reddish-brown ; its texture exhibits all the
varieties mentioned. There are imbedded in it: crystals of sanidin, or of both sani-
din and oligoclase, besides mica and hornblende ; the latter, however, is frequently
wanting. To this family belong the rocks which compose the trachytic ranges of
Washoe and Esmeralda, that of the Drachenfels, and many others.

Fam. 2d. Oligoclase-trachyte.—The paste is of the same color as in the rocks of
the first family, though darker shades prevail, and presents a similar variety of text-
ure. Imbedded are, chiefly, crystals of oligoclase and hornblende, the former being
frequently of the vitreous variety; the latter having usually the shape of broad
needles, with a black color and bright cleavage-planes. Besides those minerals,
black mica is of frequent occurrence. The rocks of this family are ordinarily associ-
ated with those of the first subdivision, but in some localities are not accompanied by
them. At Lassen’s Peak there is but one limited space where rocks of both families
intermingle ; it is near the place where the lava has been ejected. Apart from it, the
grand currents of lava, extending to from ten to twelve miles distance from the place
of ejection, consist merely of oligoclase-trachyte. To this family belong those varie-
ties of trachyte which were called ‘‘ domite” by L. v. Buch.

The fact that the occurrence of the rocks of either one of these two families
will frequently exclude those of the other, and that, even in those localities where
they are associated together, they will occupy separate places in regard to geological
superposition, appears to indicate that the distinction of these two subdivisions forms
an approach to the requirements of the natural system.

OrperR THIRD—PROPYLITE.

The rocks of this order have hitherto occupied a very undecided position in
the different classifications of rocks proposed, and just as various has been their nomen-
clature when they had to be mentioned in geological descriptions. The fact that they
bear close resemblance in mineral character to ancient diorite, while, geologically, they
are intimately allied to voleanic rocks, has been the principal cause of this uncertainty
of their position. In Hungary and Transylvania, they occur quite extensively, and,
being of practical importance as the bearers of rich metallic veins, have had to be
noticed frequently in treatises on the mines of those countries. Beudant applied for

them the name ‘‘ porphyric greenstone,” and classified them, along with syenite, among
fo) ’ ’ to) ? Oo

they are both chiefly composed of oligoclase and hornblende, would render it indeed, practically, a very difficult and compli-
cated task to compare the geological relations of different volcanic countries on the strength of written descriptions. If we
now follow the classification of J. Roth, we find the application of the name trachyte limited to those voleanie rocks which
contain sanidin, but are devoid of quartz. All those numerous varieties which do not contain sanidin, but are intimately
allied to sanidin-trachyte, by their mode of geological occurence as well as by their physical characters, are excluded from
the denomination, and, together with all volcanic rocks composed chiefly of hornblende and oligoclase, are united into one
subdivision of andesite. The reasons which justify the separation of the compounds of oligoclase and hornblende into three
different groups, (oligoclase-trachyte, hornblendie propylite, and hornblendic andesite) will be detailed in the following pages.
It may then be understood why the adoption of the different systems of the current nomenclature would render the concise
geological description of volcanic countries extremely difficult, and would conceal the harmony really existing in the rela-
tions which they present in different countries.

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OF VOLCANIC ROCKS. 7) \

the ‘‘transition rocks.” Since then, the names greenstone, greenstone-porphyry,
diorite, dioritic porphyry, and others, have frequently been applied for the same rocks
of the Carpathians. Similar names have been used for them when they were mentioned
as occurring in other countries, as for instance Mexico, where still oftener they have
been simply styled ‘‘ porphyry.” In 1860, having had sufficient evidence of the Ter-
tiary age of these rocks and their close connection with the voleanic rocks of that
period, I separated them, in a treatise on some voleanic countries in Hungary, already
referred to, by the name of ‘‘ greenstone-trachyte,” from the remainder of those rocks
which then were usually comprehended by the name trachyte. That designation has
since that time been frequently applied in geological descriptions. The attempts made
to classify the rocks included in it have, however, been rather unsuccessful. J. Roth
combines them into one group with amphibol-andesite, from which they are quite dis-
tinct as regards their petrographical as well as their geological properties ; while others
considered them as belonging to the dioritic rocks. The grounds upon which they
have been united with the latter are purely artificial, since diorite is, from a geological
greenstone-trachyte.” Breithaupt established

ue

point of view, widely separated from
a new name, ‘‘Timacite,” for a variety of our propylite, which is of very limited oceur-
rence, and in which he discovered a new variety of hornblende, called by him ‘“ gamsi-
gradite.” Most valuable contributions for the knowledge of our “ propylite,” were
recently given by Dr. Guido Stache, from observations made in Transylvania. (Hauer
and Stache, Geologie Siebenbiirgens, Vienna, 1865.) He discovered the occurrence,
in that country, of quartz-bearing varieties in greater extent than they have been
found hitherto at any other place. Stache retains the name ‘ greenstone-trachyte,”
for those varieties which contain no quartz, and proposes the name “ Dacite” (from
the Roman province of Dacia to which Transylvania belonged) for those of which
quartz is a common ingredient.

These statements will show the discrepancy of the views which have been
entertained in regard to the systematic place and the nomenclature of the rocks
under consideration. It is owing, partly to their twofold affinity with other rocks,
(mineralogically to diorite, and geologically to voleanic rocks) and partly to the fact
that they had, until lately, not been made the object of study. All observations made
during the last few years concur in this, that those rocks, wherever they have been
encountered, form a distinct link in the range of Tertiary and Post-tertiary eruptive
rocks, being everywhere the first of them in age, while they are, in regard to their
mineral character, no less distinct from any other eruptive rocks originated in those
periods. They constitute, indeed, a more natural and more distinct group than any
of the other volcanic rocks, and it has become desirable to unite them under a com-
mon designation. As no prominent property distinguishes them from diorite, and the
derivation of the name from one certain locality did not appear proper for the desig-
nation of rocks of wide distribution, geological relations alone could be used as a basis for
the nomenclature. The rocks under consideration, as we shall hereafter more fully
develop, give evidence, in all localities where they have been met with of having re-
opened the eruptive activity after ages of comparative repose. It is since then only,

WW (59)

Ay) RICHTHOFEN—NATURAL SYSTEM

that this activity has continued with extreme violence over all parts of the globe,
through the remaining part of the Tertiary and the Post-tertiary periods, growing,
however, more and more faint during the latter. Propylite was, in fact, the precursor
of all other volcanic rocks, and its appearance on the surface inaugurated a grand
revolutionary activity on the globe. It is this position, at the entrance as it were to a
new era in the history of the earth, which has given rise to the name ‘‘ propylite.”

Mode of Geological Occurrence.—The position of propylite at the bottom of all
voleanic rocks is its most important geological feature. Its age has in no instance been
ascertained with exactness. The nearest approach to its determination was made in
Northern Transylvania, where I found, in several localities, nummulitic strata inter-
sected by dykes and large intrusive masses of propylite, and covered by accumulations
of it. As the greater part of the volcanic rocks in that country are of Miocene age,
the ejection of propylite must have taken place either in the latter part of the Hocene
or in the earlier part of the Miocene epoch. In view of the facts, that volcanic rocks
have nowhere been observed to be anterior to the Hocene (probably even not prior to
the Miocene) epoch, and that propylite is always allied to them as the first link in the
order of succession, it may be inferred that propylite is, in general, of Tertiary age,
until proofs to the contrary may be found.

The forms of propylitic mountains can be observed only in rare instances, since
they are usually covered by other volcanic rocks, especially by andesite. This cireum-
stance may also explain the fact of the comparatively rare occurrence of propylite as a
surface-rock. In the environs of Bisztritz, in Northern Transylvania, it forms several
high, isolated cones with steep slopes, resting on Eocene strata; their fine, dome-
shaped appearance is scarcely surpassed in beauty by that of any other kind of rock.
No estimate can be made in regard to the relative bulk of propylite which has been
ejected, on account of its being overlaid by andesite and trachyte. It occurs, as far
as exposed to view, in quite considerable accumulations, at Nagybanya and Kapnik,
in Hungary, in Washoe, and at Silver Mountain, and covers large areas in Mexico,
where it overlies the Cretaceous. It appears to have been profusely ejected through
fissures, and its emission not to have been accompanied by volcanic action proper ; no
distinct traces, at least, have been found of propylitic voleanoes. Massive eruptions
‘are known to have occurred, besides the places mentioned, in several other parts of
the southern slope of the Carpathians, on the highlands of Armenia, and in Mexico,
while the occurrence of propylite in Bolivia, the Altai Mountains, Northern China,
and some other countries, may be inferred from the description of their geology.

Propylite has been repeatedly considered to be a sedimentary rock, metamor-
phosed iz situ. The cause of this opinion was probably the fact that there are rocks
which undoubtedly have had that mode of origin, and share with propylite the resem-
blance in mineral character to diorite. The eruptive origin of propylite is, however,
evident, as it intersects stratified rocks in the shape of dykes. In Washoe and Silver
Mountain, moreover, there are extensive accumulations of propylitic breecia and
stratified tufa. The latter consists of alternate layers of coarse conglomerate, fine-

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OF VOLCANIC ROCKS. 93

grained propylitie detritus and massive propylite, and the entire system is intersected
by dykes of the latter.

A particular interest, which has a practical bearing of some importance,
attaches to propylite, inasmuch as, notwithstanding its limited occurrence, it yields
probably a larger amount of silver than any other rock. Several of the principal
silver-bearing veins, as the Comstock vein in Washoe, the celebrated veins in Hungary
and Transylvania, as well as some of those in Mexico, and probably too in Bolivia, are
enclosed in propylite.

Mineral Composition —In propylite are united the petrographical properties of
ancient dioritic rocks with those of andesite and oligoclase-trachyte. Varieties are
numerous, but produced more frequently by the difference of the component minerals in
size and relative proportion than by the texture, which remains porphyritic in every in-
stance. The most common varieties consist of a fine-grained, microcrystalline paste of
dark green or greenish brown, more rarely of reddish and dark gray colors, in which
are imbedded crystals of oligoclase and hornblende ; the former is of whitish or light
green color, the latter ordinarily dark green and fibrous, seldom black with bright
cleavage-planes (as is the case with gamsigradite). The paste appears to be a fine-
grained aggregation of the same two minerals (the feldspathic ingredient prevailing),
with the admixture of titanic iron, and to owe its ordinarily green color to the profuse
dissemination of small particles of green, fibrous hornblende. Sporadic crystals of
augite are occasionally met with in some of the common varieties, while others contain
rounded grains of quartz sparsely enclosed. Recent observations have led to the dis-
covery of two series of varieties deviating from the usual composition as explained.
One of them is produced by the increasing proportion of the grains of quartz; the
other by the more profuse occurrence of augite. The former was found by Stache, in
Western Transylvania ; the latter observed by me at Silver Mountain. Both appear
to be of a limited geographical distribution, but are of great interest, as they extend
the limits of the order of propylitic rocks, without rendering either their petro-
graphical character, or their geological relations less distinct. In regard to the latter,
it may be stated that the entire range of the quartzose to augitic varieties was anterior
in age to andesite.

Difference of Propylite from other Volcanic Rochs nearly related to it—The rocks of
two other orders have an affinity to propylite, namely oligoclase-trachyte and ande-
site. The principal ingredients of either are hornblende and oligoclase, and either of
them contains crystals of both these minerals imbedded in a paste. Yet there is a
notable difference in character between these two, as well as between either of them
and propylite. It is so conspicuous to the eye that the respective rocks have ordi-
narily been distinguished in geological descriptions, even in those of older time ; and
even the unprofessional eye would be able to distinguish the three groups in a collec-
tion of specimens belonging to them. Yet, it escapes description. It may, at this
present time, safely be founded on what the botanist would call ‘ habitus,” a certain
general character which it is as easy to recognize by the eye as it is difficult to describe
it in words. and impossible to define its causes. It is probable that observations, such

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24 RICHTHOFEN—-NATURAL SYSTEM

as Sorby has made in reference to those minute differences of texture, which can only
be detected with the aid of the microscope, and H. Rose in regard to the modifieations
of silica and their causes, aided by exact chemical analysis and experiments made
with the view of enquiring into the differences of origin of such eruptive rocks as
differ from each other in texture, will, if further prosecuted, reveal the true nature
and cause of the properties which distinguish the rocks of these three different
orders.

A few of the more palpable differences may here be noticed. Propylite is
essentially of greenish color, and some of its varieties resemble diorite in composition
and texture ; andesite is of blackish color and approaches basalt in aspect; trachyte is
of various colors and shades, among which green and black are rarest of occurrence,
and in regard to its external characters resembles rhyolite more than any other rock.
Oligoclase, in trachyte, is frequently of the vitreous variety, scarcely ever so in pro-
pylite and andesite. Hornblende is an essential ingredient in these two, not so in the
former ; it is of fibrous texture and green color in most varieties of propylite, black in
andesite and trachyte. Mica is seldom wanting in the latter, while it is not of com-
mon occurrence in propylite and andesite. Titanic iron enters largely into the compo-
sition of these, and is contained in smaller proportion in trachytic rocks. The latter
excel by having the greatest variety of texture, while propylite has among the
three the most perfect porphyritic texture, which has given rise to its frequent
popular designation ‘‘ porphyry ;’ this name has never been applied to andesite or
trachyte.

The enumeration of all these trifling differences is, however, insufficient to
express the marked distinction which exists in thes external characters of propylite,
trachyte and andesite. As a similar, and even more conspicuous, distinction manifests
itself in their geological relations, we have to consider the existe.ce of those three
natural orders as a fact founded on observations, although we may be utterly unable
to explain, and even to express it in words.

Subdivisions —The remarks made in regard to the mineral composition of pro-
pylite have shown that the range of its varieties may conveniently, and in harmony
with geological occurrence, be subdivided into three parts :

Fam. 1st. Quartzose Propylite or Dacite—This embraces rocks which, though
having in general a similar composition to those of the following family, contain be-
sides, rounded grains of quartz, sometimes in considerable proportion. They occur in
the western part of Transylvania, where their outbreaks succeeded those of rocks of
the second family, and preceded those of andesite.® Similar rocks have been observed
in-Sinaloa (Mexico).

Fam. 2d. Hornblendic Propylite——Rocks composed chiefly of hornblende and
oligoclase, as described above. This family embraces vastly the majority of all pro-
pylitie rocks observed, among others those of Washoe. Breithaupt’s “‘ timacite ”
is one of its varieties.

» Fully described by G. Stache, loc. cit.

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OF VOLCANIC ROCKS. 95

Fam. 3d. Augitic Propylite—Rocks distinguished by the accession of augite
among the ingredients of the rocks of the foregoing family. It is present in greater
or less quantity, sometimes predominating over the hornblende. To this family be-
longs the propylite of Silver Mountain, which contains augite in larger proportion
than any other known variety.

Orper FourtH—ANDESITE.

The history of the name ‘andesite’ has been noticed conjointly with that of
“trachyte.” At the time when the former name was first proposed for certain rocks
of the Andes, a few specimens of which had been brought to Europe, it was intended
as a designation of an accidental variety, proper to that mountain range. But those
same specimens have proved since to be the type of one of the most important groups
of voleanic rocks, which is distinct from others in character, and has a wide distribu-
tion.

Mode of Geological Occurrence.—Andesite vies with basalt in regard to the
quantity of matter ejected to the surface, and probably excels the same in this respect.
In most of those parts of the Andes, in regard to the geology of which we possess
reliable information, it forms the chief bulk among voleanie rocks. The same is the
case on the southern slopes of the Carpathians, at Nangasaki in Japan, and on the
islands of Luzon and Java. Andesite succeeded the ejection of propylite, and pre-
pared the way to that of trachyte. Preéminently the greater part of it has, to all
appearance, been ejected through extensive fissures ; that is, it has been produced by
what we styled massive eruptions ; though andesitic voleanoes, too, are not of rare
occurrence, and, including those which are extinct, appear to have been particularly
grand in their activity. It may be supposed that many of the former craters have been
destroyed. Andesitic mountains are characterized by monotony in scenery. They
form continuous ranges, which are often of considerable elevation and extent, but ex-
hibit gentle outlines in their summits as well as in their slopes. Breccias only, which
accompany the solid rock ordinarily in vast quantities, cause local interruptions of the
monotony by their more rugged forms. They appear in castle-shaped rocks on the
crests of andesitie mountains, and form high walls, naked and steep, along their slopes.
Being more liable to destruction by the erosive action of water than solid andesite,
they frequently compose the sides of steep ravines and camions.

Mineral Composition —Andesite is always of dark color, mostly blackish,
though frequently reddish-brown on the weathered surface. Its mineral composition
varies, though it is confined within more narrow limits than that of propylite. We
may distinguish, in regard to it, two

Subdivisions. which are connected by gradual passage in composition and, in
a greater measure, by geological relations.

‘am. 1. Hornblendie Andesite-—Paste of bluish-black to dark-gray color, and
of microcrystalline texture, which passes by gradual steps into that of obsidian. In
the former case it is frequently vesicular like trachyte. There are imbedded in it

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26 RICHTHOFEN—NATURAL SYSTEM

small but distinct tabular crystals of oligoclase, fine grains of titanic iron, small,
elongated columns of hornblende, sometimes mica, and in most varieties a few isolated
crystals or rounded grains of augite. Frequently these different crystals are so small
as to be no longer recognizable, except by the aid of the microscope. The rock is
then similar in character to certain varieties of melaphyr, but may be distinguished
by its vesicular texture, which is scarcely ever wanting in the hornblendic varieties.
This family comprises vastly the greater portion of all andesitic rocks.

Fam. 2. Augitic Andesite-—Two groups of varieties may be distinguished
among the rocks of this family. Those of the first have a paste of an oil-brown color
passing into black, a compact appearance, and ordinarily a microcrystalline tex-
ture which passes into that of porcelain. Vesicular inflation does ordinarily not
occur. Crystals of a monoclinohedric feldspar (probably labrador) are almost invari-
ably enclosed, and frequently accompanied by hornblende and augite. For the rocks
of this group the name ‘ trachydoleritte” has particularly been applied. The other
group comprises certain varieties for which the name ‘‘ anamesite”’ has not rarely been
used, and which are of dark-gray colors. Their texture is in most cases vesicular.
The enclosed minerals are the same as with the other group; but, while for this
the feldspathic ingredient is more characteristic, and frequently alone present in large
crystals, the rocks of the second group are distinguished by the predominance of well
formed crystals of augite and hornblende, while those of feldspar are often so small
that they cannot be distinguished by the eye. The rocks of both these groups contain
titanic iron in larger quantity than those of the first family, and are of greater specific
gravity. Olivine enters occasionally into their composition, in very subordinate quan-
tity. Geologically, the rocks of this family are closely allied to hornblendic andesite.
They have succeeded it in age, and are limited in their geographical distribution to those
places where hornblendic varieties had been ejected before, frequently intersecting and
overlying them. Notwithstanding the resemblance which they bear to certain varie-
ties of basaltic rocks, they appear to be never associated with them geologically.

OrpER FirrrH—Basatr.

No one of the names applied to volcanic rocks is of equal antiquity with that of
“basalt,” and with none there has ever been less change of opinion, and uncertainty in
respect to its application. It may be inferred from this fact that the rocks comprised
by that name are very distinct in character and little liable to variation, which is indeed
true for the typical rocks of the order. But there have to be associated in the same
order with them certain other rocks, which, though nearly related to basalt in regard to
their chemical and mineral composition, differ from it in the more conspicuous external
characters.

Mode of Geological Occurrence.—Basaltic rocks are more independent than those
of any of the foregoing orders (perhaps with the exception of propylite), in respect
to their epoch of ejection as well as to their geographical distribution. In the order
of time they succeeded next to rhyolite, but locally, both are usually separated.
Basalt occurs always in the neighborhood of more ancient volcanic rocks, but the cases

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OF VOLCANIC ROCKS.

Lo

7

are rare when it intersects them and expands over them. Almost invariably it is found
accompanying their ranges at some distance, forming itself extensive ranges ; more

frequently, however

, it occurs in groups or lines of isolated outliers. It may some-

times appear to have no connection with the distribution of older volcanic rocks. But
closer investigation will always show it to be within or in the neighborhood of the
limits of former eruptive activity. The belts along which this had taken place may
be locally interrupted for quite a distance, and the gap be filled by some isolated out-
breaks of basalt, or the latter may extend the limits of those belts, in length as well
as in width. It is a strange phenomenon that these isolated outbreaks of basalt occur
particularly in connection with granite. It intersects this rock very frequently in small
dykes, and expands over it in thin sheets. This connection of granite and basalt is
very conspicuous on the eastern slope of the Sierra Nevada, and along the belt of
basaltic eruptions which traverses the middle part of Germany. Such places are often
the seat of basaltic voleanoes, which, though now mostly extinct, are also otherwise of
very frequent occurrence.

Mineral Composition —Basalt is the representative of what Bunsen has called
the ‘“‘normal pyroxenic type,” and is thereby, as well as by its specific gravity, mineral
composition, and little variety of texture, the reverse of rhyolite, which represents the
‘normal trachytic type.” <All the essential ingredients of basalt, and most of the
minerals which enter accidentally into its composition, are different from those which
are characteristic of rhyolite. Augite, labrador and titanic iron, imbedded in a paste
consisting essentially of the same minerals, constitute generally the rocks of this order ;
or labrador may be replaced by leucite, nepheline or a zeolitic substance ; or other min-
erals may enter into the composition, such as olivine, basaltic hornblende, hauyne,
apatite and black mica; and in many instances basaltic rocks consist merely of a
fine-grained aggregation of the different minerals mentioned. Ordinarily, the paste
has a microcrystalline texture, and may either contain crystallized minerals imbedded,
or may be devoid of them. This mode of texture passes by gradual steps into that of
obsidian, which is the only hyaline variety occurring. Among the peculiarities of these
rocks, and which may also in some measure be observed with augitic andesite, is the
occurrence of rounded cavities filling the rock and giving it a cellular, sometimes
spongy appearance. It is different in nature from pumice-stone, which is a modifica-
tion of hyaline texture, while those cavities occur in microcrystalline rocks.

Subdivisions —Fam. 1st. Dolerite (including nepheline-dolerite and the greater
part of anamesite).—A crystalline aggregation of augite and labrador with titanic iron.
Labrador is usually replaced by nepheline, either in part or totally. Accessory min-
erals are: olivine, hornblende, apatite, black mica.

tam, 2d. Basalt.—Paste of dark-gray or black color, and of the varieties of text-

ure mentioned ; it constitutes either the mass of the rock by itself, or encloses olivine,

augite and labrador, in crystals or crystalline grains. Besides these are frequently

enclosed : titanic iron, black mica, rubellan, zircon, apatite and other minerals. This

family comprises nearly the whole bulk of the rocks which constitute the basaltic
order, the two other families being of rare occurrence.

(65)

28 RICHTHOFEN—NATURAL SYSTEM

Fam. 3d. Leuettophyre—Rocks of porphyritic texture, crystals of augite and
leucite being imbedded in the paste.

It will be noticed that no place has been assigned in this classification to phon-
olite. As regards those rocks for which this name was first proposed, our knowledge
is still quite limited. It appears that, notwithstanding their comparatively rare occur-
rence, they form a distinct natural group closely allied to basalt; but they are so dif-
ferent from true basalt as regards lithological characters, that they should not be
classified with it before further observations will have determined their real position
in reference to the natural families of volcanic rocks. The name phonolite has, how-
ever, been so much extended in its application, that this task is not so easy to accom-
plish. Some external properties, easy of recognition to superficial observation, such
as a certain tabular structure, lithoid texture of the paste, with small, bright crystals
of feldspar enclosed, and the peculiarity of ringing by a blow of the hammer, which
have been often considered as the characteristic features of phonolite, are just as com-
mon with certain varieties of trachyte and rhyolite, and even with some of propylite.
It must be ascribed to this reason, that the name phonolite has been used for the
designation of rocks which bear an accidental resemblance to true phonolite, but are
distinct from it in nature. It has probably been oftener applied to rocks belonging to
the trachytic order than to such as have the distinguishing features of those varieties
for which the name has been first used.

CorRELATION OF THE Five Orpers oF Votcanic Rocks.

In the foregoing pages I have attempted to lay down the outlines of a classifi-
cation of the volcanic rocks by natural principles, and to apply a nomenclature which
should be appropriate to these, and embrace, at the same time, the most current of
existing names. It is the next object of this paper to prove that these rocks are
mutually connected by definite relations, and that their totality, in virtue of this
property, forms actually what may be called a system in nature, and that the form into
which we have tried to bring it, imperfect though it must be, is an approach towards
its expression. In order, therefore, to fully realize the philosophy of the natural
system, we have to contemplate the relations which, firstly, the rocks of the different
orders offer mutually among themselves, and by which, secondly, they are connected
as an entire class with ancient eruptive rocks ; while we will have, thirdly, to examine
into the mode of origin of volcanic, and of eruptive rocks in general, in order to
establish the nature of their fundamental difference from sedimentary and metamor-
phic rocks. Our task is thus three-fold. The present chapter will be devoted to the
first order of relations. They may be considered from several points of view, the
more important of which are: chemical and mineral composition, geographical distri-
bution, and all those complex relations which may be comprehended in the term
‘‘mode of geological occurrence.” We will contine ourselves to the last point of view.
But even with this restriction, we can only trace general outlines.

(66)

OF VOLCANIC ROCKS, 29

Laws relative to the Age of Massive Eruptions.

The succession of massive eruptions during the Tertiary and Post-tertiary ages
has taken place in the following order :
Ist. Propylite.
2d. Andesite.
dd. Trachyte.
4th. Rhyolite.
5th. Basalt.

This singular mode of succession, in which no regularity (as to increase or
decrease of silica or specific gravity, or as to a gradual change of mineral composition)
can be discovered at first sight, and which might indeed appear to be devoid of order,
and to bear the character of such a succession as might have been occasioned by the
codperation of accidental circumstances in one single country, can nevertheless be
proved to exist in widely separated parts of the globe. It may justly be objected to
this assertion, that observations in regard to the relative age of different volcanic rocks
are scarce, and hardly sufficient to establish definitely such a law. But hitherto no
deviation from it has been discovered,” and it appears to be true for all volcanic regions
on the globe, though with this restriction, that the epochs marked in each country by
the ejection of rocks of any certain orders have not been contemporaneous in different
countries. The commencement of eruptive activity in the Tertiary epoch has been ear-
lier at one place than at another ; it is its further mode of development in regard to the
nature of the matter ejected which has everywhere been regulated by the same definite
relations, though it has been independent, in some measure, at each place, or, to use
a more correct expression, over the area of each belt of eruptive activity. We have
to mention another restriction. Abrupt passage may be said to be an almost unknown
conception in geological matters, where the order in time is concerned ; nor has it to
be applied to the order of succession of volcanic rocks. The eruptions of propylite
appear not to have been interrupted by the ejection of any other rocks. This may
too be said, though less strictly, of the andesitic epoch. But some, as it were, retarded
eruptions of andesite, which, however, have always been insignificant, may occasion-
ally be traced during the first part of the trachytic epoch, and similar relations exist
between trachyte and rhyolite. Basalt, however, appears to have had everywhere its
own epoch of ejection, uninterrupted by the massive eruptions of any other rock of
the orders just mentioned.

We proceed to a short review of some salient facts observed in those coun-
tries where the mutual relations of volcanic rocks in regard to their age have been
made an object of study.

In Hungary and Transylvania, propylite, as we have had occasion to mention,
was ejected first of all voleanic rocks, and may be said to have maugurated all sub-

10 Tt must be borne in mind that we are speaking of massive eruptions. Apparent exceptions are known to occur
with rocks ejected from volcanoes, of which mention will be made hereafter.

I (67)

30 RICHTHOFEN — NATURAL SYSTEM

sequent eruptions of those belonging to other orders. At several places it has
been observed to intersect Eocene strata, and to expand above them. The coun-
try of Nagybdnya, Felsébdnya and Kapnik, the celebrated silver-bearing veins of
which are enclosed in propylite, offers especially conspicuous illustrations ; not less,
from the descriptions given by G. Stache, the ‘‘ Erzgebirge ” of Transylvania, where
the same kind of rock is rich in mineral veins. In this country, hornblendic propylite
forms an older series, followed by eruptions of highly quartziferous varieties. In
both countries, but chiefly in that first named, and at several other places along the
southern slope of the Carpathians, andesite may be seen intersecting propylite in large
massive dykes, and towering up above it in mountain ranges. Andesite composes
entirely the Hargitta-range, which extends over one hundred miles in length, and
twenty-five in width; the Vihorlat-Gutin-range, which is of still larger dimensions,
and the Hperies-Kaschau-range ; all of which are densely wooded, and of a gloomy,
monotonous aspect. The only change observable on their summit ranges is a more or
less dark color of the rock, occasioned by the predominance of augite or hornblende in
its composition, (the augitic varieties being invariably of more recent age than the
hornblendic), while their slopes, and particularly their ends, present a much greater
variety in rocks as well as inscenery. It is here that the more silicious volcanic rocks
are encountered. Trachyte is of rare occurrence ; but it forms several isolated cones,
some of which, on account of their prominent position, were crowned by castles in the
middle ages. Rhyolite is much more frequently met with, bursting forth on the
flanks, and skirting the foot of the andesitic ranges, particularly where they verge
towards the Hungarian plains, which in the rhyolitic epoch were still covered by a
shallow and slowly retiring sea. It projects against this in promontories, which are
now covered by the most celebrated of the Hungarian vineyards, those of Tokay
among others. The boundaries of these vineyards towards the adjoming beech forests
mark the dividing line between rhyolite and andesite. An interesting mode of occur-
rence of the former may be witnessed in large circular or amphitheatrical basins which
are surrounded by andesite. Such places are the theater, especially, of the volcanic
activity connected with the outbreak of rhyolite, and abound in endless hyaline vari-
eties of the same. Telkibanya is the most interesting among the localities of this
description. There is no lack of evidence to prove that trachyte and rhyolite are both
of more recent age than andesite, while the assertion that trachyte preceded rhyolite
in age, rests only on a few though conclusive observations. Basalt occupies a singular
position in the geology of Hungary. It keepsaltogether aloof from the places occupied
by the other four orders of voleanic rocks, and forms extensive, though isolated, hills at
some distance from them, scattered over a wide range of country. It would be difficult
to determine its relative age, but for the voleanic sediments which were of formation
contemporaneous with the ejection of the different volcanic rocks, and have been spread
over wide areas at the bottom of the then existing sea, with fossils occasionally imbedded.
In several localities, especially in the neighborhood of Kaschau, basaltic sediments may
be seen, covering those composed of rhyolitic matter, while at Gleichenberg in Styria,
Mr. Franz von Hauer has observed fragments of rhyolite enclosed in basalt.

The observation of these relations in Hungary and Transylvania has first given
(68)

OF VOLCANIC ROCKS. 31

rise to the establishment of the above mentioned law of the periodical succession of
voleanic rocks." It has since been corroborated by observations made in other coun-
tries far remote from the Carpathians. Near Nangasaki, in Japan, andesite preceded
trachyte in age, and of the former there are two varieties represented, one augitic
and one hornblendic, of which the former is of more recent origin than the latter:
thus, even the more minute relations observed in the Carpathians are repeated in
other countries. The islands of Java and Luzon are too intensely voleanic and
covered with lava to aid in establishing the laws in respect to massive eruptions
without very close observation. But another region offering copious evidence is that
of the Sierra Nevada, together with the adjoining parts of the Great Basin. Observa-
tions in these countries are still limited as regards our present subject. The great
part, however, which volcanic rocks take in their composition, as well as in that of the
highlands of Mexico, and of the entire range of the Andes, promises to make the
Western Coast of America the most prolific source of observations necessary for the
definite establishment of geological laws of which, at the present day, we can only
trace the first foundation.

In Washoe, a country adjoining the Sierra Nevada immediately to the east, propy-
lite forms extensively the foundation for all other volcanic rocks, which fact proves
clearly its priority inage. It composes the plateau of Virginia City and Gold Hill, and
derives a practical interest from the fact that the Comstock vein is enclosed between
propylite and syenite, though in some parts of it both walls consist of the former rock.
Andesite is Insignificant in bulk in that region. It composes a few small hillocks on
the propylitic plateau, and in some cuts and tunnels andesitic dykes may be seen, which
appear to have been the feeding channels of the surface accumulations. Trachyte, on
the contrary, is among the prominent rocks of Washoe. It forms a high and rugged
crest, encircling the plateau of Virginia and Gold Hill to the east, and extending for
miles to the north, while to the south it reappears across the Carson River. No evi-
dence is afforded, in Washoe, for the establishment of the mutual relations of andesite
and trachyte, while it is conspicuous that the latter was of later origin than propylite.
Besides the evidence offered by intersection and superposition, another fact may be
noticed which is suggestive for the length of the period that elapsed between the
eruptions of both rocks. It is this, that propylitic sediments occur, at least, at one
thousand feet more elevation than those consisting of trachytic matter, which fact
appears to indicate that the former have been deposited at a much earlier part of the
period marked by the gradual subsidence of the inland seas of the Great Basin than
has been the case with trachytic sediments.

At Silver Mountain, propylite, of the augitic variety, fills the bottom of a deep
basin encircled to the west by granitic walls several thousand feet in height. Its
massive accumulations are intersected by andesitic and trachytic rocks, which latter
appear to compose the summit of the high peak of Silver Mountain itself, while rhyo-
lite occurs in such a position as to make it probable that it has arrived at the surface
last of all eruptive rocks. Basalt appears to occur only to a limited extent at Silver

11 Richthofen, loc. cit.
(69 )

32 RICHTHOFEN — NATURAL SYSTEM

Mountain. But this rock is largely distributed in the regions adjoining the Sierra
Nevada to the east, and bears evidence of its recent origin. It is the only volcanic
rock. which covers in places the sand of the deserts, and the belts distinguished by its
eruptions are still marked by the occurrence of hot springs and other post-vol-
canic phenomena. Most of these are contiguous to those places where basalt and
granite are in close contact, as is very conspicuously the case at Steamboat Springs,
near Washoe, and in the Coso Mountains. No phenomena of similar nature appear
to be connected at the present time with any other volcanic rock east of the Sierra
Nevada. There are not a few instances where basalt may be seen covering propylite
or andesite ; but I met with only one case where it comes in contact with rhyolite,
close enough to establish their mutual relations. This is in Esmeralda, on the eastern
slope of the Sierra Nevada, a country of unusual interest for the study of volcanic
rocks in general. Propylite encloses the silver-bearing veins of that place. It is
overlain by trachyte and rhyolite, both of which occur in very great variety. To
the east of the place, basalt has not only flowed over rhyolite, but contains numerous
fragments of it enclosed, which fact confirms also for this country the more recent
origin of basalt.

Many other examples might be added to this short list, partly of positive obser-
vations made in the countries already mentioned, and partly of facts described in
treatises on the geology of other countries, such as Armenia, the Caucasus, Central
France, the Eifel, Bolivia, Mexico. As these descriptions, however, have not directly
in view the illustration of our subject, great care should be used in drawing from
them conclusions in regard to it. Let it suffice to remark, that every observation on
record which bears on our subject, appears to confirm the proposed law, while none
can be found giving evidence against it. This may justify the assumption that the
periodical succession of volcanic rocks, in the order above mentioned, is a general law,
true for all parts of our planet.

Laws regarding the Mutual Relations of Massive Eruptions and Voleanic Activity.

It has been mentioned in this chapter, that the law of the periodical succession of
voleanic rocks regards those outpourings of large volumes of matter not resulting
from voleanic activity proper, and which we called massive eruptions. For conven-
ience’ sake, we make use of the following expressions: propyliie epoch ; andesitic epoch ;
trachytic epoch ; rhyolite epoch ; basaltie epoch — designating thereby those epochs
in which the massive eruptions of rocks belonging to each of these orders, have taken
place in every different country. If we now direct our attention to the other mode
of manifestation of subterranean energy, the volcanic eruptions, a cursory review of
active volcanoes in regard to the nature of the rocks which they eject, shows that the
same law is no longer true for them, as their lavas belong to several different orders
of volcanic rocks. It is, however, known that each voleano ejects lava or scoria
belonging petrographically only to one distinct order, and the examination of the
material accumulated by former activity will show that with most volcanoes the
nature of the rocks ejected has never materially varied, while with some of the

(70)

OF VOLCANIC ROCKS. 33

grander vents it has undergone a periodical change. Volcanoes, whether active or
extinct, may be classified from this point of view. We shall distinguish: andesitic,
trachytie, rhyolitie and basaltic volcanoes, according to the nature of the mineral matter
which each voleano has ejected in the first epoch of its activity, regardless of any later
changes. Two noteworthy relations may be traced between these different orders
of voleanoes and the massive eruptions of the synonymous orders of voleanie rocks.
The first of them is the alliance of both in regard to geographical distribution, the
voleanoes of each order being limited, in this respect, to the immediate neighborhood
of massive accumulations of rocks similar in nature to their first lavas. From this
may partly be inferred the second relation, that the massive eruptions of each order
have been succeeded by volcanic activity, which occasioned the ejection of lava corre-
sponding in nature to their own rocks, and continued for long after-time, in many instan-
ces to the present day. This dependence of volcanoes upon massive eruptions explains
why the number of active voleanoes is so small when compared with those which
are extinct, and why the present activity even of those which are still in operation, ap-
pears to be only a faint remnant of that which the same vents exhibited in former time.
It will further explain why no vestige can be found of a rhyolitie voleano having been
active before the rhyolitic epoch, or of a basaltic voleano having originated before the
basaltic epoch, while geological observation goes to show that during, and immediately
after those epochs, the voleanoes of either order have been most intense, numerous
and extensive, and their activity has, from that epoch of culmination, gradually relaxed,
in most cases to perfect extinction.

An instructive instance of one of those grander volcanoes which have undergone
a periodical change in regard to the nature of the matter ejected from them, is afforded
by the extinct voleano Lassen’s Peak, in Northern California, which Professor J.D.
Whitney and I visited in 1866. We found it to have been originally an andesitic
voleano, and it has to be ranked as such in our proposed classification. The enormous
bulk of the ancient voleano is totally built up of stratified layers of andesitie tufa
and rapilli, which, in the steep gorge issuing from its lower crater, are exposed in a
thickness of nearly four thousand feet, notwithstanding the total destruction which
the upper part of the former cone has undergone, and the fact of its lower parts
extending down far beneath the present surface, and being therefore concealed to view.
Besides these stupendous accumulations of loose matter, currents of andesitic lava
appear to have been emitted from the crater, extending at least twenty miles from
the place of ejection. Ata later epoch, the activity of the same volcano has been
distinguished by the emission of trachytic lava from the northeastern part of the
wall of the crater; its currents have expanded to elongated and sloping tables.
bounded by abrupt descents. A third epoch is marked by the outbreak of rhyolite
at the same place whence the trachytic rocks had issued. Rhyolite composes the
present summit of Lassen’s Peak, on which it is accumulated in a thickness of more
than fifteen hundred feet, also some other summits of less altitude, and at least one
prominent current of lava of great volume.” The noteworthy fact illustrated by these
observations on Lassen’s Peak, and corroborated in numerous other instances, is this :

12 Mr. Gineree Kine has preeed the occurrence of cae ult of Poe very recent origin Metal) nari of
(71)

34 RICHTHOFEN —- NATURAL SYSTEM

that the same law of periodical succession which has been established in regard to
massive eruptions, is true for volcanic action, particularly when this happened to
assume such unusual intensity and dimensions, and has been of as long duration as
was the case at that volcano.”

Summing up these considerations on the correlation of the different orders of
voleanic rocks in respect to the age of their emission through volcanic vents, we arrive
at the following conclusion: The commencement of the activity of the volcanoes of
each separate order has been nearly coincident with, though in every instance success-
ive to, the main phase of the corresponding massive eruptions. Thence it has, by each
separate vent, either continued emitting similar material to that first ejected, until its
extinction, or it continued in the same way to the present day, or it has been subjected
to a periodical change in regard to the nature of its lavas, and this change is analogous
to that exhibited by the succession of massive eruptions. In this case, as in the for-
mer, the voleano has either become extinct when in a certain phase, or it is still active.
We are thus furnished with a natural cause of the fact, that most active voleanoes are
emitting basaltic, a smaller number of them rhyolitic or trachytic rocks, while an-
desitic lava is peculiar only to a few of them, especially to some of the prominent vol-
canoes of South America (Chimborazo, Cotopaxi, Antisana, Tungurahua, also Popoca-
tepetl, Colima, and Teneriffe) which appear never to have changed in mineral char-
acter. It will, too, be self-evident, why generally no material change in the nature
of their lava should have been observed in regard to those volcanoes which have orig-
inally emitted basalt and constitute our order of basaltic volcanoes.

A few more instances may here be mentioned in support of our propositions.
The interest attaching to volcanoes has furnished us with a much greater number
of facts in regard to voleanic rocks when occurring as lavas, than we possess in regard
to the grander and more frequent instances when similar rocks have been produced
by the comparatively neglected action of massive eruptions. Among those observa-
tions, none will be better evidence than such as prove the abrupt succession, by ejec-—
tion from the same volcano, of two rocks so dissimilar in composition as rhyolite and
basalt. On the other hand, the nature of volcanic action will explain why we should
meet among lavas, more frequently than among massive eruptions, with the fact of
two successive epochs blending into each other by the alternation of the two kinds of
rock peculiar to them separately, and it cannot be surprising if instances are occasion-
ally observed exhibiting, at least partly, a reversed order of succession.

Lassen’s Peak. It is probable that it indicates the existence of a fourth epoch in the activity of that voleano. The ejection
of basalt has been so frequently connected with the opening of vents in the neighborhood of, but not coinciding with, channels
through which its predecessors had ascended, that its local separation cannot be an argument against its belonging, in our
case, to the system of Lassen’s Peak.

13 The fact that trachytic lavas are frequently followed by such of basaltic character has been known since long
time, and was till now the only law of succession observed. Mr. Scrope has suggested the hypothesis indorsed by Mr. Dar-
win, Sir Charles Lyell, and other distinguished geologists, that in the subterranean reservoirs of voleanic matter, the heavier
particles will occupy the lower part, and the lighter ones be nearer the earth’s crust. It will easily be seen how totally
inadmissible this theory is in the ease of Lassen’s Peak. It is not less so in those cases where rhyolite was succeeded by
basalt, since the process of liquation can certainly not be supposed to have produced an abrupt passage under ground from
one mass to the other, and it would be much more natural to suppose a gradual transition to take place, there as well as in
the succession of the rocks emitted to the surface, if liquation had really taken place.

(72)

Cr

OF VOLCANIC ROCKS. 3

An instance, which is instructive on account of its simplicity, is furnished by
the island of St. Paul, in the Indian Ocean. F. von Hochstetter found its foundation
to consist of rhyolitic rocks. These are intersected by basaltic dykes. Khyolite over-
lays the first basaltic formation, and is itself superposed, first by dolerite and then
again by basalt. These two rocks of the basaltic order constitute the main body of
the island, and encircle its crater. Similar relations, though on a much grander scale,
have been observed by the same eminent geologist on New Zealand. More frequently
than this order of succession between rhyolitic and basaltic lava, has been observed
the sequence of basalt to trachyte, with the omission of rhyolite, or immediately to
andesite, when both those rocks are absent. Vesuvius is built up of rocks of the
basaltic order, and still emits lava corresponding in mineral character to its predeces-
sors, while the rocks of its surroundings (Campi Phlegrai), on the prior origin of
which geologists agree, are trachytic. The industrious explorer of Mount Etna, Sarto-
rius von Waltershausen, has described its foundation as being composed of white and
reddish colored trachytic rocks, which contain hornblende as a characteristic ingred-
ient, while among those rocks which build up the summit, as also in all modern lava of
the volcano, no hornblende but, in its place, augite is visible. This mineral and labra-
dor compose the recent lava, which belongs to the basaltic order. The much more
extensive recurrence of a similar order of succession in the Eifel and in Auvergne,
is too well known from the accurate descriptions of the geology of those regions, to
require to be here more fully mentioned. It contributes especially to confirm our
proposition, that the volcanoes of the different orders, as regards their origin, have
been nearly contemporaneous with the correlated massive eruptions. The classical
descriptions of the Hifel, by Mr. von Dechen, give conclusive evidence thereof.

Among those voleanoes the lava of which has never undergone a material
change and is, at the same time, similar in nature over the area of larger volcanic
districts, may be mentioned, besides numerous basaltic voleanoes, those of the trachytic
order in Central, and those of the andesitic order in South America, as far as may be
seen from the descriptions given of them.

We might greatly enlarge this enumeration of observations confirming our
propositions ; but, as by most authors only a ‘‘ trachytic” or a ‘‘ basaltic” character
of lava have been mentioned in a general way, they would only furnish evidence in
favor of the general tenor of the law, but would fail to give it in regard to any of its
details.

RELATION oF Voucantc Rocks to AnciENT Hruptive Rocks.

All rocks which, bearing evidence of an intrusive or eruptive origin, preceded
in age the Tertiary period, may, by principles similar to those which we applied in
tracing the natural system of volcanic rocks, be divided into two great classes, for
which we may use the terms “ granitic rocks” and ‘‘ porphyritic roeks,” derived from
the mode of texture predominating in either class. Granitic rocks are, besides,
geologically associated with granite, which is their principal type, while quartzose
porphyry occupies a similar position among porphyritic rocks. The annexed table

(73)

NATURAL SYSTEM

RICHTHOFEN

36

“ad0-U0IT
DOUBvPY snosojsruvyLy,
oyny
TOPVIGL'y

*90-U0dT
DYoUTR, SNOLoJULITT,
‘apuayquaoyy ‘ayany
‘opuaquy ‘asvpoosyo

‘910-W0aT
OYOUSVAY SHOLasUVyzLy,
‘opua[quaoyy ‘AsupodO,
(‘010-W0aT
dyousry_y snosojiuvsty)
‘opua[quaoyy ‘osepoos[g

(-zqanny)
{(oyorg) ‘opuoyquso FT
‘osnpooyyg ‘osepoosyg

(opuarqui0 fy)
‘OMOLEL
OsupOs ITO,
asepouyg
*Z.cun?)

————

qqesvg, “pe “WNT

‘aya[o(d] “4ST “Wey
1pspg—YL A P4O
oyyAdorg onisny “pg “wey ‘oHSepUY ONLINY "ps “Wey
‘aYISOPUY OIpus|quAo;T “YS “WR

‘ayyAdorg orpuce;quioyy “pe “wey ‘opsopup—ypnoy LapaQ

‘oyAtOVAT-osvpoosyg "ps ‘Wey

‘oyyAdoag-osozjaenty 4ST “Wey oyAYoua-UIpluBy “4YST “Wey

yyhdonJ~—Puy,T, PLO ‘ayliyood [, —puoogy Lape

‘sodoad oyoAqyy “pe “wey
(2zyaenty) ynoyyIA soar A )
oyuedry “pg “Wey

‘OUPCAON “YST “WUT (‘solouv A snosazizqaen?)

agyohyy—psluy Lop

“SINTIGHUONT TWILNASSOT

“SNOOY OINVOTOA + SSVI) GUINY,

adydaog onsny + Apr [UO
hahyduog opbhny—yjenoy pso
“Aad ydaog

omsny pue adydepoyy
UOOAJOd O}VIPOUMI}UL SyooY “"pZ “Ule,T

adda 4ST “Wey

whydopy Penh PO

opdydaog + Ayre 4pUO

‘apuhiydlog —puovrg “apo

‘ZWAENY) MOUILA SAONBA "pZ “WRT

“£akyduog esozywngy 4ST “Wey

hahiy duo, 2s] —BU P40

‘SUOOY OMIMAUAIOg + SSVI ANooUg

—

‘oSuquiq “PZ “Wey
‘oyuorysuod AFT puv oaqquy 4s] “WT
aspqnu—Yjenog LAPLO
‘asnqri(y puv aMol(y
WOIMJOq OPBIPSMAIOJUL SYOOY “PS “WY
‘OPMOT “AST “Wey

apo — PL, P40

‘oqmakg + Ayre ATCC

‘apiualigy—puoragy oplC

‘ouluvay ontuedg “pg “Wey
‘AUBIN “ps “UB
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(74)

OF VOLCANIC ROCKS. 37

will show the mutual relation of these two classes and their subdivisions, and of
either of them to volcanic rocks."

It appears that this general classification is based upon as natural principles as
are within reach of our still limited knowledge of eruptive rocks, and therefore may
at least approach the natural system. The following are the systematical principles
chiefly involved :

Ist. Chemical Composition.—Hach class contains all possible compounds inter-
mediate between those which Bunsen styled the normal trachytic and normal pyroxe-
nic types, and may, therefore, be represented by a numerical series of infinite grada-
tions within two certain limits, and progressing according to a definite arithmetical
law. If this law applied to the composition of rocks with mathematical precision, it
would be sufficient to know the relative quantity in which any one single ingredient
enters into the same, in order to find by calculation the relative proportion in which
every other ingredient should be present. It is, however, well known, that analysis
shows ordinarily a slight deviation from the composition as required by theory ; and,
considering the various influences to which the rocky masses must have been exposed
before and after their consolidation, we should naturally presuppose that such devia-
tions would be the rule, and may indeed be astonished to see how slight they generally
are. Silica has been found to be not only the most convenient, but also, on account
of its predominance over other ingredients, the safest element by which to determine
the place any rock occupiesin the series. In the classification as proposed in the pre-
ceding page, each class of eruptive rocks commences with those varieties of granite,
quartzose porphyry and rhyolite, which contain the highest amount of silica as found
by analysis, and descends to the most basic varieties of diabase, augitie porphyry and
basalt. In a chemical respect, therefore, the three classes are identical.”

M4 This table is only designed to show the mutual relations of the subdivisions of the three great classes of erup-
tive rocks in their most general outlines. I have purposely avoided to detail them any more, since it appears that great
progress in regard to their knowledge will be made in the next years, and considerable changes in the details of classi-
fication may have to be looked for. The composition of eruptive rocks is just at the present time being made an object of
more careful study than it ever was before. Contributions of high value in regard to the chemical and mineral com-
position of volcanic rocks are being furnished by the members of the Geological Institute of Vienna; while H. Abich
continues with untiring energy his fruitful researches on the same rocks of Armenia, the Caucasus, and the borders of the
Caspian Sea. The examination of the granitic rocks by G. vom Rath, A. Streng, Th. Scheerer and many others; the
more general labors of G. Rose, Robert Bunsen, A. Delesse, G. Bishoff, F. Zirkel; the microscopic investigations which
have been commenced with so much success by A. Sorby, and are being extended by his numerous followers—all these
labors pursued zealously by those named and a great many other workers in the same branch of geological science, whom it
would be too lengthy to mention, promise a rapid adyance of our knowledge of the properties of eruptive rocks. In
respect to their mineral composition, the discovery of some new species of feldspar (such as the plagioclase of Rath and the
microtine of G. Tschermak) which appear to be of wide distribution among the components of eruptive rocks, promises
alone to enlarge quite considerably the basis of classification, as it appears to give a clue to the causes of the differences in
outward appearance (frequently referred to in this paper) of rocks which are alike in regard to their chemical composition,
and among the mineral ingredients of which no difference could be recognized heretofore.

15 Objections have been raised against the validity of the law of Bunsen, partly on the ground of the frequent
discrepancy of the figures obtained by chemical analysis from those which would be required by the theory, and partly on
account of the highest and lowest amounts of silica occurring in eruptive rocks having as yet by no means been fully
ascertained. We referred above to the first objection. As regards the second, it is evidently very triflivg in importance.
The limits of the series may, and probably will, be somewhat extended, and the figures representing the compounds which
form those limits may have to be slightly changed, yet the series itself will remain essentially the same. The law of Bun-
sen will have to be revised and corrected when increased experience shall have established a broader basis for it ; but no
change of its principles may ever be expected, as an overwhelming amount of evidence has accumulated in support of its
essential tenor. J (75)

58 RICHTHOFEN — NATURAL SYSTEM

2d. Mineral composition, in respect to which each class represents a series of
gradations essentially dependent on the chemical composition, and therefore chiefly
coincident with the chemical series, but differing from it inasmuch as it is more articu-
late. Certain types, corresponding to certain steps in the chemical scale, and consist-
ing in distinct aggregations of a few minerals, are the centers for which the petro-
graphical names have been applied in the first place ; around them other members are
grouped which connect every two types by gradual passages, and are ordinarily com-
posed of an aggregation of all the minerals peculiar to either of them. Expansion of
the series, as it were, in a lateral sense, occasioned by the accession of minerals inferior
in importance, are not unfrequent, but do not affect in a great measure the definite
character of mineralogical gradation, as they are of local occurrence, and probably less
dependent upon any material differences in chemical composition, than upon certain
influences which acted upon the mass of the rock, either before its ejection, and
then by the admixture of matter differing from it in nature, or after its solidification,
and then by chemical metamorphism.

3d. Specific gravity, which increases in the reverse proportion of silica. In this
respect, too, each class represents a series of infinite gradations from the lowest to the
highest value.

Taking the three foregoing principles exclusively as the basis for classification,
all eruptive rocks would have to be united into one class. This union, made regard-
less of any other relations, may be traced as the leading feature of nearly all systemat-
ical arrangements proposed. There are, however, other points of view which must be
considered if a more perfect classification is aspired. They lead to the establishment
of further separations, by principles similar to those which we had to apply above in
defining the different orders of the compounds of hornblende and oligoclase among
voleanic rocks. These points of view are the following :

4th. Mode of Texture —Eruptive rocks exhibit, in regard to their modes of text-
ure, very peculiar differences, which are little capable of explanation with our present
state of knowledge. They are especially conspicuous with the most silicious rocks.
Free silica, in granite, has probably been solidified cotemporaneously with the other
component minerals ; but its solidification appears often to have been completed last
of all, as the quartz does envelop the aggregation of the other crystals. In quartzose
porphyry and rhyolite, on the other hand, free silica, at least the greater part of it,
has been solidified first, which is obvious from the fact that its perfect crystals are
imbedded in an ordinarily microcrystalline aggregation of the other ingredients, which
however contains, in most cases, some surplus of free silica that had not entered into
the composition of the crystals. This fundamental difference from granite points to
some difference in the mode of origin. Quartzose porphyry and rhyolite differ in regard
to the texture of their paste, which has a compact aspect in the first, while it is more or
less vesicular throughout the mass in most varieties of the latter. This difference, like
the former, appears to indicate, that the molecular condition of the liquid mass, at the
time when it was ejected, was different in either case. The variety of texture diminishes
with the decrease of silica, and the more basic rocks of the three classes bear a much

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OF VOLCANIC ROCKS. 39

closer resemblance to each other than the silicious rocks. The cause must be, either
that the conditions of the mass before cooling were less varied with basic than with
silicious rocks, or that the differences were alike, but manifested themselves less con-
spicuously in the character of the solidified rocks ; the latter cause is the most probable
of the two, since even our cotemporaneous basaltic and andesitie lavas offer but a small
number of varieties comparing with those of rhyolite or trachyte, although it is not
likely that there is any more variety among the circumstances modifying the latter than
among those which are acting upon the former. It is chiefly the similarity in texture
which has occasioned the general application of such names as “‘ trap,” ‘‘ greenstone,”
‘‘aphanite,” and others, for basic rocks of all ages, while those containing a large
amount of silica have scarcely ever been similarly confounded. We may, however, note
this difference among the former, that granitic texture—that is, a erystalline aggrega-
tion of the component minerals—is peculiar to those basic rocks associated with granite,
while the vesicular or trachytic texture is only proper to voleanic rocks. Occasionally,
though very rarely, rocks of granitic texture are geologically associated with porphy-
ritic rocks, as is the case near Predazzo, in southern Tyrol.

5th. Geological age ; granitic rocks being generally the most ancient in origin,
volcanic rocks the most recent, while those of porphyritic structure are intermediate in
age between both.

6th. Other geological conditions resulting from the correlation of the different
foregoing principles, the outlines of which I will attempt to trace in the following
pages.

Correlation of Age and Texture.

It is scarcely possible to treat of one of the foregoing principles in its appli-
cation to eruptive rocks singly, without constantly encountering points of intimate
connection with others, so thoroughly are they intertwined and mutually dependent.
It is the object of petrology to develop these correlations. We contemplate them
here only in their purely geological bearings, in order to try to establish the true rela-
tion of volcanic rocks to their ancient predecessors. We have, while occupied with
these considerations, constantly to keep in view how few are the observations upon
the strength of which we have to base conclusions. The area on the globe whose
special geology has been made the object of investigation, though extending every
year, is still very limited; and even in the best explored countries, little attention
has, in most instances, been paid to the distinction of eruptive rocks. Observations in
regard to them are abundant in some parts of Europe, fortunately in such countries
as are especially capable of giving a clue to their general knowledge. Distinct
conclusions may be arrived at in regard to portions of that continent, but little scope
is afforded for giving them latitude by comparison with the relations presented in other
countries. We have, therefore, to keep well separate in general petrology, those posi-
tive conclusions which are founded upon sufficient observations, and are applicable to
the relations in those countries where the latter were made, and the realm of theories,
which are arrived at, partly by the generalization of those conclusions, and partly by
making deductions from hypothetical suppositions; because, the premises being founded

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40) RICHTHOFEN—NATURAL SYSTEM

on local occurrences, and their general validity not being proved, the theories must
necessarily have a great deal of uncertainty, which will only gradually be dispelled by
the advancing knowledge of the geology of the globe.

The correlation of age and texture, as resulting from observations made in
Burope, will occupy us first. Granitic rocks are widely distributed on that conti-
nent. Their great eruptive masses are of Azoic and Paleozoic age. The rocks
have almost throughout granitic texture, though the distinguishing features of
porphyritic rocks are proper to some subordinate varieties of diorite and diabase.
The eruptive activity exhibited in the granitic era gradually relaxed. It appears
to have been insignificant, though it was by no means extinct, in the latter part
of the Devonian and the first part of the Carboniferous periods. But it recom-
menced about the time of the deposition of the coal measures, thence increased in in-
tensity, and was most violent in the Permian age, though it was, during all that
time, much more limited in extent than it had been during the granitic era. The mid-
dle portions of Germany were then its principal theater, until it changed, in the com-
mencement of the Triassic age, to the southern slope of the Alps and Carpathians.
The rocks produced during this era possess, almost exclusively, porphyritic texture.
The Jurassic, Cretaceous, and the first part of the Tertiary ages, were distinguished
by an almost complete repose of eruptive action, in Hurope. It was only after the
commencement of the Tertiary period when that violent resumption of eruptive ac-
tivity took place to which we have repeatedly called attention in the foregoing pages,
and which thereafter continued, gradually relaxing, down to our present era, the mani-
festations of volcanic action in which are its last faint remnant. Vesicular inflation,
which is the characteristic feature of trachytic texture, is peculiar to the rocks of this
class ; all of them possess it more or less, though there are varieties resembling por-
phyritic rocks closely in aspect. A combination of the trachytic and porphyritic modes
of texture is of more common occurrence than either of them singly, and is indeed
the distinguishing feature of volcanic rocks.

These relations of age and texture have been conclusively proved to exist in
Europe, and appeared to justify the conclusion, that the three classes of eruptive rocks
are geologically separated, and represent three successive and distinct phases of the
manifestation of subterranean agencies. Considering in their generality the facts ob-
served in other countries, they appeared to confirm these views, since granitic rocks
are known to be generally very ancient ; volcanic rocks to have been generated during
the Tertiary and Post-tertiary periods ; while, in regard to porphyritic rocks, observations
have been scarce, and descriptions lack distinctness, yet no relations have been recorded
in reference to them which would be contradictory to those observed in Europe. On the
western coast of North America, however, eruptive rocks exhibit some peculiar rela-
tions—diflering to some extent from those which they offer in Hurope. Positive facts
are known sparsely outside of California, but conditions similar to those observed in
that country appear to be repeated through large portions of the range of the Andes.
Prof. J. D. Whitney’s admirable researches on the age of the metamorphic rocks of
the Sierra Nevada have clearly demonstrated that the granitic rocks, which partake to

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OF VOLCANIC ROCKS. 4]

ereat extent in the structure of that mountain range, cannot possibly have been
ejected prior to the Jurassic epoch. The texture of these rocks is, notwithstanding
this comparatively recent origin, that of all true granite, and the prominent varieties
cannot be distinguished from some European kinds of granite, as for instance those of
the Adamello and the Cima d’Asta in the Southern Alps, which are among the most
recent in age on that continent. Volcanic rocks are widely distributed in the Sierra
Nevada, and are of the same or similar age as in Europe. Quartzose porphyry occurs
to some extent in Washoe, under circumstances which make the exact determination
of its age difficult, but render it certain that it is intermediate in this respect between
granitic and voleanic rocks. These relations would appear to be an exact counterpart
of those observed in Europe, with the one prominent difference, that the commencement
of the eruptive action was much later in America. Very recently, however, additional
observations have been made, which give a somewhat different aspect to these relations.
Mr. Clarence King observed granite, covered by Paleozoic rocks and antecedent to
them in age, near the Colorado River ; while Prof. Whitney and myself discovered true
quartzose porphyry in the county of Plumas, in northern California, associated with
rocks proved by the former to be of Triassic and Liassic age, in such way as to leave
little doubt about its cotemporaneous origin. Farther east, in the Great Basin,
Palwozoic granite is of no rare occurrence, and it is among the prominent features in
the geology of the Rocky Mountains ; while the discovery of porphyritic rocks may
have to await further examination, they having been in most countries the last erup-
tive rocks to be detected.

However these facts may affect the theoretical conclusions in regard to the
origin and mutual relations of granitic and porphyritic rocks, which had been made on
the strength of former observations, they appear to confirm the separation, from a geolo-
gical point of view, of both classes of rocks. There has been in the Sierra Nevada and
adjacent countries, it appears, an ancient granitic era corresponding to that of Europe,
followed by a porphyritie era which was nearly or quite coincident with the European.
But, while the manifestations of subterranean agencies almost ceased in Europe during
the following ages, they recommenced with great intensity on the western coast of
North America, and gave rise to a second granitic, followed by a second porphyritic era.
The voleanic era commenced in both countries in the Tertiary epoch, but it appears to
have been in an advanced stage in Europe while it was still in its birth-throes in
America.

Correlation of Age and Composition.

This point of view is not inferior in interest to the foregoing. The most
noteworthy fact is this, that quartzose, and in general highly silicious rocks prevail
among those of ancient origin, basic compounds among those of later ages. Granite
and syenite are overwhelmingly predominant among ancient eruptive rocks. Diorite
and diabase are generally associated with them, but remain always quite subordinate
in bulk. The relative proportion is different with porphyritic rocks. So little
attention has been paid to these, outside of Europe, that general conclusions in

: (79)

42 RICHTHOFEN—NATURAL SYSTEM

regard to them should be drawn with care. In the middle part of Germany,
and in southern Tyrol, where they have been repeatedly studied, subjected to
chemical analysis, and described in numberless treatises, quartzose porphyry is
rather predominant. But porphyrite, melaphyr, and augitic porphyry, are, in the
aggregate, little subordinate in bulk. The volcanic offers the complete reverse
of the granitic era, respecting the proportionate quantity in which the different com-
pounds have come to the surface. Andesite and basalt compose as large a proportion
of the aggregate bulk of volcanic rocks, as granite and syenite do of those of the
granitic era.

Some minor differences in age may be noticed among the different orders com-
posing the three classes of eruptive rocks. Extrusions of granite and syenite appear
to have been almost the exclusive feature of the eruptive activity during granitic eras,
and to have been succeeded only towards their close by the emission of diorite and
diabase, or of other rocks of limited occurrence, such as gabbro and hypersthenite,
Such at least has been observed to be the case in several countries, in regard to the
chief outbreaks ; but if we enter into the details of the mode of succession of the
rocks belonging to the different orders, we perceive that it has not been so definite
as with volcanic rocks—granite and syenite bearing evidence, in many localities, of a
more recent age than some neighboring masses of basic rocks. Yet, in keeping only
the main features in view, we may easily see, that the general order of succession of
granitic rocks has been conformable to a gradual decrease in silica. Syenite is usually
more recent in origin than granite; and even among the different varieties of the lat-
ter, true granite, containing the highest ratio of silica, has generally been anterior in
age to G. Rose’s granitite. There may be some connection between these relations
as they are exhibited in any single granitic district, and the fact that the granitic rocks
of the Sierra Nevada, belonging altogether to a more recent era, contain no true granite—
their chief bulk consisting of rocks intermediate in composition between granitite and
syenite. The porphyritic era, in Germany and on the southern slope of the Alps, was
inaugurated by eruptions of quartzose porphyry, and has terminated in the Alps by
those of augitic porphyry. The intermediate epoch has been distinguished by rocks
intermediate in composition. The mode of succession of the different orders is more
distinct than with granitic, but less so than with volcanic rocks. Melaphyr and por-
phyrite interchange frequently ; but, at many places, the former appears to have pre-
ceded the latter in age, in a similar way as andesite preceded trachyte. They form’
together one epoch, which, in the commencement, was occasionally interrupted by
an outbreak of quartzose porphyry.—The laws of the periodical succession of the five
orders of volcanic rocks have been developed in the foregoing pages. Their epochs
are much more distinctly separated than those of the ancient rocks, but the mode of
their succession is more complicated. With granitic rocks, silica as a component part
decreases with the age; with porphyritic rocks the same is true in a broad sense—the
precedence in age of melaphyr and porphyrite forming the only conspicuous devia-
tion ; while in reference to the voleanic era, no rule at all may be discovered at first
sight. Considering, however, the predominant rocks of that era, which are propylite,

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OF VOLCANIC ROCKS. 43

andesite, and basalt, we perceive that the former two were the precursors of the latter,
containing at the same time silica in a proportion superior to basalt. The singular
phenomenon of trachyte and rhyolite being ejected at a time intermediate between
the epochs of those rocks, is a notable deviation from the mode of succession peculiar
to granitic and porphyritic rocks. We will try to consider its probable causes in another
chapter.

Correlation of Eruptive Rocks in regard to their Geographical Distribution.

Another point of view offering in the contemplation of eruptive rocks, and which
has a close bearing to their natural system, is the correlation among them in regard to
their geographical distribution. This is, however, a vast subject, and volumes might
be written in collecting evidence for final argument of general value. At this place
we can give it only a cursory notice. We will consider, first, what is the mode of
distribution peculiar to each class of rocks, and then trace the correlations perceptible
among them in regard to their different modes of distribution.

Granitic rocks are scattered widely over the globe. Wherever its surface is
composed of ancient sedimentary rocks, and these give evidence of disturbances of
some intensity, by the plication of their strata, we may be almost certain to find granite
entering into the geological composition to some extent. In the diversified structure
of the European continent, geological maps show the existence of granite in nearly
every prominent mountain-range. Considering among them the range of the Alps and
Carpathians, we find granite scattered over its whole extent, from Savoy to Transyl-
vania, particularly on the southern slope. It forms some prominent summits, but
occurs also in subordinate positions. The geological relations of several of these
places have been examined, and careful investigations made of the mineral and chem-
ical composition of the rocks. They have resulted in proving an individuality of
granite such as is peculiar to no other kind of eruptive rocks; different eranitic
masses are of different age, and exhibit a corresponding diversity in regard to the
composition of the predominating rocks. Sometimes, it is true, several neighboring
masses are similar in respect to their petrographical character, and bear evidence of
being nearly of the same age (for instance those of the Adamello, the Cima d’ Asta,
and Brixen), but others, next adjoining, will be found differmg from them in nature
and in age. The length of geological time during which they have been ejected, has
never been established ; but the period appears to have been one of immense dura-
tion. <A similar individuality as to age and mineral character may be noticed in
respect to the granite of other mountain ranges on the European continent. Alto-
gether, the mode of distribution of the ancient granitic rocks, as far as they enter into
the structure of the surface, may be said to be in numerous small districts, which are
independent of each other in regard to their epoch of ejection and petrographical char-
acter. With reference to the latter, each separate district shows a great preponder-
ance of rocks belonging to one of the families of the granitic order, which are usually
accompanied by syenite in smaller proportion, and by some subordinate eruptions of dior-
ite and diabase. These districts are principally scattered along the present lines of eleva-

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44 RICHTHOFEN—NATURAL SYSTEM

tion. But this may be due in part to their concealment, in the spaces between those
lines, by sedimentary rocks. More signs of the separation of distinct ‘‘ regions of eruptive
action,” as they may be called, are exhibited in the porphyritic era. Eruptive activ-
ity has been intense within them, but appears not to have spread beyond certain boun-
daries. Each of these regions embraces a number of the former granitic districts,
while it leaves others excluded. Some resemblance with the peculiar features of the
granitic era is afforded, inasmuch as each porphyritic region has been independent
from others in regard to the epoch of its eruptive activity. We mentioned before
that one of the porphyritic regions comprises the middle part of Germany, while
another stretches along the southern slope of the Alps and Carpathians.—If we
proceed to the volcanic era, it presents to us the reverse of the individuality
peculiar to granitic districts, in the wonderful unity exhibited in regard to time
and space over the whole area of extensive belts. In reference to unity of time
alone, we might call the greater part of the continent of Europe, and even the
entire surface of the globe, one great region of eruptive action, during the vol-
canic era, since the first emission of rocky matter has been nearly cotemporaneous
in widely separated countries, while its culminating epochs have probably varied but
little in them separately, and the rocks have been ejected everywhere in a similar
order of succession. In regard to local distribution, however, we have to distinguish
certain belts, far exceeding in area the porphyritic regions. Each of them extends
over a number of preéxisting mountain ranges, and the eruptions in each have fol-
lowed, in their distribution, chiefly the lines of former elevation and ancient sea-
coasts. But there is unmistakably to be recognized a tendency of the agencies which
caused the eruptions, to connect these separated ancient belts of elevation ; either lon-
gitudinally, when ranges superior in extent to the preceding ones would be formed, as
appears to have been the case in the Andes; or, as it were, ina lateral sense, when
the connection of neighboring mountain ranges into table lands would be either
initiated or promoted. One of these belts, consisting of parts which had previously
been disconnected, may be traced from Armenia to the Rhine, though I will try to
show in the sequel that it is only a part of another belt which is of far greater extent.

We mentioned before, that porphyritic rocks are encountered chiefly in those
places where granitic rocks had preceded them. As regards the voleanic belts, eruptive
activity has been particularly violent in certain portions of them. It is worthy of
note that, wherever this has been the case, either granite or both granitic and por-
phyritic rocks had been ejected before. This fact leads us to consider the correlation of
the three classes of eruptive rocks in reference to their peculiar modes of distribution.
Little information exists in regard to this subject. Only one instance” shall be related,
which is highly suggestive for the existence of such a correlation, though it is of slight
value as long as it is not corroborated by corresponding facts observed in other coun-
tries. A survey, on a geological map of the Alps and Carpathians, of the southern
boundary-line of those highly metamorphosed formations which preceded the Trias in

16 Referred to in Richthofen, Geognostische Bechreibung der Umgegend von Predazzo, Sanct Cassian und der
Seisser Alp in Siid-Tyrol : Gotha, 1860.

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OF VOLCANIC ROCKS. 45

age, and chiefly compose the central portion of the Alps, shows that it is directed from
west to east in Lombardy, but in the vicinity of Lugano bends suddenly to the north-
sast, then turns as abruptly back to its former direction from west to east. After
having passed the granitic mass of the Adamello, the same change is repeated on a
grander scale. The boundary is turned again in a perfectly straight line to the north-
east, and then resumes its former course, which it follows in an equally direct maun-
ner, and in which it continues, exceedingly distinct at first—less distinct, by the
encroachment on it of more recent formations, farther east—until it turns a third time
to the northeast at the sudden termination of the Alps near Vienna, and continues in
this direction for a long distance. Finally it re-assumes, in the Carpathians, a similar
course to that which it bad on the southern slope of the Alps. Three very distinct
reéntering angles are thereby formed. The first of them encloses the country of Lu-
gano ; the second comprises the vicinity of Predazzo and Fassa in southeastern Tyrol,
and of Belluno and Vicenza in Venetia; while the third, which is by far the most
extensive, comprehends all northwestern Hungary. Hach of them has been a center
of eruptive activity, commencing with the granitic, and continuing through the por-
phyritie down to the voleanic eras, and all three are among the most classical coun-
tries for the study of eruptive rocks. There is, however, a conspicuous difference in
the mode of manifestation of the eruptive activity in each of the three eras. Little
connection exists apparently between the granitic masses of the three countries. They
are portions of the generally scattered granitic outbreaks, and differ among themselves
probably as much in age as they do in regard to the nature of their rocks. In the
porphyritie era, eruptive activity was contemporaneous in the three localities, but
scarcely extended beyond them. In the volcanic era, when the southern slope of the
Alps and Carpathians formed only a portion of a much more extensive. belt, the
countries adjoining those three places were chiefly distinguished by the intensity of
eruptive activity.

Great as have been the interruptions between the different eras, the continu-
ance of the selection of those three nooks at the foot of a prominent mountain range
for the manifestation of subterranean energy, from Palaeozoic down to modern time,
is evident. Similar instances, though less striking, might be mentioned from other
parts of Europe, such as the porphyritic region of middle Germany. Reverting to
other parts of the globe, it appears to be a general experience, though it is far from
being absolutely proved, that all the principal accumulations of volcanic rocks are
encountered in the neighborhood or immediate vicinity of granitic masses. These are
scattered over areas where no voleanic rocks occur ; but the distribution of the latter
within any of the volcanic belts appears to have been dependent, in a great measure,
upon the vicinity of the channels which had in preceding time afforded vent to granite.

The general law deduceable from these relations is this: that, with the growing
thickness of the earth’s crust, the systems of fractures which were formed in it at cer-
tain epochs, and partly gave vent to the emission of rocky matter, increased in depth
as well as in length, and were more and more concentrated to definite portions of the
crust, which are recognizable upon the earth’s surface by the partly coincident areas of

K (83)

46 RICHTHOFEN—NATURAL SYSTEM

eruption ; that further, simultaneously with the increase in extent, each area manifested
a growing complexity in regard to the distribution of eruptive activity within it, and
that this distribution depended in a great measure upon that of the outlets of more
ancient eruptive matter. The first recognizable stage of this development, which,
however, was probably a far advanced one, is the individualization of those nu-
merous districts of fractures, the narrow limits of which are made manifest by the
mode of occurrence of Azoic and Paleozoic granite. A growing development from that
stage, in the different directions mentioned, is conspicuous in the porphyritic era in
Europe ; when, besides the greater extent of the regions in which subterranean agencies
manifested themselves by the fracturing of the crust, every such area had become
more definite concerning its boundaries towards those which were not fractured during
the same era. On the other hand, however, each porphyritic region was more complex
than the granitic districts had been, inasmuch as the former were composed of certain
areas of greatest activity, which, as we had occasion to remark, appear to have been
chiefly dependent upon the distribution of the granitic districts enclosed within each
porphyritic region, while intermediate portions were contemporaneously affected by
disturbances merely, but not by any eruptions. A more advanced stage in this gradual
development seems to be exhibited by the Jurassic granite of the Andes. The belt
distinguished by its eruption appears to have been distinctly bounded, and the area
over which contemporaneous manifestations of like character took place, to have been
even more extensive than the porphyritic regions of Europe. All these features, how-
ever, are conspicuous on a much grander scale in the voleanic era. The growing
definiteness of the boundaries of the volcanic belts towards large areas which were
entirely free of eruptive activity, the increasing complexity of their interior arrangement,
and its dependency upon preéxisting lines of elevation, and particularly on the
distribution of granitic and porphyritic rocks, are evident from what has been said before.
We may even trace a development from the andesitic to the basaltic epoch. It is
known how far superior in extent is the range of basaltic outbreaks to the area occupied
by other volcanic rocks. If considered with attention, it will be found that there is a
tendency in the former to connect within each belt the subordinate ranges of the latter,
longitudinally as well as laterally. The only connection, for instance, between the
voleanic districts of Hungary and those on the lower Rhine, is occasioned by the chain
of isolated basaltic hills which extends through the central part of Germany.

On tHE OriGiIn oF Votcanic Rocks.

The facts established in the foregoing chapters, and the inferences drawn there-
from, may assist us in considering some questions of wider bearing. We have seen that
the volcanic rocks of several, and probably of all parts of the globe, are connected by
simple and definite relations, which together comprehend the main features of their
natural system; and we found that correlations of a similar nature ally the volcanic
with all the ancient eruptive rocks, justifying, to some extent, the supposition that the
eruptive rocks of all ages and places form one harmonious whole, and that we may be
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OF VOLCANIC ROCKS. AT

able to discover the laws of the natural system embracing their totality. In order to
arrive at a more perfect understanding of this system, we must now attempt to exam-
ine into the causes of those mutual relations. It would naturally occur to us that they
would be implied in the circumstances which attended the generation of the eruptive
rocks, and in the conditions in which they have been before arriving at the places they
occupy at present. We have, therefore, to investigate the following questions : What
was the nature of volcanic rocks before they arrived at the places which we now
see them occupying? Where did they originate? By what agencies did rocks con-
nected in widely separated places by simple and definite relations come to their
positions among others which bear no such relations either to them or among each
other? The importance of these questions for our subject, the attention which they
have attracted through the whole history of geological science, and the great diversity
of opinion prevailing in regard to them, will make it necessary to treat them more
fully than might otherwise appear consistent with the objects of this paper.

Voleanic action and massive eruptions, notwithstanding the similarity of the
material produced by both, would appear, from the most cursory review of the
phenomena connected with either of them, to differ to some extent, not only in regard
to the causes to which they owe their origin, but also in regard to the position the
matter occupied before its ejection. We have for this reason to keep again distinctly
separated these two modes of manifestation of subterranean energy. In regard to
massive eruptions, which will first occupy our attention, we can scarcely draw up any
argumentation without enlarging on the entire range of eruptive rocks, and extending,
at least partially, our views to them.”

1. On the Origin of Massive Eruptions.

In order to establish some positive premises available for drawing conclusions
in regard to the origin of the massive eruptions of volcanic rocks, we reiterate the
following facts, of which mention has partly been made in the foregoing pages:
Ist. The eruptive (including the voleanic) rocks offer a great diversity of chemical
composition ; but all the compounds represented by them are mutually connect-
ed by simple and definite arithmetical relations as regards the figures which

17 The following considerations are given notw ithout some hesitation, partly on account of the uncertain ground on
which they have to move, and partly because some of their main features are, of necessity, only well known theories repro-
duced, though, perhaps, under a somewhat different form. Yet, the establishment of the relations detailed in the preceding
chapters, and other observations made of late years, may allow us to arrive at more satisfactory conclusions in regard to
some weighty problems than could be done before, or, at least, to determine more precisely the only direction in which we
have to look for their solution. It should be borne in mind that among the theories recently proposed upon the subjects
specified above, there is not one which has not already had its prototype in the phantasmagorias of the time of the dawn of
geological science, and that it is these which have been constantly reproduced, enlarged, diversified, remodeled according to
the advance of science, and supported by continuous accumulation of evidence. Propositions which had been accepted as
being beyond the necessity of proof, and which are still occasionally reproduced as axioms in popular works on geology, have
been weakened, and not unfrequently overthrown, when facts newly revealed would withdraw their chief supports, but have,
after some time, revived under new forms. In treating on a topie where the degree in which the results of our speculation
appear satisfactory to us depends upon the degree of probability which we think we see in the theories arrived at, and of
their faculty of explaining observed facts, and where we are in constant danger of making incorrect deductions from imper-
fect premises, not enough can be done in the way of weighing the evidence by which different doctrines are supported ; and
this is particularly necessary in reference to those theories which we are too much accustomed to consider as matters of fact,
upon which further conclusions may be safely built. (85)

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48 RICHTHOFEN — NATURAL SYSTEM

express the value of the different ingredients entering into their composition. The
law of Bunsen, in which are embodied these relations, though true for all eruptive
rocks, has never been applicable to those of sedimentary origin, nor to the met-
amorphie sedimentary rocks. 2d. The order in which massive eruptions of vol-
canic rocks have taken place in different countries, is by no means conformable to
a regular succession of gradations in chemical or mineral composition, but shows
the existence of several distinct groups, each of which comprises, chemically, a certain
portion of the entire range of compounds intermediate between the two extreme types,
and has, besides, its own peculiarities of texture and mineral composition. — It is for
the latter reason that rocks, belonging geologically to two different groups, may be
identical as chemical compounds, and yet differ petrographically. We may repeat
here that these groups, wherever their mutual relations have been made an object of
study, occur in the same order of time, and that the rocks belonging to each of them
present generally a great similarity in character wherever they are encountered.
3d. The massive eruptions of volcanic rocks are distributed over the globe in certain
belts, differing in extent and width as well as in direction. ;

There are, among those of a general bearing, two manifest conclusions at which
we may arrive by the aid of these facts. The first relates to the origin of the matter
ejected. It may be said that equality of physical and chemical properties, and of the
mode of occurrence, will commonly involve equality of origin. We have, therefore,
to infer, that the source from which volcanic rocks have derived their origin has
been similar in nature in every locality, that the definite numerical relations, existing
in the chemical composition of the matter ejected must exist similarly at that source,
and that they must there pervade matter equally in different parts of the globe; and
we may further add, that at the same source the different kinds of matter which cor-
respond to the different passages in composition among volcanic rocks, must be ar-
ranged, at every locality alike, in a definite order of position in reference to the
distance from the center of the earth, since such relation in space can alone explain
the definite order of succession in time in which those rocks have been ejected. If
we consider the relations existing between voleanic and ancient eruptive rocks, there
can be no doubt that all of these have derived their origin from that same source, and
that it is of general distribution under the surface of the globe. The second general

conclusion relates to the cause and mode of ejection. As like effects imply like

causes, and the similarity of the phenomena connected with the massive eruptions of
voleanic rocks is conspicuous, we may infer that the agencies to which they have been
due, were, in the main, similar in all cases. However accidental and local circum-
stances may have caused minor differences, the prominent features in the mode of
geological occurrence and geographical distribution cannot be due to them. In how
far we may be justified to ascribe the ejection of the granitic and porphyritic rocks to
similar agencies, will depend upon the degree of similarity of their part in the struct-
ure of the surface of the globe with that of the voleanie rocks. The conclusion must
be drawn from those correlatious which have already been mentioned, that the funda-
mental causes of all eruptive activity are alike, and implied in general planetary pro-
cesses which are closely connected with the evolution of the globe. These general
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OF VOLCANIC ROCKS. 49

conclusions appear to be the only ones which are fully justified. If we venture to
inquire into the nature of the agencies which caused eruptive activity, and to explore
their fundamental causes, we have to transgress the limits of legitimate speculation
and enter the realm of hypothesis, the only way of proceeding in which consists in
weighing probabilities. Further experience, it may be hoped, will extend the limits
of induction, in the same measure as it will furnish increased evidence.

I will attempt, in the first place, to consider, what must be the nature, and
what the position of the source at which the voleanic and all eruptive rocks have
originated, in order that their physical and chemical properties and their mutual rela-
tions may be explained, and then to examine what have been the agencies that have
had, in all probability, the most immediate influence upon the facts connected with
the mode of distribution and succession of eruptive rocks in general. We have to
start in these considerations by a well-known argument.

If no changes had ever taken place on the surface of the globe, and no sedi-
ments had ever been deposited on it, but that state was still*preserved which the
globe must have exhibited when the substance of the rocks was liquid on its surface,
then the matter nearest to this would probably be of a nearly uniform chemical com-
position, and, consequently, of a uniform specific gravity. If we imagine the globe to
consist of concentric layers, it is exceedingly probable, from physical laws, that then the
substance of each successive layer would be, too, of a nearly uniform composition
throughout its extent. Proceeding, however, from those layers nearest to the surface
to those at greater distance from it, the specific gravity of the matter composing them
severally would necessarily increase gradually towards the interior. This would be
effected by the tendency of the heavier elements to predominate as the depth became
greater, while the lighter would increase in as gradual a ratio towards the surface.
If the layers were infinite in number, then the passage, in chemical composition as well
as in specific gravity, from the surface towards the center of the earth would be one of
infinite gradations. This condition is eminently the most probable, if physical laws
are taken into consideration. If the crust was then allowed to solidify, to any depth
required, the same interior arrangements would continue to exist unaltered, regardless
of any changes on the surface. That a condition similar to this actually exists at the
present time, is rendered evident by the well-known fact, that the specific gravity of
the mass of the globe greatly exceeds that of the average of the matter which com-
poses its exterior crust. Hruptive rocks have been carried to the surface from places
beneath it. The depth from which their material has been derived, the way in which
it was rendered liquid, and the cause and mode of its ejection, these are the principal
points of conjecture in regard to them. The most probable place of its derivation
are those imaginary layers which are beneath the theater of external changes, and still
occupy their primeval position. This appears to be manifest from the two facts to
which we have constantly to refer: the ejection of identical chemical compounds in all
countries and ages, and the exhibition by eruptive rocks, in each country, as well as
over the entire globe, of a series of compounds the specific gravity of which increases
in inverse ratio with the amount of silica. The suggestion that eruptive rocks, in
virtue of these facts, represent the arrangement of matter in the interior of the globe,

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50 RICHTHOFEN — NATURAL SYSTEM

those which are rich in silica and of little specific gravity bemg derived from places
nearer to the surface than those in which a smaller proportion of silica is attended by
a higher specific gravity, was first distinctly made by Sartorius von Waltershausen,
who attempted to prove this theory by mathematical calculation, and has at least suc-
ceeded in demonstrating its adaptation to the nature of eruptive rocks. The assump-
tion, hypothetical as it may still appear, has an eminent degree of probability, and is of
the highest importance for the geology of these rocks, for the reason that it is capable
of explaining numerous observed facts which have failed to be satisfactorily explained
in any other way. ;

We may still draw a line of negative argument in corroboration of this theory.
It is perfectly clear that the source of eruptive rocks must either have been below the
lowest depth at which rocks of sedimentary origin occur, or above it. We found that
the former assumption gave a satisfactory explanation of prominent facts, and it is
just as conspicuous that the second fails to give any explanation at all. If eruptive
rocks had had their original seat within the crust of sedimentary rocks, and had been
generated from their substance, they must be analogous to them in chemical compo-
sition, that is to say, they would vary in this respect within very wide and complex
limits in any single country, and we should encounter no lesser differences when com-
paring the totality of eruptive rocks in one part of the globe with that in other parts.
The fact that no arithmetical relations can be recognized as connecting the various
sedimentary rocks in their composition, would hold equally good among eruptive
rocks, since it is impossible to conceive a progress from the indefinite to the definite,
from that which is void of any recognizable relations in respect to the composition of
matter to that in which such definite relations are plainly evident, by the mere
influence of such agencies as would cause fluidity, (that is, the combined action of
heat, pressure, and water.) As no agency, indeed, is known to which such a result
may be ascribed, this argument removes beyond the limits of probability the
assumption, that the original seat of eruptive rocks has been above the foundation of
the gradual accumulation of those rocks which have been produced by external
changes, and therefore allows only the other alternative, that it has been below that
boundary.

While the theory of Sartorius will explain the causes of the prominent mutual
relations existing among eruptive rocks in regard to their composition and general
similarity in all countries, we have still to trace those causes which effected their ejec-
tion from a deep-seated original place to the surface. This question is more abstruse
than the foregoing, inasmuch as the manner in which forces have formerly been acting
beneath the earth’s crust, is a subject more involved, and allowing a larger range of
hypothetical explanation, than we met on a field where the observed properties of
matter offered a comparatively safe guide. Complicated as the processes appear to
have been, to the codperation of which massive eruptions were due, it seems that
they may severally be traced back to one common cause of a higher order, of which
all, or most, of them are but different modes of manifestation. Probabilities accumu-
late to point out as this fundamental cause, the process of the gradual cooling of the
earth and the solidifying of its crust toward its interior.

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OF VOLCANIC ROCKS. 5]

Since the early day of the ingenious speculations of Descartes, this great and
general cause has been considered as the main agency to which the disturbances
on the surface of the globe are due; and, though having been more or less in
favor at different times, the doctrine has at no time been completely abandoned.
The various aspects which it has periodically assumed, the latitude given to it
on one side, and the objections raised against it on the other, mark one feature
of the phases of the gradual progress of science. It has been applied in different
forms to explain the mode of origin of ancient eruptive rocks; and, since Dolo-
mieu and, in a more elaborate way, his pupil Cordier, have assigned the same
original source to voleanic activity, the contraction of the interior of the globe, by
radiation of its heat into space, has been considered as offering a sufficient explana-
tion for the majority of the phenomena which are often united under the term
‘‘vuleanism.” But, giving all due consideration to the vast effects of which it has
undoubtedly been the cause, the conception of the modus operandi of this agency
(contraction) alone meets with considerable difficulty. An outward tension might,
indeed, result in the formation of fractures on elevated places; and, supposing for a
moment that these fractures would descend into regions where matter was in a liquid
state, then the latter might possibly be ejected, and caused to accumulate on the sur-
face. But such would hardly be the effect of an inward tension. It may, too, cause
the rending of the crust and, possibly, the filling by liquid matter of those rents
which are in the lowest places ; but it does not explain the extrusion of this matter
to the surface, nor the fact of its particular accumulation on elevated parts of the
same. It would take too much of our space fully to detail the numerous mechanical
difficulties which occur, if one attempts to explain all the phenomena comprised in the
name ‘‘ vuleanism” by the exclusive assumption of the contraction of the interior of
the globe. Some of the more obvious objections against this theory will be briefly
mentioned in the course of the following considerations.

A number of facts point towards the existence of some unknown force below
the earth’s solid crust, which counteracts in a considerable measure the permanent
subsidence of the latter by contraction. It is perfectly evident that the secular rising
of parts of the surface of the globe above the level of the sea cannot be merely the
apparent effect of the different degree of its general subsidence, as has been maintained
by very distinguished geologists; but that elevation, that is, the periodical increase of
the distance of parts of the surface of the globe from its center, must be a reality.
Considering the amplitudes of the changes of level that have taken place in historical
time, they will be found to sum up to such figures as, if reduced altogether to subsi-
dence, would indicate a far greater shortening of the radius of the globe within that
time, than is compatible with astronomical calculation. It is true that the retardation
of the rotation of the earth by the tidal wave must counteract, in some measure, the
acceleration caused by any shortening of the radius of the planet ; but this retardation
is insignificant if compared with the amount of the changes of level. The reality of
elevation is forced more directly upon the mind, if those cases are taken into consider-
ation where certain portions of continents are rising above the level of the sea at a
more accelerated rate than neighboring regions, which is of very frequent occurrence

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Hy) RICHTHOFEN — NATURAL SYSTEM

in yoleanic countries, and conspicuous in those numerous instances where elevation is,
or has been in former times, proceeding more rapidly along the crests of mountain
ranges than at either foot of them. Adherents of the theory, that all oscillations of
the earth’s crust are only due to its contraction, consider elevation as real in these
cases, and have tried to explain it by the assumption, that folds must be formed by the
subsidence of the ample shell on the contracted nucleus ; that these folds would increase
in amplitude in consequence of a lateral pressure caused by further subsidence, and
thus an absolute rise would be effected. It is, indeed, very probable that this process
is of vast importance in the formation of mountain ranges ; but it cannot explain the
total amount of the changes of level. If Fourier’s calculation, that, taking the present
loss of heat by the globe as standard, the radius of the latter should have shortened
seventeen centimeters in twenty-five centuries, is correct, the diminution would have
been 1,700 centimeters in 250,000 years, and about six hundred feet in one million
years. It is manifest how utterly insignificant would be the corresponding diminution
of the earth’s cireumference, when compared with the vast changes of level which the
surface must be supposed to have undergone during such a length of time.

We are bound, for these reasons, to consider, not alone the process of elevation
of mountain ranges, but also the secular rise of extensive regions, as realities which
cannot be exclusively explained by the contraction of the globe. But if so, there must
exist another force, the effects of which oppose those of contraction. The united action
of both, and the periodical prevalence of either of them, would then be capable of
explaining the alternation of elevation and subsidence at every single place, and the
contemporaneous action of both in neighboring regions. This antagonistic force con-
sists, probably, in an increase of volume attending the slow and perfect crystallization
of matter, by cooling down, during immense periods, from a viscous state. With a
great number of bodies, contraction by loss of heat appears to continue to the very
moment of solidification, and to increase during the latter when no time is allowed for
crystallization, but to be diminished, and finally reversed, in the same measure as oppor-
tunity is given for a slow and perfect crystallization. In the special case of rocks
made up of silicates, this must remain a supposition which is not proved, but is emi-
nently probable.*

18 An increase in yolume by crystallization has been found, by experiment, to take place in numerous instances, in fact
in most cases when a perfect crystallization from a molten state has been obtained, provided that the volume of the substance
experimented on could be determined immediately before the act of crystallizing. In respect to those silicates which make
up crystalline rocks (not taking into account the water entering into their composition), experiments could hitherto not
be made, because it is impossible, with our present means, to allow the viscous mass sufficient time for crystallizing. Yet,
there are some suggestive facts which have, in some measure, the value of experiments. We mention among them par-
ticularly one which was observed by Ferd. Zirkel in analyzing, with the aid of the microscope, the texture of the
minerals participating in the composition of eruptive rocks. He found that “glass cavities” contain ordinarily several
of those vacuities which are also exhibited by the cavities filled with water, and have been supposed by Sorby
and others to have originated from the contraction, by cooling, of the enclosed matter. If the substance contained
in the glass cavities bears signs of an incipient crystallization, by having partly a “lithoid” texture, the vacuities are
of rarer occurrence, while they are entirely wanting in the so-called stone-cavities, when the lithoid texture pervades the
whole mass filling the cavity. It is scarcely possible to make an experiment more convincive than this natural occurrence,
which is more open to subtle observation and measurement than is ordinarily the case with experiments on cognate subjects.
It needs hardly to be mentioned, in connection with this question, that those experiments which have been made by Bishof

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OF VOLCANIC ROCKS. 53

We may then distinguish as eminently probable, the following immediate effects
of the cooling of the globe: 1. Contraction of the liquid portion of the interior, by
cooling down to the temperature required for solidification under the respective press-
ure. 2. Expansion by slow and perfect crystallization. 3. Contraction of the erys-
tallized masses by further cooling. We may add to these as a secondary effect of
greatly inferior importance, the flow of heat or change of chthonisothermal planes, at
those places where subsidence causes an accumulation, or elevation an abrasion of
matter on the surface. This process, to which a truly wonderful importance has been
ascribed by some, must be going on continually and everywhere. Being itself the
result of motions of the crust, its proximate cause must be the processes attending the
cooling of the globe, while in its totality, it may have a further, though very insignifi-
cant, effect in shifting the areas of elevation and subsidence.

Other agencies, with which we are yet unacquainted, may probably result from
the same common cause. But taking into consideration only those already mentioned,
it may appear difficult to form a clear conception of the mode in which they must
codperate, in order not to counteract each other, and thereby to result in a general
movement of the crust in one direction only, but to bring about that variety of
effects which manifests itself chiefly in the change of elevation and subsidence, and in
other modes of dislocation of the crust. It is perfectly evident, that the aggregate
effect of those agencies is different under different portions of the crust of the globe,
contraction prevailing in some parts, and expansion in others; the former causing
subsidence and deposition of sedimentary matter, the other elevation and denudation.
But if we consider how gradual is the passage in the state of aggregation from the liquid
interior of a current of lava to its solid crust, and if we bear in mind how immensely
vaster in volume are those masses which constitute the interior of the globe, and what
immensely longer time is given for their cooling and crystallization, the conclusion is
irresistible, that the passage in the states of aggregation from solid to liquid must
extend there over an immensely greater space. Crystallization will very probably
take place next to those portions of the crust which are already solid. If it is attended
by an increase of volume, and this increase produces tension, it is very likely that the
next adjoining masses, though not yet crystallized, will offer too much resistance to
allow this tendency for expansion to find immediate relief by yielding to the gen-
eral tendency to contraction which may prevail in liquid masses at greater depths.
It may thus be explained why, in different parts of the crust, a motion, independent
and in opposite directions, may result from the two-fold tension attending changes
of volume which take place at different distances beneath the surface, and which

and others with a view of ascertaining the increase in volume which different rocks undergo by melting, do not affect our
supposition. In the first place, the molten state may be quite different from that in which the same substance would be
immediately before crystallizing, and the passage from one to the other may be attended by a decrease in volume which is
not counterbalanced by the subsequent increase taking place by the passage of the substance into the crystallized state.
Then, the rocks on which the experiments were made had been solidified under a great pressure, while, when in a molten
state, they were only exposed to the pressure of the atmosphere. Finally, it has not been ascertained whether all the water
which is contained in the rocks does escape on melting or not. If not, then it will probably have the effect of inflating the
molten mass.

G (91)

54 RICHTHOFEN —- NATURAL SYSTEM

would oppose, and, to a certain degree, destroy each other, if the state of aggregation
of subterranean matter allowed of a free conduct of motion.

; These slow and continuous agencies, chiefly those among them which give origin
to the process of elevation, must have been, too, instrumental in causing the massive
eruptions of rocks. At least, no other force supposed to act beneath the surface can
account as fully for certain facts connected with them, such as the gradual changes
in the mode of their geographical distribution, or the connection of the manifesta-
tions in any certain system of fractures, or the fact that ages of comparative
repose have been interrupted by paroxysmal actions of great violence which have
taken place during certain eras in the history of each separate country. The
latter circumstance points clearly to the assumption, that during the eras of repose,
or at least their later portion, a constantly increasing amount of potential energy
must have been accumulated under the crust of the globe; since the subterra.
nean agencies called into existence by the cooling of the globe did of course never
rest, nor can they be reasonably supposed to have been more intense in the Tertiary
than they were in preceding periods. It is self-evident that the increase of tension
must have been much more considerable under areas of elevation than under the
more extensive regions of subsidence. In the latter case, when a downward tendency
is caused by contraction, the weight of the crust must come in its aid ; the essence of
the resistance to that tendency may therefore be concisely expressed as cohesion minus
weight ; while weight added to the cohesion will give approximately the resistance to
elevation by expansion. The tension under a crust of great thickness must therefore,
in the latter case, increase to a stupendous intensity, until it is sufficient to overcome
the resistance, and will then be able to result in such paroxysms as that by which
the volcanic era has been inaugurated, and the main feature of which consists in the
formation of fractures which, by their mode of association, constitute distinct systems
or belts, separated by areas in which the existence of fissures in the crust cannot be
recognized on the surface.

Although the processes suggested may furnish us the cause for the prime condi-
tion to the emission of rocky matter from below, namely, the periodical opening in the
earth’s crust of such fissures as widened with the approach to the surface; yet
they fail completely to explain, by themselves alone, the more immediate causes and
the mode of that emission. It may be presumptive to extend speculation upon this
topic beyond those limits which we have reached ; but a safer guide to conjecture on
the same than had been known at any former time was given during the last few
years by the experiments of Daubrée, and the microscopic examinations of the texture
of rocks by Sorby. It has been held quite generally till of late, that the opening of
a fissure to a liquid mass below, would be sufficient by itself to cause the ascending
of the liquid through it to the surface ;” but this supposition is utterly irreconcilable

19 Tt must here be remarked that some of the most eminent writers on the subject of voleanoes, chiefly Pou-
lett Scrope, Prof. Dana and others, have suggested long ago, that the ascending of lava in a voleanie channel mast be
due to both the fluidity and expansion imparted to it by the globular state of the water which finds ingress to the chan-
nels of the lava and enters into its composition. They have anticipated, by this suggestion, in some measure, the results of

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OF VOLCANIC ROCKS. a)

with physical laws. Supposing for a moment that theory to be correct which consid-
ers al the changes of level as originating in contraction alone, then it is quite proba-
ble that fissures would be formed in the center of the areas of subsidence. But as
they must be closed near the surface, and open in their lower parts, it is difficult to
see in what manner the liquid matter could ascend through them to the surface, while
no better account could be given for the occurrence of eruptive rocks on high table-
lands. In order to explain it, recourse has been had to the most arbitrary assump-
tions. It may even be read in our time in various geological books, that the crust below
might give way from the overload, and the whole be “plunged” into the semi-fluid
mass beneath, causing it to overflow. Mountain ranges of a thousand miles in extent
splash it out”

have been assumed to subside suddenly upon the liquid mass, and to ‘
through fissures. Leaving out of consideration these fantastical theories, there remains
a number of others, according to which the weight of portions of the crust would
cause the ejection of liquid matter. If the substance composing the crust exceeded
in specific gravity the liquid matter below, then this mode of ejection would be prob-
able, and we should be indeed surprised that large portions of our globe were not
flooded over repeatedly by molten masses from below. If the specific gravity of the
crust and the fluid matter below was the same, then it would require the most exten-
sive fracturing of the crust and uplifting of its fragments, in order to make the liquid
mass overflow the latter. But all that we know in regard to the subject goes to show
that the masses below the crust are of greater specific gravity than those composing
it. To suppose the weight of the crust to cause the protrusion of liquid matter
through fissures, is therefore to suppose an action which is mechanically impossible.
We arrive at no more satisfactory conclusion in regard to our present problem,
if, besides the contracting forces, we assume the existence, beneath the crust of the
earth, of others which have their origin in the increase of volume by erystallization,
and thus produce expansion. I attempted to show that they furnish the most probable
agent to which may be ascribed the opening of the systems of fissures, which partly
served as the channels for the extrusion of rocky matter. These fissures would have
to be open at the surface and to decrease in width below, because formed by an out-
ward tension on areas of elevation. It cannot be assumed that they would descend to
any greater depth than the lowest limits of solid rocks; they would, therefore, not
reach down to any matter sufficiently liquid as to be capable of being forced up
through them. And if it should be able to ascend, then it would solidify within the
fissure, almost instantly, by loss of heat, and long before reaching the surface. There
is, however, even a more forcible argument to demonstrate why it should not have
been capable of ever entering the fissures. For, if our supposition that the silicious
masses beneath the crust increase in volume by crystallization is correct, the relief
from pressure by the formation of fissures must have the immediate effect of rapidly pro-

experiment established by Daubrée. But the supposition was only made for the case of lava, and not for that of rocks which
were ejected without volcanic action proper. Yet even in regard to volcanoes it did only explain the extrusion of lava to
the surface from a place at a limited distance below it, and failed to give a clue to the manner in which the constant supply
of matter to those places was kept up.

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56 RICHTHOFEN — NATURAL SYSTEM

moting the crystallization of vast masses which had been held in a viscous state before,
by the existence of the tension itself. It needed, for all these reasons, another agency
which would not alone force up molten matter through the fissures, but also cause it to
arrive at the surface in that particular state of aggregation which it has had, according
to the observations of Sorby. This agency is indicated, by the experiments of Daubrée,
to have been water, the descending of which into the fractures is indeed a necessary
consequence of their formation. There it would convert the state of aggregation of
the masses surrounding the lowest parts of the fracture into that called by Daubrée
‘‘ aqueous fusion,” which appears, indeed, to have been the state in which all eruptive
rocks have been immediately before their consolidation. The.process of aqueous fusion,
as has been shown by the same eminent geologist, is attended by a very considerable
increase in volume of the masses affected. It would, therefore, give rise to processes
totally different from those which had preceded. For this expansion would imme-
diately cause a motion of the masses rendered liquid, in the direction of least resist-
ance, that is, upwards in the fissure, and would, if continued for a sufficient time,
make the same overflow on the surface of the crust, even if unassisted by other eject-
ing agents, such as the vapor of water.”

We may carry these deductions still further, if we revert to our previous con-
clusion, that the relief from pressure by the fracturing of the crust would have caused the
crystallization of masses below it, which had been held before in a viscous state by the
tension itself. This process would extend in depth as well as laterally, and gradually
affect the viscid masses beneath an entire belt of fissures. It would have had again to be
attended by an increase of volume. But those crystallizing masses not being in a state
of aqueous fusion, the resistance could in this case not be overcome by the extrusion
of that portion of them by which they were increased in volume, and the effect would
be, as in the ease first mentioned, accumulation of potential energy. I will attempt to

20 If we consider the geological features of all large accumulations of eruptive rocks, such as the great andesitic
ranges of Hungary, or the quartzose porphyry composing a plateau of great dimensions in southern Tyrol, or those granitic
masses which, by overlying the edges of stratified rocks, give evidence of having been ejected to the surface in a liquid state—
it would appear that their emission has been a slow and mainly a quiet process of long duration, hardly attended by those con-
yulsions and paroxysms which form the prominent features of volcanic action, and should have been no less characteristic of
massive eruptions if they had been due in any large measure to the expansive force of vapor. The process of the emission
of the rocky matter has, it is true, been evidently intermittent in most cases, as may be inferred from the occurrence of vast
accumulations of breccia, and it appears that an extensive solfatarie action has frequently taken place through neighboring
fissures ; but the manner in which the matter was protruded through the main fissures and deposited on the surrounding parts
of the surface, had evidently no similarity to the mode of ejection of scoria, ashes, and lava from most of the active volea-
noes. The mode of action described, which may be inferred from geological observation, is perfectly in accordance with what
we should expect it to have been by reasoning a prior? on the basis of our previous suppositions. For, if G. Bishof’s cal-
culation is correct, that the elastic force of steam is at its maximum when it has the same density as water, which it would
acquire under a pressure of 8,300 atmospheres, the loftiest column of lava (taking its mean specific gravity to be 3) which
should be supported by it, would be, according to Jukes, 88,747 feet. The original seat of those eruptive rocks which were
protruded without being accompanied by volcanic action, must necessarily have been at a much greater depth, and we should
therefore, also from this point of view, be led to suppose that the expansive force of steam has had only an insignificant part
in their protrusion from greater to Jesser depth. It must, of course, have come into action when the liquid masses arrived
near the surface, but will have caused hardly more than an ebullition, even in viscid masses, on account of the extent of the
openings. The formation of conglomerates could thereby be vastly promoted, but their final deposition and consolidation
must have been quite different from the manner in whieh similarly subdivided matter would be deposited around a volcanic
orifice.

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OF VOLCANIC ROCKS. 57

demonstrate in another chapter that the changes of level which have been connected
with the ejection of rocks, appear to confirm these suppositions. Suffice it here to
draw the necessary consequence, that the increasing tension must finally have had the
effect of rupturing the newly consolidated masses. If in the intermediate time the
masses filling the fissures of the first epoch had, at least partially, been consolidated,
then a new system of fractures would be opened, within the limits of, though not coin-
ciding with, the first. These fractures of the second epoch would descend to greater
depth than those of the first, and in allowing the access of water to masses situated in
lower regions and being of a more basic composition, would open for these the way to
the surface. By the repeated occurrence of this or similar processes, the theater of
action from which the rocky masses were conducted to the surface, might descend, by
steps, into considerable depth within a comparatively short period, and thus there
could be produced a great diversity among the rocks emitted through one system of
fractures, though this diversity would be regulated by definite relations in regard to
the nature and succession of the rocks ejected. The process would come to an end
when the solidification of matter and the formation of fractures had descended to those
masses, the state of aggregation of which was such as no longer to allow them to crys-
tallize when the pressure was diminished, and in this way any further increase of
volume would be prevented. The matter filling the fractures would now solidify, and
the communication of the interior with the surface be cut off, with the exception of
the voleanic channels. The resistance offered by the crust of the globe would hence
be greater than it had been before, and there would follow another era of repose,
longer than that which had preceded the era of eruptive activity.

The application which may be made of these processes suggested by theory, to
the explanation of the actual correlations of eruptive rocks in regard to their age,
chemical composition and geographical distribution, is obvious. We may, indeed,
venture to deduce @ priori the history of eruptive action, in its main features, from the
hypothesis of Sartorius, and the assumption that silicates will increase in volume by
perfect crystallization. We should have to conclude that ma remote period, when
the crystallized crust and the sedimentary shell of the globe were inferior in aggregate
thickness, fractures and eruptions of rocky matter would have been of frequent occur-
rence, and that highly silicious compounds should have prevailed among the ejected
masses. Little, if anything of them, is probably visible at the present surface, as the
rocks of the Azoic and Paleozoic formations are probably the monuments of an already
far advanced stage of the development of our planet. All the distinguishing features
of the eruptive rocks of these periods (including of the Palsozoic age only the Silurian
and the first part of the Devonian), such as the great number and individualization of
the granitie districts, the independence of each of them in the subtler differences
regarding the commencement and further development of the eruptive activity as well
as the peculiar nature of the rocks ejected, the great preponderance of highly silicious
compounds, the common association with them of small quantities of basic rocks, and
the slight increase of the proportion of the latter in the Devonian period—all these
phenomena are easily understood, without further explanation, in the hght of our
hy pothesis.

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58 RICHTHOFEN —NATURAL SYSTEM

Let us now direct our attention at once to the voleanic era. The conditions
of the globe must have been very different in the Tertiary from what they had
been in the Paleozoic period. A longer time of comparative repose had in most parts
of the globe preceded the inauguration of the violent manifestations of vulecanism in
the Tertiary period than had ever before elapsed between any two eras of eruptive
activity. The globe had cooled down. Volumes of sedimentary matter had accumu-
lated, and added externally to the thickness of its crust, while it had increased in a
vastly greater measure by the crystallization of liquid matter below. Those silicious
compounds especially, of low specific gravity, which had formerly yielded the material
of the vast accumulations of quartziferous eruptive rocks, would have been consoli-
dated, and the limit as it were between the solid and the viscous state of aggregation
receded into regions where the matter would be of a less silicious composition and of
greater specific gravity. The similarity in distant countries of the rocks first ejected
(propylite and andesite) goes to show that the recession of that limit into greater
depth must have proceeded in a nearly equal ratio in all those regions where volcanic
rocks are distributed. When the tension below had increased sufficiently to overcome
the resistance, it would now no longer manifest itself in the formation of small and
differentiated systems of ruptures. In the direct ratio of the increase of the resist-
ance the fractures would have to be of greater extent, and those elongated belts of
them would be formed which even now are partially distinguished as the belts of vol-
canic activity. The first rocks ejected would necessarily be of a more basic composition
than the predominant rocks of the granitic era, while the repetition, at a later
epoch, of the process of fracturing would give rise to the ejection of rocks in
which silica would be contained in a still lower proportion. The greater portion
indeed of the ejected rocks consisted of propylite and andesite, in the first, and of
basalt in the second half of the voleanic era. A notable but only apparent anomaly
in the regular order of succession has been the emission of trachyte and rhyolite
between the andesitic and basaltic epochs. But if it is considered that these rocks were
ejected partly from the same fractures through which andesite had ascended, and
partly from others in their immediate vicinity, while the distribution of basalt has
been independent, to a certain extent, of all foregoing eruptions, it is evident that
the occurrence of trachyte and rhyolite is closely dependent on that of andesite, and
bears only a very remote relation to basalt. It appears that after the ejection of the
chief bulk of andesite, when other processes ending in the opening. of fractures into
the basaltic region were being slowly prepared in depth, the seat of eruptive activity
ascended gradually toregions at less distance from the surface. There is, within the
limits of conjecture based on physical laws, no lack of processes which could codperate
to that effect. The consolidation of the ejected masses within the fissures would prob-
ably proceed simultaneously, by loss of heat, from the surface downwards and, by
pressure, from below upwards. The opening of new branches from the main frac-
tures, the remelting (by the aid of the heat of the molten mass within the latter, and
of water finding access to it) of solidified matter adjoining the fracture, the emission
of that remelted matter through those branches: all these are secondary processes
depending on the first almost necessarily. The supposition that to these is due the

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OF VOLCANIC ROCKS. 59

order of time in which trachyte and rhyolite have been ejected to the surface, is
corroborated by the fact that these rocks occupy generally a subordinate position in
regard to quantity, and have had, to a great extent, their origin in voleanie action.
When treating about the latter, we will come back upon this subject. There remain
some peculiar features of volcanic rocks which cannot be satisfactorily explained at
the present time. We mention, among them, the fact that the three modes of texture
of rhyolitic rocks are often severally limited to certain localities ; the mode of forma-
tion of the laminated structure of rhyolitie and trachytic rocks ; the occurrence of the
compounds of hornblende and oligoclase in that threefold form to whieh we have
repeatedly referred; the fact that basalt has been followed only to a very limited
extent by rocks bearing to it a similar relation, as trachyte and rhyolite do to andesite.
As regards the long lapse of time intermediate between the Devonian and the
Tertiary periods, the mode of occurrence of eruptive rocks in the same shows in
nearly every respect a gradual transition from that which was peculiar to the granitic to
that which we just described as being characteristic of the voleanic era. This interme-
diate period may be designated as the porphyritic era, though this name appears to
apply more properly to its first part only. Quartziferous rocks were not so predom-
inant in it as in the granitic era, porphyrite and melaphyr having nearly equaled
quartzose porphyry, in point of quantity. Augitic porphyry was ejected in a
much larger proportion to the aggregate bulk of the porphyritic, than diabase to
that of the granitic rocks. Where it occurs, it was the last in the order of rocks
erupted, while quartzose porphyry was generally the first among them, though this
place is sometimes occupied by porphyrite. In reference to the general features of
their geographical distribution, porphyritic rocks occupy no less distinctly an inter-
mediate position, as may be seen by what we have said on this topic on another page.
There are exceptions to the order of general development as here specified.

They regard chiefly the texture, and are almost exclusively to be found among the
rocks of the porphyritic era, though the recurrence, in propylite, of the properties of
ancient diorite, is a phenomenon of a no less exceptional nature. Leaving this rock
(propylite), or rather only some of its varieties, out of consideration, the voleanie rocks
have their peculiar characters, by which even the most basic rocks are to be recog-
nized when seen in large accumulation. In the rocks of the granitic era, if we
consider its end to be within the Devonian period, the characteristic features of por-
phyritie and voleanic rocks are probably never to be observed. Among the excep-
tional occurrences within the porphyritic era, may first be noticed the fact that perfect
granitic texture is still occasionally encountered, as, for instance, near Predazzo and
on the Monzoni in southern Tyrol, where some subordinate masses of rock resembling
granite and syenite have been ejected in the Triassic age, together with the well-known
porphyritic rocks of that region. There are similar instances known from other places
on the European continent, but they are scattered, and the respective rocks always
quite limited in extent. The grandest exceptional instance that is known up to this
time, is the recurrence in the Jurassic period of perfect granitic texture in the erup-
tive rocks of the Sierra Nevada. But as regards their mineral composition, these rocks
belong to the family of syenitic granite, containing hornblende as a very characteris-
@)

RICHTHOFEN —NATURAL SYSTEM

tic ingredient, while silica enters into their average composition probably in a
much lower proportion than it is contained in ordinary granite. The varieties known
as granitite are of rare occurrence. In another respect, namely, the mode of geograph-
ical distribution, the Jurassic granite of the Andes marks an advanced stage, if com-
pared with the Permian and Triassic porphyries of Europe, inasmuch as the area of
its distribution excels the porphyritic regions of the latter continent in regard to their
extent and the unity of their inner relations, while it does not come up, in these
respects, to the properties of the great volcanic belts.

2. Origin of Volcanic Action.

I have tried to demonstrate in another chapter, that volcanic activity is inti-
mately connected with massive eruptions in a three-fold way, namely, in respect to
the epoch of its commencement, the mode of its distribution, and the nature of the
rocks ejected. Basaltic volcanoes occur in ranges built up by the massive eruptions of
basalt, or in their vicinity, in such connection as to make obvious the nearly contem-
poraneous origin of both. Andesitic voleanoes are found in the neighborhood of those
masses of andesite which had been ejected without being attended by volcanic action ;
while voleanoes which have emitted rhyolitic or trachytic lava are so situated as to
justify the inference of a close connection between the first opening of their vents
and the origin of neighboring accumulations, either of rocks of similar character or of
andesite, but which are due to massive eruptions.”

No distinct line of demarcation can be drawn between the two modes of mani-

21 This affinity in regard to petrographical character and geographical distribution is probably the cause why both
agencies have ordinarily been confounded. It has been a current notion, strongly advocated at times, that all volcanic
rocks have come to the surface in a similar way to that in which lava is being ejected from volcanic vents, and that the
cause of the ejection has been the same in all cases. In regard to the fact that extensive mountain ranges are completely
built up of voleanic rocks, it was argued that it is by no means necessary, and in fact erroneous, to suppose them to have
originated in events surpassing in magnitude those of the present day, as the length of geological time would explain how
they could be built up by the gradual accumulation of innumerable currents of lava. Where traces of former volcanic
vents could not be found, the easy destructibility of the matter of which the sides of craters are usually composed, afforded
a convenient and apparently just argument for explaining their absence by denudation. The same doctrine was applied for
explaining the mode of formation of ancient eruptive rocks where we see mountain masses made up of them, while others
have arrived at the conclusion that they were the “ roots” or “cores” of volcanoes; granite itself is by them considered
to have had formerly, and to have now this function, even in the case of those volcanoes the lava of which consists of basalt.
Both these doctrines have been adopted the more readily, as they appear to be in harmony with the favorite hypothesis, that
at no time have any changes on the surface of the globe been more violent than those going on at the present day, nor
different from them in mode. However ably this theory, which contributed so much to the advancement of science by
checking the phantasmagorias of former time, has been advocated, an unbiased comparison of the grand manifestations
recorded in the geological structure of an andesitic mountain range, with the mode and degree of activity of present vol-
canoes, must lead to different conclusions. In the endeavor to sustain the @ priori assumption of the equality of force in all
ages, too little stress has been laid on the circumstance, that, however nearly equal the aggregate amount of force acting on
the globe may have been, its modes must have undergone a change, chiefly by the partial conversion of the heat of the
globe into other forces. At the same time, the degree of the intensity of the manifestations of subterranean forces must
haye varied in a two-fold way. With the continuous increase of resistance, the aggregate motions of the crust must have de-
creased, since a growing amount of these forces*was required for overcoming the resistance. On the other hand, the manifest-
ations of the same forces had to become periodical and paroxysmal, and periods of violent action had to be separated by
others of repose. The violent activity of the Tertiary period has passed, and our present volcanoes appear to mark the transi-
tion into another period of repose. I will try to demonstrate that the causes of their action are utterly inadequate for ex-
plaining the grander phenomena and correlations of massive eruptions.

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OF VOLCANIC ROCKS. 6]

festation of subterranean energy, as some of the monuments of the grand voleanie
action of the past indicate the former existence, at certain places, of a stage intermedi-
ate between both. But in those numerous instances where the nature of either of
them can be distinctly recognized, some conspicuous differences may be noted between
them. Volcanoes are provided with a channel connecting the seat of volcanic action
with the surface. The matter which they eject consists either of stratified layers of
ashes and scoria, or of currents of lava in the shape of flat sheets superimposed over
one another, or of alternating layers of both kinds of material, the latter structure
being most frequent. A low dip, verging on horizontality, of all planes dividing sub-
stances of dissimilar nature, or marking successive deposition, and homogeneity of
material parallel to those planes, together with a structure of the mountain masses
distinguished by the radiation of all mineral matter from one common center, may be
considered as those features. by which even extinct volcanoes, or fragments of them,
may unmistakably be recognized. The center may shift, or have shifted, within nar-
row limits, or a series of centers may follow each other in close succession, but this
will obliterate the true character only in a slight degree. Similar rocks, when they
came to the surface by massive eruptions, do not present these distinguishing features.
They usually compose ranges of small width in proportion to their length, and in the
place of one or more distinct centers an elongated axis may be detected, from which
the structural and morphological features originate. As regards the interior structure,
there may be observed a certain massive character of the rocks. which is partly pro-
duced by the prevalence of their, compact. varieties, and partly by the circumstance
that homogeneity of lithological character may be traced to a great distance in a verti-
cal direction. In sections, masses are frequently found thousands of feet in height,
which do not ‘vary perceptibly in character, and show no horizontal structure. If the
slopes of the ranges are examined, the rocks will be found preserving a homogeneous
character chiefly in a direction parallel to the axis, while it is less persistent at right
angles to the latter. The planes dividing dissimilar rocks are inclined at all angles,
and have very frequently a steep or nearly vertical position. Breccias even, which
sometimes occur in very large masses, are bounded in this way towards the adjoining
compact varieties; they are of irregular shape, and do not often occur in stratified
layers or in elongated currents, as is the case when they are produced by volcanic
action. The ranges made up by massive eruptions show no signs of craters. Yet
they are frequently the foundation of voleanoes. Oftener still do voleanoes occur on
the lower portions of their slopes, or they may form a series parallel to the axis of
the main range, and even greatly exceed it in elevation.

Notwithstanding these points of difference, there are not only stages of transi-
tion between the distinct geological features resulting from either mode of action, but
a similarity in character may be produced under certain circumstances, which makes it
difficult to decide what was the mode of origin of-an accumulation of volcanic rocks.
In the first place, the matter extruded through fissures may have been so liquid as to
expand at once in thin sheets. This is very frequently the case with basalt, in the
great accumulations of which, whether they be due to massive eruptions or to voleanic
activity, the prominent differences wrought by both modes of action in the external

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62 RICHTHOFEN—NATURAL SYSTEM

features are generally least conspicuous. This basic compound is not only more fusi-
ble than the more silicious rocks; but, it appears that the admixture of superheated
water will increase its fluidity more than that of the other compounds, while those
unknown influences which cause the mass to solidify in the particular form of basalt
must be still more potent in increasing its fluidity, since lavas consisting of dolerite,
leucitophyre, and trachydolerite, which are quite or nearly identical with the former
in chemical composition, are never so liquid as those of basalt, and are not unfrequently
quite viscous. These rocks, together with all those of a more silicious composition, ex-
hibit more distinctly the differences in origin, and this is probably due, in a great meas-
ure, to the peculiarity just mentioned. The great fluidity of basalt, which is also made
evident by its frequent occurrence in very thin and yet very extended dykes, causes
it, even when confined in the narrow space of a volcanic vent, to let the vapors of
water escape, in quiet ebullition, in its orifice, as Dana has so beautifully illustrated in
his description of Kilauea, while, at intervals, it will break out and cover the sur-
rounding country with flat sheets of lava. The action connected with the ejection of
the same rock from larger fissures, in former time, appears to have been similar to
this. Yet, the numerous instances of the occurrence of basaltic cinder cones, and, on
the other hand, of large accumulations of solid basalt with no perceptible horizontal
structure, go to show that also in the case of this rock the modes of occurrence may
be different when they are the result of different modes of ejection. Other instances
of a similarity of the manner in which the matter has been deposited, when due to
either mode of ejection, are frequent on the flanks and at the ends of the andesitic
ranges of Hungary, where currents of andesite as well as of rhyolite have been emitted
through fissures in andesite, at little elevation above the foot of those ranges. They
appear to be due to processes intermediate in kind between both modes of ejection.
We may, finally, mention those cases where massive eruptions were sub-aqueous, and
layers of fine-grained tufa formed, alternating with coarser conglomerates, between
which may be intercalated solid layers of the same kind of rock of the fragments of
which those are composed. The similarity of this kind of depositions with the sedi-
ments of submarine voleanoes is often very great.

The principal point of difference between massive eruptions and voleanic action
appears to be the depth of their source under the surface, and all the minor differences
are probably dependent upon that. The region from which the former have derived
their material, is, as we tried to prove, at a great depth beneath the deepest sedimen-
tary rocks. The seat of volcanic action appears to vary within wide limits in regard to
its distance from the surface, but to be, on an average, at much less depth than that of
the massive eruptions ; though there are circumstances which render it probable that
it is in all cases beneath the shell composed of sediments. Hvidence has been gathered
by Prevost, Dana, Scrope, Hopkins, and others, in favor of the assumption that vol-
canoes are not connected with the molten interior of the globe, and are therefore not
to be considered as safety-valves. The comparatively little distance of the seat of vol-
canoes beneath the surface is rendered particularly evident by the small area of the
earthquakes attending their activity, when compared with the wide extent of others
which must be dependent on some deep-seated action, but have no recognizable con-
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}

OF VOLCANIC ROCKS. 3
nection with any particular voleanic vent. It is no less obvious from the fact that two
neighboring voleanoes may not only eject, contemporaneously, different kinds of lava,
but also be, to some extent, independent of each other in their manifestations.

We may base further conclusions in regard to the origin of volcanoes upon the
following premises, which we repeat from the foregoing pages: First, several facts
appear to indicate that the source of volcanic action is at a comparatively limited
depth ; second, all voleanoes, whether active or extinct, are intimately connected with
massive eruptions ; third, this connection is of such a character as to establish, in the
majority of cases, the close similarity and chemical identity of the mineral matter
ejected by the volcano in its first epoch of activity with that of neighboring hills or
of its own foundation, which had been accumulated by massive eruptions; while in
other cases these neighboring hills or the foundation of the voleano are composed of
voleanic rocks not identical with the lava, and then the latter will belong lithologically to
a kind of rock which, according to the order of succession of massive eruptions, would be
of a more recent origin than the former; that is, trachytic or rhyolitic voleanoes were
frequently opened at those places where only andesite had been accumulated before,
and basalt where either andesite or trachyte had preceded ; but the reversed order
appears not to occur, no basaltic voleano having been succeeded, in its own neighbor-
hood, by massive eruptions of trachyte or andesite ; fourth, many of those volcanoes
which have been active through a long period, have undergone a periodical change
in regard to the character of the mineral matter ejected by them, and this change is
in general (though with exceptions) conformable to the order of succession observed
in regard to massive eruptions.

From these facts may be inferred the complete dependency of volcanoes upon
massive eruptions. The latter, as we attempted to show, were due, firstly, to the open-
ing of systems of fissures which extended throughout the solid crust ; and secondly, to a
quiet outflow, which was caused by the expansion attending the change of aggregation
of solid or highly viscous matter around the lowest part of the fissures into that of
aqueous fusion. The relations of volcanic activity to massive eruptions are indicative
of a process by which the elongated and extensive vents of the latter were gradually
differentiated into isolated and narrow channels feeding isolated orifices on the surface.
It has been observed that cinder eruptions mark generally the last stage of volcanic
action. We may go a step further back, and say that volcanic action is the last stage
of massive eruptions.

In order to arrive at a conception of the manner in which the change from one
mode of action to the other could be effected, let us suppose that a main fissure was
filled with matter from below, and mountains of volcanic rocks accumulated above it
by the long continued overflow. Solidification would at once set in, and proceed
downward whenever a cessation of the extrusive action occurred, independent of
the question whether it would not simultaneously proceed upwards from the depth.
Its progress would not be equal in all parts of the fissure, since this must be
wider at some places, and more contracted at others. In this condition we should
have one of the causes for the isolation of centers of action, for the length of time

during which different portions of the matter would remain in a fluid condition, must, of
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64 RICHTHOFEN —NATURAL SYSTEM

course, depend in a great measure upon the width they would severally occupy within
the channels, and those filling its wider parts must for a longer time remain susceptible
of renewed expansion by accidental circumstances. Another cause which would have
the same general effect, is the localization of the ingress of water. Some of its chan-
nels would, to all probability, become obstructed, and the rate at which new ones
would be opened in their place would probably diminish in nearly equal ratio with the
total amount of the manifestations of energy connected with the phenomenon of ejection
in its different stages. As it appears that the supply of water, which may be either
constant or intermittent, is, next to an elevated temperature, the chief condition for
entertaining voleanic action, it may be inferred that the cause mentioned would contrib-
ute greatly towards the isolation of certain portions within the main fissure, by helping
to keep the matter within them in a liquid state. If this second cause was coincident
with the first, by the restriction of the ingress of water to a place where the fissure
expanded, then both circumstances would combine to prolong the state of liquidity at
that point. The connection between the bottom and the surface may have been
kept open in a certain part of the mouth of the fissure, while solidification was proceed-
ing over the rest of it. An isolated vent would then gradually be formed, and nar-
rowed down to the size of a voleanie orifice. Obstructions of the outflow, by periodical
consolidation, would become more frequent, and thus would proceed a slow change of
the mode of action of massive eruptions to that which is peculiar to volcanoes.

This is probably the simplest manner in which volcanoes can originate. It will
apply particularly to a number of those which have undergone no change in regard to
the character of their lava, and the lofty cones of which rise over mountain ranges
consisting of the same material with their own lava and cinders, though owing their
origin to massive eruptions. We must now consider a third cause which would aid in
promoting volcanic action, and probably come very often into play. It is indicated by
the frequent occurrence of series of voleanoes extending in lines parallel to the axis of
the main outbursts. Their only possible cause is the formation of fractures, parallel to
the main fissure, and branching off from it. In order, however, that these could be
formed, solidification must have proceeded downward in the main fissure, without any
communication with the liquid portion in depth having been kept open. This process
would necessarily imply a temporary cessation of the process of extrusion. That this
could take place may be the more readily understood, if it is taken into consideration,
that the liquid masses filling the fissure in its whole extent must contract considerably
by constant loss of heat, and that any additional expansion produced by the promotion
of aqueous fusion at certain places had first to equalize this loss of volume, before
it could manifest itself in a rise of the whole mass. This twofold action, which is
probably one of the main causes of the intermittent character of volcanic ac-
tivity, must produce an alternating motion of matter within the fissure, and
there would be given ample opportunity, during a period of its subsidence, for
the consolidation of the upper portion to a great distance down from the surface.

If a period then followed in which, by dislocations of some kind, a change took
place in the conditions subterranean, and expansion began again to prevail within the
fissure, the new supply of force would manifest itself at the upper limit of the liquid
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OF VOLCANIC ROCKS. 65
portion, where the least resistance was offered, and new fractures would be opened
branching off at that depth from the main fissure. The liquid matter would ascend
through these secondary fractures, and, if these were of sufficient dimensions, give rise
first to the formation of parallel ranges of volcanic rocks by massive eruptions, and
then only to a gradual isolation of channels of voleanic action, as in the case first ex-
plained; or, if the fracture consisted of a series of smaller ruptures, cause at once the
formation of a series of voleanoes. The activity in these secondary fissures could con-
tinue long after any manifestations had ceased over the main fissure, and even after the
consolidation of the matter contained in this, with the exception of the volcanic hearths,
had proceeded into far greater depth. The formation of secondary fractures, branch-
ing off from the main fissure, might be repeated at different depths, and fissures of a
third order be formed, branching off from those of the second. Subterranean reser-
voirs of liquid matter, which may either be isolated or connected, would thus be formed
at different depths, and be arranged after a similar plan below ground, as we notice
among the active and extinct volcanic orifices above ground. The hypothesis of
the existence of such subterranean seas of melted matter, as they have been called, has
also, though in a very different meaning, been arrived at by the adherents of the theory
of a metamorphic origin of voleanic rocks. This coincidence increases the degree of its
probability. But unless the nature and distribution of those reservoirs is made de-
pendent on grander phenomena having connection with the interior of the globe, they
will not be capable of explaining the harmony prevailing either in one volcanic region
or between all these regions. We have been led, by arguing on our suppositions, to
the same conclusion at which we arrived before by induction from observed facts, namely,
that the seat of volcanic action must be at a comparatively limited depth. Yet it ap-
pears that this depth is in all cases below the shell of sedimentary rocks. Among the
reasons supporting this assumption, we mention only one. This relates to the chemical
composition of lava. The volcanic is typically the era of andesitic and basaltic com-
pounds. The occurrence of trachyte and rhyolite among the ejected rocks goes to
show, that the seat of action receded at certain places from the andesitic regions to
those of the compounds corresponding in chemical composition to the two kinds of rock
named. But if it had been partially above those regions where matter is still in its
primeval position, then we should expect that there would be volcanoes the lavas of
which, being derived from sedimentary rocks, would, as a whole, deviate in composition
from the law of Bunsen. No such voleanoes are known, and it is, therefore, not prob-
able that the seat of any of those the lava of which has been analyzed is within the
shell of sedimentary rocks. It is true that subordinate deviations from the composition
as required by theory occur; but it has been found sufficient to ascribe them, as in the
ease of Vesuvius, to a mechanical destruction of the rocks surrounding the channel of
ejection by the ascending lava.

Active voleanoes themselves furnish an illustration in evidence of their own
origin as here advocated. It is well known that small cones are frequently met with
on the slopes of larger volcanoes. If they occur in larger number, as on Mount Etna,
they are usually situated in lines which radiate from the crater. Each of them is built
up of layers of scoria and ashes sloping away from the center, where a crater is im-

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66 RICHTHOFEN — NATURAL SYSTEM

mersed, and such cones will occasionally emit currents of lava, and be in fact the rep-
etition on a small scale of the mother voleano. The usual and probably correct ex-
planation of their mode of occurrence is this: That, through crevices or by passage
through porous rocks, water gets access to glowing lava, and, by its action on the same,
causes the opening of a fracture, and, in immediate succession, the repetition of the
same phenomena which the mother-volecano presents when active. Just as these par-
asitic voleanoes have their roots in the glowing lava, volcanoes in general must, as is
demonstrated by their mode of occurrence, be considered as parasites on certain sub-
terranean portions of the material of massive eruptions, which still possess a high tem-
perature and are kept in a liquid state by the molecular combination with water which
finds access to them. This mode of origin of voleanoes, however, is only a repetition
on a small scale of the manner in which massive eruptions themselves originated, inas-
much as voleanoes bear a similar relation to the latter as these do to the primeval sub-
stance composing the interior of the globe, to which the fractures descend. These
main fissures, which are probably very few in number in every volcanic belt, form the
great arteries in this harmonious system. Their common origin will furnish an ex-
planation of the general similarity of the phenomena presented by different volcanic
belts ; while the varied and possibly very intricate mode of their ramifications towards
the surface, together with the different conditions of the rocks which they intersect, the
various rate and local diversity of the access of water, the different circumstances which
may determine the depth in which the expansive force of vapor can be brought into
action (among these may be the relative proportion in which chlorine, fluorine and sul-
phur are present), and other influences unknown, would give ample means for explain-
ing the diversity of all the phenomena of vulcanism within each separate volcanic belt:
such as the apparently intricate, and yet to a certain degree harmonious, mode of dis-
tribution of the rocks ejected ; the correlations existing among the latter in regard to
their composition ; the dependency of volcanic action on massive eruptions ; the mode
of distribution of hot springs, solfataras, geysers, and other phenomena which were ap-
parently associated with both modes of action ; the different phenomena connected with
the occurrence of earthquakes ; the small size of their area of disturbance when con-
nected with voleanic action, its varying, and sometimes very great extent in other
‘ases Where no connection with any one distinct volcano can be discovered ; the singu-
lar correlations, finally, which have been observed to exist between different volcanic
vents situated on the same belt. Some hints may even be got in regard to the remoter
correlations which apparently exist between the phases of volcanic action on neighbor-
ing belts, though it must be conceded that in respect to the latter numerous facts have
been observed which cannot be satisfactorily explained in the way here proposed, and
rather appear to indicate the influence of the phases of magnetic currents on the
manifestations of vuleanism.”

22 An extremely valuable contribution to the elucidation of these correlations respecting the phases of volcanic ac-
tivity was lately given by Dr. Emil Kluge, in his work, Ueber den Synchronismus und Antagonismus von vulcanischen
Eruptionen, Leipzig, 1863. The clear and able compilation of facts will be of lasting interest, though grave objections may
be raised against the author’s views on the origin of volcanic action, which are not based on any inferences drawn from the
correlations demonstrated in the same book.

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OF VOLCANIC ROCKS. 67
The foregoing considerations may explain what appears to have been the most
probable way in which voleanic activity was developed at and near the places of mass-
ive eruptions. They apply without difficulty to those cases where voleanoes have un-
dergone no change in regard to the character of their lava. We have still to consider
those more intricate cases where the latter has periodically changed. There are volea-
noes which exhibit a regular succession of andesitic, trachytic, and rhyolitic lavas, fol-
lowed in a later period by the outpouring of basaltic lava, either through the same, or
through other vents in the immediate vicinity. There are others in which only a part
of this series can be observed, such as the succession of basalt to trachyte or rhyolite,
which appears to be of the most frequent occurrence, or of rhyolite only to andesite.
As the order of succession is generally the same as that exhibited by massive eruptions,
it would appear that it must be due to the same causes in both cases. If we take it
for granted, that those extensive reservoirs of melted matter, from which was either
continued its quiet protrusion, or voleanoes were fed, have had their seat in those con-
centric layers of the crust composed of silicious masses which correspond to trachyte
or rhyolite in composition, then there is little difficulty in explaining the protrusion
of portions of them. For there must be a limit to the expansion of a substance such
as a given mass of andesite, by aqueous fusion, and thereby a limit to its ejection.
The eruptive activity might then either come to rest or continue. As it is very im-
probable that water takes originally a part in the composition of those masses below
the shell of sediments which have crystallized from a molten state, its access to them
at places contiguous to a source of heat, such as must be given by a fissure filled with
molten matter from below, must be attended by a powerful influence on them. It
would exert itself in aqueous fusion and expansion. But the viscidity peculiar to these
highly silicious substances would not allow their extrusion until after the more liquid
andesitic masses had been ejected. Supposing the reservoir in which this first
change was effected to have been in the trachytic region, further action from the same
could be cut off by the cessation of the ingress of water to it. Another reservoir, sit-
uated in the rhyolitic region, might then be isolated within the educting channel, filled
now with trachytic matter, and the same process repeated, as before, ending with the
change of trachytic into rhyolitic rocks. As regards the succession of basalt to these
silicious rocks, we refer to the fact established before, that the fissures which gave vent
to basalt, have all been formed at a much later epoch than those through which andesite
had ascended, and that they were only partly coinciding with them. It would appear
that subterranean reservoirs of liquid matter, connected with the surface by channels of
ejection, should have offered, in many cases, the places of least resistance. It may,
therefore, be inferred, that basalt, the great comparative liquidity of which is a well-
known fact, would enter many of those reservoirs, and be emitted through the same,
or through newly-formed channels, in preference to any matter of a more viscous con-
sistency ; and it will not be difficult to understand why basalt should, in many instances,
have again been followed by, or alternated during long epochs with, lavas of a rhyo-
litic composition.
These considerations, which may be equally applied to the order of succession
of massive eruptions and to that of volcanic lavas, are not given with a view of explain-
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68 RICHTHOFEN — NATURAL SYSTEM

ing this intricate subject, but only to show that natural occurrences may, with the aid
of the theory here advocated, be explained without having recourse to any forced as-
sumptions. There is no one of the processes pointed out which is not within the limits
of those we are accustomed to consider as highly probable in regard to that part of
voleanie action which is removed only a little way beyond immediate observation, and
therefore more accessible to well-founded speculation than is the remoter connection
between volcanic action and the fissures through which the massive eruptions of vol-
eanic rocks took place.

3. Other Theories respecting the Origin of Eruptive Rocks.

The various theories which have been proposed in regard to the origin, not
only of the volcanic but of all those non-foliated crystalline rocks which are made up
of silicates, diverge in different directions. Most of them, however, leave unnoticed
the most essential features of those rocks, such as their nature in regard to the details
of chemical composition, their similarity in character in distant countries and different
ages, the laws of their mode of succession and distribution, and the fact of their peri-
odical emission after long periods of repose ; and no one undertakes to account for
all of them. There may be distinguished two classes of these theories : the first com-
prehends those which assume the original seat of eruptive rocks to have been beneath
the sedimentary rocks, while the second embraces those which would have it to be
within the shell composed of the latter. It was the purpose of our foregoing theoret-
ical considerations to point out, that it is exclusively in the direction followed by the
theories of the first class that we may at all look for a satisfactory explanation of the:
relations presented by the eruptive rocks. But, though the views here advocated
belong altogether to this class, the leading theories embraced in it have a very different
scope. That form of them which was held by Buch, Humboldt and others of the most
prominent geologists, and is still quite largely adopted, starts from the assumption that all
eruptive rocks were ejected in the same condition in which they are supposed to have
been when at their original place in the earth’s interior, that is, molten by dry heat ;
while the contraction of the globe by loss of heat is regarded as the sole cause of their
ejection. Among the weighty objections which may be raised against these theories,
may be mentioned: that the eruptive rocks on their arrival at the surface have evi-
dently not had a temperature which would be sufficient for their dry melting ; that
they contain a certain proportion of water enclosed, which was formerly not brought
into account ; that the ejection from the depth to the surface of masses molten by dry
heat is a process impossible of explanation, and that, if it was possible, the rocks
should have a different texture from that exhibited by granite, diorite or propylite ;
that, finally, contraction alone is as little capable of furnishing an agent for the rending
of fissures opening towards the surface, which is the prime condition of eruptive
activity, as is the cooling of the globe of giving the conditions requisite for the process
of ejection itself. The theories mentioned have been longest maintained on the
European continent, where they are even now advocated by many. Though approach-
ing nearest of all to the most probable mode of origin of eruptive rocks, the reasons

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OF VOLCANIC ROCKS, 69

why they are untenable, with our present state of knowledge, are too obvious to be
made an object of a more detailed explanation.

More numerous and more obvious objections may be raised, chiefly from a geo-
logical point of view, against those theories of the second class, according to which all
the rocks under consideration, with the exception of those lavas which we actually see
being ejected from volcanoes, would have derived their origin exclusively from meta-
morphism z stu; be it that the change is supposed to have been mainly of a chemical
nature, and effected on or beneath the surface, by the action of water alone containing
certain substances in solution, or that refuge be taken to the codperation of super-
heated water and pressure at a great depth. A large amount of positive facts as well
as of sagacious reasoning have been applied in their defense ; but viewed in the light
of those observations which present themselves continually to the geologist in the
field, in evidence of an essentially intrusive and extrusive nature of those rocks, the
premises on which argumentation is based must appear extremely deficient. Also
will the reasons which we shall adduce against the leading theory of our day, be appli-
eable a fortior? against the assumption of an origin of our ‘‘ eruptive” rocks only by
metamorphism in sttw. Before entering upon that leading theory, we have still to
mention the existence of a number of others, which, though acknowledging the proba-
bility of an origin of all lava and ‘ trap-rocks” from the liquid interior of the globe,
assume that granite, syenite, diabase, diorite and certain porphyries are so-called hypo-
gene rocks, that is, have originated by metamorphism of sediments iz situ. Against
these theories may be raised the collective objections which apply severally to the
others.

The obvious objections which may be made to the theories hitherto mentioned
have given more and more ascendency to another doctrine, which we may designate
as the metamorphic theory of eruptive rocks, and which owes its great influence upon
modern geology to the fact of its starting from a certain number of established geo-
logical facts and lithological observations, and from the results of experiments. It is
eminently a theory of the second class. No arguments against it will be more potent
than those which prove the fallacy of the basis of all the theories of this class, as
they will show that we must look for the origin of eruptive rocks altogether in a dif-
ferent direction from that followed by them.

The metamorphic theory is essentially to the purport, that all eruptive rocks,
whether of recent or of ancient age, whether ejected by volcanic action or carried into
their present position without any sign of the latter, were originally sedimentary rocks
rendered liquid by the codperation of heat, pressure, and water. It is supposed that
these rocks, by the continued superposition of immense masses of sediment, had ar-
rived at a great depth under the surface, where the agencies mentioned would codper-
ate to modify and transform their state of aggregation, resulting either in a molecular
change alone, or in their conduction into a state of fusion. In the former case, the
sediments would be simply metamorphosed, while in the latter, they would either
crystallize in depth, with a total loss of their original structure, and form ‘‘ plutonic,”
or ‘‘hypogene,” or ‘‘indigenous ” rocks, or be forced upwards through fractures, and
solidify partly in the conducting channels and partly on the surface, when they would

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70 RICHTHOFEN — NATURAL SYSTEM

form ‘‘ trap rocks” and ‘‘lava.’’ One common cause is thus assigned to both meta-
morphic and eruptive (including volcanic) action, the latter being considered an ad-
vanced stage and ultimate result of the former. The immense action of metamorphism
is an undeniable fact. But while formerly, on account of the violent agencies which
were supposed to have been required for it, and the growing conviction that heat alone
could not produce effects on such an enormous scale as had been suggested, the sub-
ject had to be treated with caution, and any extreme assumption was received with
doubt, if not with a certain repulsion, the condition of things, in this respect, has of
late undergone a great change, as scientific experiments, and especially those of Dau-
brée, have demonstrated the extent of the influence of water and pressure in producing
metamorphic action. They have proved the remarkable fact that, at a comparatively
low temperature, and with the aid of pressure, the effects of water, if continued for a
sufficient time, particularly when it is charged with alkaline substances, will be able
to produce changes in the nature of rocks which surpass even the most audacious
assumptions of former time. An apparently safe foundation was now given to the
widest generalizations, and the consequence is, that an almost unlimited action is at the
present time brought to the account of metamorphism. The least founded concep-
tion, however, of the faculties ascribed to it, we consider to be its supposed sole in-
strumentality in the production of volcanoes and the eruptive action of former ages
from portions of the shell of sedimentary rocks. It is true that a great additional
degree of apparent probability has been given to the metamorphic theory of eruptive
rocks by the results of the microscopic examination of rocks so successfully instituted
by Sorby, since they prove a remarkable similarity, in the minutest texture, of the
minerals constituting granite and some cognate non-foliated rocks, with the main in-
gredients of certain foliated rocks made up of silicates, such as gneiss and micaschist.
These had been observed long before to form the last link of a series which may be
traced, by slow gradations, from unmetamorphosed sedimentary rocks, through those
which are undoubtedly metamorphosed, up to the crystalline foliated rocks mentioned.
The newly discovered facts appeared to indicate that granitic texture is the most ad-
vanced stage in a progressive series of molecular changes, and, carrying the argumenta-
tion still farther, the assumption was apparently justified that a corresponding origin
must be ascribed to other rocks, such as those of volcanic origin, which are connected
with granite by another long and gradual line of passage.

In drawing up our argument against this theory, (including all those doctrines

23 It may be frequently noticed, that those eminent men who, by creating a firm basis for induction, have pointed
out the path through regions in the field of science the knowledge of which had consisted before of a confused accumula-
tion of facts and suggestions, did themselves apply the newly acquired views within moderate limits, while sweeping general-
izations on the same basis were usually made by others. It is so in the present case. The limits within which Daubrée
himself has applied his ingenious conclusions from his experiments on the action of superheated water and pressure upon
silicates, will probably never be drawn any closer. No discovery, however, could have been more opportune to those who
had advocated before the origin of eruptive by the remelting of sedimentary rocks. Against the form of this theory which
was first proposed by Hutton, and enlarged by Lyell, Babbage, Herschel and others, similar objections could be raised as
against that form of the theories of the first class which was held by Buch. With the aid of the new views acquired,
however, it was remodeled and brought into its present shape. A far greater extent is thus being given to the conclusions
from Mr. Daubrée’s experiments than their author ever intended.

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OF VOLCANIC ROCKS. 71

which assume either all or a part of the massive crystalline rocks composed of silicates
to have originated by the metamorphism of sediments) we will first point ont some
reasons against its general tenor, which may be conclusive in regard to those rocks
the eruptive nature of which is almost generally conceded, while we will have to
bring some additional arguments against its special application to those which are often
designated as ‘‘ hypogene ” or ‘ plutonic.”

The metamorphic theory starts from the assumption of an alternating progres-
sion from the center of the earth, and recession towards it, of the chthonisothermal
planes, the former being caused by sedimentary deposition, the latter by denudation.
It is then argued that, by the progression of these planes, sedimentary strata would
acquire a more and more elevated temperature, and, being permeated by water, would
be metamorphosed, and finally rendered liquid. It is demonstrated that heat, generated
by the plication of the strata in the lowest and central part of an area of subsidence,
would aid in promoting these changes, which would end in the rupturing of the crust
and the protrusion of liquid matter. Eruptive activity should, according to these
views, be confined to areas of subsidence, in particular to their central portions. Geo-
logical observation does not favor this conclusion, since the emission, at least of the
voleanic rocks, has taken place on the borders of those areas, on high table lands, and,
in general, in places which have undergone elevation before and since the time of the
first commencement of the eruptive activity.

It would appear that the experiments of Daubrée should not be too freely
applied to reasoning on processes within the shell of sedimentary rocks. They have
been made with relatively large quantities of water, such as can scarcely be expected
to be present in solid rock at some distance below the ground. Supposing that that
quantity naturally enclosed in it would, under great pressure, cause its fusion, then
no reason can be adduced why there should not prevail a liquid state of all matter at a
limited depth below the surface, and over extensive regions, if not over the whole
globe. That such is not the case, is evident from the want of any signs of subterranean
tides. Leaving this difficulty out of consideration, another presents itself concerning
the periodicity of eruptive activity. If processes such as those suggested by the
adherents of the metamorphic doctrine were its cause, then it would be impossible to
give an explanation, why there have been long eras of rest intervening between others
of violent eruptive activity, or why the latter was of general distribution over the globe
during the Tertiary era, and preceded by a period of rest which was probably no less
general. The phenomena of vulcanism might have manifested themselves at a certain
time more in one country than in another, if metamorphism had been their cause, since
their principal theater would be constantly shifted to the places of the most violent
metamorphic action ; yet they should, at least in the aggregate, have been continuous.

These objections, however, against the metamorphic theory of eruptive rocks
are of little weight when compared with another, respecting their nature as chemical

24 Tt is needless to enter here upon a discussion of the causes and effects attributed to these phases in the “ flow of
heat” by the adherents of the metamorphic theory, since this topic has been made lately the subject of a paper by the skillful
hand of Professor Dana.

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72 RICHTHOFEN —— NATURAL SYSTEM

compounds. It is from this point of view, as we have repeatedly remarked, that are
offered the chief and most deeply-founded differences between the eruptive and the
stratified and foliated rocks. Definite numerical relations on the one side, complete ab-
sence of them on the other, and a transition between both, marked by the gradual dis-
appearance of those relations with the passage from granite, in which they are very
distinct, to gneiss and micaschist—these are in short the prominent characters of the
great divisions of those rocks which are accessible to the observations of the geologist.
We alluded at another place to the corollary of this distinction, namely: that it is
impossible that eruptive rocks are remelted sediments, because, if they were, they
would necessarily have to participate in the varied and indefinite chemical composition
of these. This important argument appears to have been completely overlooked by
the adherents of the metamorphic doctrine, and it would be sufficient by itself to
make the latter appear to be in contradiction with the true state of facts.”

If we descend from this general tie embracing the totality of eruptive rocks,
to those relations which either separate or connect distinct groups of them, in regard
to the time at which they came to the surface, we are unable to find any explanation
of the peculiar uniformity of these relations, if we adopt the metamorphic doctrine.
Rocks of great mutual similarity might have been occasionally ejected at different
places; but the repetition of the same order of succession in distant regions would be
just as inexplicable, and contradictory to the nature of sedimentary rocks, as the per-
fect chemical identity of rocks which are widely separated in space, but occupy a
similar position in the order of succession.

Another geological consideration may be mentioned which appears to weaken the
metamorphic theory. It is known that the most ancient rocks are distinguished, in
general, by a much greater similarity among each other in respect to chemical compo-
sition than is the case with those of more recent origin, and that one prominent feat-
ure of the majority of them is the presence of silica in a similar proportion to that in
which it is contained in granite. The formation of sedimentary rocks having been
due, at all times, to the disintegration of rocky matter antecedent to them in age, it is
obvious that ancient sediments would have participated in the silicious nature of the

25 Tt can hardly be comprehended that it should have been maintained, and be believed, with our present state of knowl-
edge, that clayslates were, by metamorphic action, converted into granite and syenite, and sandstones into porphyry ; since
granite and (quartzose) porphyry are chemically identical, while clayslate and sandstone differ in this respect not only among
themselves, but each of them represents a large number of different accidental compounds, without any law of mutual con-
nection. The supposition that ‘“ the presence of the sandstone formations affords the conditions required for the occurrence
of the great porphyry-masses,” gives evidence of an interpretation of natural occurrences by preconceived ideas. ‘There are, it
is true, numerous instances of sandstones having aequired, by metamorphic action, a more or less perfect porphyritic texture,
so as to be hardly capable of being distinguished, in specimens, from true eruptive porphyries; but geological observation
will seldom leave any doubt in regard to their true mode of occurrence ; and, as a generality, it is not difficult to distinguish
these metamorphic rocks with porphyritie texture from “the great porphyry-masses,” such as those of southern Norway, or
middle Germany, to which the above-mentioned supposition has been especially applied. It is evident that the eruptions of
the porphyritie rocks were there subaqueous, and that the true relation is exactly the reverse of that suggested: the ejec-
tion of great masses of quartzose porphyry afforded all the conditions required for the formation of sandstone beds. Quartzose
porphyry did overflow vast regions, and, by its immediate disintegration into tufaceous matter, gave origin to those red
sandstones by which it is so often accompanied. These cover the porphyry often in horizontal beds, and bear to it similar
relations of gradual passage and interstratification, as trachytie tufa does to trachyte, where the eruptions of the latter were
subaqueous.

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OF VOLCANIC ROCKS. 73

rocks by the destruction of which they originated, and should have had a tendeney
towards an average between them in regard to chemical composition. If such sedi-
ments were afterwards rendered liquid by metamorphic processes, and either carried to
the surface as eruptive rocks, or solidified beneath it, so as to form part of it only after
ages of denudation—if they were then again partially disintegrated, re-deposited and
reérupted, and so forth, down to our time, the final result should be an ever-increasing
uniformity in composition of both eruptive matter and sediments. It may be objected,
that this tendency towards uniformity must have been checked by the formation
of chemical sediments, as by them a portion of certain elements was eliminated, and
must have become almost irrecoverably lost for further’ participation in the supposed
revolving process. The substances prevailing in chemical deposits are lime, soda and
magnesia. Recent eruptive rocks should, therefore, besides showing a uniformity of
composition, be somewhat deficient in those substances, and contain in proportion
more of silica, alumina and potassa. The facts are exactly reverse. Not alone has the
variety of eruptive rocks rather been constantly increasing ; but, among those most
recent in origin, such are greatly predominant as contain lime, soda and magnesia
in larger proportion than ancient rocks, while they are relatively deficient in silica,
alumina and potassa. We arrive by this last line of argument at the conclusion, in
the first place, that the present variety of rocks, especially of those of eruptive origin,
can in no way be explained by, but is contradictory to, the assumption that they orig-
inated by the repeated destruction and reformation of the material which constituted
the surface of the globe in the most ancient time. From this it follows, in'the second
place, that, even supposing that the theory should be admissible for the majority of
eruptive rocks, entirely new matter must have come repeatedly to the surface, in order
to replace the constant loss, as it were, of substances such as lime and soda. Eruptive
rocks ascending from sources beneath the deepest sediments are the only means imag-
inable for the performance of this function. The enormous extent to which matter
must have been supplied from that source is especially evident, if it is considered that
there must have been a time when no sediments existed on the face of the globe, and
that the disintegration of preéxisting matter is the prime condition to their formation.
The entire mass of the sediments which make up the exterior shell must, therefore,
have been derived from the destruction, partly of the original crust which solidified on
the globe, and partly of those rocky masses which protruded through that crust to the
surface. It is eminently probable that the mass of matter derived from the latter
source exceeds very far that which is due to the first. This consideration shows
that it is hardly possible for us to form an adequate idea of the vast importance which
the periodical emission of rocky matter from places beneath the primordial surface must
have had in the history of the formation of the crust of the globe.

We may thus start from any point of view within the range of positive
knowledge, and we shall find that there is not one line of argumentation which does
not go to show that the doctrine of the origin of eruptive rocks by the metamorphism
of sedimentary matter is irreconcilable with the most prominent established facts. Not
only does it fail completely to explain them, but with most of them it is in obvious
contradiction. But as a metamorphic origin has been proved to evidence in regard to

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7A RICHTHOFEN— NATURAL SYSTEM

certain foliated rocks, and as it appears no less safe to conclude that those rocks, the
intrusive or extrusive character of which is generally admitted, originated from be-
low the sedimentary rocks, we have still to examine where is the limit to which the
theory of such an origin may be safely applied. A line of gradual passages connects
the foliated rocks with gneiss, and through it with granite, and another line of
gradual passages can be traced from the volcanic through the porphyritic to the
granitic, and through them again to the gneissoid rocks. The affinity of granite and
gneiss may therefore be said to be the central point from which all other lines of
passage in the nature of rocks diverge ; and it must be ascribed to this relation,
that many geologists, while admitting the origin from below the crust of the globe
of “trap-rocks” and “lava,” still consider granite, syenite and cognate rocks to
have been generated by the metamorphism of sediments, and to occupy now the
same position which these sediments had before being metamorphosed. The pe-
culiar character of granitic texture, which is nearer allied to that of foliated crystal-
line than to that of voleanic rocks, the geological occurrence of granite in intimate
connection with those and its apparent independence of position in reference to the
latter, the singular order of solidification of the minerals constituting granitic
rocks—these and other reasons of a similar kind would indeed appear to support
the assumption that granitic and volcanic rocks differ in regard to their origin ; while,
on the other hand, additional probability is apparently given to the connection of
granite and foliated rocks in regard to their mode of origin, if it is taken into con-
sideration that many varieties of gneiss resemble granite closely in chemical com-
position, although others deviate from it in this respect and do not conform to the
law of Bunsen.

According to the theory here advocated, all those rocks will be the true repre-
sentatives of the primordial mass of the globe, which are subject, in regard to their
composition, to the law of Bunsen, and occur, besides, in positions which place
beyond doubt their intrusive or extrusive origin, so far as they no longer occupy
their original place. As regards the first distinguishing mark, it is known that granite,
syenite, diorite, diabase, and nearly all those porphyritic rocks which have been con-
sidered as of hypogene origin are subject to the law of Bunsen, as far as their composi-
tion has been ascertained. In this respect, therefore, they are fundamentally different
from sedimentary and metamorphic rocks. Although believing this to be conclusive
evidence of their origin from below the shell of sediments, we must enter into a short
discussion of the chief objections which have been raised against the necessary corollary
from this mode of origin, which is the eruptive character of those portions of granite
which we see on the surface, and the assumption of the existence of this as the foun-
dation of all sedimentary rocks.

It is argued by the adherents of the metamorphic theory of eruptive rocks, in
the first place, that granite cannot be the fundamental rock in the crust of the globe,
on which those of sedimentary origin are resting, since even very ancient masses of it
are frequently found to overlie stratified rocks. But this fact, which was known long
ago, shows only that it is impossible, on account of the great thickness of sedimentary
rocks, to know by ocular observation of what nature is the fundamental rock. It is
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OF VOLCANIC ROCKS. 75

logically absurd to assume the non-existence of a foundation consisting of rocks differ-
ing in origin from sedimentary rocks ; and the conclusion is inevitable, that it must be
composed of rocks which were generated by the solidification of such masses as formed
part of the primordial substance of the globe, next to the surface. Probabilities
accumulate to point towards the assumption that very silicious granite composes that
foundation, together with such gneissoid rocks as must be supposed to have been formed
in an incipient sea of very high temperature, and under the pressure of the superin-
cumbent atmosphere of aqueous vapors, at a time when the cooling of the globe had
advanced far enough to allow of their first condensation, and when pressure, water,
and heat could codperate to produce on its surface similar effects to those which the
same agencies are supposed to have wrought in later periods at an ever-growing dis-
tance from it. The most probable mode of these ancient processes has been pointed
out by Daubrée. The true nature of the foundation rock can, of course, not be
positively known, but must remain a matter of conjecture. No solution of this
problem will, however, be more satisfactory than that which is based on the hypoth-
esis of Sartorius, because it is in harmony with all the phenomena of vulcanism.
The probabilities in favor of a granitic foundation are, therefore, very great, while
no valid objection has yet been raised against its having the nature indicated.

It is further argued that, the position of granite being always in the midst of
sedimentary, or of such foliated crystalline rocks as are connected with the former by
gradual passage, and often conspicuously above such rocks, the only material from
which it could be generated are the sedimentary rocks themselves. The arguments
brought in evidence of these assertions are among the most potent which can be
adduced against them. Granite, in Norway, is superposed on Laurentian rocks turned
upon their edges, and, as no channel can be found by which it ascended, it is argued
that it must have been generated by metamorphism 7m situ. But as, on the other hand,
it has justly been remarked, that no position is more favorable for metamorphic action
than that of upturned strata, it is difficult to comprehend why the Laurentian rocks,
which were evidently nearer to the source of heat, should not have been metamor-
phosed in a much higher degree than the overlying granite. The same objection may
be made in other instances, as in the case of the Huronian and Laurentian rocks in
Canada, which were found to be overlain by granite thousands of feet in thickness.
The negative evidence, apparently afforded by the fact that no channels have been dis-
covered through which the granite could have protruded, is of very slight value, if it
is taken into consideration how rare are the instances where the conducting channels
can be seen, in the case of overflows of volcanic rocks, or even in that of currents of
lava. There are a few more instances known where granite can be distinctly observed
to overlie the upturned edges of stratified rocks, and must, therefore, have overflowed
the same in a liquid state. In some other cases, as in those of the extensive granitic
areas of Bessarabia and Western Australia, the true geological position of granite
escapes observation ; while, ina number of others, it has the appearance of an intrusive
mass.

The strongest objection which may be made to the eruptive nature of
granite relates to its lithological characters, which are different from those of the

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76 RICHTHOFEN — NATURAL SYSTEM

eruptive rocks of the present era. The metamorphic theory suggests that enormous
volumes of those masses which it supposes to have been rendered liquid by meta-
morphism, and small portions of which were occasionally emitted to the surface, did
solidify below ground with a total loss of their previous structure, and may arrive at
the surface by elevation and denudation. Granite is supposed to have been in all
cases a hypogene rock, generated by metamorphism in those same positions in which
it is found at the present time, and to be still formed continually in the same way ;
though it is admitted that the tension which may have attended its former state of
liquidity, may have caused it to ramify into rents and fissures of the overlying rocks.
We have, of course, to assume the existence of an immensely greater amount of hypo-
gene matter corresponding in composition to all the different eruptive rocks, but buried
so deep beneath the present surface, that, possibly with the exception of a few granite
masses, it never does, nor ever will, form part of it. We have, for this reason, to con-
sider all non-foliated crystalline rocks made up of silicates, and which are visible on the
surface, as having been removed far from their original seat, and therefore as being
eruptive. It is a matter of course that only a very small portion of all matter protrud-
ed from the depth would be ejected to the surface, as by far the greater portion would
solidify within the channels of ejection. Granite occupies frequently the latter posi-
tion quite distinctly. But if we consider those large accumulations of the same rock
covering hundreds of square miles, we can only explain them in two ways. Hither
they must occupy their primeval position, as is supposed by the adherents of the
metamorphic doctrine, or they must have been ejected and spread in a liquid state
over the underlying rocks. The objections against the first supposition are, the adapta-
tion of granite to the law of Bunsen, and the fact already noticed, that it overlies
the upturned edges of strata ; the objection against the other is the peculiar mineral
character of granite. But we should always bear in mind that our knowledge in re-
gard to the conditions required for producing any certain kind of mineral character
within a compound of silicates of any certain composition is very limited, as is illus-
trated, among others, in the cases of propylite or dolerite ; and the inferences based
on subjects which are imperfectly known to us should, in geology, always be made
subordinate to those which we can positively establish by observations. It is not on
the ground of the latter that a hypogene origin is ascribed to all granite, but on the
ground of its mineral characters. It is said that granitic and voleanic rocks offer quite
generally a different appearance, and that the texture of the former indicates crystal-
lization under great pressure, while the volcanic rocks were evidently solidified on the
surface. As regards the first assertion, such rocks only should be compared together
as have a similar composition, that is, granite with rhyolite, or diabase and augitiec
porphyry with basalt ; not, as is ordinarily done, granite with phonolite or basalt. We
referred already to the similarity in character between certain varieties of rhyolite and
granite. It is corroborated by the microscopic examination of these rocks by Ferd.
Zirkel, from whose memoir® I quote the following passages: ‘ Quartziferous trachytie

26 Dr. Ferdinand Zirkel, Mikroskopische Gesteinsstudien, in Sitzungsberichte der Kais. Acad. der Wissenschaflen zu
Wien, vol. 47, 1863.

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OF VOLCANIC ROCKS. 77
porphyry (rhyolite) cannot be distinguished under the microscope in any respect what-
ever from felsitic porphyry. If one dare not doubt the eruptive origin of the former,
one cannot but acknowledge the same mode of origin in regard to the latter.” And
again: ‘Let us here remark, that the microscopic structure (texture) of trachytic
quartz does not differ in any respect from that of granitic quartz ; there is no difference
between these two families of rocks so widely separated in time, either as regards the
number or the appearance of the water and glass cavities.” Considering the second of
the before-mentioned assertions, it is true that Mr. Sorby has, by his sagacious obser-
vations, come to the conclusion that granite solidified at a temperature of about 600°
Fahr., and that the various granites have been formed under a pressure equivalent to
a depth of from 40,000 to 69,000 feet. The calculation by which these figures were
obtained, is made upon the basis of the proportion between the bulk of the water
contained in the microscopic cavities, and the size of the vacuity, the latter being sup-
posed to indicate that the water had formerly filled the entire cavity and contracted
within it after the solidification of the surrounding rock. The laws of hydrostatic
pressure, however, as Daubrée justly remarks, are in such a case not applicable in the
same way as they would be in a column of water ascending through a fissure ; temper-
ature and pressure may, in a mass of rock solidifying from a viscous state, be pre-
served, as in a closed vessel, to within a few feet from the surface ; ‘‘it is, therefore,
possible that many processes, such as the crystallization of granite, may have been
going on under pressure, though at a very limited depth.” Considering that a mass
of granite, after the solidification of its exterior portion, will indeed be enclosed within
walls that may offer a strong resistance equivalent to a great pressure, it may be in-
ferred that it will assume, in crystallizing, a similar texture to that which it would
have obtained if the same process had been going on at a great depth below the
surface. It can hardly be suggested what must have been the texture of the crust of
a granitic mass. But the age of the granite is sufficient to justify the conclusion that
the crust of those masses which solidified on the surface of the globe must have been
completely abraded, and only those portions be preserved to observation which con-
solidated under considerable pressure.

Though these arguments may show that the difference in the character of a
large mass of rock need not necessarily be proportionate to the depth under the sur-
face of the globe at which it solidified, and that granitic and volcanic rocks are nearly
related as regards their microscopic texture ; yet, the conspicuous external differences
between these two classes of rocks must remain a problem difficult of solution. The
proportionate rapidity with which volcanic rocks, on account of their usually small
volume, must have cooled, may be among the reasons. But it is not the only one. There
are differences in the nature of rocks, which escape our present means of explanation.
Hornblende-propylite, though no doubt at all can be entertained in regard to its sub-
aérial solidification, has completely the character of the so-called Plutonic rocks, and
yet lacks their supposed principal distinguishing mark: crystallization in depth. Con-
sidering that certain varieties of propylite, andesite, and trachyte are modifications of
the same group of chemical compounds; and, though having been solidified on the sur-

face of the globe within a short period, yet exhibit marked external differences, we can-
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78 RICHTHOFEN—NATURAL SYSTEM

not be surprised to see, that other rocks composed of silicates, which are so widely sep-
arated in time as is the case with granite and rhyolite, should offer even greater differ-
ences. Some further clue to a better knowledge of this subject may be expected from
the study of the nature of volcanic rocks. We see the same chemical compound
forming highly viscid lava in one voleanic crater, while it is quite liquid in another.
There, it solidifies to dolerite or leucitophyre; here, to basalt. Similar influences
appear to work greater differences in more silicious compounds. We should therefore
put little value on the doubts entertained in regard to the solidification of granite, as of
a rock ejected on the surface and expanded over it, until we are better acquainted
with the causes of the lithological differences among volcanic rocks; and the evidence
regarding the mode of origin of granite should, till then, be mainly taken from its
geological occurrence and its chemical composition.

RELATION OF THE DISTRIBUTION OF THE VOLCANIC Rocks TO THE CONFIGURATION OF
THE SURFACE OF THE GLOBE.

If we embrace in a broad review all those relations mentioned in the foregoing
chapters which regard the history of eruptive action, we arrive at the general conclu-
sion that, in reference to the entire globe, it is one homogeneous and harmonious
whole, and has been attended by such gradual changes only as were necessarily
occasioned by the progress of the physical development of the globe itself, while, in
regard to every different part, it presents a series of distinct phases intimately con-
nected by mutual relations. It would be a subject worthy of the closest investigation,
to trace the effect which the events of these phases have had severally upon the strue-
ture and the configuration of any separate country. A comparison of these effects
as they are manifested in different regions would then aid in establishing some of the
chief causes of the differences of structure peculiar to each; and the knowledge of
these would make us better acquainted with the laws governing the evolution of the
globe, and prepare the way for a more thorough understanding of its physical geog-
raphy. I will only attempt at this place to trace the effects referred to in regard to
the last of the phases of eruptive activity, the only one which was nearly contempo-
raneous in all countries. It occurred at a comparatively recent date, and we can view
the results in a clearer light than we can those which are remote in time and partly
obliterated by the vast changes which since then have revolutionized the surface.

We have, in the first place, to trace more in detail than we have done before,
the peculiarities of the distribution of volcanic rocks. It has been observed that ac-
tive voleanoes are chiefly situated along the lines of the present sea-coasts, especially
at the foot of mountain ranges parallel to them; or that they follow elevated sub-
marine ranges, when they will either remain submerged beneath the sea, or protrude
above it, forming chains of islands. Some of these appear to mark the lines of high
mountain ranges bounding submarine continents, if we may use this expression for
those areas of a shallow sea-bottom which are separated from others of great depth
by wall-like elevations, as are indicated for instance in the main ranges of the vol-

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OF VOLCANIC ROCKS. 79

canoes of the Indian Archipelago. Active voleanoes have also been found to be par-
ticularly numerous in those regions where the narrow terminations of two continents
verge towards connection, as is the case in Central America, between Alaska and
Kamtschatka, and between Australia and Farther India. The mode of distribution of
extinct voleanoes has been little investigated. But as they are far more numerous
than active vents, and as volcanic action has completely ceased in extensive regions, a
perfect understanding of the distribution of volcanic activity on the globe can only be
acquired when they shall be fully taken into consideration. The general laws of their
distribution appear not to differ from those relating to active volcanoes. They, too,
have been evidently dependent upon the course of the sea-coasts which existed at the
time of their activity, and were in many places quite different from those of the pres-
ent epoch. But we must note, besides, this additional feature of extinct volcanoes,
that among the theaters of their grandest and most extensive displays are certain
countries which were formerly covered with large inland basins filled with salt water,
and form now mostly elevated table-lands on which may still be seen the remnants of
those former inland seas. Volcanoes so situated form often crowded groups occurring
at distances of more than five hundred miles from the sea-coasts which existed con-
temporaneously with their activity. To them belong the numerous extinct volcanoes
on the plateau between the Sierra Nevada and the Rocky Mountains, a region which
has probably had no equal in regard to the extent in which volcanic activity has taken
place, as it comprises the States and Territories of Oregon, Idaho, Montana, Utah, Col-
orado, Nevada, California, Arizona, New Mexico, and continues southerly to the high
lands of Mexico. Other volcanoes of high table-lands are those of the Mongolian
Desert, the Thian-Shan, Armenia and, probably, eastern Africa. The volcanoes of all
these regions are extinct, with the exception of a few in the southern part of the pla-
teau of Mexico, which are in action, and some others which still exhibit solfataric phe-
nomena.

Interesting as would be a map embracing all active and extinct volcanoes, it
would give only the outlines of the distribution of those larger accumulations of volcanic
rocks which were produced by massive eruptions, and have ordinarily attracted little
attention.” Geological maps being often imperfect as regards them, it is not yet pos-
sible to distinguish any other features of their distribution than those mentioned in
reference to voleanoes, as they can always be found in the vicinity of these. We may
however add, that massive accumulations may eventually be found near the summits
of mountain ranges, where volcanoes do ordinarily not occur.

2 This may be exemplified in regard to the Andes of North and South America. Humboldt (Cosmos, vol. iv)
distinguishes five groups of volcanoes, separated by non-voleanic regions. They are mentioned by him as follows: Ist, vol-
canic group of Mexico, nearly five hundred miles in length, but occupying scarcely more than one degree of latitude, on
account of its direction from east to west; 2d, three hundred and fifty miles free from volcanoes; 3d, group of Central
America, having a length of over eight hundred miles; 4th, seven hundred and thirty-five miles non-voleanic ; 5th, gronp of
New Granada and Quito, five hundred and fifty-five miles in extent; 6th, the longest interval without volcanoes, being 1,133
miles; 7th, group of voleahoes of Peru and Bolivia, extending over five hundred miles ; 8th, a non-voleanie space of six
hundred and twenty miles 9th, the group of Chile, the most extensive range of voleanoes of America. being 1,143 miles in
length. Summing up, Humboldt takes about 3,000 miles to be the aggregate extent in length of the voleanic regions, 2,838
miles that of the intermediate spaces which are free from yoleanoes. If all the voleanic rocks occurring in the Andes

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80 RICHTHOFEN — NATURAL SYSTEM

If we turn our attention from the mode of the geographical to that of the geo-
logical occurrence of voleanic rocks, we may first consider from this point of view the
relations just mentioned. The two modes of distribution which are most conspicuous,
namely, on the foot of mountain ranges and along sea-coasts, are nearly identical in a
geological aspect. For most, if not all of those ranges the flanks or vicinity of which
are distinguished by the occurrence of volcanic rocks on a large scale, have been contig-
uous to sea-coasts in the Tertiary period, or they are so at present, or they have been
so in the intermediate time. This is true of active and extinct volcanoes, as well as for
massive eruptions, in the Carpathians, on the southern foot of the Alps, on the borders
of the plains of northern Germany, on the slopes of the central plateau of Asia, in their
whole extent from Armenia and the Caucasus, passing Lake Issikul and Lake Baikal,
to the vicinity of Pekin, in the coast ranges of California, in the Cascade Mountains of
Oregon, and in numerous other countries. As regards the prominent occurrence of
voleanic rocks on high table-lands, it is analogous to their situation on sea-coasts, imas-
much as the salt lakes occurring on them, which were of much larger size in the Ter-
tiary period, would be an equivalent of the vicinity of the sea. Those larger regions
not covered by salt lakes, which are situated in the interior of continents, and have
been so since the commencement of the Tertiary period, were generally not the theater
of outbreaks of voleanic rocks. This circumstance renders almost certain the influence
which the neighborhood of large quantities of salt water has had upon the commence-
ment and main phases of the eruptive activity, though its continuation in the latter
phases, which are those of volcanic action proper, may in numerous instances have
been maintained by fresh water.

The relations of the distribution of the volcanic to that of the granitic and por-
phyritic rocks have been considered in another chapter. There is probably no place dis-
tinguished by the accumulation of the first where the previous eruption either of granitic
or of both granitic and porphyritic rocks may not be observed, if the conditions are such
that observations in this direction are possible. But as there are porphyritic regions out-
side of the volcanic belts (for instance, the quartzose porphyries of southern Norway, or
the Triassic trap-rocks of the Connecticut Valley), and as the same is true of a great
number of known granitic districts, and probably of a far greater number of others
which are not accessible to observation, it is evident that the existence of ancient chan-
nels of ejection did not necessarily imply their reopening in the Tertiary period, while
there can scarcely be any doubt that the once shattered places where they were situ-
ated offered less resistance to the formation of new fractures, than other portions of
those regions in which the greatest disturbances took place in the Tertiary period.

We have, finally, to mention the connection which apparently exists between the
occurrence of volcanic rocks and the regions where ancient formations have been

were laid down ona map, it would hardly show such great intermissions. The extensive fields of lava found by Capt.
Fitzroy, in southern Patagonia, the voleanic rocks of the Desert of Atacama, and of northern Peru, are instances of their
extensive distribution in the non-voleanic spaces of South America, while on the northern continent they are one of the
main features in the structure of the great western mountain ranges, probably throughout their whole extent from Panama
to the peninsula of Alaska, and are accompanied probably by thousands of extinct craters.

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OF VOLCANIC ROCKS. 81

highly disturbed and subjected to an extensive metamorphism. It appears that the
eruptive action was in a great measure, perhaps absolutely, limited to regions which
offered this condition. But this connection by no means justifies the conclusion that
the existence of metamorphic foliated rocks was among the first causes of the ejection
of voleanic rocks. There are not only countries, such as the mountains of Scandinavia,
the Ural Mountains, the Appalachians, and other ranges distinguished by the records
of a very ancient metamorphism, where no trace of volcanoes has been found ; but this
applies even to a few, though particularly grand instances of mountain ranges which
were the theater of metamorphic action on a grand scale within the most recent periods,
such as the Alps, the Himalaya, and the Pyrenees.

The two last named relations, both conspicuous, yet both secondary in import-
ance, are evidently nearly identical, since granite enters, probably in all cases, into the
structure of those same countries which have been the theater of an extensive meta-
morphic action. It is, however, worthy of note that, of the regions so distinguished,
part of those only appear to have become the theater of eruptive activity in the Ter-
tiary period, where granite must, by the mode of its occurrence, be assumed to have
formerly been extruded to the surface, as is the case in the Sierra Nevada, on the
southern foot of the Alps, in middle Germany, and in central France, while no volcanic
rocks arrived at the surface in other mountain ranges where granite occurs only in the
shape of wedges surrounded by foliated rocks, and bears a merely intrusive character,
as in the central range of the Alps.¥

28 Another peculiarity in the distribution of volcanic rocks may be noticed. It is their outbreak at places of dis-
location or faulting of mountain ranges, if we may apply these expressions for the immense displacements which some of the
latter have undergone, along lines either at angles with or parallel to their direction. A remarkable instance of the first
kind is afforded in Hungary, where a fracture, along the line Eperies-Kaschau, crosses the Carpathians in a nearly meridional
direction. ‘To the west of it, ancient formations, along with granitic, porphyritic, and voleanic rocks, make up the surface,
and are elevated to high mountain ranges culminating in the High Tatra. ‘They extend eastward close to the fracture, and
there terminate abruptly. East of it, the main chain of the Carpathians continues as a range of little elevation, and con-
sisting chiefly of Cretaceous and Eocene rocks, while the high foothills of the western part are replaced by the Hungarian
plains. Immediately out of the fracture rises the Eperies-Kaschau range, of nearly one hundred miles in length, which
consists exclusively of volcanic rocks. The subsidence of the land on the eastern side of the fracture must have amounted
to several thousand feet in this case. A transverse dislocation of similar magnitude, by which the western side was sunk
thousands of feet, is observable on the eastern bank of the Rhine, along the boundary of Switzerland and Vorarlberg. No
volcanic rocks however are visible at that place, which fact appears to be in accordance with the absence of any ancient
eruptive rocks in the neighborhood, and the wedge-like shape of granite in the nearest places of its occurrence. A peculiar
instance is afforded by the three porphyritic regions on the southern slope of the Alps, each of which indicates a deep de-
pression, bounded by longitudinal and transverse fractures, of that area which has in each case been the seat of eruptive
action. Prof. J. D. Whitney and I had occasion to observe another instance, more forcibly striking for its grandeur than
any of the foregoing. Following the crest of the unbroken range of the Sierra Nevada, from south-southeast to north-
northwest, one arrives in the northern part of the County of Plumas at a sudden change of scenery and rocks. ‘lhe meta-
morphic rocks of the Sierra Nevada are broken off and terminate in a ]jne apparently at about right angles with the diree-
tion of the crest of the former. The rugged and wild scenery which they present, particularly in this county, gives way
abruptly to one equally remarkable for its beauty and geological interest. The high voleano, Lassen’s Peak, rises in the dis-
tance, amidst an intensely volcanic group of hills. Low forest-clad ridges built of lava, and separated by meandering plains
covered by meadows, stretch southward from it, gradually sloping down towards the line of dislocation, and there abut
against the wall of metamorphic rocks. The whole surface, though lower than the summit range, is entirely voleanic. Sev-
enty miles northwest from Lassen’s Peak rises Mount Shasta, more sublime even than the other. Between both is a deep
depression, through which the Pit River takes its course from northeast to southwest, at little altitude above the sea.
This depression is situated at the very place where the crest of the Sierra Nevada should pass, and is directed transversely
to it. Northwest of Mount Shasta, Prof. W. H. Brewer found the continuation of the Sierra Nevada, with similar char-

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82 RICHTHOFEN — NATURAL SYSTEM

These appear to be the most essential geographical and geological relations
regarding the mode of distribution and occurrence of volcanic rocks. It is perfectly
evident that no one of them singly was the chief cause of their occurrence, nor were
all of them together ; but each of them had a marked influence, either upon the direc-
tion and the location of the orifices of the fractures, or upon the mode of ejection.
Keeping in view these different relations, I shall dwell, in the rest of this chapter, more
particularly on the connection between the occurrence of volcanic rocks and the con-
figuration of the surface. This subject may appear to be beyond the scope of this
paper. But the examination of every question which relates to the inner connection
between the phenomena attending the ejection of volcanic rocks will aid in disclosing
the true nature of these, and promote the knowledge of the principles of their natural
system. We will endeavor to answer the following questions: Was the particular
structure of certain portions of the crust of the globe, which is indicated by the situa-
tion of elevated regions on its surface, among the causes of the eruptions of volcanic
rocks? or were the inequalities of level on the surface, in the volcanic regions, due to
the processes attending and the agencies causing the eruptions?

The answer to both questions must be in the affirmative. The peculiar structure
of the earth’s crust, at those places where mountain ranges and highlands rise on its
surface, appears to have influenced in a great measure the distribution of the volcanic
rocks, because those among them which are accompanied by the latter, had been
elevated before the time of the eruptions, and the adjoining lowlands were not the
theater of eruptive activity. But, on the other hand, it is evident that the ejection of
voleanic rocks, or rather those subterranean processes of which they were one of the

acter in rocks and scenery to that which that range has in other parts. It appears that a gap of more than one hundred
miles in length has been formed in the region of the two volcanoes, by the displacement of a portion of the Sierra Nevada,
which was probably bounded by two lines of fracture transverse to the direction of the mountain range, and has subsided
thousands of feet, and that then an immense accumulation of volcanic rocks filled up the gap, and closed in building up the
two giant volcanoes. Other lines of dislocation which have given vent to voleanic rocks, and which have more frequently
been noticed, are directed parallel to mountain ranges. Of such nature appears to be the abrupt descent of the Sierra
Nevada towards the Great Basin, which has been the theater of violent eruptive action ; and probably the relations on the
western slope of the Rocky Mountains are of a similar nature. The Vihorlat-Gutin Range, in Hungary, offers a striking
illustration of an extensive accumulation of volcanic rocks along the foot of a preéxisting mountain range, though the dis-
location is not conspicuous in that country, on account of the deposition of recent sediments which filled up the Hungarian
Basin. ‘There may be some affinity between these modes of occurrence of volcanic rocks and the manner in which they are
met with in certain areas of flat or hilly countries, surrounded by ranges composed of ancient rocks. The best illustration
is afforded by the Basin of Transylvania. The undulating country of the interior is encircled by high ranges consisting of
ancient formations, which are lined on their inner side with volcanic rocks. Hungary itself affords a similar instance, though
less regularity is perceptible ; and the same structure is somewhat approached in the geological relations of Bohemia. We
may also mention, as recalling that mode of occurrence, certain depressions between the two summit ranges of the Sierra
Nevada, such as the Basins of Sierra Valley and Lake Tahoe, which are encircled, first by a ring of volcanic rocks, and
then only by the metamorphie and granitie rocks which form the bulk of the Sierra Nevada. Orthe “ Parks” of the
Rocky Mountains. heir geology, it is true, is almost unknown. An interesting description of the San Luis Park, the
greatest among them, recently published (see American Journal of Science and Arts, November, 1867), shows that the
elevated rim of its basin, which is estimated at eighteen thousand square miles, is made up of ancient formations, and that
voleanie rocks encircle more immediately the extensive plain forming its bottom, which is itself composed of volcanic
sediments, thus completing a structure that reminds of that of Transylvania, in more than one respect. If we look for
instances on a grander scale, we may find some analogy with the mode of occurrence of volcanic rocks as just described,
in the voleanic ranges encircling the Basin of the Pacific Ocean. And it may not be out of place if we call attention to the
similarity with these circular basins which is presented by the configuration of the surface of the moon.

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OF VOLCANIC ROCKS. 83

resulting events, exerted a powerful reaction upon the further promotion and accelera-
tion of those motions of the crust by which the irregularities of level of the surface were
produced. This may be inferred, in the first place, from the fact, that in and along
volcanic belts, elevation and plication of sediments have taken place, probably in every
instance ; both modes of disturbance have affected, partly sediments anterior in age
to the eruptive action, and partly such as were contemporaneous with and posterior to
it; and there may be frequently noticed a diminution of the intensity of these dis-
turbances, in an inverse ratio with the distance from the voleanic belt.22 But besides
this fact, observation has placed it beyond doubt, that in the second half of the Ter-
tiary period, and after it, a greater aggregate amount of relative, and probably too of
absolute elevation has taken place generally over the globe than had been the case for
ages before. The altitude of the rocks composing the summits of the highest moun-
tain ranges and table-lands of the present time, exceeds that which they had in the

——

*9 This evident connection, in voleanic regions, of elevation and plication, as cause and effect, appears to be one of
the gravest objections against the general validity of the theory of Mr. Hall on the mode of formation of mountain ranges,
though the incongruity of an explanation of the granitic wedges by metamorphism in situ with the chemical nature of
granitic rocks, the total insufficiency of a gradual sedimentary deposition for accounting for a foot-for-foot subsidence of
large areas, and the difficulty of finding any natural cause of the assumed fact that the belts of greatest amplitude within
each area of subsidence should have been elevated to mountain ranges in preference to other neighboring parts, are objec-
tions of scarcely less weight. It is a well-established fact, that the Alps existed as a mountain range, at least from the time
of the Jurassic period (though probably long before it), and have gained in height in later periods. Yet the Jurassic and
Cretaceous strata are as much contorted as are those of more ancient age, and the same is true in a still greater measure
of the Tertiary Flysch. If, therefore, Mr. Vose, in his Orographie Geology (p. 68), arrives at the conclusion: “So far as
we have examined the facts, no other hypothesis than that of the slow sinking of vast masses of yielding sediments can at
all satisfactorily account for the plication and other evident effects of a compressive force so invariably exhibited in moun-
tain districts,” the more recent formations of the Alps confirm this view as little as the above-mentioned example regarding
the voleanic belts. It may be mentioned, in this connection, that, while Mr. Hall’s theory would necessarily imply an
increase in the intensity of plication towards the axis of greatest accumulation, which is considered by him to have been
coincident, after the elevation, with the axis of the mountain ranges, this conclusion is not confirmed by the structure of
the Alps. In Vorarlberg and northern Tyrol, which offer probably, of all parts of the Alps, the most normal and regular
structure, all strata, commencing with those of Triassic age, are least disturbed where they are nearest to the axis, and the
plication increases towards the foot of the range. Superposition, by contortion, of older formations on those of more recent
origin, is encountered with increasing frequency of instances and growing distinctness, in crossing the parallel waves from
those nearest to the axis, towards the northern foot of the range. Another objection, nearly related to the foregoing, is
this, that, granting that a trough-like depression would be able to cause plication by the settling of the yielding strata
towards the axis of greatest subsidence, the waves thereby formed would have their steeper inclination towards that axis,
and their flatter slopes directed outwardly, since a greater resistance would oppose the motion of the lower strata than
would be offered to the higher ones. The analogous instance, to which reference has been made, of a wave rolling towards
the beach, shows plainly the effect of the retardation suffered by the lower portion of the water by friction, in the steepening
of that side nearest to the place towards which motion is directed, and the final throwing over of the top in the same diree-
tion. A familiar instance, which repeats more exactly, thongh on a small scale, the conditions suggested by Mr. Hall’s
theory, is afforded when a viscous mass, such as resin or pitch, moves downward on a slightly inclined plane. If the neces-
sary conditions are given for the formation of waves, their steeper sides will always be on the lower end, or, on that which
would be nearest to the axis of a trough-like depression. It is obvious that this shape is the reverse of that of the waves
of plication which Mr. Rogers and others have observed, and on which even Mr. Hall’s theory is partially founded. This
theory, which was first proposed for a mountain range in which recent formations do not occur, and where, therefore,
not all of these objections could be raised, may, if somewhat modified, have its limited applicability for partially explaining
the mode of formation of certain mountain ranges; but numerous arguments appear to preclude its general applicability,
especially so in the case of the European Alps, and probably not less in the case of the Himalaya. It appears, too, to be
quite inadmissible in the case of the Rocky Mountains ; and as regards the Sierra Nevada, only a chain of the most arbi-
trary assumptions would be able to give it an apparent validity for the explanation of any features in the structure of that
range.

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84 RICHTHOFEN

NATURAL SYSTEM

Cretaceous period by thousands of feet. This is not only true for those ranges which,
like the Andes and Rocky Mountains, have been intensely volcanic, but also for such
as show a less immediate connection with eruptive activity in the voleanic era, as is
the case with the Pyrenees, the Alps and the Himalaya. The elevation which sue-
ceeded the Cretaceous period appears to have everywhere been slow, intermittent, and
partly retrograde, in the Eocene epoch, while its chief phase is traceable to the Miocene
and subsequent epochs, by the relative altitude above the level of the sea to which
sediments of different ages have been raised.

There can scarcely be any doubt that this acceleration of the changes of level,
and the extrusion of voleanic rocks, have had an intimate connection. But the mode
of this connection has by no means been ascertained, and appears to be quite an intri-
cate subject. It was formerly believed that the eruptive rocks were themselves the
upheaving agents, their intrusion and ejection having been the mechanical means of
the elevation of mountain ranges. But geological observation and argumentation have
conclusively demonstrated that such could not be the case, and made it probable that
both elevation and eruption are the most conspicuous symptoms of different agencies
which were dependent on one common cause. When that first theory had to be aban-
doned, another was put in its place, to the effect that the ejection of rocks must have
been attended by subsidence ; and, although we shall show in the following pages
that the rise of the crust greatly predominated in the vicinity of the eruptions, there
can be no doubt that subsidence was probably in all cases among their intricate effects.
But the cause generally ascribed to it is not at all capable of explaining it. The
assumption that eruption must be attended by subsidence is usually made on the ground,
that the removal of a certain volume of matter by ejection, from a deep-seated place,
would cause the formation of a vacuity, corresponding in extent to the volume
removed, and that consequently the overlying portions of the crust would settle down.
This assumption appeared to be corroborated by the observation that the region sur-
rounding a volcano subsides during its activity, while it rises in periods of rest. But
it is evident that the size of a current of lava is not proportionate to the settling of an
area of hundreds of square miles to the amount of several feet. If, moreover, the
views here advocated, regarding the cause and mode of eruptive activity, are correct,
that is, if the ejection of rocky matter is the effect of the increase of volume which it
undergoes on passing into the state of aqueous fusion, then the eruption is only the
discharge of the surplus matter which has no room within the space of the fissure ; no
vacuity could therefore be formed, and the commonly adopted cause of the subsidence
would not exist. There are, however, two other causes which would produce sub-
sidence, and have probably been acting in every case of massive and volcanic erup-
tions. The first of them is the contraction of the liquid mass in the conduits, by cool-
ing. Its amount must be considerable in proportion to the space of the fissure, but
small when compared with the volume of a mountain range on the surface. Its effects
will be local and abrupt. In the case of volcanoes, it is the probable cause of the
familiar phenomenon of the subsideuce of the bottom of craters, and of those less fre-
quent cases where whole portions of the cone are suddenly engulfed. As regards
extensive accumulations of rocks by massive eruptions, there are certain features of
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OF VOLCANIC ROCKS. 85
the configuration of their surface which cannot be more plausibly explained in any
other way. To these belong the sudden breaks in the continuity of their surface,
which consist sometimes in elongated and steep walls, thousands of feet in height, and
miles in length, or in crateriform or semi-circular basins, and in other more or less
abrupt depressions, which may be chiefly noticed where large regions are uniformly
covered with granite, porphyry or volcanic rocks. This cause will affect the mass of
the eruptive rock itself, but not perceptibly the surrounding country. Yet, this is
subsiding when a voleano is active, and it can be definitely proved in the case of the
andesitic ranges of Hungary, that during the epochs of the massive eruptions, the gen-
eral rise has been repeatedly interrupted bythe subsidence of the suroundings of the
theater of activity. To these changes applies probably the second of the causes alluded
to. Whenever a fissure is filled by liquid matter injected from below, its surroundings
must necessarily become heated, and, by their expansion, produce a slight increase of
the rise of the surface. This heat escapes, in the case of a volcano, chiefly in the epochs
of its activity, by the emission of laya, vapors and boiling water, and other phenom-
ena associated with voleanic action. Massive eruptions will have been attended by the
escape of heat on a much larger scale. They appear to have been often accompanied
by an extremely violent emission of hot water, as may be inferred from the great
accumulation of deposits of silica in some volcanic countries, or from the immense
overflows of extensive regions by voleanic mud: this occurs in Hungary and on the
western slope of the Sierra Nevada on so grand a scale as to almost exclude the possi-
bility of its having originated merely from volcanic action, especially as no volcano is
visible from which they could have escaped. These processes must of course have had
the effect of lowering the temperature of the masses surrounding the fissure to some
distance from it, and of producing subsidence during the eras of activity. Yet they
are not sufficient to explain the extent to which it has often taken place, though it had
in no ease, in the vicinity of the theaters of eruptive action, more effect than to reduce
locally the amount of elevation.

A few examples will suffice to show how vast are the changes of level which
have taken place since the commencement of the volcanic era, and to demonstrate their
connection with the other manifestations of vulcanism during the same era. An in-
structive instance is furnished by the country situated between the Pacific coast and
the Rocky Mountains. The labors of several distinguished geologists have made us
acquainted with some of the main features of its complicated structure ; but it is only
by the detailed examination of some of the most important portions, together with the
accurate determination of the age of several of the sedimentary and metamorphic forma-
tions, made by and under the direction of Professor Whitney, that the foundation has
been laid for an exact exploration of the entire western part of North America, which is
now proceeding with rapid steps, and promises to give important contributions towards
the solution of the questions discussed in this essay. There appears to have existed, as
we mentioned before, an ancient granitic era in that region. But it is as yet impos-
sible to recognize the relation of this ancient granite and the metamorphic rocks by
which it is accompanied to the ancient or to the present configuration of the surface.
In the western countries, those formations are concealed by the immense overlying

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86 RICHTHOFEN — NATURAL SYSTEM

accumulation of Paleozoic and Mesozoic sediments, and the subsequent denudation of
these, though grand in the extreme, has only been sufficient to expose to view the
ancient granite in some localities, among which we mentioned that on the Colorado
River. Other masses of ancient granite are visible at the surface in numerous places in
the Great Basin, but their exact relations still remain to be determined, while in the
Rocky Mountains they are one of the prominent features, partaking largely in their
structure.

The next period of interest, in regard to the occurrence of eruptive rocks, is that
of the deposition of Triassic and Liassic sediments in great aggregate thickness, which
were found by Whitney to extend from the Pacific coast far into the Great Basin. They
prove that all this country was then still submerged beneath the sea, while the Rocky
Mountains formed probably a broad belt elevated above it. It is in this period that
were ejected the quartzose porphyries of the County of Plumas, in northern California,
almost contemporaneously with those of the southern Alps. Whether this event, which
was probably not limited to the region mentioned, was attended by any changes in the
configuration of the surface, cannot yet be decided. After it, however, they must have
taken place on a grand scale, reminding one of the emergence of the central body of the
Alps from the sea after the deposition of the infra-Liassic limestones. The ejection of
granite found, it appears, almost the whole country embraced between the western
slope of the Sierra Nevada and the eastern slope of the Rocky Mountains lifted out of
the sea. But changes greater than those which had preceded them, appear to have
attended and followed those gigantic outbreaks. They were probably due, in a great
measure, to the intense metamorphism which was connected with them, and has com-
pletely changed the petrographical character of the preceding sedimentary deposits.
When the volcanic era commenced, which was probably in the Miocene epoch, all the
ancient formations of the Sierra Nevada were nearly turned upon their edges, and the
depressions of the surface in the Great Basin were filled with salt water. But the alti-
tude of the entire plateau above the level of the sea was probably insignificant at that
time compared with what it is now, as may be inferred from the fact, that rivers did then
flow on the present western slope of the Sierra Nevada, parallel to its crest, which they
could not have done if that slope had had its present inclination. We must imagine
that where the great mountain range raises now its lofty summits, a hilly country ex-
tended then, ascending slightly to the east. The first outbreaks of voleanic rocks found
those rivers still flowing in their beds, as is proved by the higher sediments in the old
river-channels, which consist of voleanic tufas. But great changes occurred after the first
commencement of the voleanic era, changes which contributed probably more towards
imparting tothe western portion of North America its present features than any which
had preceded them. The voleanic belt extending along the entire western coast of
America, had its greatest breadth between the coast of California and the Rocky Moun-
tains, and the eruptive activity has been violent over that vast region. It appears that
to that era is due the principal part of the elevation of the high table-lands. Such, at
least, is the case in their western portion. The crest of the Sierra Nevada must have
been elevated at a quicker rate than its western foot, as distinc. traces are left in the
gravel deposits that those ancient rivers which were flowing parallel to its crest have
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OF VOLCANIC ROCKS. 87

been gradually turned from their channels, overflowing their banks at several places in
succession, and taking a course down the slope, until an entirely new system of water-
courses was created, at right angles to the crest, the steep ravines and gorges of which are
now one of the characteristic features of the Sierra Nevada. It appears that the changes
of level since the inauguration of the voleanic era have also been progressing on a grand
scale in the eastern part of the Great Basin, where the eruptive activity had probably
even more gigantic proportions than in the west. The central portion of the Rocky
Mountains was raised, according to the estimate of Dana, about seven thousand feet since
the Cretaceous ; and its eastern part from one to two thousand feet, since the Miocene,
according to the observations of Hayden.

These figures render evident the great elevation which the mountain mass
between the Sacramento and the Missouri must have undergone since the commence-
ment of the volcanic era. ‘Twice in the history of that country, since the Triassic
period, may there be recognized an extraordinary intensity of all those changes which
we must ascribe to subterranean agencies. The first instance was in or about the
Jurassic epoch, when strata, which appear to have been quietly deposited during pre-
ceding eras, were elevated, and a gigantic intrusion and ejection of granitic masses was
attended by an intense and wide-spread metamorphism, by which disturbances and
plications were promoted; the second, in the volcanic era. It is probable that similar
phases to those mentioned will be recognized throughout the entire range of the Andes,
the geological structure of which appears to offer much similarity in different parts. To
the events of the volcanic era, chiefly, will have to be ascribed the connection of both
parts of the continent, though it may have been prepared by that preceding era of in-
tensified actions, which manifested itself in the ejection of the granite of the Sierré
Nevada, and appears to have left no less distinct traces in other portions of the range
of the Andes. We may still note a peculiar difference in the mode of the changes of
level if we proceed across the continent from west to east. It appears that the narrow
strip of land adjoining immediately the western coast has been subjected rather to
periodical oscillations than to any lasting changes of level, while the great elevation of
the mountain ranges and highlands must be chiefly ascribed to the circumstance that
all changes have acted there essentially in the same direction, producing the elevation
of extensive regions. This would explain why the western descent of the Andes has
been periodically increasing in steepness and the strip between them and the coast is of
little width. The latter appears to correspond to the boundary between an area of
elevation to the east and an area of subsidence to the west, one of which was especially
subjected to the manifestations of vulcanism, while in the other it has left no recogniz-
able signs. It is quite different on the eastern side of the range of the Andes, where,
in both parts of the continent, a slow rise has taken place, which has connected, during
the volcanic era, that mountain range with other ranges farther east, by those exten-
sive low-lands which are so important a feature as regards the extraordinary produc-
tivity of both parts of the continent.

Similar to these are the relations presented by the European continent, in regard
to which we will only mention a few prominent facts. During the porphyritic era and
the time immediately succeeding it, great changes of level had taken place on that

io)

Ss RICHTHOFEN —— NATURAL SYSTEM

continent. It appears that in Middle Germany the Triassic period designates the com-
mencement of an era of repose, while in the Alps, where the porphyritic outbursts took
place in the Triassic age, the Liassic strata still participated in the disturbances and
elevations, which partly attended the porphyritic era and partly succeeded it. There-
after followed another period distinguished by the comparatively small amount of dis-
turbances which took place in it. Subsidence appears to have prevailed in its first part,
while towards its end, during the second half of the Eocene epoch, that renewed rise
commenced which contributed so much towards imparting to that mountain range its
present configuration. In surveying the general features of the geology of Europe, it
will be found that the changes of the boundaries of the continent and the sea, as pro-
duced by rise and subsidence, have, in the period intermediate between the porphyritic
and the volcanic era, been inconsiderable in proportion to the length of time; and,
though grand in the totality of their results, they are probably far surpassed by those
which have been produced since propylite reopened the eruptive activity. None of
the prominent mountain ranges, it is true, have been called into existence since then,
Some of them, of little importance and consisting completely of volcanic rocks, have
been formed, and sedimentary formations have been folded quite extensively, so as to
form hilly regions ; but the main ranges had existed before, and only underwent an in-
crease in volume, though one of considerable magnitude, and the rate of elevation appears
to have been greatest with those mountain ranges, during the period indicated, which
have the greatest altitude at present. The amount of elevation which Eocene and
Miocene strata have experienced in the Alps, Pyrenees, and others of the prominent
mountain ranges justifies this conclusion, while the mode of occurrence of these forma-
tions in the Alps allows us to infer that their central portion was elevated at a more
rapid rate than either the northerly or the southerly part. It is worthy of note that
not alone in the Alps, but quite generally in Europe, the Jurassic and Cretaceous
strata have undergone only little more disturbances than those of Hocene age. In the
belts elevated during the voleanic era, all three formations participate ordinarily at
nearly equal rates in the structure of the low ranges of the hills stretching between
the main ranges and girting their foot. The conclusion appears therefore to be justi-
fied that in the volcanic belts, at least, the elevation of the Jurassic and Cretaceous
has chiefly taken place during the volcanic era.

There may be distinguished a twofold mode of elevation in the volcanic era: it
manifested itself either in a secular rise of continental areas, or in the elevation of
mountain ranges, as may be exemplified by referring to the difference in the mode of
elevation between the Andes and the countries adjoining them to the east.

The distinction is not so definite if we refer to the Huropean continent. Among
the regions elevated during the voleanic era may be chiefly noticed a broad belt ex-
tending from the Alps through the Turkish Peninsula, Asia Minor, Armenia, and
Persia to the Himalaya, and continuing less distinct westward to the Pyrenees. The
three main ranges have experienced the greatest amount of elevation within this belt.
The rest of it, which is distinguished by the folding of the Nummulitic strata over its
entire area, marks the region of the next greatest intensity of the elevating forces within
a much more extensive area of continental rise. The increase of altitude has been com-
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OF VOLCANIC ROCKS. 89

paratively inconsiderable over this area ; yet it is, in its totality, of greater import for
the configuration of the surface of the planet than the prodtiction of the mountainous
and hilly regions mentioned. For it had the effect of connecting regions which were
before separated by the sea, and of increasing thereby the area of continents in cer-
tain directions. It is well known that the sea, which in the beginning of the volcanic
era extended in the valley of the Danube to above Vienna, bathed both slopes of the
Carpathians, and reached eastward to the foot of the highlands of Central Asia, has
since then considerably retired, and that the secular rise of northern Africa, Arabia,
and Asia Minor has, also in these countries, enlarged the continental areas.

So much time has elapsed, in most countries, since the main phases of the vol-
canic era, that the influence of the events connected with them upon elevation and
subsidence has probably relaxed. It appears that portions of the areas of elevation
have suffered a periodical subsidence, not only since that time, but also during the
voleanic era. But its aggregate amount having been in most of them less than that of
elevation, the final effect manifests itself generally as a rise; though it appears that
in recent times some regions, chiefly those which are situated on the northern and
southern boundaries of the great area of continental elevation, are undergoing a more
marked subsidence. Even now, however, those countries chiefly are rising which
were distinguished by eruptive activity in the voleanic era.

These few instances will suffice to show how great have been the changes in the
configuration of some portions of the surface of the globe during the relatively short
geological time which has elapsed since the commencement of the volcanic era, and how
much superior they appear to have been to those which had been going on during
preceding periods of much longer duration. Those facts which are known in relation
to this subject, allow to infer, that the changes of level were proceeding at an acceler-
ated rate, during, and probably also in the time next preceding, the volcanic era, and
that the elevation was limited to certain belts of great extent which were, in the
inajority of cases, distinguished from neighboring regions by eruptive activity. But
this is not true for all cases, since there are some regions the elevation of which was
especially grand, and in which very few rocks, or none at all, were ejected. In ex-
amining into the causes of the connection of elevation with the other events of the
voleanic era, we must therefore keep in view the distinction of those cases where it was
attended, and those where it was not attended, by eruptive activity.

As regards the first cases, we may refer for their explanation to our previous
theoretical considerations of the causes and effects of the formation of fractures. If
the imerease of volume by the slow and perfect crystallization of viscous masses be-
low the crust was the cause of the rending of fissures, a slow rise must have preceded
this process. That this was so on the European continent, is verified by the fact, that
the Eocene strata had been elevated quite considerably when the first of those sedi-
ments, which can be proved to have been contemporaneous with eruptive activity,
were deposited, If then the relief from pressure caused the extensive crystallization,
at the depth to which the fissures reached, of masses which had been held before in a
viscous condition by the pressure itself, an accelerated rise would be the effect ; and if

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90 RICHTHOFEN — NATURAL SYSTEM

the repeated rupturing of the crust caused a repetition of this process, in the way in-
dicated before, epochs of ‘an accelerated rise would alternate with those of a retarded
elevation. This theory will explain why a very considerable rise, and of greater ex-
tent than that which had preceded, attended the phenomena of the basaltic epoch.

Although we must consider this subterranean process as the chief cause of con-
tinental elevations, so far as they attended the phenomena of eruptive activity, it was
certainly not their only cause. There can be no doubt that the process of metamor-
phism must have promoted those changes. As far as we understand the conditions
required for its production, it appears that they could not be created by any other
known processes in a degree similar to that in which we must necessarily suppose them
to have attended the ejection of rocks, whether its mode be that of massive eruption
or of voleanic activity. Even on the surface there are no places where we can observe,
at the present time, metamorphic processes of greater intensity than near the orifices
of voleanoes, or in solfataras, or at the theaters of other processes attending or succeed-
ing volcanic activity ; nor is there at any other places more evidence afforded of subter-
ranean metamorphic processes. The latter will exceed those visible on the surface in
nearly equal proportion with the increase of temperature and pressure in depth ;
while we may conclude theoretically, that the neighborhood of fissures filled with
heated substances from below would not only have a higher temperature than
other portions of the crust, but also allow of a comparatively free circulation of water,
on account of its shattered condition. In respect to metamorphic action, as in all
other respects, the grandeur of the phenomena of massive eruptions would make us
presuppose that they would have given rise to a far more extensive and intense meta-
morphism than the insignificant processes of volcanic activity. The occurrence of
ancient eruptive rocks in belts of highly metamorphosed sediments removes it almost
beyond doubt that they are connected in the relation of cause and effect. It is assumed
that metamorphic processes, on account of the entering of water into the composition
of rocks, and their crystallization, must be attended by an increase of volume, and
will have the mechanical effect of elevating the surface.

We may thus obtain a few hints as to the causes of the changes of level which
those portions of the surface of the globe which were the theater of eruptive activity
in the voleanic era, have undergone since its inauguration. The agencies to which
they were due must be even more intricate than the motions of the crust, ‘because the
movement in one direction may be the resultant of several forces working in opposite
directions. Increase of volume by crystallization below the crust, in those regions
where the formation of fractures relieved the masses from pressure, is probably
the most potent of these agencies. Metamorphism would work in the same direction,
while loss of heat would cause subsidence. All these agencies have their first and
common cause in the formation of fractures by an upward tension, and all other
phenomena of vulcanism are the immediate or mediate effects of that same cause,
which itself results from the gradual cooling of the globe.

There remain those cases to be considered where some of the most prominent
mountain ranges, in the structure of which volcanic rocks take no part, were elevated
during the voleanic era, at a much greater ratio even than those in which these rocks
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OF VOLCANIC ROCKS. 9]

occur. The Alps and the Himalaya are the most striking instances of this kind. The
first of these, and probably, too, the other, had, as we mentioned before, undergone a
considerable elevation in and after the porphyritic era, but have probably changed their
altitude very little in the next following periods. The acceleration of their elevation
during the voleanie era appears, however, to have greatly exceeded that of the Andes.
The coincidence in time of these events, and of all the phenomena characteristic of the
volcanic era, justifies the supposition that they are connected in their origin.

’ If we contemplate, in its relations to the volcanic era, the great elevated belt of
which the Alps and Himalaya form the axis, we find it to consist of three parallel
zones. ‘The central one comprises those two mountain ranges and the broad moun-
tainous country which connects them, and which owes its configuration chiefly to the
events of the volcanic era. The Alps and Himalaya are free from volcanic rocks. This
is also true as regards the central portions of those chains which branch off from the
Alps in a southeasterly direction, and of those (as far as their geology has been ex-
plored) which extend westerly from the Himalaya. The farther one proceeds in the
Turkish peninsula towards the east, the more frequent are the monuments of eruptive
activity of the volcanic era; they are known in Servia and Bulgaria, in the north, and
in Macedonia, Thracia and Epirus, in the south. They increase in similar ratio as
one proceeds westerly from the Himalaya, through the highlands of Armenia to Asia
Minor. If it is considered that active volcanoes are particularly numerous in those
regions where the elongated portions of two different continents have, as it were, a
tendency to connection, the analogy with the mode of distribution of volcanic rocks
in the regions mentioned is conspicuous. The system of the Alps with its southeastern,
and that of the Himalaya with its western branches, were disconnected before the Ter-
tiary period, and were united into one mountainous belt during the same. The eruptive
activity contemporaneous with this slow process was remote from the main axis where
this had existed as an elevated range before, and approached it more and more, from
both sides, increasing, at the same time, in intensity, until it culminated in that
region where the connection of both systems was effected. Massive eruptions and
voleanic activity have ceased completely in this central zone.

North of this, is another zone, which was distinguished in its entire length by
the intensity of eruptive action in the volcanic era. It stretches from Central Asia by
the Caspian Sea, the Caucasus, the Crimea, the Carpathians and, in branches, through
central Germany to central France. In the European portion of this zone, there are
only recognizable in thermal and mineral springs the last feeble remnants of former
volcanic activity, while they are somewhat more energetic in that portion which is
situated on the Asiatic continent. A third zone, not less distinguished for the in-
tensity of eruptive action in the volcanic era, accompanies the main axis to the south.
It traverses India, and, in the Bay of Bengal, is connected with the volcanic belts of the
Indian Archipelago, while it continues to the west through Arabia, Syria, Palestine, to
the Mediterranean, where it comprises the active voleanoes of the Grecian Archipelago
and Italy, and is connected by Sardinia and southern Spain with the Azores. Volcanic
activity still continues in this belt, but the period of the massive eruptions has passed

long ago.
(129)

Ne)
bo

RICHTHOFEN — NATURAL SYSTEM

The entire belt, composed of these three zones, has to be considered as one great
area of elevation, characterized, at the same time, over its greater portion, by the rem-
nants of the eruptive activity of the volcanic era. No portion of it, however, has, as
we mentioned before, experienced an amount of additional elevation during that period
equal to that of those mountain ranges which existed before it, and among which the
Alps and Himalaya are the most prominent. In these cases there cannot have been any
connection between the elevation and the ejection of the rocks to the surface, because
these do not occur. But this does not preclude the connection between the elevation
and those agencies which are the causes of eruption. It has been often demon-
strated that the changes of level must have been accompanied by the formation of
such fractures as would be closed next to the surface, and, though allowing of an in-
trusion of plastic matter from below, would prevent its extrusion to the surface. It is
probable that fissures of this kind may have been chiefly formed where high mountain
chains are composed of metamorphic rocks, since they frequently exhibit vertical disloca-
tions, partly parallel to their axis, and partly at right angles to it, by which the strata
on one side have been moved thousands of feet above the other, and which, notwith-
standing, did not give vent to the ejection of rocks. These faulting fissures appear to
form frequently the lateral boundaries of mountain ranges and the limits of the mani-
festation of vulcanism, and both these circumstances allow us to infer that they extend
downward to great depth. Though one of the main features in the geological
structure of the Alps, they have hitherto been little examined. But itis probable that
their formation was coincident in time with the main phase of elevation, that is,
with the volcanic era. if, then, in that part of the crust over which the Alps are ele-
vated, such fractures were formed as did not open on the surface, then all the agen-
cies below would codperate towards elevation alone. The expansive force produced
by aqueous fusion would find no vent for the discharge of the masses by which the
volume had increased, nor would there be any opportunity given for the escape of
heat by hot water and other means; while, on the other hand, the masses in depth
would be relieved from pressure by the formation of this kind of fractures in the same
way as by those which would allow the passage of liquid matter to the surface. In the
case of the latter, an immense amount of force is spent in other modes of action, while
in the first case it could be applied almost exclusively to elevation. Besides this direct
action, however, we must also keep in view, that in those regions where fissures would
not be open at the surface, the conditions required for metamorphic action would be
given on a particularly grand scale. Gases and water would not reach the surface, but
be employed in depth in promoting metamorphic action over vast regions, and imcreas-
ing the rate” of elevation. It may be by processes of this kind that already in ancient
times, contemporaneously with an accelerated elevation, were formed those granitic
wedges surrounded by broad belts of foliated metamorphic rocks, which are peculiar
to certain mountain ranges, particularly to those which also in the voleanic era have not
been the theater of eruptive activity. The intrusion, from below, of heated masses into
fissures closed towards the surface, would give all the conditions required for the exhibi-
tion, on a grand seale, of those processes of hydropyric metamorphism which Daubrée
has rendered probable by his experiments and theoretical deductions.

(150)

OF VOLCANIC ROCKS. 93

Leaving aside these theoretical considerations, and comparing only the differ-
ent facts mentioned in regard to the changes of level which have taken place on the
surface of the globe since the inauguration of the volcanic era, we are forced to the
conclusion that they must be intimately connected, as regards their causes, with the
other phenomena by which this era was distinguished from preceding periods. Eleva-
tion and eruptive activity, even when locally not quite coincident, are codrdinate effects
of the cooling of the globe ; but while the one isits immediate effect, the other results
from it only by the concurrence of other agencies, which by themselves alone would
have been incapable of producing results of such magnitude. As there are distinct
phases in the history of eruptive action, dependent on and marking the evolution of
the globe, so we may recognize different stages in the mode of manifestation of the
elevating forces. In those elevations which took place during the volcanic era, there
are certain peculiar features evident, among which may be mentioned : the increase in
altitude of those mountain ranges included in the volcanic belts ; the union into one
main chain of several smaller ridges, the axes of which are situated on a line; the
lateral connection of parallel ranges into highlands; the oblique connection of the
ends of main chains, the axes of which are situated on parallel lines, but are remote
from each other longitudinally, by broad belts of mountainous regions (of this
character is the connection of the Alps and the Himalaya); the elevation, final-
ly, above the level of the sea, of large areas which had been submerged before,
whereby distant mountain ranges were connected by extensive lowlands, the size
of the continents increased, and their outlines rendered more uniform. The mode
of these great changes of the configuration of the surface of the globe was probably
a repetition on a large scale of similar changes which had been going on in former
eras of eruptive activity. It is not our object here to develop them. But throughout
the course of these changes there appears to be conspicuous a tendency to connect, in
certain directions, what was before disconnected, to increase in size and render more
definite certain areas of elevation, and to separate them from other areas of subsidence,
which appear to have been likewise increasing in extent. We may even recognize an
increase of this tendency during the volcanic era, as the phenomena connected with
the basaltic epoch have been far more widely and generally distributed over the areas of
volcanic belts than those of the preceding (propylitic and andesitic) epochs. There must
be certain general laws which regulate the directions in which the connection of discon-
nected parts takes place. Their knowledge is still very limited. The admirable manner
in which Dana has laid out the grand outlines of the arrangement of the islands of the Pa-
cific, has indicated the way by which we may look for the solution of this problem. Ifthe
structure of the crystallized crust of the globe gives the most probable cause of the
definite directions which may be recognized in the outlines of continents and chains of
islands, it should be borne in mind, that the direction of structural planes, if they
exist, must vary gradually in depth, in the same ratio with the chemical composition,
and that they may at the depth of andesitic compounds be different from what they
would be in the granitic depths. The singular way in which the connections by ele-
vation are effected, and which has been already partly pointed out by Dana, appears to

Q (151)

94 RICHTHOFEN—NATURAL SYSTEM OF VOLCANIC ROCKS.

find only an explanation when such changes of the direction of the structural planes
are assumed to take place in depth.

The tendency to simplify beyond their present conditions the general outlines
of the configuration of the surface of the globe, continues probably without intermis-
sion, though an era of comparative repose has followed the violent exertions of vulean-
ism of the voleanic era. The great backbone of the Eastern continent, comprising the
Alps and the Himalaya, and the second great belt which comprises the Andes, and,
more properly, encircles the whole Pacific Ocean, are at the present epoch the main
features in the orography of the globe. Both have derived their prominent position
from the events of the volcanic era. The conclusions at which we have arrived in
these pages have been drawn chiefly from observations made in regions of prominent
interest in either belt, and it is for this reason especially that we believe that at least
some of them will be found to be susceptible of general application.

T am fully aware how imperfect is this attempt to develop, in the correlations of
the volcanic rocks, the principles of their natural system. This system should be, as
we remarked before, not a classification of objects by certain conspicuous properties,
but a classification of objects by their definite mutual relations. We have not to make
the divisions, but to find them. We shall be able to do this with greater or less per-
fection, in the same measure as those relations are studied and firmly established.
Much is needed to arrive at this end. It can only be-achieved after evidence has been
accumulated by the combined and harmonious labors, in different countries, of geolo-
gists in the field as well as in the laboratory. The range over which both modes of
observation extend from year to year, and the field on which their results meet, is in-
creasing in a surprising degree. Yet eruptive rocks form ordinarily an object of less
exact observation than those of sedimentary origin; and it cannot be denied that har-
monious observation, by which alone can be established the principles of comparative
petrology, is rendered nearly impossible, as long as no uniform system of nomenclature
is applied. These pages are the result of an attempt to add one more to the many
contributions towards establishing the foundation of both, and to show the value which
the study of eruptive rocks and of those mutual relations which comprise their natural
system promises to have for approaching the solution of some of the highest problems
of geological science.

TABLE OF CONTENTS.

PAGE
HINTERODUCTORNC ETL: SETS Herd cols See MER a ene ee Ne ree te nib teem rs il
Tony NATORAD SYSTEM) OF VOLCANIC: IROCGKS)..2...5..-.0.-6----.--+.++-0 foereres 5
Oxpury HURST) RAW OLEH sere. crcleiersierss eters Ate nO Oke ee CI Renee OE Tee ae 12
ORDER SECOND: “TRACHYTH),..5 052-2422. eee <1 ee Sree ae aS tare ne he ei Mee F 17
ENDER BE LHTRD my RO PIII ME ey scarr sie ssts ein oe so la ei Pare eae tS os aoe lap lal see 20
Orper FourtH: ANDESITE............... Rohe ere SEA DHE eee SRO ERS UCB OS ante COI 25
ORDERBHURTHE PDA SAINTS er rrarare eri ane vocidir< stots eet ede oe re eee Saunier lc
CoRRELATION OF THE FivE ORDERS OF VOLCANIC ROCKS.............+......-.-- eee 28
Laws Relative to the Age of Massive Eruptions............ si efeloye is ays ase areheye 29
Laws regarding the Mutual Relations of Massive Eruptions and Voleanie Activity 32
RELATION oF VoLcanic Rocks to ANcreENT Eruptive RocKS..............-.-eeceeeee 35
Chmelennnvare Age anda Recline reer Soe eee ono ee ad & Soe Sek een a ees ihe
Correintion0f Age and, ‘Composuiwre <2... onc deste yxeci eee ciet ween ean 41
Correlation of Eruptive Rocks in regard to their Geographical Distribution... ... 45
On PrriegORIGUNG OL MV OLGANTO SOCKS aay es iicy sce. 1c its oie wae tae Serae austere ars ites hares 2 = 46
is OCOD LTC: OEY OUT 107 fe eR <r a 47
MO MeGiINy AMV OLA ACEO crane tiva a 2 as wats BT OS lowes ee Ge Lee 60
3. Other Theories regarding the Origin of Volcanic Rocks..... > MOOR eee 6 3de 68

RELATION OF THE DISTRIBUTION OF VoLcANIc Rocks TO THE CONFIGURATION OF THE SURFACE
Ors Gri Cane! F oe couaeceeedsoc ROBO COL ODA och CON een oo tp Bee Ons CSTE he:

tat

ran.

" e -
a 7 <
ae et Sh

MEMOTRS

OF THE

\ . :
Gatirornta Acapemy or ScrENCES

Volume I]

SAN FRANCISCO
PUBLISHED BY THE ACADEMY

1888 — 1896

Il.

Ill.

CONTENTS OF VOLUME II.

PAGE
ON THE ANATOMY OF SUTROA ROSTRATA, A NEW ANNELID OF THE SUB-FAMILY LUM-
BRICULINA. By Gustav Hisen

Meee vetatwrasea et on cel arevarose) BE oNa(APn aierage aie atey si nia¥ay stacese lavampemiees-Vev co's ehcraieteare.sa alata l
(Published January, 1888.)
ON SOME ANCIENT SCULPTURES FROM THE PACIFIC SLOPE OF GUATEMALA. By Gustav
JON reat Haale CMA BER ACO UG COR BOD DIC Cc OCI COCOA TOT TORT ACCOR ereC Orn Or Hartree ene rama 9
(Published July, 1888.)
ONGOAGIRORNTASHUDREIMED A, (By Gustan Misen «=<. costes sles coc eisieeele\sivcissele c vleeoc/ aoa ceos ses 21
(Published February, 1894.)
PAGLBIC COAST OETIGO CH AIWAN vie SBynGustan: Hager crt) se aie ieele -(eiais a/</areteisctelsis se be see sees ola 63
(Published March, 1895.)
PAOTETOIOOAST ORIGOCH AIDA Ghee ByiGustan! Hise. .e cleieeislciernieieeiciciers + ole eiei</e/c\scte a.cicieieeive viz cisia'e 123

(Published February, 1896.)

INDEX OF SCIENTIFIC NAMES.......

'
aig!

25

GR Fo bell) a Lh LS)

Oa

we

lf

MU

(feo ew See ye Ween S yee vs

Gustav Eisen.del

I. On toe ANATOMY OF SutrRoa RosTRATA, A NEW ANNELID OF THE FAMILY OF
LUMBRICULINA.

By Gustav EIsEn.

The oligochetous annelid, which is the subject of this monograph, was first
found by me some ten years ago in the small body of water known as Mountain
Lake, situated close to the Marine Hospital, on the Reservation west of San
Francisco, California. During a recent visit to the same locality, in May, 1887,
I found the same worm in exactly the same part of the lake, but, as before,
in very few specimens. Perceiving the necessity of creating a new genus for this
oligochete, I have done myself the honor to dedicate the same to Mr. Adolph
Sutro, whose magnificent public scientific library, marine aquarium and zodlogical
station on the Pacifie Coast will in the near future become one of the most promi-

nent and valuable institutions of learning in this country.
OLIGOCHALTA.
Family. Lumpbriculide.
A. Sub-family Trichoprittva Vejd.

No solitary albuminous gland opening on the ventral side of the body.
B. Sub-family Lumpricocina Vejd.

A solitary albuminous gland opening on the ventral side of one of the ante-

rior segments. Of this sub-family three genera are now known:
1. Lumbriculus Grule

Four pairs of seminal receptacles in segments 8, 10, ll and 12. A solitary
albuminous gland in the ninth segment. The free secondary perigastric vessels
begin in the thirteenth segment, and are all pulsating. The preseptal secondary
vessels not branching, the postseptal one branching. Spines not distinctly forked.*

Cephalic lobe not filiform.

" Vejdov-ky, System, ete., der Olig chaeten, Prague, 1554, pag. 52.

Mem. Von. I. 1. January, 1888.

3 CALIFORNIA ACADEMY OF SCIENCES.
2. Rhynchelmis Hoffin.

One pair receptacles in the eighth segment. A solitary albuminous gland
in the ninth segment. The postseptal secondary vessels are simple, the preseptal

ones are feathered and branched. Spines are entire. Cephalic lobe filiform.*
3. Sutroa n. gen.

Seminal receptacles consist of several pairs of lobes, entire or forked, which
all open in the so-called albuminous gland in the eighth segment. A solitary
albuminous gland in the eighth segment. Preseptal and interseptal secondary dor-
sal vessel are branching and feathered. Postseptal vessels are gastric, not feathered
nor branching. Spines simple, not forked. Cephalic lobe filiform.

It will thus be seen that Sutroa is nearly allied to the two other genera of
the sub-family, the most prominent characteristic being the concentration of the
receptacles and their opening into the albuminous gland. But the vascular system
also shows some great peculiarities, which will soon be fully described.

As only one species is known of the genus Swtroa, the generic and specific

characteristics will here necessarily be considered together.
Sutroa rostrata n. gen. et sp

Habitat—l\ have found this species only in Mountain Lake, near the Marine
Hospital. west of San Francisco, California. It occurs here only on the north
shore, Just at the mouth of a small spring, which empties into the lake. It
lives in the richest mud, between the roots of aquatic plants, probably with the tail
vibrating in the water. When captured and set free in the water, the worms swim
rapidly through a peculiar wavy motion of the body, showing to advantage its
really magnificent iridescent colors, found to the same degree in no other fresh
water annelide. The color of the body is very similar to that of Rhynchelmis limo-
sella, a lively fleshy ved.

The worm appears to be quite rare, as 1 did not succeed in getting over a
dozen worms during each excursion. It also seems quite restricted to a certain
part of the lake, where it lives in the rich mud at the margin of the bank. On
the south, east or west ends of the lake I have looked for it in vain.

The egg capsule or cocoons of Sutroa are in size nearly similar to those of
Rhynchelmis, but in shape different. They can nearest be compared to the hanging
nests of some birds—small, bag-shaped bodies, with a flat, somewhat concave top—

suspended by a long semicircular membrane. Figure 15 gives an enlarged repre-

*Vejdovsky, 1. c, pag. 52.

ANATOMY OF SUTROA ROSTRATA. 4

sentation of the same. I generally found three eggs or young worms in the
sapsule.

The worms appear fully developed in July and August, at which time I
found mature specimens. In May only immature ones were found, but as some
of those were quite small it is likely that the worms remain mature for a long

period, and that perhaps the proper breeding season is during the winter months.

In size and form Sutroa rostrata much resembles Rhynchelmis limosella, Hoflin.
Mature specimen reach a length of three inches and a width of one-eighth of an
inch, but generally are considerably smaller. The form is also similar to that of
the above worm, The body is round or quadilateral; in many specimens the sides
are even concave (Fig. 3). The posterior part of the body is again very much flat-
tened out, quite transparent, and so brittle that care is necessary to yet any entire

specimen.

The anterior part of the worm, or the cephalic lobe proper, is elongated and
filiform, its length exceeding the width of the body. This characteristic is
also found in Rhynchelmis limosella, and indeed so similar are the two worms exte-
riorly that one may easily be mistaken for the other. Fig. 1 represents a Sutroa,
nat. size; Fig. 2, the anterior portion of the body; and Fig. 3, a transverse section

of a quadilateral specimen.

Vascular System.—The dorsal vessel (Figs. 4 and 5, v. d.) is pulsating. The
ventral vessel (Wig. 4, v. v.) is not pulsating. In the figures the pulsating vessel is
represented, red, the non-pulsating one as blue. The ventral and dorsal primary
vessels are connected in the cephalic lobe. where at the apex one vessel connects
directly with the other (Fig. 4, I). Between this cephalic plexus and segment
XIII no direct connection exists between the two primary vessels. But from
segment XIII toward the posterior end of the body, we find in every one
a secondary gastric vessel which undoubtedly connects the dorsal and ventral pri-

mary trunks.

The dorsal primary vessel (Figs. 4 and 5, d. v.) is entire, not forked, as is
the case in the other genera of Lumbriculina. In every segment it emits secondary
vessels, which are of two kinds, gastric and perigastric. The perigastric vessels
occur in all the segments of the body; the gastric vessels only in the posterior,

beginning with segment VIII.

In each of the six anterior segments we find only one pair of perigastrie ves-

sels, but each vessel shows a distinet forking, less so in segment I, but more dis-

4 CALIFORNIA ACADEMY OF SCIENCES.

tinct in the others. In segment VI the forking is so deep that it nearly
divides the vessel in two (Fig. 4, I to VI). In segment VII the forking is
perfect, and from that toward the end of the body we find in every segment

two distinct pairs of perigastric vessels. Each of these vessels again are forked at

the apex, and feathered—that is, besides the forking at the apex, it emits two pairs
of side branches. All these secondary perigastric vessels end blindly in the peri-

gastric cavity and do not connect directly with the ventral vessel (Fig. 5).

The ventral vessel is not pulsating. It is forked in segment VIII, the
forks again uniting with the dorsal vessel in the cephalic lobe (Fig. 4). A similar
forking of the ventral vessel is known in the other genera of Lumbriculide. The
two forks of the dorsal vessel are again connected by secondary perigastric vessels
(Fig. 4, v. pr. v.), which, however, in no way connect with the dorsal vessel or its
secondary perigastric vessels. In hynchelmis limosella* as well as in Phreatothrix +
the contrary takes place. Here the secondary ventral vessels connect the ventral
forks with the dorsal pulsating vessel. My observations on the transparent Svtroa
satisfy me positively that my above description is correct, and that the dorsal and
veutral vessels are not united in those segments. The blood in all the vessels is

lively yellowish-red.

The alimentary canal is extremely simple, consisting of a simple duct. In the
twelve anterior segments this duct is narrow and quite pellucid, but in the thir-
teenth segment it is considerably enlarged and continues so toward the posterior

part of the body. In the same segment we first meet with gastric vessels.

The nervous system resembles that of Lhynchelmis. The two ventral nerve
cords are, as in Lelipidrilus, connected by numerous anastomosing commissures.
The cephalic ganglion is rather long and narrow. No lateral nerves project from the

ventral nerve cord.

Generative System.—Vhe sexual organs are of two kinds—generative and recep-

tive. The generative include:
Testes, ovaries aud albuminiferous gland.

The receptive are: Efferent ducts, with atriwn and seminal vesicle; oviducts and
receptacles,

“Vespovsky: Anatomische studien. Rhynchelmis, Zeitsch, f. w. Zool. Ba. XXVII, Taf. XXI, Fig. 1.
+The sane. Ueber Phrcatothrix. Same, Bd. XXVII, Taf. XXXIX, Fig. 2.

ANATOMY OF SUTROA ROSTRATA. 5

The testes consist of one pair, situated in segment IX, and attached to
the dissepiment between segments VII] and IX. They present small bodies

of rather irregular shape (Fig. 14, test.).

The ovaries also consist of one pair, situated in segment X, and attached to
the dissepiment between segments IX and X.+ They are somewhat smaller than
the testes, and of a more regular form (Fig. 14, ov.).

The oviducts consist of one pair of small cup or funnel-shaped organs (Fig.
14, ovd., and Fig. 11). The exterior porus is on the ventral side of the body be-
tween segments XI and XI]. The interior funnel-shaped part is extremely
delicate and translucent. I found the eggs quite frequently in the act of passing
out. The oviduct of Sutroa differs from that of ARhynchelmis, in being somewhat

more elongated, with a longer and narrower neck.

The efferent ducts are four, and their funnel-shaped interior openings are
all found in segment XI, freely projecting from the dissepiment between segments
X and XI (Fig. 17). These efferent ducts are extremely long, extending through
some twenty segments, or from XI to XXX or XXXI. They here enter
the atrium, which is similarly elongated, extending from segment X to XXXI.
The exterior porus of the atrium is situated very near the center of the
ventral side of the line between segments X and XI. This enormous devel-
opment of the efferent ducts and atrium was previously only found in

Ocnerodrilus.*

The efferent funnels are comparatively small (Figs. 6, 7 and 14).

*I will here take occasion to correct an error in my former description of this latter genus, for which see
“On the Anato ny of Ocnerodrilus. Upsala 1878." The organs which I have there describe] as seminal receptacles are
undoubtedly nothing but the atrium. During a visit to Central America, I found four new species of Ocnerodrilus, and
a cursory micruscopic inve tigation showed me immediately thit s-minal receptacles existed in several pairs in some of
the anterior segments, which makes it evident that the large bodies which open in the same porus as the efferent
ducts must be considered as atrium. In the Californian species which I described as Ocnerodrilus occidentalis, these
small seminal receptacles were evidently overlooked. Professor Fr. Vejdovsky, in his admirable work, ‘‘System der
Oligochzten,” page 144, takes the same view of these organs, and refers to them correctly as the atrium. I now beg to
append a correct diagnosis of the genus Ocnerodrilus: Dorsal vessel weakly pulsating, in segments IX and [X;
Jurnished with two pairs of strongly pulsating h-arls. In the eighth segment it emits two side branches which continue
toward the cepha ic lobe. VENTRAL VESSEL ts nut forked, but continucs u.divided to the bucealic s gment. The sccondary
vessels are of two kinds, GAsTRIc and PERIGASTRIC, The perigastric oxues only connect with the ventral vessel. The «ferent
ducts are not united with the alrium, but bith open in the same porus. SEMINAL RECEPTACLES occur in puirs in several of
the anterior segments. TEstES—Two pairs in ninth and tenth segments. T'wo ovipucrs in fourteenth segment. Ovariss—

One pair in twel/th segment. The genus Ocnerodrilus constitutes undoubtedly a distinct fam'ly.

6 CALIFORNIA ACADEMY OF SCIENCES.

The exterior porus of the atrium is in Sutroa surrounded by strong muscu-
lar swellings, and the atrium itself forms a kind of penis, which probably is to some
degree projective. The exterior surface of the atrium is nowhere covered with
clands, as is the case in Rhynchelmis, but consists of a long, narrow duct of a

uniform width (Fig. 6, atr., and Fig. 14, atr.). No basal glands.

The seminal vesicle consists of an enormous sac-like body, entirely filling
the twenty segments occupied by the atrium and the efferent ducts. 1 did not suc-

ceed in finding the placé at which it was attached to the atrium.

The most characteristic interior organ, however, is the solitary body, for
which I here adopt the name given by Vejdovsky, glandula albuminifera or albumi-
nous gland. Iam, however, by no means fully satisfied of the functions of this
organ, and its structure seems somewhat different from that of the albuminous
glands of Rhynchelmis and Lumbriculus. The solitary albuminous gland of Sutroa
opens in the center of the ventral part of the eighth segment (Fig. 14, gl. alb., and
Fig. 10). It has the shape of a globulous body, connecting by a narrow neck
with the body wall and opening through an external large porus. But the
histological construction of this gland is quite different from the gland described

by Vejdovsky in Lumbriculus* and in Rhynchelmis.+

In the latter two genera this organ is distinctly glandular, but in Suéroa it is
of nearly the same construction as the receptacles, being covered by smooth epithe-
lium, under which are found numerous long and narrow cells (Fig. 10). I have,
however, no reason to doubt but that these organs are analogous in all the three

genera.

The receptacles in Sutroa consist of three pairs. In Rhynchelmis and Lumbricu-
lus we find the receptacles open each in a separate porus; in Sutroa, however, there
exists the great anomaly that all the six seminal receptacles open into the narrow
neck of the above described solitary gland (Fig. 10, re. sem.). These receptacles
consist of long narrow sacks, containing fully developed spermatozoa (lig. 8).
Generally there are three pairs of receptacles, but in some specimens I found three
receptacles on one side and only two on the other. Sometimes the interior end of
a receptacle is forked; generally, however, they are entire. That these elongated

bodies are real receptacles is proved by their being full of spermatozoa fully devel-

*Vejdovsky, System Cer Oligochwten. Taf. XII, 16. g. alb.

{Same. An. stud. RKhynchelwis. ‘Taf, XXIII, Fig. 17.

ANATOMY OF SUTROA ROSTRATA. T
oped. In the albuminous gland, however, I have searched in vain for any, though

I must confess the number of specimens at my disposal has been very small.

The segmental organs are found in all the segments posterior to XII. They
consist of long slender organs, the body of which is destitute of any glandular
appendices or cells similar to those found in Rhynchelmis. The interior opening of
the organ is surrounded by brown glands, behind which there is a larger head-like

enlargement of similar structure to the duct proper (Fig. 12).

The spines are eight in every segment and occur in four pairs. They are
J
slightly S-shaped, entire, not forked (Fig. 13).

EXPLANATION OF THE FIGURES.
Puate I.

Fig. 1. Sutroa rostrata, natural size.

Fig. 2. Front part of the worm, with the cephalic lobe, showing its filiform
elongation,

Fig. 3. Transverse section of the body of a quadrilateral specimen.

Fig. 4. The front part of the worm, showing the vascular or circulatory
system. The dorsal vessel is represented red; the ventral
vessel, blue.

d. pr. v.: dorsal perigastric vessel.
v. pr. v.: ventral perigastric vessel.

The twelfth to fourteenth segments, showing the vascular system

Or

Fig.
in those segments.

ac.: alimentary canal.

dv.: dorsal vessel.

d. pr. v.: dorsal perigastric vessel.

d. gr. v.: dorsal gastric vessel.
Fig. 6. The efferent ducts and atrium.

ps.: penis.

eff. d.: efferent ducts.

atr.: atrium.

Fig.

SI

Efferent funnel more magnified and seen in different aspect.

CALIFORNIA ACADEMY OF SCIENCES.

[o,0)

Prats II.

Fig. 8. Ventral nerve trunk and cephalic ganglion.
Kig. 9. Seminal receptacles and albuminous gland.
re. sem.: seminal receptacles.
gl. alb.: albuminous gland.
Fig. 10. Interior extremity of one of the receptacles, showing the sperma-
tozoa accumulated in the same.
Fig. 11. One of the oviducts, with eggs in the act of descending.
Fig. 12. The interior extremity of one of the segmental organs.
Fig. 13. One pair of spines.
Fig. 14. Interior view of the ventral part of the worm.
gl. alb.: albuminous gland.
re. sem.: seminal receptacles.
test.: testes.
OV.: ovaries.
ovd.: oviduets.
atr.: atrium.

eff. d.: efferent ducts.

Fig. 15. One of the egg capsules or cocoons. In the interior are seen three

young worms, magnified about five times.

The Roman numerals everywhere indicate the segments. In numbering the
same I have everywhere in this paper referred to the first segment as segment
I, thus following Vejdovsky and others. In former papers I have always referred
to the first setigerous segment as segment 1, and believe yet this way to be the least
confusing. But as uniformity in numerating the segments is highly desirable,
and as every investigator seems to have his pet way, leading to endless misunder-
standing, I shall for the future accept the system used in this paper, hoping all

other investigators will do the same.

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On some Ancient Scutprures From THE Paciric Store or GUATEMALA.
By Gustav EIsen.

During my visit to Guatemala, Honduras and Salvador, in 1882, I made the
archeological remains of the Mayas and the Aztees in those countries my particu-
lar study. These researches were embodied in a paper presented to the Smithso-
nian Institute in 1883. This paper treated principally of the ruins of Copsn and
Quirigua, in Honduras and Guatemala. Through an unfortunate mishap a portion
of this paper was lost in transferring from the Smithsonian Institute to the
Bureau of Ethnology in Washington, D. C. This delayed the printing of the
same four years. In the meantime Mr. Alfred Maudslay, of London, carried on
extensive excavations in both the above places. His researches being so much
more extensive than my own, and now soon to be published, will in a measure
supersede them, and make their publication less desirable or entirely superfluous.

The following paper contains only those portions of my former researches
which refer to parts not visited by Mr. Maudslay. The historical traditions of
these localities are entirely lost, and instead of indulging in vague and at present
highly unsatisfactory speculations about possible relations, life, characteristics, ete.,
of the people who produced the sculptures found there, I shall mainly confine
myself to describing what I have seen, leaving future explorers, with more
material, to generalize and speculate upon the, of late, rather fashionable myths of
Tlaloes, Quetzalcoatle, Toltecs, ete.; but this much I will say in regard to the people
who originated these sculptures—that they were, undoubtedly, of Aztec origin; the
worn gylphs yet to be distinguished show similarity to Aztec and not to Maya

writings.

EL PORTAL, PANTALEON, LOS TARROS, SANTA RITA, SANTA LUCIA COTZUMALGUAPA AND
AGUNA.

In the above named places as well as in many others along the Pacific Coast
of Guatemala, from the very slope of the voleanoes to the shores of the Pacific
Ocean, are found numerous traces of ancient civilization. So numerous seems the
population of the country to have once been that there is now scarcely a single
farm upon which is not found ruins or relics of some kind. But the monuments

Mem. Vou, II.7 2. July, 1888.

10 CALIFORNIA ACADEMY OF SCIENCES.

on this coast are of an entirely different type from those of Guirigua and Copan,
and it is perfectly evident that they were constructed by entirely different people,
who resembled each other, neither in regard to physiognomy, dress or ornaments,
nor in regard to the glyphs used for expressing their ideas, etc. The monu-
ments of the Pacific Coast are of several kinds, viz.:

1. Larger or smaller hills or mounds made of soil, or soil and rough or
rounded stones. Generally three or four of these mounds are grouped together
forming between them a kind of inclosed courtyard. Some hills are several thou-
sand feet in length and some twenty feet high; others only fifty to sixty feet long;
the height being the same as the former.

_ 2. Smaller mounds, in which are seen a low foundation of rough stone, not
fixed with mortar.

3. Bridges and aqueducts made of cut stones, beautifully laid without mor-
tar, and wonderfully withstanding the destruction of time. Such are found
around Santa Lucia and Los Tarros.

4. Sculptured stones, representing heads of men or animals, in all grades of
skillful manufacture, some most beautifully made and designed, such as the large
head from Los Tarros; others again very primitive, like those of Aguna.

5. Hollow stones or fonts, richly ornamented, used either for baptism or
for sacrifices.

6. Sculptures in low relief, representing mythological or historical events,
or records of some kind.

7. Pottery of different kinds, such as dishes, vases, and musical instru-
ments with heads of beasts and man; also smaller idols of burnt clay. This pottery
is evidently from different epochs. The finest made wares are found near or on the
surface, while the poorest kinds are found deeper in the ground, as far down as
nine feet. I have this assertion from several gentlemen owning farms on the coast,
especially from Don Joaquin Durand, who has repeatedly found ancient pottery at
different depths on his farm — “‘ Sapote,”’ at the base of Volcan de Fuego. In our
present time no such wares are worked by the Indians anywhere on this coast, nor
can the present crockery of the Guatemala Indians, in skillful workmanship or
even in the fineness of the clay used, compare with the ancient pottery found in
the ground. Only in the wild and entirely unconquered country of the Lacandres,
between Guatemala and Yucatan, have the natives to this day preserved the art of
finer pottery making. Edward Rockstroh, who made an expedition to this country

ANCIENT SCULPTURES OF GUATEMALA, 1]

a few years ago, and who succeeded in penetrating far enough in the same to visit
several hamlets or ranchos, brought from there several idols well made. The
general form was a small dish, furnished at one side with a large head. These
objects were both idols and sacrificiary places. In the hollow of the pots was
placed incense or libation, and the god for which it was destined, was represented
by the head attached to the dish. These heads, or faces, were in many regards
exactly similar to those found on the Pacifie Coast, and bore also a remarkable
resemblance to the faces and heads of the Peruvian pottery. The ground color of
the pots was also a dark grayish-black, while the pottery from the Pacific Coast, as
well as that from Copin and other places of Maya origin, is yellowish-red, and
sometimes ornamented with painted glyphs.

In the “Instituto Nacional” of Guatemala, is a fine and interesting collection
of pottery from different places of that Republic; but like everything else in that
country, it is in a deplorable state, the catalogue being missing, so that the nume-
rals on the different objects cannot be referred to. Nobody has charge of the
“museum,” or takes any interest inthe same. Mixed with the ancient Indian pottery
are some Egyptian idols, as well as pieces of unburnt EHuropean pottery, most
likely believed by the “authorities,” to be of the same origin, and having as
much or as little interest.

The sculptured stone heads found all over the coast are all furnished with a
large conical projection, as can be seen by referring to the different figures appended.
This projection was evidently formed for insertion into the adobe or stone walls of
the respective temples, and served to keep the objects in position. They have no
connection with phallic symbols or worship. Most likely, when the sculptures
were in their original places, these projections were not seen, but were entirely
hidden in the walls.

In producing drawings of different sculptnres, my intention was not, neither
did my time permit me, to again draw stones which had once been drawn and pho-
tographed by former explorers. My intention was only to bring to notice what
was not previously known, and thus, as much as I could, complete the work so ably
begun by others. This especially refers to the sculptures of Santa Lucia, where
Dr. Habel and Prof. Bastian have already done considerable work. In regard to
the sculptures of the other places here referred to, I believe so far no accounts of
the same have been published, and I drew there as many as I could find, within the
limited time at my disposal.

12 CALIFORNIA ACADEMY OF SCIENCES.

EL PORTAL.

The above is the name of a coffee plantation and farm, owned by Don Manuel
Herrera, at the time of my visit Minister of Agriculture in the Government of
Guatemala. The plantation is situated only a few minutes’ walk from “Antigua,”
the old capital of Guatemala. The monuments of this place comprise a large natural
hill or mound several hundred feet high, and perhaps five hundred feet in diame-
ter. The top is covered by three smaller mounds, artificially made. From the top
of one of these smaller mounds, and down the steep south side of the larger one, is
seen the very much ruined remains of a roughly made stairway, the stones once
composing its steps having neither been cut nor well fitted. Along the lower part
of this stairway, and also between the three small mounds at its top, were found all
the sculptures known from this place. At the base of the stairway is yet seen a
large basalt block, in which can barely be made out the figure in low relief of a
large tiger of natural size. A little higher up are seen several very rough heads of
animals, as well asa deer head, but they can hardly be called sculptures, the original
stones having somewhat resembled heads, and only been helped by the artist by the
addition of a line for the mouth or a hollow for the eye. All the better sculptures
were some time since removed to the near dwelling-house and here sheltered from
rain and destruction; Sefior Herrera being one of the few native gentleman in Gua-
temala who takes interest in the ancient relics of the country. The following sculp-
tures are all which are at present known from this place. The numerals refer to
the figures appended only where “Fig.” is put in front of them.

Figure 1. Large block of basalt with the upper surface covered with a tiger
in bas-relief. The tongue is stretched out, and on the same is seen perched a bird

with long bill, very similar to an Alcedo. Size 53 by 3% feet.

Fics. 2 and 8. Head of a male personage—Fig. 2 representing the same
seen from the side, and Fig. 3 from the front. The face is serene and well made,
furnished with a beard. On the head is seen a kind of helmet, the original of
which was perhaps made of wood. In this helmet is seen a smaller face, and on
the side of it the two arms bent upward. The back is furnished with a projection.
Size of Fig. 2, 19 inches long by 143 inches high. Size of Fig. 3, 145 inches high
by 83 wide.

Fig. 4. Head of some animal, perhaps a snake, seen from the front. Rectan-
gular posterior projection. Size, 11$ inches high by 13 inches wide. Roughly

made sculpture.

]
=?
le

ANCIENT SCULPTURES OF GUATEMALA. 13

Fie. 5. Head of another similar animal, but seen from the side. The up-
per surface of the skull has inscribed upon it two C’s,a similar ornament often

being found in snakes. Size, 19 inches long by 114 inches high.

Fies. 6 and 7. Head of some fictitious animal, perhaps modeled after a
tiger. Fig. 6 represents it seen from the front. Size, 1334 inches broad by 19 inches
high. Fig. 7 is the same seen from the side. Size, 25} inches long by 19 inches

high.

Fig. 8. Head of a fictitious animal. Behind is seen the remains of a hollow,

perhaps a font. Front view. Size, 13 inches broad by 83 inches high.
SANTA RITA OR CARMEN,

This place is situated between Santa Lucia Cotzamalguapa and Pantaledn.
So far only one sculpture is found in this place, represented in Figs. 9 and 10. The
lower part of the skull is wanting. Fig. 9 represents it seen from the front and
Fig. 10 from the side. This skull, as well as the heads from Pantaleén, have the

angle of the face very large. The sculpture is at present at Pantaledn.
PANTALEON.

This place is situated one league southeast of San Lucia, and belongs at
present to Don Manuel Herrera. The sculptures found on this place are removed
from their original site, and now to be seen in a yard behind the main dwelling-
house. Originally, however, they were found on the ground between four small
mounds, enclosing between them a kind of courtyard, having the entrance on the
south side. The sculptures consist only of heads of male and female personages,
originally six in number, but one of them has been taken to Berlin, Germany, by
Dr. Bastian. Iam, however, not aware of it having been described. The remain-
ing sculptures are all very well made, cut out of a dark. basalt from Volean de Fuego,
some thirty miles distant. The rocks, however, were most likely found on the
place, as rounded stones of similar nature are in some places abundant on the
ground. All the sculptures are furnished with a large posterior projection for
insertion in the wall.

Figs. 11 and 12. Head of an old woman, the face with very deep
wrinkles all over. Remarkable in all these heads is the head-dress, which is just the
opposite of the head-dresses worn by the idols of Quirigua and Copin. Instead

of being enormously large, it is extremely sinall, and evidently used more as

14 CALIFORNIA ACADEMY OF SCIENCES.

an ornament than as a protection. In this head it consists of a small central bow,
from the sides of which extend two wings of feathers. The whole seems attached
to some kind of frame, on each side of which hangs down two smaller ribbons and

one larger one. Fig. 12 shows this head-dress as seen from above.

Fig. 15. A head of a young woman, of handsome features. The hair is
abundant and seems to hang below the ears. These, as in all the rest of the sculp-
tures, are unnaturally large, most likely enlarged by some process. The head-dress
is in the form of a crown, or at least ornamented as such. The figure shows the

head from the front.

Fies. 14, 15 and 16. Head of a very old woman, seen respectively from the
side, front and from above. The hair is laid in thick tresses and covered by an
exceedingly small hat. The ears are enormously large, their lower lobes extend-
ing and projecting forward. The most remarkable things to be seen in this face are
the eyeballs, which are hanging down outside on the cheeks. The upper lip is
small, while the lower one is very large and projecting. The nose is broken, but

enough is left to show its original large size.

Fics. 17, 18 and 19. Head of an old woman, seen respectively from the
side, front and from above. The hair is ornamentally laid and covered by a very
small hat; by the same is seen a bow. The right eyeball is hanging on the cheek.
The nose is large and aquiline, not broken. Below the face is seen a kind of collar

extending from ear to ear.

Figs. 20 and 21. Head of an old man, seen from the side and from above.
The nose is partially broken. Forehead and face deeply wrinkled. Head covered
by a cloth and a very small hat. The ears have large ornaments.

The size of these five sculptures is nearly the same. Length, 32 inches,
hight 24, and width 16 inches.

What these heads represent will at present remain only a conjecture. In
Yucatan criminals were punished by the extraction of their eyes, but I am doubtful
about these heads representing criminals, as they then would most likely not have
been ornamented, as it seems, in so very fashionable away. Besides, the expressions
of their faces are always noble and serene, which would not have been the case if a
criminal had been represented. Even those having their eyes extracted show no
signs of pain whatever. The faces represent evidently very old people, at least
judging by the very deep wrinkles which cover their foreheads and cheeks. May

an) ie

ANCIENT SCULPTURES OF GUATEMALA. 15

not these heads represent saints, some of whom had martyred themselves and whose

images had since been set up for adoration ?

I was informed that similar sculptures are not uncommon in the neighbor-
hood, but the above ones are said to be much superior in execution. The following
localities are said to contain similar sculptures: Santa Ana or Mistan, about nine
leagues south of Pantaleén; Gomera, about ten leagues south of Pantaleén. Also
in the vicinity of Santa Lucia six heads were to be seen, which were said to exhibit
the peculiarity of having animals and bars projecting from the mouths. Even in
a place north of Pantaledn is said to be a large head more than six feet high, in
similar style with those removed to the house and described above.

LOS TARROS.

This place is situated northwest of Pantaleén a few leagues distant. I did
not visit the place, the time and weather not permitting it. One of the two sculp-
tures found on this place is removed to Pantaleén, and arrangements have already
been made to remove the other. Both sculptures are said to be nearly exactly
alike, the one remaining in its place only being a little larger. Originally both
sculptures stood on a plain facing each other, and 20 to 30 yards apart. Fig. 22
represents the smaller of the sculptures, seen from the front, and Fig. 23 the same
seen from the side. The size is: hight, 68 inches; width, 44 inches; thickness, 24
inches. The sculpture represents the bust of some prominent male personage,
standing out in bold relief from a stone slab, slightly ornamented. Above the slab
is seen the head of a large fictitious animal, with a large hole perforating the base of
the same. The face of the personage is most serene and noble, and expresses great
kindness. The head-dress is very large, a kind of turban, very much inflated, in
front furnished with an ornamental projection, in which is seen a face lying on the
side, and in front of this a smaller ornament like a gem. Behind the face
are seen three leaves, and behind them, extending towards the margin of the stone
slab on each side, a bunch of very long and rather narrow leaves, with a nervation
similar to that of Melastoma. The ears of the personage are large, furnished with
ornaments, in the middle of which is seen a hanging boss. On the breast is an
ornament with five small holes in the margin, and a human face in the center. The
fantastic head, hanging ominously above the slab, is of a wonderful effect, and the
perforation of its side breaks the heaviness of the sculpture. In elegance of design
and in workmanship this stone is undoubtedly one of the very finest ever found in
Central America; and it is much to be regretted that it is not protected from the-

16 CALIFORNIA ACADEMY OF SCIENCES.

injurious influences of the tropical rains and sun. Originally the stone had a large

posterior projection, but this was cut off so as to facilitate transportation.
SANTA LUCIA COTZUMALGUAPA.

The sculptures from this place are pretty well known through the works of
Dr. Habel* and Dr. Bastian.** Some of the slabs have been removed to Berlin, but
the largest part is yet to been seen at the original place, but exposed to rain and
other bad treatment. Many of the sculptures are so faint that it is necessary to

draw up the lines with chalk before any complete idea can be had of the same.

The sculptures, when originally found, were covered by soil, the same being
thrown up into hills or mounds. At present the whole country around is covered
with these mounds of smaller and larger size, and as only a few of them have been
opened, it is not unlikely that the number of sculptures will be considerably in-
creased when a thorough search has been made. The way the sculptures were, or
are, hidden in the ground, is very peculiar. Some of them are lying down flat on
or near the top of the mound; others are standing upright, but covered with soil
up to their very tops. It is evident that the soil has not accumulated slowly from
decayed vegetation, etc., but it seems rather as if the sculptures have once pur-
posely been covered up, to be hidden, perhaps, at the approach of some enemy.
At the time of the Spanish conquest the whole of the Pacific Slope was here densely
populated, especially around and near Hscuintla, from which Santa Lucia is only

distant some thirty miles.

Through the kindness of Mr. Edward Rockstroh, I came into posession of
some lead pencil and ink drawings of sculptures from Santa Lucia, made by
the late Dr. Carl Herman Berendt, and received also permission to publish the
sume. One of these drawings I had already a copy of, made by myself at Santa
Lucia; but as my drawing was wanting in the lower part—this part at my visit
being covered with soil—lI prefer to give the drawing of Dr. Berendt as being more
perfect. As this as well as his other drawings, were not exactly in a state to
be lithographed, very often having corrections of minor details on the margins,
I recopied them, inserting the corrections, etc. It is to be regretted that no notes

as to size accompanied the drawings, and only meager ones as to localities.

*S. Habel.—The Sculptures of Santi Lucia Cotzumalguapa, ete. Smithsonian Contribution to Kuo wledge.
269. Apr. 1879.

** The work of Dr. Bastian, in quarto, 3 plates, printed in Berlin? has no title, at least not the copy that I
lave seen.

~!I

ANCIENT SCULPTURES OF GUATEMALA. 1

Fig. 24. Large stone slab, about nine feet high by six wide. The sculpture
represents some religious ceremony. ‘Two men are carrying between each other a
standard, and appear to deliver the same to a third person. The standard is orna-
mented by several appendices, one being especially large and circular, and having
in its midst the face of a monkey. In the margin of the stone are seen several
round rings with glyphs or signs, but all are so effaced that no correct delineation
is possible. This slab is yet standing upright in a mound, and was originally entirely
covered with earth, but at present the larger part of the face is uncovered.

Fig. 25. Stone slab found in the grounds of Virgilio Pais, on the plain of
Quayabo, near Santa Lucia. Drawing made after a sketch by Dr. OC. H. Berendt.
The central figure represents a chief richly dressed, with a girdle round the waist,
one extremity of the bow ends in a snake-head extending toward the woman stand-
ing infront. One of the legs of the chief is ornamented; the other bare. The head-
dress, behind which is seen a large bird, perhaps the common black vulture of
Central America, is large and ornamented with many feathers. In front of the man
stands a woman, evidently emaciated, as her ribs are plainly seen. She appears
entirely naked, except a girdle round the loins. Her hair is elegantly dressed and
tied with ribbons. Behind her is seen an evil spirit, or other mythical personage,
with claws on hands and feet, and a very pronounced male member. The chief has
his mouth covered with a bandage.

Fig. 26. Original from the grounds of Virgilio Pais; the d rawing after a
sketch by Dr. C. H. Berendt. Represents a sepulchral tablet, on which are seen
the portraits of perhaps man and wife, their different head-dresses, ete., indicating
decidedly their different sexes. From the mouths of the respective portraits extend
as usual curved figures with notes or nodes. A similar speech writing is found in
all Mexican manuscripts, as can be seen by a reference to Lord Kingsborough’s Mex-
ican Antiquities, for instance, Tom. II, p. 26. This writing is entirely unknown in
Maya sculptures and manuscripts, but is seen on some sculptures at Chichen,
Yucatan.

Fig. 27. From the same locality, and after a sketch by Dr. C. H. Berendt.
A stone tablet, most likely, like the former one, a sepulchral tablet, having in its
center a forced deadhead, with outstretched tongue. Above the same are seen two
crossed bars, perhaps meant to represent two crossed bones.

Figs. 28 and 29. The two sides of a sacrificial font or altar. The front is
twenty-four inches high, ornamented with a face like that of a dead man. The ears

18 CALIFORNIA ACADEMY OF SCIENGES.

are enormously large. Above is seen a cavity in the margin, in which perhaps the
victims were made to place their heads at decapitation. The hollow posterior cav-
ity of the font, used for the reception of the cut-off head, is twenty-one inches
broad by twenty inches deep. Locality: Xata. (After a drawing by Dr. Berendt.)

AGUNA.

Situated some twenty miles northwest of Santa Lucia and owned by Dr.
Guillermo Rodiguez of Guatemala. The whole place is covered by artificial mounds.
About nine miles south of the main dwelling-house are seen, among other hills,
three smaller ones made of soil and rough stones. These mounds, perhaps fifteen
feet high and forty feet long, are placed so as to form a small courtyard. In this
courtyard were found the sculptures here represented, viz.:

Figs. 30 and 31. A stone sculptured as a head, with a posterior projection
for insertion in a wall. The sculpture is represented both from the side and from
the front.

Figs. 52 and 33. <A similar stone, with a forced deadhead in front and sev-
eral ornaments. In execution these heads do not compare with the sculptures of
Pantaledn, being very much inferior, both in design and workmanship. Their size
is about four feet long by two high.

EXPLANATION OF THE FIGURES.

FROM EL PORTAL.

Fig. 1. Sculptured tiger in low relief. The tongue is outstretched and on the
same is sitting a bird.
2. Head of male personage, seen from the side.
Head of the same, seen from the front.
4. Head of fictitious animal, front view.
5. Head of an animal similar in size and shape to the above, but side view.
6. Head of a fictitious animal, seen from the front.
The same, side view.

8. Fictitious animal or deadhead, remains of a font, front view.
FROM SANTA RITA OR CARMEN.

9. Sculptured deadhead of man, seen from the front.
10. The same, side view.

24.

26.

ANCIENT SCULPTURES OF GUATEMALA. 19

FROM PANTALEON.

Sculptured head of woman, side view.

The former, seen from above, to show the head-dress.
Sculptured head of woman, front view.

Sculptured head of woman, side view.

The same, front view.

The same, seen from above, to show the head-dress.

Sculptured head of woman, seen from the side.

The same, front view.
The same, seen from above, to show the head-dress.
Sculptured head of woman, seen from the side.

The same, seen from above, to show the kead-dress.
FROM LOS TARROS.

Large, sculptured stone slab, in the middle the bust of a male personage;
above, a fictitious animal's head.

The same, seen from the side.
FROM SANTA LUCIA COTZUMALGUAPA.

Large, sculptured stone slab, in very low relief, representing some relig-
ious or civil ceremony. Two male personages are seen carrying a
standard, and presenting it to a third.

A chief or other personage, in rich dress, and head-dress or helmet with
plumes. Behind the same is seen a bird; in front, a female figure,
and behind her a fictitious personage. The sculpture is in low relief.

Smaller sculptured slab, perhaps originally the cover of a tomb or made
in memory of the two personages whose portraits are seen on the
stone.

Sepulchral stone slab; in center the image of a deadhead surrounded
by many ornaments.

Sacrificial font, seen from the side. The font has a shield with a sculp-
tured face.

The same as above, front view. The small cavity in the top was evidently
used for holding the victim’s neck; the font behind receiving the
head when decapitated.

20

30.
dl.
32.
33.

CALIFORNIA ACADEMY OF SCIENCES.

FROM AGUNA.

Sculptured stone head, with posterior projection; seen from the side.
The same; front view. )

Sculptured stone head—a forced deadhead, with ornaments; front view.
The same; side view. aa |

PL

Mem, Cal Acad. II,

Gustav Eisen, del.
9/1882.

Sculptures of El Portal, Guatemala

ARTO, BRITTON & REY, S F

Gustav E1sen, del.

9/1882.

Sculptures of Portal, Antiqua;
and Santa Rita or Carmen, Pantaledn, Guatemala,

ARTO.

BRITTON & REY SF

a

ARTO. BRITTON & REY, S F

Gustav Etsen, del,

9/1882.

Sculptures of

at

Mem. Cal. Acad, Il PLES

Gusrav Eisen, del. ARTO. BRITTON & REY, S F
9/1882.
Sculptures of Pantaledn, Guatemala,

a

Mem, Cal, Acad. II, P], Vi

Gustav Eisen, del. ARTO, SRITTON & REY SF

Sculptures of Los Tarras, Pantaledn, Guatemala.

ah | Ae

egal)

Mem. Cal, Acad, Il

Pl. ITIL,

Gustav Eisen, in part, after sketch by Dr. Berendt. : ARTO. BRITTON & REY, SF

a il _
Sculptures of Santa Lucia Cotzumalguapa, Guatemala,

Mem. Cal. Acad, II Sd hgh

.

Gustay Eisen, after sketch by Dr.
8/1882,

Berendt.

ARTO. BRITTON & REY, SF

Mem. Cal, Acad. II, FI, X,

Gustav Etsen, after sketch by Dr. Berendt.
9/1882.

ARTO. SRITTON & REY, S F

Sculpture of Santa Lucia Cotzumalguapa, Guatemala.
B g

| <..

Mem, Cal. Acad, II, PI. XI,

Gustay Eisen, del. ARTO. BRITTON & REY, SF

4] 1882.

Sculptures of Aguad, Guatemala,

ON CALIFORNIA EUDRILIDZ.
BY GUSTAV EISEN.

The California Oligochets, to be described below, are all related to the well-
known genera Microscolex, Pontodrilus, Plutellus, ete., referred by Benham, Rosa
and others to the general family of Eudrilidee, and by Beddard to his family Crypto-
drilidx. I will not here discuss the relative merits of the respective families, as the
number of species belonging to them is hardly sufficient to enable us to as yet come to
any definite conclusion. For the present, at least, I retain these genera in the family
of Eudrilide. But I must concede that this family, as defined by the above inyesti-
gators, contains two distinet main types—one represented by Eudrilus and the other
by Microscolex—and a separation in families should be made in reference to them,
though I am uot prepared to make the separation as wide as Beddard has done. Our
California species ally themselves to the Microscolex group, and especially to Micro-
scolex and Plutellus. A species of Pontodrilus was found too late to be included. I
believe this family may be conveniently divided into several subfamilies based upon
the openings of the spermduets, prostates, penial sacs, as well as upon the location of
the nephridio-pores. Our California forms may thus be grouped as follows:

PONTODRILINI.

The eight set are separate though in four couples on eyery somite.

Clitellum complete.

Male pore in somite xviii.

Nephridia—mega-nephridia only—commence posteriorly to somite xiii; the pores are in line with setw 2’,
not alternating in position.

The prostate receives the spermduct previous to its entering the male pore.

No gizzard, no typhlosole, no subneural vessel. PonTopRILus E. P., ISS1.

PHoTopRILuS Girard, 1887.

MICROSCOLECINT,

Eight set separate in four couples; the sets of the inner couples not always parallel.
Clitellum complete. -
Male pore in somite xyii.
The prostate receives the spermduct previous to its entering the male pore; or opens separately in the same
somite.
The nephridia commence anterior to somite vy; the nephridia in line with one of the setw 3 or 4. The first
few anterior pores may open in front of a different sets from the posterior ones.
No gizzard, no typhlosole, no subueural yessel. Ruopoprites Beddard, 1889.
Microscouex Rosa, 1887.
DELTANIA Eisen, 1893.

PLUTELLINI.

Sete eight, in couples or equidistant.

Clitellum perfect or imperfect.

Male pores in xviii.

The prostate either receives the spermduet or it opens separately, but in the same pore.

The nephridio-pores show a systematic alternation in position either in front of or outside of the sete.

A gizzard; a small typhlosole may be present PLUTELLts Perrier, 1873.
ARGILOPHTLUS Eisen, 1893.

22 CALIFORNIA ACADEMY OF SCIENCES.

The genera related to Microscolex may be arranged as follows:

I. Male pore in somite xviii. Nephridia commences posterior to somite xiii.
PoONTODRILUS and PHOTODRILUS, etc.
II. Male pore in somite xvii. Nephridia commences in the anterior somites ii to y.
a, Spermducts open independently of the prostates. RHODODRILUS.

Sp.: minutus; nove-zelandie.
b. Spermducts open in the prostate, close to the body-wall.

1. The set of the inner couples (set 1 and 2) in the vicinity of the clitellum are parallel with

each other and do not converge toward any one of the genital pores. Nephridia commence
in ii to iy. MICROSCOLEX.
Sp.: modestus; algeriensis.

The sets of the inner couples (set 1 and 2) in the vicinity of the clitellum converge toward
one or both of the genital pores. Nephridia commence in somite ii to vy. DeLranta.
Sp.: elegans; Troyeri; Benhami; Poultoni; dubia.

bo

DELTANIA.
Deltania Eisen, Zoe, iv, 250, October, 1893.

Prostomium encroaches on somite i. Eight setee in four couples ventral and
lateral, beginning in somite 11. The sets of the inner couples in the genital region
converging toward the male pore. The sete of the outer couples are further apart
than those of the inner couples. There are a buccal cavity, pharynx, cesophagus and
sacculated intestine, but no gizzard, thyphlosole, or cesophageal pouches. Clitel-
lum, which is perfect, contains three large somites and two outside smaller ones,
extending from xiii to xvil. No dorsal grooves. Testes in x and xi, free. Sperm-saes
present, free. Spermatheca present or absent, variable in position and shape.
Ovary one pair in xiii. Oviduct one pair in xiv. Two pair of ciliated rosettes in
x and xi. Spermduets open in xvii together with a large prostate which is placed
parallel to the segmental grooves. The spermducts join the muscular part of the
prostate at the point where it enters the body-wall. The glandular part of the prostate
consists of two layers of cells. Penial setee open in the same duct as the prostate.

The dorsal vessel with three pair of lateral hearts in x, xi and xii. No sub-
neural vessel. Only very few blood vessels on the nephridia.

Nephridia are of two slightly different kinds. Those in the first few anterior
somites, commencing in somite ii, open in front of and a little interior to the 4th seta,
while those posterior to the former open in front of and a little lateral to the 5d seta.
The nephridia begin generally in ii, rarelyin y. All nephridia furnished with a large
terminal bladder which in the posterior nephridia develop a ccecal prolongation.

Small, transparent-glassy, more or less colorless worms with orange colored
clitellum, and living in moist, especially sandy soil.

Systematic position. Under the genus Deltania I group all species which
would otherwise be referred to Microscolex Rosa, but which agree in haying a deltoid
arrangement of the ventral sete surrounding the generative and especially the male
pores. With this character I believe a closer investigation of the species of Microscolex
will join others, principally in regard to the nephridia which have not been sufficiently
studied, probably on account of the scarcity of specimens for research.

Among species which must be referred to the genus besides those described

to

CALIFORNIA EUDRILIDA®,

below are Microscolex dubius of Rosa and HLudrilus dubius of Fletcher, I will first
refer to the latter. Fletcher’s description is sufficiently minute to allow us with cer-
tainty to refer it to the genus, but the details are wanting to such an extent, that it is
difficult to understand its further relationship. There are three points in the deserip-
tion which are of special interest.

1. Absence of spermathece.

2. The beginning of the nephridia in v.

3. The junction of the spermduct and the prostate half-way between the
glandular part and the body-wall.

As to the first of these the spermatheee may be really wanting or it may have
a substitute similar to the one found in Deltania elegans as described below. At any
rate this character brings the species Hudrilus dubius close to Deltania elegans as well
as to Beddard’s Wicroscolex Poultoni.

The beginning of the nephridia in somite y brings 2. dubius close to Beddard’s
species but separates it distinctly from De/tania elegans in which the nephridia com-
mence in i, as will be shown below.

The third character requires to be reaffirmed and described more in detail.
The joining of the spermduct and the prostate is always of the utmost importance and
interest and a mere general statement will not suffice for properly characterizing a
species, especially when the group is little known.

Rosa at first considered #. dubius to be identical with his JJicroscolex mod-
estus, but a later investigation of new material convinced him of the distinct char-
acter of the species and he then describes both as two different species of Microscolex.
It must therefore be considered certain that the deltoid arrangement of the ventral
setee does not occur in JMicroscolex modestus. In regard to the respective species of
E. dubius and M. dubius described by Rosa and Fletcher, I am not fully persuaded
that both actually belong to the same species, and I believe that nothing short of an
actual comparison of the specimens can decide if they do so.

Beddard has at two different times described species of the genus Microscolex,
but which differ from each other in several important points. Wicroscolex novwe-zelan-
die resembles the old genus Rhododrilus in the independent opening of the spermduct.
Instead of referring the above species to Microscolex, and merge Rhododrilus in the
latter genus, I consider it more proper and convenient to retain Rhododrilus and re-
fer JL. nove-zelandic to it, as the independent opening of the spermduet appears to
me of sufficient importance to be considered a generic character. Another species,
Microscolex algeriensis, also described by Beddard, ean, I believe, best be retained in
the genus Microscolex, as it evidently possesses the sete parallel throughout the
ventral side of the clitellum.

Microscolex Poultoni however is probably a true Deltania and Beddard’s excellent
description leaves no important characters in doubt.

In his description of Microscolex Poultoni, Beddard refers especially to the
deltoid arrangement of the sete in the clitellial somites. He says: “ From segment

24 CALIFORNIA ACADEMY OF SCIENCES.

xix backward and from segment xiii forward, the distance between the two ventral
setee of each side gradually increases.”

Distribution and habitat. The species of the genus Deltania described below
are undoubtedly natives of the Pacific Coast of North America, and more particularly
ef California. The species are found not only in the Golden Gate Park of San Fran-
cisco, where they might have been introduced, but also in localities distant from
gardens, and in which no cultivation has ever taken place. Thus I have species
from Berkeley, Santa Rosa, Lake Chabot, Mount Diablo, ete. Moreover, since this
paper was finished in MS., I have found species of Deltania in localities so far from
civilization that there can be no doubt as to the native habitat. I refer to species
found in the high mountains of the Cape region (in the vicinity of Cape San Lucas)
of Baja California, at an altitude of 4,000 feet, in almost inaccessible localities. A
species has also been found in a gulch or river-bed at Ensenada, Baja California,
thus proving the extensive territory over which the genus extends. The description
of these species must be deferred to another paper.

As regards Deltania dubia (?Eudrilus dubius Fletcher) and Deltania Poulton
(Microscolex Poultoni, Beddard) it may be possible that the former, at least, is an
introduction from abroad. The locality where the latter is found shows the genus to
possess a wide neotropic extension.

While it is thus certain that species of Deltania are natives of the new world
and the Atlantic islands, and especially of the Pacific Coast of North America and
Mexico, the distribution of Microscolex must yet remain undecided. Whether it
was imported from the Argentine Republic to the Mediterranean region, or vice
versa, must remain an open question, though the finding of two species in the Medi-
terranean countries seems to point to the probability of that region being its real
habitat.

Species. Of the genus Deltania, California now possesses at least three species
with the very great probability of a discovery of new species, as soon as a wider area may
have been explored. All the species appear very sensitive to dryness and heat and
disappear with the first warm and dry weather, hence the difficulty of finding them
except at their proper season which appears to be limited to February to April in the
vicinity of San Francisco. The species in Baja California were found in September
and October.

Deltania elegans.
Figs. 1-20, 49-58.

Deltania elegans Eisen, Zoe, iv, 248, October, 1898.

Size about 2 to 4 inches by from ,', to } inch wide. Septal glands very small, the
posterior one the smallest. Spermathec very pellucid, minute, and irregular in their
position both as regards somite and the place in the somite, but generally opening be-

tween viii and ix. Sperm-sacs comparatively small, deeply lobed, one pair in xi and

CALIFORNIA EUDRILID®, Zo

one in xil. Prostate is at the top helix-like folded. Penial papill:e prominent, with
two or more penial sete in each sac.

The worm is very pale, glassy, semi-transparent with reddish-orange clitellum,
and with the dorsal vessel showing prominently through the body-wall. It is very deli-
cate, succumbs readily to heat and can only with difficulty be kept alive. It is the
largest of the species belonging to this genus as far as known.

Habitat. Deltania elegans is so far only found around San Franeisco, Califor-
nia, and at Santa Rosa and Mount Diablo, especially under decaying manure in natura]
hollows in the Golden Gate Park. It is common immediately north of Strawberry
Hill in the sandy hollow where rain water collects and keeps the soil sufficiently moist.
The worm is not found in the water, but at the water’s edge, the water, however, being
only temporary during the rainy season. A few specimens also found in Berkeley
along the creek. In the locality in Golden Gate Park the worms congregate in large
masses, always in the sandy soil. At Santa Rosa and Mount Diablo I found only few
specimens, the season being at my visit in May far advanced and the soil was drying
fast. Mature in April and May.

DETAILED DESCRIPTION,

Exterior characteristics. The worm varies considerably in size, but averages
about three inches in length, by a width of from two to three lines, tapering consid-
erably towards the tail. The color is pale flesh, with a bluish cast, and quite trans-
parent, glassy, with a yellowish clitellum. Altogether, the exterior appearance of the
worm is one of great deticacy, greatly heightened by its semi-transparency.

The cephalic prostomium is prominent, and divides the peristomium about ?
of its width (fig. 3). This, the first somite, is wider than any of the following.
Somites ii and iii are next in size, and about equal in width. Somites iv to xi are
smaller, about equal in width, slightly decreasing backward. Somites xii and xiii are
smaller than any of the other anterior somites, and of about the same respective width
(fig. 2). No dorsal pores.

The clitellum (figs. 2 and 15) commences with somite xiv, though xiii is gen-
erally somewhat thickened. The clitellial somites xiv, xv and xvi are very wide,
about as wide as somites ii and iii, and of the same size. Somite xvii is much smaller.
This somite carries the male pores and papille (fig. 15). These are situated in the
posterior part of the somite, almost on the edge of the intersegmental groove (fig. 15).
The papillie are slightly raised around the opening of the sac in the penial sete. The
papillee are situate close together very near the median line of the body, a short dis-
tance only from the ventral ganglion. The papilla is oblong, sigmoid, with a pore
for the sete at inner end. Between it and the intersegmental groove is seen the slit
in which open the prostate and the spermduct. It is situated slightly to the outside
of the penial papilla (fig. 14).

The oviducal pores are in the anterior part of xiy, generally in a depression
anterior to and more ventrally located than the inner couple of sete. Spermatheeal
pores are variable, not perceptible. The nephridio-pores are in front of the third set,

26 CALIFORNIA ACADEMY OF SCIENCES.

while the three anterior pepto-nephridio-pores are in front of the fourth sete (fig 2).
Clitellum is conspicious, occupying somites xiy to xvii, encroaching slightly on xiii
(fig. 2 and 15).

In viewing the caudal end of the live worm a number of irregular white spots
are seen in mature specimens. ‘These are rows of mature ova which agglomerate there
sometimes in large quantities. How they finally find their way through the oviduct
is not readily explained.

Sete (figs. 2 and 4). As usual the sete begin in somite 1. All except
those in the penial sacs, are sigmoid and arranged in couples of two, eight in each
somite. The ventral setee are about 4 closer than the lateral ones. Thus if we con-
sider the distance between the inner and outer couples to be 50, that between the
sete of the outer couple is 50, that between the setee of the inner couple 22, that be-
tween the inner couples 40. This being the distance in somite xxvii, where it may
be said to be normal. In the region of the clitellum, and posterior to it, the inner
setee are unequally distant in the respective somites. Thus we may consider the inner
setee 1 and 2, to have the normal distance from each other in somites xii to xxvii.
From these two somites the sete in the inner couples conyerge toward somite
xviii in which the sete are about 4 as far apart as in xii and xxvii. In xyii the sete
of the inner couples are wanting.

The innermost or row No. 1, forms a continuous line from one end of the body
to the other, while the row No. 2, forms an angle with somites xvii and xviii at the
apex. The sete in rows 3 and 4 are parallel and normal. The seta 1 in xviii is pre-
sent (pl. xii, fig. 4).

This arrangement of the sets appears very constant, and is characteristic of
the species, the details being somewhat different in the other species of the genus,
while the general characteristics are the same. The normal sete (fig. 16) of the
clitellum are not smaller than those of other parts of the body. The penial setie are,
however, very much the largest.

Penial setw (fig. 17). The two pairs of sacs containing the penial sete are situated
in front of the spermidueal pore in somite xvii. They open immediately in front of
that pore, in a slit, at either end of which is situated a pore, each pore being the
outlet for the respective fork of the penial sac. The two sacs are connected at the
upper margin as usual by arciform muscles. Each sae contains not less than two, and
sometimes three or four setie, straight or slightly curved, but not sigmoid. The sete
vary considerably as regards shape, but resemble each other in not being sculptured,
and are only marked by rings. The penial sac reaches to the upper part of the mus-
cular part of the prostate.

INTERIOR CHARACTERS.

The Septa begin between somites iv and y. Those between vii and viii, and
following as far back as somite xiv, are slightly thickened, all, however, being of the
ame general thickness,

The body wall contains the usual layers. The hypodermis is slightly thinner

CATTIFORNIA EUDRILIDA,

te

than the circular muscular layer, and this again is thinner than the longitudinal
muscular layer. The clitellum is perfect, but somewhat thicker on the upper part.
The glandular layer is very thick, being about ten times thicker than the two mus-
cular layers combined. The longitudinal fibres are nowhere arranged in such
feather-like: bunches as in Lumbricus, but much more irregular, with a faint ten-
dency to feather-like bunching. This is the rule both in the clitellum and elsewhere.
This stratum of longitudinal fibres is, as usual, interrupted in four places by the setal
grooves, though there are also fibres between the sete. In the center, between
these grooves, the layer is thickest, from there gradually tapering towards the setie.
The under side, or the ventral part, of the layer is somewhat thicker than the dorsal
part.

The clitellar glands are less regularly paired than in Lumbricus though
the general arrangement is that of two or three rows of glandular cells together. The
clitellum is very thick and developed all around the body.

Alimentary canal (fig. 1). The buceal cavity is eversible to a remarkable de-
gree, so much so that it is often projected like a large sae or bladder, covering more
or less perfectly the prostomium and part of the peristomium. Its walls are very
thin and transparent. The pharynx commences with the prostomium and covers
somites i, ii, iil, but is only dorsally developed. It is much and irregularly folded, the
sinuses being sac-like and not parallel, the largest ones being in somites ii and iii.
The muscles of the pharnyx are thickly covered with salivary glands. Superiorly
these glands project along the under side of the muscular bands, which extend back-
ward, thus forming three rows of parallel projections tapering from base to apex. ‘The
anterior of these salivary glandular masses is the largest, the third, or the most posterior
one, the shortest (fig6). The posterior half of the salivary glandular mass forms one
single projection equal to all the anterior ones together. This gland is connected with
and partially rests on two muscular bands attached to the body-wall between somites
vii and yiil.

(Hsophagus commences between ii and iy, extending backward to somite xvi,
being slightly differentiated in xiv, xv, and part of xvi, as a tubular intestine (fig. 1).
(Esophagus is much sacculated, first rising upward and forming a sigmoid plexus in
somite vii, after which it lowers itself somewhat in viii and then extends gradually
backward to the sacculated intestine, at the same time gradually diminishing in size.
The tubular part in xv or between xy and xvi is the narrowest part of the cesophagus.
The glandular epithelium of the cesophagus is yery narrow in the anterior somites, or
those in front of the clitellum, and the blood sinuses in them are narrow. In the
anterior part of the clitellum the epithelial villi become greatly elongated with in-
creased blood supply, while in the central part of the clitellum these blood sinuses be-
come very large, occupying the largest part of the epithelial lobes. The nuclei in those
epethelial cells are everywhere round.

The sacculated intestine, tig. 1, s. i., commences in xvi, is generally about four or
five times wider than the tubular intestine. It does not increase gradually, but at

28 CALIFORNIA ACADEMY OF SCIENCES.
once, in which it differs from the corresponding organ of the two other species de-
scribed here.

Septal glands (fig. 1). The septal glands of the esophagus are found paired in
somites yi, vii, vill and ix. They are all comparatively small and very thin and
flat. The posterior ones are smaller, but the difference between the glands is not
so great as for instance in some species of Oenerodrilus, or as in Deltania Troyeri.
They are more nearly of the same size, and they extend all around the cesophagus
and are more or less divided in several separate lobes. We can, however, always
distinguish one pair in each somite connected with the septum below the alimentary
canal, extending upward with its free upper lobes, which do not connect.

Nephridia (figs. 19 and 20). The nephridia commence in somite ii, and are
found in all posterior somites. The three anterior nephridia—in ii, iii and iv—are
slightly different as to size and outward form, and may be considered as a kind of
pepto-nephridia. They open, also, differently from other nephridia, their pores
being in front of and a little interior (nearer the ventral ganglion) to seta 4, while
all other nephridia open in front of and interior to the third seta. The nephridio-
pores of the clitellum are considerably larger than the pores in the other somites, and
appear like large, transparent dises when viewed from the outside. All nephridia
possess a vesicle or bladder next to the body-wall. A small collar of tubular cells
surround the nephridio-pore. The vesicle is smaller in the anterior pepto-nephridia,
gradually increasing in size backward, being largest in the post-clitellar nephridia,
where it is several times larger than in the pepto-nephridia. In the three pairs of
pepto-nephridia, the vesicles are almost circular, and of the same size in the three ne-
phridia. That of the first common nephridium in somite y is of about the same size
as the pepto-nephridial vesicles, but from this on the bladders increase gradually, but
slowly, in size to the end of the clitellum. But in somite xviii and following to the
end of the body, the vesicles are much larger, about twice as large as those in the
clitellum. Thus the nephridio-yesicles in the clitellum are about three to four times
shorter than the tubular part, while the post-clitellar vesicles are half, or more than
half, as long as the tubular part or duct, when ordinarily folded. In the pepto-
nephridia, the vesicle is about five times shorter than the folded tube, and the tubular
duct extends more backward than in the other nephridia—especially so in the first
nephridium

encroaching on the next posterior somite, reaching diagonally across the
somite, while all the other nephridia run parallel with the intersegmental grooves.
This diagonal position of the nephridia is, however, not always constant, except in
the most anterior nephridium. The vesicle in the posterior nephridia consists of two
more or less distinct lobes, the posterior one (to the duct), which is more rounded
and bladder-like, forming a ececum, and the anterior, which is elongated or deltoid.
This difference is more pronounced in the posterior than in the anterior nephridia,
most so in the nephridium in sonite xviii, which nephridium is generally the largest
of all. From this somite the nephridia diminish somewhat in size, both anteriorly
and posteriorly. The posterior margin of the vesicle is considerably lobed, and in

CALIFORNIA EUDRILIDA. 29

general outline convex. ‘The anterior margin is convex-deltoid, with the apex at the
exterior pore.

The single duct from the vesicle leaves the vesicle in a different manner in
the respective nephridia. In the pepto-nephridia it ascends from the center of the
vesicle; in the anteclitellar vesicles it leaves from the apex of the deltoid longer part
of the vesicle, while in the postclitellar nephridia it leaves from the side of the vesicle
below the apex. The inner structure of the nephridia corresponds almost exactly
with that of the nephridium of Argilophilus, which genus, however, does not possess a
vesicle. With this exception, the nephridia of the two genera might be considered as
almost identical. The most characteristic feature is that the short single duct from
the nephridio-stome after joining the nephridial body, does not enter it as a single tube,
but as a spongy duct full of irregular and connecting ductules, which later on join
into one larger branching canal, the ductules, or arms of which enclose the two reg-
ular underneath-lying ducts. At the inner bend of the duct the ductules disappear and
the returning lobe contains three main canals, one of which is ciliated. This arrange-
ment reminds us greatly of the one observed in certain leeches and described by
Bourne. The vesicles consist of an outer muscular layer, which extends all around
the bladder, and from it, along the duct through the clitellum or body-wall, to the
exterior pore. Between it and the ccecal epithelium there is a continuous row of
connecting chambers probably analogous with the ductules of the ducts. This epithe-
lium is furnished with some few blood-vessels. The duct from the bladder to the
nephridio-pore is not otherwise differentiated, the glandular cells of the clitellum joining
directly on the muscular duct. Before reaching the pore but while in the body-wall,
the duct is enlarged, forming a small pear-shaped urinary bladder, which again opens
into a narrow duct surrounded by a row of long tubular cells, which open directly
into the duct. These cells form a veritable collar, the upper cells lining the inner
surface of the nephridio-pore (fig. 49). This in the elitellar nephridia. In the
nephridia posterior to the clittellum, the long tube between the vesicle and the pore
is entirely wanting, the bladder connecting directly with the collar of tubular cells
(fig. 50). The tubules, or yacuols, in the vesicle collect into at least two tubes, which
run downward between the muscular and glandular layers of the vesicle and appar-
ently open on either side at the beginning of the collar at the nephridio-pore (figs. 49
and 50). The secretion from this glandular layer of the bladder may be of such
nature as to facilitate the ejection of coarse matter such as calculi which are found
often in enormous quantities in the vesicle or seen as just ejected through the pores.
The single duct which leads from the bladder to the nephridial body proper or the
folded canals, apparently does not connect directly with the ductules or vacuols of
the bladder, but opens directly into the large central chamber of the bladder, which
again connects directly with the collar of the nephridio-pore. For a more detailed
account of the canals of the nephridia in these two genera see the description of next
genus, Argilophilus. Below the nephridial collar in De/tania elegans is found a large
branching body, probably a ganglion. It sends out branches to the nephridial collar,
though in my sections I haye not seen their actual connection with the collar. In

50 CALIFORNIA ACADEMY OF SCIENCES.

fiz. 50)., the section is made through the junction of the nephridio-vesicle and the
collar, and the ganglion (x. g/.) below is small, not yet branched. In fig. 50c, the
section is made close to the pore where the ganglion (x. g/.) is branched. These
branches always extend towards the collar, never the other way. Both these sections
from which the drawings were made are not quite in right angle to the longitudinal
muscular layer, but slightly slanting.

The nephridio-stome is rather large, but very transparent and quite difficult to
detect. It is flat, rosette-like and placed half-way between sete 1 and 2. The duet
descending from it is remarkably thin, straight and never winding. ‘The nephridio-
stome is normally situated, piercing the anterior septum as usual. There is no central
cell and nucleus larger than the other, but seyeral smaller cell nuclei are visible in
the center below the large marginal cells, which are about 16 to 18 in number and as
usual placed in a crescent. Between these cells and the real margin of the lip there
is a row of smaller cells and nuelei, the latter being slightly lower than the inner
large nuclei, but sufficiently situated in the same plane to be seen at the same time.

7Zestes consist of two pairs of small bodies, as usually situated in somites x and
xi, attached to the septum close to the body-wall in the anterior part of the somite.
They are deeply lobed and the lobes are very narrow, parallel and generally three or
four in number. The testes are free, not enclosed in the sperm-sacs.

Sperm-sacs (figs. 5 and 6). There are two pair of sperm-sacs, one pair in
somite xiand one pair in xii, attached to the anterior septum in the somite, rather
high up, but below the cesophagus. The four sperm-sacs are of about the same size,
comparatively small, and deeply lobed very much like the posterior sperm-sac in
Ocnerodrilus Beddardi, ete., none of the lobes being very much larger than the other.
It must be remarked that the sperm-sacs are not situated in the same somites as the
testes, somite X possessing testes, but mo sperm-sacs, and somite X11 possessing sperm-
sacs, but no testes. The respective sperm-sacs are not connected with each other and
there is no median sperm pouch. The trabecula or muscular divisions of the sperm-sacs
are numerous and the chambers enclosed by them are comparatively small.

os.7,8, 11). There are two pair of ciliated rosettes,
oD >

one in xiand one in xii,as usual. They are very large and folded, deeply crumpled and

very prominent, occupying about + of the somite. The funnels are wide, with a

Spermducts and rosettes (fi

slight posterior swelling. The spermducts run backward and sideways, connect in
xii, and continue to somite xvii, where they leave the body-wall, ascend slightly and
connect into one single duct, which opens out jointly with the prostate, in the
posterior part of the somite. The duct does not enter the prostate, but both open
jointly in one slit, running parallel to the intersegmental furrow. The spermduct is
throughout of the same size, only widening out just previous to entering the rosette.
The lower descending part of the duct, between the spermiducal pore and the bend
where it starts forward, is slightly narrower than the anterior part.

CALIFORNIA BUDRILIDA. 51

Prostate (fig. 8). There is one pair of prostates of rather prominent size ex-
tending parallel with the intersegmental groove reaching almost across the somite.
The shape of the prostate resembles at the top somewhat a curved feather, the inner
apex being helix-like, curving backwards. This form appears quite constant and while
I found the length and the width of the prostate to vary, I never found one, which did
not show the helix-like, convolution. The thickness of the prostate varies consider-
ably. In some specimens it was almost twice as thick as in others, the increased
thickness being caused by a gradual widening toward the inner apex. In some speci-
mens the prostate was longer, more slender and its longitudinal sides almost parallel,
but the convolution was generally always thickened. In most instances the convolu-
tion could be considerably straightened out by a pushing with a needle, but it would
when released assume its natural helix-like form. The spermduct connects with the
muscular part of the prostate in the muscular layer of the body-wall. The sac con-
taining the penial sets is situated immediately anterior to the prostate and somewhat
closer to the ventral ganglion in line with the regular sete, but opens in the same pore
with the prostate and spermduct. (Figs. 8, 51, 52, 53.)

A eross section of the glandular part of the prostate shows that it is composed
of two layers of cells, the outside one containing large cells of flask-like shape, the
inner are narrower rectangular cells. The contents of both layers resemble each
other greatly and are difficult to discern. Both layers of the prostate contain nu-
merous blood vessels arranged like radii ina circle, penetrating both of the cellular
layers. But the inner layer is seen to also possess a vascular system of its own with
many smaller vessels similarly arranged. These vessels are generally wider at the
periphery of the prostate and narrow toward the center, many if not all collecting in a
network of capillaries spreading on the inner surface of the prostate (figs. 55 and 56).
Otherwise these vessels do not anastomose. All these vessels are fed by a branch from
the ventral vessel of the body, which divides on the prostate into two or three large
branches which again fork toward the apex of the prostate in many smaller ones, as
in Deitania Troyer (fig. 45).

This junction of the various male organs is affected in this manner. The two
spermducts run jointly on the top of the inner longitudinal muscular layer of the
body-wall. When reaching the lower or muscular part of the prostate they turn and
run parallel to it. Immediately before reaching the place where the prostate enters
the wall, the two spermducts fuse into one duct, the lumen of which then is wider
than the adjoining part of the prostate.. This duct joins the prostate in the iongitudi-
nal museular layer of the body-wall. After reaching the transverse muscular layer
this duct joins the pore of the penial setze (figs. 51, 52, 55).

Spermatheca (fig. 13). The spermatheci consist of very minute bodies, pear-
shaped in outline, and of extremely delicate structure, without any differentiation of
the wall in a muscular and glandular layer. In size, the spermatheca is not much,
if any, larger than one-half the width of the somite, when contracted in alcohol. But
the most peculiar feature of this organ is that it is variable in number and_ position.

a2 CALIFORNIA ACADEMY OF SCIENCES.

In one specimen I found only one spermatheca, situated in the center of the ventral
line, between somites ix and x, but in x. Another specimen had two spermathece,
one in ix /x, another on the left side, between x and xi. Another had two sperma-
theez betweeen x and xi, one on each side. Another had two spermathece between
ix and x, one in center of median line, and one on the right side, always attached to
the intersegmental wall. This arrangement and variability of the spermathece
reminds us of the spermathece in Microcheeta, where the number varies on either
side; but in other respects there is no similarity between the two. The exterior
spermathecal pores are not conspicuous, and not perceptible when viewing the out-
side of the body, mounted, for instance, in glycerine. As will be seen, the sperma-
theea in this species differs very much from those in Deltania Troyeri and Benhami,
in which two species this organ is constant, and furnished with two diverticula each.
The spermatheca in Deitania elegans resembles greatly in structure a sperm-sae, from
which it only differs in size and in position. It reminds me greatly of the peculiar
organ described by me in Ocnerodrilus occidentalis, where, apparently, the posterior
testes have become modified, and assumed the function of spermathecee, with a dis-
tinct and ciliated duct perforating the body-wall.

The spermatozoa are always found agglomerated in spherical masses in the
spermatheea, hardly regular enough to be designated as spermatophores. The tails are
lone, either extending straight out, or arranged screw-like in the same direction

aD fo} fo) oa
around the sperm-ball. These balls vary greatly in size, some being twice as large as
others, but they are always round and apparently globular (fig. 15).
« © J dD D

Ovary and Oviduct (fig. 9). As usual the ovary is found in xiii. It offers no
great peculiarities. It is rather deeply lobed and very large. The oviduct opens in
xiv, with its funnel in xiii. The oviducal funnel is very thick, substantial and round,
with a circular and very regular outline. The figures (5 and 9) give correctly its
outline, but the depressed folds have been too distinctly marked. There is no ovisae,
and the oyary and oyiduets are entirely free.

Blood vessels. The dorsal vessel emits three pair of hearts in x, xi, xii, and the
ventral vessel is forked between somites ix and x. The blood is yellowish-red, more
decidedly yellow than red. There are but few blood vessels on the nephridio-tubes and
none on the nephridio-vesicle.

The ventral nerve-cord is considerably wider in the posterior part of the
somites where it emits the customary pair of nerve fibers. In the anterior part
where the single septal nerve pair is emitted, the nerve-cord is quite narrow. In
Deltania Troyeri the nerye-cord is quite uniform without any nodular enlargements,
as wide at the anterior as at the posterior end of the somite. The brain is narrow,
slightly curved and the posterior sinus shallow. — It is situated in somite ii (fig. 6).

CALIFORNIA EUDRILIDA.

Deltania Troyeri.

Figs. 21 to 39.

Deltania Troyeri Eisen, Zoe, iv, 251, Oetober, 1893.
Size about 14 inch by $ line. Septal glands comparatively large, the one in
vi the largest. One pair of large, opaque spermatheca, furnished with one pair of
diverticula, which are about $ or more longer than the spermatheca proper. Sperm-
sacs in x and xi, not lobed. One developed seta in each sac of penial setee. Pros-
tate is tubular, not helix-like, with the top either straight or bent at right angle, pro-
jecting backward. The exterior penial papillee not as prominent as in the preced-

ing species. The inner couples of sete are further apart than in the following species.

Habitat. This species was first brought to my attention by Professor Carlos
Troyer, of San Francisco, who found it, together with the preceding species, in the
Golden Gate Park, in San Francisco, immediately north of Strawberry Hill. It
occurred there in sandy depressions, where the rain and drainage water had moist-
ened the soil in March and April. As the soil dried up the worms disappeared.
The worm is very searce at any time, and not one specimen is found to every hun-
dred of Deltania elegans.

Evterior characteristics. Fixteriorly this species is characterized at once from
Deltania elegans by being very much smaller, as much so as an Enchytreeus is
smaller than an average medium-sized Lumbricus. The length in the largest speci-
mens is about two inches, when stretched to its full capacity, though the average ones
hardly reach one inch. The width is less than one line at the clitellum and less than
! line at the tail end. The first somite is much longer than any of the following.
The second somite is next in size, while all the others are smaller and of very much
the same proportions as in Deltania elegans. Thus iii, iv and y, are larger than the
following, and those between v and xiii are smaller and of about the same size.

The clitellum occupies the same somites as in Deltania elegans, or from xiy
to xvii, with the two outside somites smaller than the central ones. The body tapers
towards the tail end, the last somites being somewhat larger than the others and
rather obtuse.

The color is pale flesh, with a darker, yellowish clitellam. The whole body
is very transparent, just as the former species, but much less so than in the following.
It is a very tender worm indeed, and can only be brought home alive with great
care, as the least increase of temperature is apt to kill it. In no instance did I sue-
ceed in keeping it alive more than a couple of days. In this respect, however, all
the species of the genus are very much alike, and if there is any difference the larger
species is the most tender.

Sete (figs. 21, 24 and 39). The general arrangement of the setz is similar to
that of Deltania elegans, but the two inner sete are much less close together, in compari-
son with the two outer ones, than in the latter species, but not as close by one-half as in

.

34 CALIFORNIA ACADEMY OF SCIENCES.

Deliania Benhami. Tn the genital region, the distance between the two inner sete
diminishes toward the male pore, almost in the same way as in De/tania elegans, with,
however, a slight but characteristic difference. The inner or first seta in xyili is
often, but not always, wanting, probably falling out when young, before its full devel-
opment, as more frequently a rudimentary seta is seen in place of a fully developed
one.

The first and second sete in xix and xx are closer together than normally. but
already in xxi the sete have regained their proper distance. Again, anterior to the
male pore, the setee 1 and 2 in xi to xvi are closer than normally, those in xiv to xvi
are equidistant, while those in x to xiv rapidly approach. If we thus compare with
Deltania elegans, we find that the arrangement relatively in front and behind the
male pore is reversed. While in De/tania elegans, the anterior setee quickly con-
verge, the posterior ones approach slowly. In Deltania Troyeri, the opposite is the
case.

The shape of the sete in the two species is very similar. Compared again
with the arrangement of the sete in Deltania Benhami, we find that in the present
species the sete in the ventral couples, as well as those couples themselves, are
much further apart than in Deltania Benhami. The deltoid arrangement, also, is
different in the two species, of which the figures give a better idea than any lengthy
description (figs. 24, 39, 40).

The sacs of penial setw (fig. 33) are found as usual in the vicinity of the male
pore in xvii. There are seldom more than one seta in each sae. This seta is long,
slender, almost straight, occupying the whole length of the sac. Now and then there
is a rudimentary seta in the same sac, but never more than one developed seta. In
Deltania elegans there are three or four set in each sae.

Alimentary canal (figs. 26 and 27). The buceal cavity extends superiorly to
ii, inferiorly to y. The pharynx ends in y, and is much less developed than in
Deltania elegans. The upper fold is, however, very large. There are one pair of
long and narrow salivary glands in each of iii and iy, and one pair very large com-
pact ones in iy. The cesophagus commences in y, and rises to a sigmoid plexus in
viii. It is greatly contracted at the septa. In xy and xvi it narrows down to a tubu-
lar intestine.

The sacculated intestine commences in xvii, but attains its full width first in
xix or xx. There is no gizzard, no ealciferous glands nor pouches of any kind at-
tached to the alimentary canal.

Nervous System (fig. 28). The characteristic feature of the nervous system is
the eyen width of the ventral ganglion, the two sides being nearly parallel throughout,
with almost imperceptible contractions at the septa. In Deltania elegans this contrac-
tion is very prominent, and the ganglion is almost twice as wide in the posterior part
of the segment as in the anterior one. This characteristic appears constant. In
Deltania Benhami the ganglion is narrowed somewhat at the septa, but the posterior
part in each segment is not any wider than the anterior part.

CALIFORNIA EUDRILIDA, Do

Septal Glands (figs. 26, 27, 29). When the worm is laid open and the cavity
viewed from above, it is seen that there are 4 pair of septal glands surrounding the
cesophagus in somites v, vi, viiand vii. In this view the anterior gland appears the
largest, and the posterior one in viii the smallest. This is, however, only an illusionary
appearance, caused by the position of the glands. There is in reality not any very
considerable difference in their size, as may be seen when separated and spread out.
Seen ina slightly eccentric longitudinal section, the gland in vi appears the largest both
above and below the cesophagus, though in some sections the lower part is not as large
as the lower part of the gland in vii. The anterior gland in y is short but broad. The
one in vil is larger than those in v and viii, but smaller than the one in vi. The
upper part of the gland in viii is larger than the corresponding part of y, the lower
part of the latter being the smallest. As will be seen, all the glands are developed,
both superiorly and inferiorly, as regards the csophagus, but the glands on either side
in the somite do not connect, but only touch. There is a slight, but irregular lobing
of the glands, frequently unequal on either side, as one gland may be almost entire,
while the other again is furnished with three indentations. On the under side of
each main gland there is a smaller lobe, almost entirely separated from the rest (fig.
29). The glands are furnished with blood from the subcsophageal longitudinal blood-
vessel which projects a branch to each gland, on which it again divides in two or
three parts (fig. 29). I may add that the septal glands are very large, almost filling
the respective somites, and as compared with those of De/tania elegans, about three
times as large, considering however the relative size of the two species. The glands
are nearly similar to those of Deltania Benhami, but with more unequality as to size.

Salivary or Pharyngeal Glands (fig. 26). These glands resemble those of the
preceding species, Deltania elegans, in general appearance. There are two very long
glands behind the brain, attached on the underside of the two long muscular bands
which stretch upward. The anterior one of these is the smallest and rather short. the
second in order from the brain is the longest. The posterior gland, which forms the
posterior projection of the pharynx, is much shorter, more compact and rounded
than the corresponding gland in Dedtania elegans. The whole mass of glands projects
much less posteriorly than the glandular mass of the pharynx in that species.

Spermatheca (figs. 30, 31, 32 and 39). These organs are prominent and char-
acteristic of the species. There is one pair in somite ix opening in the inter-
segmental groove between that somite and viii. The external pore is in front of the
inner couple of setze, but not interior to the sete. The organs are thick, opaque and
of the form of pointed saes, each one with two diverticula, one on each side, which
connect with the main sac close to the external pore. The outline of the sae is irreg-
ular in some places, toward the inner apex assuming the appearance of one or more
warty diverticula, which, however, never assume the size of the diverticula. Of these
latter there are one pair which are slender, cylindrical, of more or less irregular out-
line with the apex sometimes slightly wider, sometimes helix-like, turned on itself.
The lower part of the spermatheca is muscular, but this muscular part is quite small,

36 CALIFORNIA ACADEMY OF SCIENCES.

not extending above the junction with the diverticula. In general form the sperma-
theea resembles that of the following species, D. Benhami, but the outline of the main
sae is less regular and the diverticula are larger, being about one-half as long as the

main sac (figs. 30 to 32).

Testes. As usual there are two pair of small testes occupying somites x and
xi. They differ in nothing particular from those of the other two species of the

genus.

Sperm-sacs. ‘There are two pair of sperm-sacs, one each in somites x and xi.
They are long, not very opaque bodies, occupying a large part of the somite, though
not filling it closely. The spermatophoric spherules are comparatively large,
globular. The sperm-sacs are not greatly lobed, extend considerably in length from
the dorsal to the ventral side, and are of more undecided shape than the sperm-saes
of Deltania elegans and Benhami. They are also, comparatively, much larger (prin-
cipally higher) than in that species. It will be noticed that the sperm-saes occupy
somites x and xi, while in Detania elegans, Benhami and dubia they occupy somites

xi and xii.

Ovary and Oviduct (figs. 35, 36, 37, 38). As might be expected the ovary is
situated in xiii. It offers no characteristics of interest. There is no ovisac. The
oviduet is as usual funnel-shaped, either deeply cut or folded on one side, the inner
funnel in xiii, the ovipore in xiv. Close to the oviduct in xiy there is a very peculiar
sac (figs. 35 ef seq.), of the size of the oviduct or smaller. It does not open directly in the
oviduct, and it has the general shape of aseptal gland with many rounded lobes arranged
as the petals in a rose. The epidermal covering does not closely cover the inner cells
which are irregular, apparently not closely packed, of uneven size and shape, round
or conical, each with a round nucleus, and grainy cell contents. This gland does not
connect with the opening of the oviduet funnel, from which it is separated by the
septum between xiii and xiv. There is one pair of those glands, one behind each
oviduct. The gland is affixed to the posterior part of the anterior septum in xiv. A
similar gland does not exist in Deltania elegans. As to the nature of this organ IT ean
say nothing definite, and I hesitate to consider it as an ovisac, until a more extensive
material will allow other investigations.

Spermduct and Prostate (fig. 33). As in the preceding and following species,
the spermduet and prostate open in the same pore in somite xvii, The spermduet
opens slightly posterior to the prostate and also more outwardly, though both organs
closely join at the pore. The spermduct is quite wavy throughout its length to the
ciliated rosettes, which as usual are found in x and xi, The rosettes are less folded
and crenate than those in Deltania elegans. The prostate differs somewhat from the
one in the latter species. The muscular duct is not helix-like at its upper end, either
straight or bent at right angle to itself, with the distal end pointing backward, being
parallel to the longitudinal axis of the body. The relative size of the prostate is in

CALIFORNIA EBEUDRILIDA, 37

this species much larger than in Del/tania elegans, which appears to be characteristic
also with nearly all the other organs. The upper glandular part of the prostate is
about three times wider than the lower muscular part. The latter is about equal in
length to the penial setee and their sacs.

Genital or exterior male zone (fig. 25). In somite xvii there is a pair of ven-
tral papillee close to the ventral ganglion, and situated in the transyerse median line
of the somite. In these papillze open the penial setee, in the place which otherwise
would be oeceupied by the regular sete. Between these papillee and the ventral
median line of the body, somewhat nearer to the posterior margin of the somite, are
seen on either side a circular cup-shaped depression, from the center of which is spread
backward a large fan-shaped branch of muscles connecting with the posterior inter-
segmental groove. In the center of this suctorial organ, and at the very point from
which the fan-shaped muscular fascicle starts, is situated the exterior opening of the
spermduct and the prostate. In the median line between the suctorial cups is a smaller
triangular depression. The anterior part of the somite is raised and thicker than the
posterior part, or rather there are two large anterior folds, and several smaller posterior
ones in the vicinity of the male pores (fig. 25).

Deltania Benhami.
Plates xv and xvi, figs. 40-48.
Deltamia Benhami Eisen, Zoe, iv, 212, October, 1895.

Size about 1 inch by ;;. The inner couples of setee as well as the sets in the
inner couples are much closer together than in any other species. The spermatheca
are large, opaque, situated in ix, and opening between ix and viii, with two diverticula,
which are less than } as large as the main spermathecal sac. A small species, in
many respects resembling Deltania Troyeri, but very distinct by the aboye character-
istics.

Habitat. I have found this worm only in a gulch or ecaiion at the outlet of the
waterworks and dam, known as Lake Chabot, east of Alameda and San Leandro, in
Alameda Co., California. The worm is very searce and lives under damp leaves in
the very top layer of the soil around the roots of trees. The exact locality is to the
right of the gate which closes the reservation, down by the creek, not far from the
wire fence. It occurs here alone, not mixed up with any other species, and to all ap-
pearances this species is a true native and not introduced. It is an exceedingly deli-
cate worm, almost transparent, white, with yellowish clitellum, very impatient of
being handled and can only be kept alive with great care. It is much more trans-
parent than any of the other species.

Exterior characters. In general appearance, the worm resembles e/tania
Troyeri, but is slightly larger in size. The second somite is much narrower than in
that species, being larger than the third somite. But it is especially as regards
position of the sete that the greatest external difference exists (fig. 40). The ven-
tral setee in one species are much closer together than in the other species of the

to) CALIFORNIA ACADEMY OF SCIENCES.

genus, and average about twice or three times as close as in Deltania Troyert.
This is a prominent and constant characteristic. The characteristic feature of the
genus, which consists in the narrowing together of the ventral setze on both sides
and towards the male pore in xvii, is readily seen in this species. On account of
the already very close approach of the ventral setee of each couple, this narrowing
together is, however, less perceptible in our present species, but it is still consider-
able and readily seen. The figure (fig. 40) will illustrate this better than any
lengthy description. Not only does this approach of the sete exist on both sides of
the male pore, but it is also seen towards the spermathecal pore, though here in a
smaller degree. In Deltania Troyeri, there is not a trace of such an increased ap-
proach of the ventral setee towards the spermatheeal pore. Another characteristic as
regards the setae is that the ventral or inner couples are much closer together than
in the other two species described here, so close that the main bodies of the sperma-
thee almost touch.

The spermathecal pore is situated in the intersegmental grooye between vili
and ix, in front of seta 1. In Deltania Troyeri, the spermatheeal pore is in front of
the corresponding seta 1, but considerably more lateral, and on account of the
greater distance of the ventral couples of setae, the spermathece in that species are
much further apart, and do not crowd the ventral ganglion. Ovidueal pore as usual
in xiy. Penial setze on papille in xvii, opening in the male pore. The clitellum
comprises three full and wide somites, xiv, xv, xvi, and two smaller ones, xii and
XVli, as in other species.

The genital region (fig. 48) around the male papillee is much simpler than in
Deltania Troyeri. The two papille are situated much closer together, and there are
no suctorial depressions with their fan-shaped arrangement of muscles. The penial
setee are very slender, sickle-like, bent at the free apex, with a narrowed and sharp
point. There are a dozen or more external fibres of exceeding thinness stretching
from the anterior part of somite xvii to the posterior part of xvii. These
threads, however, exist only in the longitudinal groove between the papillee, all run
parallel with each other and with the long axis of the body (fig. 48). As to their
nature Tam not at all certain. They are apparently too thin to be muscles. The
clitellum proper in this as in the preceding species ends at the papillee or male pore.
Anterior to it the clitellum consists of regular flask-like cells; posterior to it again
the body-wall has assumed its regular appearance. In Deltania Troyeri the elitellar
thickening stops just before it reaches the papillee, but in Deltania Benhami it stops
just as it reaches the male pore. The papillee, however, continue to the posterior end
of the somite.

The length of the species is about 1 and 14 inch, by ;'; to $ inch wide, taper-
ing towards the caudal end. In faet the general form and size does not differ from
the preceding species. In all I found some fifteen specimens, but they were, un-
fortunately, in poor condition when I arrived home, and the part posterier to the
clitellum had already begun to decompose. The following account of the anatomy
is, therefore, not as full as desirable, but enough is known to perfeetly characterize

CALIFORNIA EUDRILIDA. ow

the species. There are four pair of septal glands in vy, vi, vii and viii. The glands,
as far as I can judge from dissections only, are of almost equal size, somewhat longer
there than in Deltania Troyert. They are equally developed on the lower and upper
side of the @sophagus. The alimentary canal offers no great peculiarities. The
specimens were all very much stretched, and I am not certain if the form of the
csophagus will prove constant. However, the contractions at the septa were much
smaller than in any other species. The cesophagus from somite ix narrowed down
toward somite xii, here it began to gradually increase in width, but the sacculated
intestine begins evidently first in xvii, increasing in width gradually backwards until
it reaches the region between xxvii and xxxy, where it suddenly narrows and con-
tinues as narrow to the end of the body.

Spermatheca (figs. 42 and 43). There is one pair in ix, opening between viii
and ix, of the same general appearance as in Deltania Troyeri with minor character-
istic details. The main sac is ovoid and somewhat lunate, pointed, with very smooth
outline and with no trace of warty excrescences. There are two diverticula affixed
halfway between the base and the glandular part. They are much smaller than
those of Deltania Troyeri, being less than one-third as long as the main spermatheca,
while in Deltania Troyert the diverticula are one-half or more as long as the main
spermatheca, and affixed to the muscular part close to the base. The spermathece
open in front of the Ist setee and are situated much closer together than those in
D. Troyeri. The opaqueness of the spermatheca is the same as in the latter species.
There is, however, a decided difference in the location of the spermatheca. In
D. Troyer they are situated so far apart that they do not touch the ventral ganglion.
In D. Benhami, however, they crowd it, this approach being caused partly by the
closer proximity of the ventral or inner couple of sete, partly by the situation of the
spermathecal pores which in our present form are more ventral to the sete.

The sperm-sacs (fig. 41) are of a very characteristic form. They are larger in
proportion than those of D. e/egans, but not quite as large as in D. Troyeri. There are
two pair, one each in ix and xii, and they do not connect with each other.
Each one consists of a very large flesh-like lobe, at the base of which are seen half a
dozen smaller and globular saes, all connected at the place of adherence to the an-
terior septum. These different lobes, the large one as well as the small ones, are
full of rounded, oblong or irregular spermatophoric sphzerules. The sperm-sacs do not
by far fill the somites. The form of the sperm-sacs varies to some extent, but the
main character is the same in all, a large, rounded or flask-like lobe, at the base of
which are several smaller ones.

Spermduct and ciliated rosettes as in D. Troyeri. The prostate offers the char-
acteristic of having the lower part of the glandular sac considerably swollen, conical
and gradually diminishing in size towards the apex, which again is slightly enlarged
like a knob. There is only one long slightly bent or almost straight seta in each of
the penial seta-sacs, both setee opening on a small papilla, and which as far as [ can
see more resembles that of D. Troyeri than D. elegans. The blood is very pale yellow,
paler even than in the other species. There are three strongly pulsating hearts in

40 CALIFORNIA ACADEMY OF SCIENCES.

somites x, xi and xii, connecting the dorsal and ventral vessels. There are also
connections between those two vessels in ix, x and xiii as in the other species, but
those connections consist of regular secondary vessels narrow and tubular, and in no
way resembling hearts. There is no subneural vessel. The ventral vessel is divided
in somite vi, and in the anterior six somites there is a heavy capillary system connect-
ing the ventral and dorsal vessels.

In the following I have endeayored to give a comparative table of the species
which I refer to Deltania. As far as concerns those species which I have not myself
investigated the table must be considered tentative, especially so in regard to the char-
acter of D. dubia, the description of which by Fletcher is somewhat insufficient. I need
here hardly add that Deltania Poultoni is the species described from Maderia by Bed-
dard as Microscolea: poultoni. The description given for D. dubia is taken from Rosa’s
description of Microscolex dubius Fletcher, and not from Fletcher’s own.

DELTANIA.

x — ===

D. elegans un. sp. D. Troyerin. sp. D. Benhamin. sp. D. Poultoni Bedd. D. dubia Rosa.

Septal glands in vi, vii, vili, ix. Vv, Vi, vil, viii. Vv, vi, vii, viii. Not present.
Sperm-sacs in nah BON Xx) oxd. xi, xi. xi, xii. xi, xii.
Spermatheca Very small and Large, apart, with Large, close togeth- Not present. Not present.
pellucid. No di- 2 diverticula each. er, with 2 diverti-
verticula. Opaque. cula each. Opaque

The ventral or inner Only a little closer Only a little closer Several times closer Several times clos- Several times clos-
sefe in each couple, together than the together than together than er together than er than the outer
exclusive.of those setw of the outer those of the outer those of the outer the outer couple.) couple.
in the genital re- couple. couple. couple.
gion, are:

Prostate, the gland- Helix-like at the Not helix-like; tub- Not helix-like; tub- Helix-like at the
ular part top. Museular ular, straight or ular and broad at top. Museular
part not very bent. the base. ‘Top en- part very short.
short. larged knob-lke.
Gizzard None. None. None. None. Rudimentary.
Diverticula of the Absent. More than }as large About 4 as large as
spermatheca as the spermathe- the spermathecal
cal pouch. pouch. ~
Penial sete Two on each side.) Two on each side. Two on each side. One on each side. Two on each side.
|
Nephridia com- ii. ii. | ii. | ii. v.
mence in
2 aa j | - ;
Nephridio-pores in/4th and 3d setw.| 4th and 3d sete. | 4th and 3d sete. 3d setie, 3d setae.
front of and little) Prominent in the) Not prominent in
interior to clitellum. the clitellum.
Spermducts Remain separate Remain separate up Remain separate up|Remain separate Unite to form one
up to the atria. | to the atria. | to the atria. | up to the atria. tube on each side
| which opens in
the muscular part
of the atrium.
Posteriorly to the xxvi and xxyii. Xe | XXxl. Xxili and xxiv.
clitellum the sete |
have regained their
normal distance in |
somite |

CALIFORNIA EUDRILID®. 4]

ARGILOPHILUS.
Argilophilus Eisen, Zoe iy, 252, October, 1893.

Prostomium encroaches on somite i. Eight setse in four couples, ventral,
lateral and dorsal, commencing in somite 11. The sete of the inner couple not con-
verging towards the male pore, but closer set than the sete of the outer couple. The
alimentary canal consists of an eversible buceal cavity, a pharynx, cesophagus, gizzard,
tubular intestine, sacculated intestine, typhlosole, but no cesophageal glands or pouches.
Clitellum not developed ventrally, occupies somites xiii to xviii.

Spermathecal pores, one pair vii /vili and one pair villi/ix. Ovipores in xiy-
Male pore in xvili. One or two rows of ventral intersegmental papille. Two pair
of spermatheex. Testes in x, xi. Sperm-sacs paired in x, xi, xii, generally enclos-
ing the ciliated funnels and testes. Two pair of ciliated funnels. Two pair of sperm-
duets, which join a pair of very large, tubular-coiled prostates in xviii, at the upper end
of the muscular duct. Two penial setee open in the same pore, but not in the same
duct as the prostate.

Dorsal vessel and ventral vessels connected by 5 pair of hearts in xiv to x. No.
subneural vessel. Blood red. Many blood vessels on the nephridia. No pepto-nephri-
dia. The nephridio-pores variable as to location, the majority open in front of or lateral
to the 4th sete, though many open interior to the 4th sete. No ccecal bladder at the
exterior pore. Large earthworms with thick round bodies, of pale flesh color, mar-
bled bluish.

Distribution and habitat. The genus Argilophilus appears to be an undoubted
native of the Pacific Coast. Specimens have been found in the San Joaquin Valley
in California, and as far north as British Columbia (Vancouver Island). In California
these worms are our most common earth worms, appearing close to the surface with
the advent of the rains in the autumn and disappearing deep in the soil with the dry
weather in May, after which time they are not any more found in even locally moist
places. During the summer months I have sometimes dug up these worms from a
depth of 5 to 6 feet or more, each worm tightly rolled up as a little ball and appear-
ently encysted in a chamber of clay, the inner surface of which is smooth and hard.
In these cysts the worms pass the dry season. ‘These worms are hardly ever found
outside of heavy adobe or clayey soil; the more clayey the soil, the better the worms
appear to thrive, provided also the soil is rich and fertile. In poor soil the worms are
seldom seen, and the best indorsement for a soil is that it contains worms of this genus:

The color of the worms of this genus is fleshy pink, thickly marbled, with steel
or slate gray, (fig. 1382). The clitellum is yellowish red, and the whole anterior part
is pinkish. The color of these worms is very handsome and distinguishes them from
the deep brown Allolobophora so common in moist or swampy places in this State.

Exterior characteristics (figs. 125-151). The most prominent exterior feature
of this genus is the color which has just been described. Another is the frequent
eversion of the lining of the buccal sae (fig. 130). As to size the worm must be

42 CALIFORNIA ACADEMY OF SCIENCES.

considered as our largest earthworm, though very variable, as might be expected.
The smallest adult worms measure about 15 inches by 3 lines, the largest again 6
inches by 4 to 45 lines, the size appears to depend greatly on the locality and richness
of the soil. The prostomium divides somite i to about 5 or % (fig. 180). Somites vii,
viii, ix, are larger than the other anterior somites. The clitellar somites (figs. 125 to
151) are large, the post-clitellar ones are very much smaller. The spermathecal
pores are more or less conspicuous (fig. 129 spth.), sometimes hardly visible, at times
again elevated and appearing as small round rings. They are situated just lateral to
setee 2, but of course in the intersegmental grooves between vii / villi and yili/ix. The
ovipores are closer together, situated a little more ventrally than sete 1, sometimes, if
not generally, connected by a depression. The clitellum is only developed dorsally
and laterally, the ventral part between the male pores being normal and appearing as
considerably depressed, in hardened specimens the depression reaching as far forward
as to the center of somite xiv or the oyipores.

The male pores which open in line with sete 2, are situated on either side on an
elongated papilla, which again is more or less surrounded by a circular depression,
outside of which is seen a high semi-circular ridge, which is thicker anteriorly and
posteriorly than laterally (fig. 25, 129). The penial setee are one pair in each
pore, and are seen protruding through the male-pore.

The regular setee are sigmoid, not greatly bent. The sete of the inner couples
are closer than those of the outer couples; all the setee are in parallel rows (fig. 24).
The nephridio-pores are difficult to view from the outside. Their arrangement is
variable, but the majority are found outside of, or more lateral than the fourth row of
setee. The three anterior nephridio-pores are seen in front of sete 4 (fig. 24). A
more detailed description will be given further on.

Setw. The ordinary sete have been already described as sigmoid. There
are two sacs of penial sete attendant to each prostate, and opening in the same pore,
but not in the same duct, as that organ. There is only one seta in each sac. This
seta is sickle-like, much more curved than those in Deltania. The point is needle-
like, and curved (figs. 122, 123). The very point is void of sculpture, but the part
back of the point and up to the sac is sculptured as in the figure 125. The largest
part of the seta is smooth, only showing the rings for the attachment of the muscles.
The inner couples of setee of the clitellum are somewhat raised, though otherwise not
differentiated.

ANATOMICAL STRUCTURE.

The body-wall (figs. 114, 115, 116, 117, 118). The body-wall outside of the
clitellum shows the usual sets of layers. The innermost vascular layer, which covers
the longitudinal muscular layer, is very thick and prominent, though not very
crowded with blood vessels (fig. 114). Under this layer, and between it and the
longitudinal muscles, passes the spermduct, almost throughout its length, from the
place where the two ducts unite to the one where they rise to join the spermduct.
This layer is less pronounced anterior to clitellum, and appears absent in the vicinity

CALIFORNIA BUDRILIDA. 15

of the spermatheea and anterior to them. In the elitellar somites this yaseular layer
is especially prominent. The muscles of the longitudinal muscular layer in this
genus, as well as in Deltania, are arranged in groups or projecting lobes, between
which pass projections of the vascular layer, as well as transverse muscles in certain
places. These lobes vary in width, and on the ventral side below the ganglion are
arranged fan-like (fig. 103, 7. p.), diverging from the median line. The longitudinal
museles are never arranged around a central axis, as is the case in so many |um-
bricides, though they show a faint trace of symmetrical arrangement. The zone of
the transverse muscles is much thinner, five or six times narrower than the longi-
tudinal zone (117).

The hypodermis is thick, with large, glandular cells of a flask-like or spindle-
like shape. In the clitellar somites these glands become irregular, club-like, and
project as far inside the layer of clitellar glands as the hypodermis is thick, or more

(fiz. 116).

The clitellum is developed only dorsally. The glandular layer is much
thicker laterally and dorsally, tapering towards the ventral side, and ceasing entirely
at a line drawn outside of the male papilla and parallel with the ventral ganglion.
The glandular layer of the clitellum is very thin, and as compared with that of
Deltania, about 4 narrower. The cells of this layer are, however, very wide and
long, there being generally eight or nine in the row. They are irregularly grouped
in twos or threes, separated by narrow blood vessels, which, at rather regular inter-
vals, are thicker. They are supplied with blood from sinuses situated between the
transverse muscles, and which connect through these branches with a capillary net-
work on the hypodermis (figs. 116 and 116).

Transverse muscles (figs. 118 and 119). There are numerous sets of trans-
verse muscular bands in the clitellar somites, quite similar to those described by
3enham in Moniligaster indicus. They are more numerous and prominent in the
somite of the oviduct than elsewhere, and form there three distinct muscular bands
the ventral ends of which terminate at the inner couple of sete (fig. 119); the lateral

>

ends again terminate on the lateral side of the body wall.

The posterior band is the smallest of the three and begins in the posterior and
ventral part of the somite, in line with, but slightly posterior to the inner setie, stretch-
ing from there diagonally aeross the somite, ending laterally at the seta 5 (fig.
IS iy We

The next band is much larger and begins in the anterior part of the somite
also in line with the inner setie and stretches diagonally backwards ending posterior to
but in line with seta 4 (fig. 119, m 2.) The third muscular band is of the same
size and runs in the same general direction as the last, begins and ends in front of it,
ending in front of the fourth seta (fig. 119, m3.) A fourth muscular band of a
somewhat similar character connects the posterior part of the oviduct with the body-
wall terminating in front of the muscular band just described as m 1, on the figure
(fig. 119, m 4.) Similar muscles as the oyiducal one, are common in all earthworms,

44 CALIFORNIA ACADEMY OF SCIENCES.

connecting the yarious organs with the body-wall and require here no particular men-
tion.

In the other clitellar somites we find bands of transverse muscles which
stretch diagonally (fig. 118) across the body wall from the ventral part, almost im-
mediately below the ventral ganglion, to the vicinity of the outer sete. The ends of
these muscles penetrate between the lobes of the longitudinal muscular layer (fig.
118, 7. m.) These muscles are not continuous through the somite but are grouped in
bands.

Ventral papille (figs. 120, 125 to 131). Argilophilus is readily distinguished
from other California Eudrilide so far known, by its ventral papillee, occupying the
ventral side of the intersegmental grooves from the spermatheca to the segments next
posterior tothe clitellum. In Argilophilus marmoratus ornatus we meet sometimes with
as many as 7 or 8 pairs, while in A. m. papillifer we find as high a number in a single
median row, under the ventral ganglion. In no instance did I find a papilla on the
segment itself, all invariably occurred rising from the groove between the somites,
being in other words intersegmental. From cross-sections it will be seen that the
papilla consists of two distinct parts, one exterior and lateral, consisting of elongated
hypodermic supporting cells, which more or less fully enclose the interior main body
of the papilla. This central part consists of larger or smaller gland-like bodies, vary-
ing in size and number according to the size and age of the papilla, ete. In some
papille these bodies fill the larger part of the papilla, in others they are confined to
the bottom or very center of the organ. As to the nature of these organs I am, as yet,
somewhat in doubt, though they certainly must be considered as sensory organs
of some kind. Stained with osmie acid the bodies present in sections a darker center,
which appears of ganglionic nature, around which are grooped larger sacs which
again are composed of smaller, light-refractive granules, giving the idea of a reticu-
lated protoplasm. Numerous nerve fibres connect with those bodies, and evidently
penetrate to the gaglionic center. Long tube-like cells butt directly on these gland-
ulous bodies, while others grooped in bundles penetrate between them connecting with
nerve ganglia. The transition between the supporting cells and the drainpipe cells
of the papilla is sudden and not gradual. In fact the line of demarcation between the
two is quite prominent. The supporting cells in the young papilla, entirely enclose
the papilla, while in the larger and full grown papilla, they are pushed towards the
side leaving the whole center to the drainpipe cells and the glands.

When stained with safranine the nuclei of the glandular cells become prominent,
less so when stained with hematoxylon. The nuclei vary greatly in size and number.
In some of the glandular bodies there are from one to three nuclei,in others none. In
sections there are always less nuclei than divisions (cells?) apparently lying on the top of
the reticulated mass. The nuclei may be seen directly above the nerve center, but
generally they are found outside of it. It seems as if these glandulous bodies are
modified agglomerations of hypodermic glands.

The arrangement of the glandulous bodies and the tube-like cells is very

CALIFORNIA EUDRILIDA. 15

regular, the former resembling small sugar-loaves standing in a row, with the fan-like
arranged tubular cells between them (fig. 120). In some sections a bunch of nerve
fibers is seen on either side touching the papilla, connecting on the other hand with
the ventral ganglion. As to the nature of the papilla Dr. Michelsen suggests that a
somewhat similar organ in Acanthodrilus georgianus is a taste or * Wollust”? organ.
It is, however, not unlikely that the minute light-refractive bodies are glandulous.

Many of the tubular cells contain a fine granulated secretion in yarying quan-
tities, which stain dark red with eosine. The fact that the whole papilla is concave in
the center speaks also strongly for the glandulous nature of the organ. The figure of
the “Augenapfela”-lke organ of Acanthodrilus, given by Michelsen, may possibly have
been taken from a young papilla in which the glandulous bodies had not yet developed.
Organs of a somewhat similar nature have also been described by Horst from Ponfo-
scolex corethrurus which, however, he does not figure in connection with nerve fibres,
It is not improbable that all the sensory organs in the genital somites of the higher Oligo-
chieta are of an analogous nature. Among such organs would be included the tuber-
cula pubertatis, the puberty grooves, as well as some other epidermal structures in
Heliodrilus, Hyperiodrilus and Eudrilus described more in detail by Beddard. How-
ever all the organs require re-examination by the aid of other methods, as one single
method of staining will not suffice to reveal their true nature.

Septa (fig. 86). The anterior septa are greatly pouched, generally to such
an extent that in cross-sections the various organs appear to lie several somites further
back than they really do. Thus the gizzard and the posterior spermatheca may be
seen in the same cross-section and this is also the case with the oviduct and the sperm-
sacs. This pouching is principally restricted to the septa 5 to 15. These septa are
also slightly thickened especially those bounding somites viii to xi.

Nephridia (figs. 59 to 77). The position of the nephridio-pores places Argilo-
philus very close to Plutellus. There are one pair of nephridia in each somite as usual.
The first pair of nephridia are found in somite ii and others follow in all the posterior
somites. The first five or six nephridia are somewhat larger than the others and open
in front of the fourth sete. All the following open irregularly in front either of the
third or fourth sete, or in the space between and anterior to them, or even outside of,
or more lateral than seta four. Those which open laterally of seta four do not even
open in the same row, as we find one nephridio-pore say as far out laterally from
the fourth seta, as that seta is distant from the third, while others are half-way or one-
third of the way between the fourth seta and the most lateral nephridio-pore. There
is no regularity as regards this succession, though rarely two successive nephridia open
in line behind each other. For instance, one nephridium opens in front of seta
3; the second as far outside of 4 as 4 is distant from 3; the third nephridio-pore is
in front of seta 4, the following half-way between seta 4 and the most lateral pore,
the following % of the distance from seta 4 to the most lateral pore, the following in
front of seta 4, the following again % of the distance from seta 4 to the most lateral
nephidio-pore, ete. (Fig. 124 np. p.)

Memoirs, Vou. II, 3. January, 1894.

46 CALIFORNIA ACADEMY OF SCIENCES.

The structure of the nephridium is considerably complicated. The nephridio-
stome, found in front of the ventral sete, is unusually minute, and leads to a com-
partively short duet, which connects with the body of the nepridium proper, at the
point where the single-tubed outlet leaves the folds. This tube is not conyoluted (fig.
59 G@)), but almost straight, very hard and solid, widening toward the base. When
it reaches the folds it does not at once enter a tube, but forms a long, cylindrical,
spongy mass (fig. 63), which extends the whole side or one-third of the length of the
nephridium, before it assumes the proper shape of a clear canal (fig. 59, @) to @),
and fig 64). In the beginning at (@), this mass shows no regular lumen, but a num-
ber of irregular pores and tubes, which might best be compared to the inner canals
of a common washing sponge (fig. 64, @),@,@)). In the center of this mass are
imbedded the two parallel folds of the main nephridial canal (fig. 64, a@ and 6). At
first the spongy tube is located principally above the two canals, but soon the mass is
shifted and the canals become imbedded in the center of the mass, or very nearly so
(fig. 64, G)toG)). The small connecting tubules at (2), which were at first so ir-
regular, soon assume a greater degree of regularity at (8), while at the same time two
longitudinal lumens are formed—one on the upper (@) and one on the under side
(@),, fig. 64) of the two central canals (a and }). At first these canals (@) and @)
are indistinct and irregular, but soon they assume the character of regular longitudinal
canals (from () toG@), @ and @). From @) to @) these canals (@) and G) are
connected by the transverse tubules, which completely surround the two central
canals (a) and @). At G) the lower canal G@) becomes narrower, and the trans-
verse ducts drop into the main canal (©), while the former lower canal @) assumes
the character of a rather thick, epithelial lining. The general effect of this arrange-
ment is, that, seen with a lower power, the spur between (¢) and () appears to con-
sist of four distinct parallel canals, while from («) to () the fold contains only three
parallel canals. At (¢) these canals become very crowded, the whole fold being nar-
rower. The length of this narrower part varies, but generally already at (), or
shortly before entering on the bent plexus at (®), the fold has regained its original
width. At @) the canals turn; that is, @) connects with @) and the central canal
folds upon itself, and for a short distance we have four almost parallel canals. At
(3) the formerly central canal (from (3) to @), 2) and @)) leaves the lobe and crosses
over to the other fold, and at () becomes the original canal @), which runs its
course all through the two folds, at @) turning downward, becoming the lower canal,
which again at @) becomes canal ©), from there on pressing backward and again
forward from (@) to @), at which point it separates itself from the fold, and, running
along the inner body-wall, forms the outlet duct opening at the exterior pore at (°).
The connecting bridge between the two main folds, or rather between one fold and
the free lobe of the other, between (@) and (®), is very narrow at (1), suddenly
increasing at (4), gradually tapering toward (5). As regards the ciliation of the
canal, it may be stated that it does not extend all through the tube. It appears that the
canal is ciliated at places where the passage of the excretions is difficult. The narrow
duct from the funnel to () is ciliated as well as the funnel itself. Again, the ciliation

CALIFORNIA EUDRILIDA, 17

begins at the point of recurrence at @); but ceases after the duct has assumed its straight
course at (8); it begins again at (@), and is with a certainty found between (©) and
(@), and possibly between (@) and (2), where, however, I am not quite certain of
its presence. The whole nephridial fold is imbedded in a wing-like cellular and
fibrous mass, which at either end becomes thicker, supporting large pellucid perito-
neal cells, such as found in the nephridia of many Oligochieta (fig. 64, per. c.), and a
complex system of blood-vessels, which branch and form capillary ioops partly on the
folds, but principally between the spongy tubes and the two central canals (figs. 59,
63, 64, 67, b/.). The blood-vessels originate from a branch of the ventral yessel as in
Lumbricus.

Scattered over the peritoneal cells are masses of free cells, generally agglom-
erated in separate heaps. Each cell is deeply crenate, as if it consisted of several in-
dividual cells, but in each such small agglomeration there is only one round nucleus
(fig. 80).

If we recapitulate, we find that the nephridum of Argilophilus consists of the
following distinct part from the nephridio-stome to the nepridio-pore:

1. Nephridio-stome, engaged in the anterior septum.

2. Narrow duct, which connects the nephridio-stome with the body of the ne-
phridium proper, especially with the spongy tubules.

3. Spongy tubules, which are at first irregular, but which soon fuse into one
main-tube with many branching tubules.

4. Main nephridial canal, which, recurring on itself several times, forms two
distinct folds and one spur. The posterior fold contains besides the tube and tubules
two turns of the canal, one of which is recurring. The anterior fold contains three
turns of the canal, two of which are recurring. Part of this anterior fold disengages
itself from the main nephridial mass and forms the

5. Spur. This spur contains four canals, two of which are recurring, the
point of recurrence for all four being at the distal end. One of the recurring canals
of the spur is connected with the posterior fold by the

6. Bridge, a part of the canal much narrower than the other, spanning the
distance between the two folds.

7. The wide duct leading to the nephridio-pore, directly connected with the
recurring canal of the posterior fold. In different nephridia this wide duct is of
varying length.

8, Nephridio-pore, apparently without urinary bladder or collar.

9. A large wing-like, but rather thin, mass of peritoneal cells.

10. A complicated and extensive system of capillary blood vessels on the va-
rious parts of the nephridium.

Compared to the nephridium of Lumbricus as described by Benham we find
a few more important differences, especially in the post-septal part. Thus the post-
septal part of the narrow tube after reaching the first loop assumes the shape of a

48 CALIFORNIA ACADEMY OF SCIENCES.

tubular network, which only gradually fuses itself into one duet with numerous
branching ductules, while in Lumbricus the single duct continues through the differ-
ent lobes and in recurring winds around itself. The tubules in Argilophilus em-
brace the two tubes but cease after leaving the first fold. The muscular duct which
is so prominent in Lumbricus is not represented in Argilophilus, but is replaced by an
elongation of the single wide tube corresponding to the “ wide tube of the 3d lobe” in
Lumbricus.

Compared again with the nephridium of Deltania, we find that the principal
difference consists in the absence of the urinary bladder and the collar at the ne-
phridio-pore as well as in the absence of a eweal bladder. The irregular or alternate
locations of the nephridio-pores distinguish Argilophilus and Plutellus from all other
earthworms, as far as known.

Alimentary canal (fig. 87 to 92). The buccal cavity is, as has been stated,
greatly eversible, and generally remains everted after the worm is dead. The
pharyne is only developed superiorly though there is a slight thickening of the lower
wall of the buceal cavity at the junction with the cesophagus, at which place numer-
ous muscles are seen to connect with the lower part of the body-wall. The pharynx
is as usual furnished with numerous salivary glands (fig. 86, s/. g/.), extending from
the vicinity of the brain to the posterior part of somite iy, the most posterior glandular
mass being the largest (fig. 86). The various glandular lobes offer some characteristics,
which if carefully noted and compared may be found to be constant enough to serve
as species characteristics.

The Qsophagus begins in iii and occupies somites iii, iv and y, (fig. 86, A. w.)
forming first a narrow tube, which widens out, and rising upwards connects with a
very large gizzard (gz.). This gizzard occupies in reality only somite vi, but its great
length causes it to push far backward to such an extent that it actually occupies the
space covered by viii and sometimes by ix. The gizzard is compressed from above,
but widened laterally which makes it appear very much larger when viewed from
above than when seen in vertical section. It connects posteriorly, in vii, with a very
long narrow tubular intestine, which extends to somite xii, but which in somites xiii
to xv is strongly nipped by the septa and considerably enlarged (fig. 86, s,) without,
however, being strictly sacculated.

The sacculated intestine proper, however, begins first in xvi, and is much wider
than the other part of the alimentary canal. The structure of the pharynx offers
nothing of unusual interest. The wall of the gizzard contains the usual layers, but they
are poor in blood vessels. The longitudinal muscular layer is thickest, and on the
widest part of the gizzard it is five or six times as thick as the epithelium, gradually
diminishing in size anteriorly and posteriorly (figs. 86, 87, 88, 89).

Pear-shaped or Chylus Chambers. Beginning immediately behind the gizzard,
and extending throughout the tubular intestine, we find imbedded between the
epithelial folds numerous pear-shaped organs of doubtful funetion. In pronouncing

CALIFORNIA BUDRILIDA. 19

these as chylus chambers, I do so with much hesitation, as I am not at all satisfied but
that instead of being organs of absorption, they may not in reality be organs of dis-
charge, or in other words, glands. TI found these chambers always empty, appearing
entirely transparent in sections of the intestine, and they neyer stain, apparently
showing a want of contents, though this may be a temporary condition, owing to
the state of the worms when killed, at which time these organs may have happened
to have been temporarily empty. As regards location, they are found principally in
somites vii to xii, and seldom extend further back than that somite. It is to be re-
membered that in somite xiii the tubular intestine changes its form and becomes
considerably sacculated. In somites vii to xii this tubular part is very poor in blood
vessels, or rather in large blood sinuses, while in xiii these sinuses begin to appear in
large number and of large size. Thus, with no blood sinus in the epithelial folds, there
are many pear-shaped chambers, while on the contrary, the cessation of pear-shaped
chambers is accompanied by numerous and large blood sinuses (fig. 93). The pear-
shaped chambers are imbedded between the epithelial cells, and probably all of them
reach the inner cavity of the alimentary canal, though, from the sections made, this is
not quite evident. Some of them, however, do, as will be seen from the figures 90 and
91. They are unequally distributed; in some places they fairly crowd out the epithelial
cells, as in fig. 91, which is drawn from a longitudinal section, showing an unusually
large number. Fig. 92 represents a surface view where they are less numerous,
and fig. 90 represents a camera drawing from a transverse section of the gut. Each
chamber consists of six, seven, or, in some instances, of only two or three cells, arranged
as the clefts in an orange, around a central pore or short tube, which, however, does not
extend all through the papilla, but ends blindly. At the lower end these individual
chambers connect with numerous smaller and more irregular cells, which join
each other close to the transverse muscular layer. The small central pore stands
frequently in connection with the alimentary cavity. In the smaller and inner
chambers this pore is less pronounced or entirely wanting. At the base of the
chambers are seen a number. of small glandular masses, with grainy and opaque
contents (fig. 90 g/.). As regards the distribution of the chambers among the epithelial
folds of the same somite, it may be noted that they are principally numerous in the
dorsal and ventral regions, disappearing in the lateral regions (fig. 133, ep. and c. 7.).
This is the case in all the somites where these chambers appear in the tubular intestine.
Chylus canals have been described by Michaelsen as present in the intestine of En-
chytrzeus, but the difference between their structure and those of Argilophilus is
quite great. In Enchytrieus these eanals are principally intercellular, and connect
with bloodyessels or sinuses, while the pear-shaped organs in Argilophilus are extra
cellular, and oceur in parts of the intestine especially poor in blood vessels. The im-
possibility to procure fresh worms will necessitate a closer study of these organs, to
be deferred to a more opportune time.

as oe
Lyphlosole. The typhlosole and typhlosolar region is small, and not especially
pronounced in front of and in the clitellar somites. Anterior to somite xix the

50 CALIFORNIA ACADEMY OF SCIENCES.

typhlosolar vessel, which is entirely dorsal, is many times smaller than the longitudi-
nal dorsal vessel, but posterior to that somite the typhlosolar vessel assumes a larger
size, almost equalling the main dorsal vessel. In the anterior somites the typhlosolar
region does not project down in the intestine, but is only somewhat wider than the
balance of the intestinal wall. The typhlosole in this part appears to be filled with a
fibrous and spongy mass, in which, however, I have seen no distinct nuclei (fig. 134,
c. ».). These cavities are more oblong at the walls, more round or angular at the
center, where several larger cavities are seen. At the upper margin of the typhlo-
sole this fibrous body gives room to two large and several smaller longitudinal canals,
one at each extremity of the typhlosole, all separated by muscular fibers or cell-like
chambers. In fig. 136 the fibrous nature of the interior of the typhlosole is somewhat
more pronounced than it should be, the fibers appearing rather more regular than in
reality, though the appearance is always as if the majority of the fibers radiated in a
fan-like way from the central spongy mass. This part of the posterior typhlosole is
in cross-section deltoid, with the point projecting into the cavity of the intestine
(136, ty). The size of the typhlosole varies with the individual specimen.

Spermatheca (figs. 81 to 86). There are two pair of spermathecze in viii and
ix opening in the intersegmental grooves between vii and viii and yili and ix. The
spermthecal pores are found in front of and slightly outside of the second sete. As
regards the size the spermathecze may be said to be very large, but unequal, as we
seldom find two of the same size. One or two are generally developed at the
expense of the others and fill all the available space in the somites, frequently pushing
the septa into the nearest somites. Seen in a transverse section of a segment the
larger spermatheca may be occupying as much as three-fourths of the cavity (fig. 86).
The spermatheca consists of two distinet parts, of which the upper is by far the largest,
rounded in outline or potato-like with comparatively thin walls (fig. 82a). This part,
though somewhat warty, carries no diverticula and there are no smaller cavities for
the storing of the spermatozoa. The lower muscular part is twisted, and set obliquely
to the former, but can in no way be said to form any kind of a diverticulum (figs. 81
and 82). The muscular layer, which is a direct continuation of the longitudinal layer
of the body-wall, is only arranged in one way forming a circular muscular stratum of
the spermatheca much thicker at the base than at the top (fig. 82 /.m). This muscular
layer extends to the upper part of the spermatheca (82 a), but is here quite narrow.
The inner glandular layer is singularly well developed (figs. 82, g/. ¢.; 84 g/l. ep. and
83), projecting inwards in large folds like the epithelial folicles of the intestine.

Secretions accumulate as a large whitish mass in the upper part of the sperm-
atheca, and are seen to be sparingly mixed or streaked with spermatozoa. But
the most characteristic part of the spermatheca is the interlacunary system for the
storage of the spermatozoa in the lower or muscular part. A section of this part shows
(figs. 82, 83 and 84, /. s.) a row of chambers imbedded between the epithelial cells,
or between them and the muscular layer, and which connect more or less directly
with the cayity of the spermatheeca by means of narrow passages (fig. 83 p.) Some

CALIFORNIA EUDRILID®. 51

of the chambers stand also in connection with each other as at 84 sp. b. and 83.
In these chambers are stored the spermballs proper, one in each, rarely oceupy-
ing the whole space of the chamber, but leaving considerable of the lacunary room
empty, it never containing any free spermatozoa, only agglomerations or spermballs.
These spermballs may also be seen as white opaque globules from the outside of the
spermatheca as represented in figs. 81 A and Lb, sp. b.

Testes (figs. 94, 95, 96.) There are two pair of testes, one in somite x, one in
xi, as usual post-septal. The anterior testes are enclosed in the sperm-sac, the pos-
terior ones are generally free. Cross-sections show the testes to be deeply multilobed,
with the lobes spreading. As regard the enclosing of the anterior testes it may be
remarked that it is more or less complete, evidently depending on the size of the
sperm-sac. In some specimens the testes were entirely enclosed, in others only the
posterior apex was invested in the sperm-sac. In most specimens the testes were
found pressed close to the body-wall and projecting backwards, the point of adherence
to the anterior septum being immediately above and adjoining the ciliated funnel of
the nephridium in line with the ventral sete.

Sperm-sacs. ‘There are three sperm-sacs, more or less but generally paired in
somites x, xi and xii, the one in the latter somite being much more lobed than the
anterior ones, and more frequently paired. The two anterior sperm-sacs are much
the largest (fig. 96 and 97). They fill the whole somites, are not always paired, but
the lobes connect all along the dorsal body-wall. The anterior sac closely inyests the
testes and ciliated rosettes in x and furthermore often encloses the ventral ganglion (fig.
96 and 97). The sperm-sae in xi is also often closed on the dorsal side, but it does not
invest the testes nor the sperm funnels of that somite (fig. 98). The sperm-sac in xii
differs from the other by being deeply lobed in the plane of the trabecule which are
arranged ina fan-like shape from the alimentary canal. This sperm-sac which is
seldom paired, does not extend below the alimentary canal as the anterior ones do,
but so to say rests entirely on the intestine (figs. 86 and 99). The anterior sperm-sacs
are also traversed by numerous trabeculee, which however are irregularly arranged
(fig. 100). The sperm-saes are frequently infested with parasitic coceidixe of round
shape and with from one to three germ cells (fig. 100, coc.) In size the sperm-saes
are so large that they push far backwards, the one pair covering the other, the
whole mass often reaching as far back as to touch the prostate, thus entirely covering
the ovary and oviducts. This is especially evident in transverse sections when the
ovary, oviduct and the three sperm-sacs may be seen at once.

The ovary is, as usual, found in xiii and offers no great peculiarities. It is
generally deeply lobed, and in cross-section shows the same projecting lobes as the
testes. ;

The oviduct is rather thin and characterized by a thick and long upper lip
(fig. 103 u. 7), which generally is bent to one side. In one instance the upper lip
was forked (as in fig. 103, c.). The muscle attaching the upper part of the oviduct to
the body-wall is unusually strong (fig. 103, m. s.).

52 CALIFORNIA ACADEMY OF SCIENCES.

Spermducts and prostates (figs. 103 to 115). There are two pair of sperm funnels
and ducts, the ducts joined together. The ciliated rosettes or sperm funnels are
found in x and xi. The anterior pair is engaged in the sperm-sac of that somite
(fig. 97), while the posterior pair is generally free (fig. 98). I have, however, seen
the sperm-saes attached to the funnels in these somites in some specimens, and there
appears in this respect to be considerable variation; generally, however, the rosettes
are free (fig. 98) in somite xi. Also with the anterior funnel there is some variation.
In some specimens the funnels were entirely enclosed in the sperm-sacs, in others the
sperm-sac was merely attached to the free surface of the funnel, while in others again
one-half of the funnel was imbedded in the sperm-sac, while the other half was free.
Figs. 97 and 98 show cross-sections with the sperm funnels or ciliated rosettes free or
imbedded.

The spermducts join in xii and continue in a direct line to the prostate, which
they enter in somite xvill. The point of junction is in the lower part of the glandular
part close to the muscular duct (fig. 86, e.). The duet runs between the very thick
vascular layer and the longitudinal muscular layer of the body (figs. 117 and 118, sp.).
The two ducts are never fused together, but continue distinct and separate, though out-
wardly joined, as to the very entrance in the muscular part of the prostate (figs. 112
and 115). While the junction of the spermduct and the prostate is, to all appearances,
in the glandular part of the prostate (figs. 110 and 113), the real point of entrance is
in the muscular duct (fig. 118, spd.). After having touched the glandular part, the
spermducts bend and cross the intervening space to the muscular prostate which they
enter in a slanting direction, then passing considerably downward enclosed in the
muscular part of the prostate, before entering the lumen proper (figs. 106, 107, 110,
111, 112, 115). Fig. 115 represents a cross-section in which the spermducts have
been cut twice. Part of the spermduct is seen free close to the glandular part of the
prostate, part again is seen just at the fusion of the ducts with the lumen of the
muscular part. The duct closest to the lumen has been partly differentiated, the cells
having lost their nuclei.

The prostate consists of a very large cylindrical, but greatly coiled, duct (figs.
106 and 107 and 1), which generally lies pressed flat to the body wall of xviii (fig. 86,
pr.) It opens outwardly in xviii in the posterior part of the somite in the same pore
as the penial setze.

There are two layers of cells in the glandular part, but apparently no muscular
layer between them. The outermost layer consists of large glandular lobes containing
glandular cells which pass between the inner cell layer, and discharge in the lumen of
the prostate (see figs. LO8 and 109). There is a large system of blood vessels which
penetrate both of the cellular layers, but which is not developed to the same extent as
in Deltania.

In cross-section of the body-wall (fig. 103 4) the prostate is seen to open
laterally to the penial sete, though in the same pore, situated at the very junction of
the glandular clitellum and the ventral zone (v. p.) of the body (fig. 103 A.¢ ).

CALIFORNIA EUDRILIDA. Hy}

Vascular system (fig. 86 Band 119). The following remarks on the yascular
system can only be considered as preliminary, a more detailed report being reserved
for a future study of living specimens. The main system consists of two longitudinal
vessels, one dorsal and one ventral. There is no subneural vessel. Three pair of
stout, oblong, thrice-contracted and sac-like hearts connect the ventral and dorsal] ves-
sels in x, xi and xii. The typhlosole has already been described.

The secondary vessels are characterized by numerous bead-like constrictions
and swellings of the smaller branches, especially of those surrounding the funnels of
the nephridia, and of the secondary vessels of the ventral longitudinal vessel.

Of the genus Argilophilus there are two rather distinet forms in California, which
however, only differ externally, and, strange enough, have not, as a rule, been found in
the same locality. Through careful dissection and sectioning I have not been able to
distinguish any anatomical differences between the two species, though I have never
seen any transitory forms as regards to the external markings. According to our
present knowledge of earthworms we always expect to find internal specific charae-
teristics of the species, and if such are not found we must hesitate to consider the re-
spective forms as different species. Under such circumstances I will here refer to
the two forms of Argilophilus only as varieties or subspecies, leaving to further inyes-
tigations, if possible, to detect any internal differences.

Argilophilus marmoratus ornatus.
Figs. 125 to 129.
Argilophilus marmoratus ornatus Eisen, Zoe, iv, 253, October, 1893.

There are two rows of ventral papillee between some of the somites in the vi-
cinity of the clitellum, interior or posterior to it, or both. These papillse are always
more or less in line with sete i, of rounded or slightly oblong form, and generally
more or less of the same size. The number of papillse varies, and frequently one
papilla in a pair is wanting.

Habitat. I have found this worm only in the vicinity of Santa Rosa, Sebasto-
pol, ete., north of San Francisco Bay. It occurs there in rich heavy soils, and con-
stitutes the most common earthworm of the district. In the vicinity of Sebastopol |
found this subspecies both in manure piles and in the wet, soggy places near the la-
goon, places which part of the year must be covered with water from the slough or
lake. Among several hundred worms collected in the month of May during one day,
I found only one single specimen with median papille; all the others possessed lateral
papille (fig. 125 to 128).

The body is thick, cylindrical, and only slightly tapering toward either end
(fig. 132). As usual, the anterior segments are much wider than the post-clitellar
ones. The prostomium frequently protrudes like an inyerted sac hanging over the
peristomium (fig. 130), as is probably the case with most genera of this family.
While the anterior somites are well set off, the clitellar somites are less distinet,

54 CALIFORNIA ACADEMY OF SCIENCES.

though they can always be made out. The clitellum comprises somites xili to xviii,
though the posterior + of xii is generally also somewhat thickened. Of the anti-
clitellar somites vii, vill, ix are thicker than the others, and as thick as the somites
of the clitellum. All the somites are deeply segmented, except the clitellar somites
and those anterior to iii. The external pores are situated as follows: Spermathecal
pores, two pairs, between vii/yiii and vili/ix, a little lateral of the seta 2. Ovidueal
pores in xiy in front of seta 1, generally in a slight depression, bordered by a swelling
of the body-wall. The male pore and pore for penial setee combined in one, open in
line with seta 2 in xviii. The male pore is characterized by being situated on an ele-
vated papilla more or less oblong. This papilla is surrounded by a circular depression
which encroaches on the two adjoining somites (xvil and xix), and this depression
is again surrounded by a lunar ridge, open ventrally, and extending from the center
of xvii to the center of xix, thus reaching slightly outside of the clitellum (fig. 129).
The clitellar glands do not extend further ventrally than toa line running from the
center of the lunar ridge parallel to the sete. The ventral part of the clitellum is
depressed.

Ventral papilla. One of the most interesting features of this genus are the inter-
segmental glands occurring on the ventral side and which in our present species are
paired. They have possibly the same general function as the tubercula pupertatis of
the true Lumbricids, or the ventral papille of Pericheta, ete., which latter they
greatly resemble, but unlike the former they are variable in position, sometimes oc-
curring to the number of as many as seven pairs. In other specimens again they are
reduced to one solitary tubercle, situated on one side of the ventral median longitud-
inal line. The most common form is as follows:

One pair between vili/ix.
One pair between ix/x.
One pair between xy/xyl.
One pair between xvi/xvii.
One pair between xix/xx.

Other specimens possessed in addition to these pairs:
One between xx/xx1.
One between xx1/xxii.

To show the variability of these papille I give their places in a few individuals
picked out at random, and I only add that other variations occur, as may be expected,
from what is shown here. All these specimens were taken the same day and at the
same locality:

No. 1. One pair, vili/ix. .
One pair, 1x/x.
One pair, xv/xvyi.
One pair, xvi/xvii.
One pair, xix/xx.

CALIFORNIA EUDRILIDA, Te)

No. 2. One pair, ix/x.
One single, xv/xvi, left side.
No. 3. One pair, ix/x.
One pair, x/x1.
One single, xiv/xv, right side.
One pair, Xv/xvi.
One pair, xvi/xvii.
One pair, xx/xxi.
One pair, xxi/xxil.
No. 4. One pair, x/xi.
One pair, xv/xyvi.
One pair, xix/xx.
No. 5. One pair, x/xi.
One pair, xix/xx.
The structure of these papillee has been referred to in connection with the
body-wall, and apparently does not differ in the two forms.

Argilophilus marmoratus papillifer.
Fig. 131, A and B.
Argilophilus marmoratus papillifer Eisen, Zoe, iy, 253, October, 1893.

The ventral side of the somites with one single median row of ventral papillie,
which are generally largest between the clitellar somites, diminishing gradually in
size toward the anterior somites. The papillie of this form are generally, but not al-
ways, more oblong than in the preceding form, where they are much more rounded.
The clitellar papillee are frequently diamond-shaped (fig. 131). The papille yary
greatly in number and size in various individuals, but they are always median, never
paired. Sometimes there are 6 to 7 papillie posterior of the clitelium.

Habitat. This worm is very common in the vicinity of San Francisco Bay,
south of Santa Rosa, where the former form begins. I have also this form from Santa
Clara County, Monterey County, Fresno County, ete. It is common in the foothills
of the Sierra Nevada, in Nevada County. Among many hundred specimens col-
lected there was only one which possessed the paired papille of the former form.

Systematic position. The peculiar variation of the nephridio-pores places
Argilophilus in undoubted proximity to Plutellus, both as described by Perrier and
Benham. The extra chietal pores in Argilophilus warrants however the forma-
tion of a new genus, even if no other important characteristics would help to make it
yet more distinct. As is well-known, Perrier’s description of the ovaries, ete., in
Plutellus have always been considered doubtful, and by Benham haye been shown to
be incorrect. This of course only in the case that Benham’s worm really belongs to
the same genus as the one described by Perrier. I do not doubt that this is so, be-
cause the positions assigned to the ovaries by Perrier is so abnormal that it is more

56 CALIFORNIA ACADEMY OF SCIENCES.

than likely that a mistake was made, as Benham has already pointed out. But on the
other hand Benham’s worm came from Queen Charlotte’s Islands in the Pacific, while
Perrier’s worm came from the Atlantic Coast, many thousand miles away, and it will
always be impossible to with certainty decide upon the position of the ovaries in
Perrier’s Plutellus until the original worm has again been investigated. That it will
be found to entirely agree with the normal type I do not doubt.

From Plutellus our Argilophilus differs in several important points besides the
location of the nephridio-pores. First, in regard to the number of the ciliated rosettes—
in Plutellus only one pair, in Argilophilus two pair. One pair of sperm-saes in Plu-
tellus, three pair in Argilophilus. One pair of testes in Plutellus, two pair in Argilo-
philus. Four pair of small spermatheca in Plutellus, two very large ones in Argilophilus.
Perfect clitellum in Plutellus in at least two somites, ventrally imperfect clitellum in
Argilophilus. In other respects the two genera resemble each other greatly, and
through the respective arrangement of the nephridio-pores, they form a distinct tribe
or sub-family for which I propose the name P/lute/lini.

The following table will show the principal characteristics of the two genera:

|
> % |
Argilophilus. Plutellus.
Prostomium dovetailing the peri-
BUODAUIN osayay- qricversitereteaeleletake set | Partly. Completely.
(CLUPICUITE rete ogd DOS OOOO COI ACTA Ventrally imperfect. Ventrally perfect in at least two
somites.
Nephridio-pores..............-...|In front of sete 3, 4, and outside of 4; gen-| In front of 3 and 4, alternating
erally the latter, the arrangement being) rather regularly.
irregular.
Qagzard tie Ree nc oe any senate Vv. vi.
Spermatheca ........+...+........| Two very large pair in vii and viii. Four or five pair in vi, vii, viii
and ix.
SPERMAPUMMEU casei ase cies hetero Two pair in x and xi. One pair in x.
Tegteg tee sate Salgceeh Manatee se Two pair in x and xi. One pair in x.
SYMP UREN 26 o.oo clbbouco tecdenigeys Three pair in x, xi, xii. One pair in xi. °
1
Ovary and oviductin ........... | xii. xiii.
Owipore lay J.cacsncines torch | xiv. | xiv.
|
Maile por erin my ctocetes sents Xvill. | xviii.

Prostate Very large, coiled, extending through several Straight, tubular, not very large,

somites. confined to one somite.

Nephridia commence in.......... ii. ili.
|

Nephridial bladder at the entrance Absent, the duct being single and straight. | Large, but of varying size in the
to the body-wall. . 2. .ccsee ae | two series of nephridia.

Ventral papille on the body-wall.| Present in pairs or in a single one, | Absent.
Present. Absent.

Copulatory seta

Typhlosole. senses. esses...) A small typhlosole present. No typhlosole.

CALIFORNIA EUDRILID®. 57

PAPERS REFERRED TO,

Benuam, W. Braxtanp. An Attempt to Classify Earthworms. Quarterly Journal of Microscopical Science, yol.
XXX, pt. li.

Benuam, W. Buaxtanp, The Nephridium of Lumbricus, ete. Quarterly Journal of Microscopical Science, xxx,
li, 1891, 293.

Benuam, W. Buaxtanp. Description of Three New Species of Earthworms. Proceedings of the Zoological Society of
London, Feb. 16, 1892.

Bepparp, Frank UW. The Classification and Distribution of Earthworms. Proceedings of the Royal Physical
Society, Edinb., 1891.

Bepparp, Frank E. On the Earthworms collected in Algeria and Tunisia. Proceedings of the Zoological Society
of London, January 5, 1892.

Brpparp, Frank FE. Some New or Little Known Oligochwta. Proceedings of the Royal Physical Society, Edinb.,
June 13. 1893.

Bourn, A.G. The Nephridia of Leaches. Quarterly Journal of Microscopical Science, vol. xxxiv, 592.

Eisen Gustay. California Earthworms of the family of Eudrilidw, Zoe, a Biological Journal, San Franciseo, Cali-
fornia, vol. iv, No. 3, 1893, pp. 248-254.

Friercner, J. J. Notes on Australian Earthworms. Proceedings Linnean Society of New South Wales, vii, 374.

Micnarsen, W. Oligochwten yon Siid Georgien. Jahrbuch der Wissenschaftlichen Anstalten zu Hamburg, 1887.
Hamburg, 1888.

Rosa, Dr. DanteLe. I Terricoli Argentini, etc. Annali del Museo Civico di Storia Naturale de Genoa, ser. 2,
vol. ix, 509, 1890.

Rosa, Dr. Daniete. Sui Generi Pontodrilus, Microscolex, Photodrilus. Bol. Musei, ete., di Torino, No. 39, vol.
iii, 1888.

Rosa, Dr. Dantete. Microseolex Modestus. Bol. Musei, ete., di Torino, No. 19, vol. ii, 1887.

EXPLANATION OF THE FIGURES.
DELTANIA ELEGANS, Figs. 1-20.

. A medium sized worm, natural size.

2. The anterior part magnified, seen from the ventral side, to show arrangement of set and the exterior pores of
various organs. p.n.p. the first three pepto-nephridio-pores in somites ii, iii and iy. n. p- nephridio-pores
of the following somites, opening in front of the 3d set~#, the pepto n. p. opening in front of the 4th seta.
0. p. ovipore. s. pd. p. spermiducal pore.

In order to show the outer sets, the body has been slightly flattened out. In reality the outer set should be
slightly more dorsal than what appears.

3. The prostomium with somites i and ii.

4. Schematic view of the arrangement of the sete on both sides of the male pore. The italic numerals indicating
the order of the setw, the roman numerals indicating the number of the somites.

5. The interior of the ventral surface of the genital somites viewed from above, showing the arrangement of
the generative organs, ete. spth. spermathece. c. 7. ciliated rosettes. sp.s. sperm-sacs. sp.d. spermducts.
ov. ovary. od. oviduct. p.s. penial set and sacs. pr. prostate. v. gl. ventral ganglion. ¢. testes.

6. Longitudinal section of the anterior part of the body. The posterior section is a little more eccentric than
the anterior. br. brain. s/. gl. salivary glands. s. gl. septal glands. phx. pharynx. s. plx. sigmoid plexus
of the esophagus. sp.s. sperm-sacs. h. hearts. dv. dorsal vessel. ¢.i. tubular intestine. s. i. sacculated
intestine. pr. prostate. s. ps. sacs with penial set. v. gi. ventral ganglion. es. esophagus. spth.
spermatheca. ¢. testes. c. r. ciliated rosettes. ov. ovary. od. oviduct.

8. Prostate and male apparatus. pr. prostate. sp. d. spermduct. p. s. penial sete and sacs. ms. longitu-
dinal muscular layer of the body wall.

9. Oviduet in xiy and xiii.

58

26.

46.

47.
48.

CALIFORNIA ACADEMY OF SCIENCES.

One of the septal glands in somites vi, vii, viii and ix.

Ciliated rosettes.

Ovary.

Spermatheca.

Exterior male-pore and penial papille. 4 male-pore. p.s. penial set and papilla.
Exterior view of the clitellum seen from the ventral side. op. ovipore. 4 male and prostate pore.
One of the regular set.

Penial set from somite xvil.

Testis from somite x.

Nephridium from somite xv.

Anterior nephridium from somite il.

A nephridio-stome from one of the posterior nephridia.

DELTANIA TROYERI, Figs. 21-38.

Diagramatie view of the regular arrangement of the set# in several somites posterior to the clitellum.

A medium size worm.

The largest specimen.

The anterior part seen from above, showing prostomium and somites i, ii, iii.

Schematic view of the arrangement of the inner sets in the region on both sides of the male-pore. ovp. ovi-
pore. p.s. penial sete. 4 male-pore.

The outside of the male genital region. f. folds in the body-wall. p.s. penialset#. p.p. penial sete papilla.
pr. and 4 prostate and male-pore. c.d. cupshaped concave region or sucker near the male-pore. c. c.
central cavity between the male-pores. m. st. place for missing setw No.1. s. st. seta No.2. p. m, penial
muscles, arranged fanshaped, connecting the male-pore with the intersegmental groove.

Longitudinal section of the anterior part of the body. b.c. buceal cavity. phnx. pharynx. si. gl. salivary or
pharyngeal glands. br. brain. s. gl. septal glands. s. ple. sigmoid plexus of the esophagus. sp. s. sperm-
sacs. @s. esophagus. ¢. testes. spth. spermatheca.

Somewhat schematic view of the alimentary canal seen from above. es. cesophagus and pharynx. - sl. gl.
salivary gland. s. gl. septal gland. d. v. dorsal vessel. A. hearts. ¢. 7. tubular intestine. s. i. sacculated
intestine.

Brain and yentral ganglion.

One pair of the septal glands (vi).

Spermatheca from ix. Side view.

Spermatheca seen from below.

Spermatheca seen from above.

Male apparatus. pr. prostate. p.s. penial sacs andset#. sp. d. spermduct. ms. muscles of the body-wall.

Nephridium from somite xy.

Oviduct and ovisae, left side.

The same from the same individual, but from the right hand side of the ventral ganglion.

The same sac seen from above.

Detail of the same.

Arrangement of the set surrounding the spermathecal pores.

DELTANIA BENHAMI, Figs. 40-48.

Arrangement of the sets surrounding the spermathecal and male-pores. The same proportion and Scale as in
the preceding figure.

Sperm-sac.

Spermatheca, left side.

Spermatheca, right side of the same individual.

Oviduct.

Prostate gland and penial set. v.v. blood vessel covering the prostate.

Alimentary canal. pha. pharynx. sl. gl. salivary glands. ss. gl. septal gland. @s. cesophagus. s. i. saccu-
lated intestine. c.i. contracted part of the intestine generally commencing somewhere between somites xxvii
to xxxyi and continuing towards the tail. In the first six somites the contraction is generally much narrower
than in the following.

The worm natural size.

The male papill on somite xvii, showing the projecting penial sete. The upper shaded part is the clitellum
proper which ends at the center of the somite in line with the male-pore.

Fr

CALIFORNIA EUDRILID. ov

DELTANIA ELEGANS, Figs. 49-56.

49, The lumen and coecal part of the nephridium in transverse section to the body. fl. c. ¢. flask-like glandular
cells of clitellum. /. v. blood-vessels in the transverse muscular layer. /. m. transyerse or circular museu-
lar layer. /. m. longitudinal muscular layer. 7. x. lumen of ccecal part of the nephridium. 7. /. interior
lining of the nephridium. g/. glandular outer layer of nephridium. cal. calculi projected by the nephridium.
gi.c. glandular cells of the clitellum. atr. urinary bladder or large contractile (?) chamber near the exterior
pore of the nephridium. ms, muscular layer of the nephidial duct joining the flask-like cells of the clitel-
lum. col. collar of the nephridium forming the exterior pore and consisting of regular layers of large, almost
rectangular cells.

SOA. Section through the body-wall and the lower part of a nephridium in somite xviii, but posterior to the clitellum.
n.d. nephridio-duct. ¢.. cocal nephridio-bladder. /. m. longitudinal muscles. c. m. circular muscles.
ep. epidermis and cuticle,

50B. Section through the body-wall and the nephridial collar in a somite posterior to the clitellum, illustrating the
large ganglion (?), below the nephridial collar. ‘The section shows the beginning of the nephridial collar and
the place where it joins the ccecal vesicle.

500, Section closely following the former nearer the nephridio-pore, showing a larger part of the ganglion. i. ¢.
interior chamber or tubule, probably opening into the nephridial caecum. ¢./. inner lining of the cecal
bladder. m. . muscular layer of the bladder. — ¢. np. collar of nephridium. 7. g/. nerve ganglion. /. m.
longitudinal muscular layer of the body-wall. ¢. m. transverse layer of the body-wall. h. hypodermis.

51. Lower part of the prostate and penial sac, section through the male-pore, showing the junction of one of the
penial sacs with the prostate. p.s.s. penis sac with set. p.s. one of the penial setw broken. pr.
and ps. prostate and male-pore. pr. d. prostate duct.

2. Section through the male-pore, cross-section through the body-wall, showing junction of the two spermducts,
and prostate, and one of the penis sacs. sp. d. spermducts. j. sp. junction of spermducts. pr. prostate.
Jj. sp. and pr. junction spermduct and prostate. s. ps. sac of penial sete. j. pr. and p.s. junction of prostate

or

and penal set. ¢ male-pore. ep. epidermis. c. m. layer of circular muscles, body-wall. 1. m. layer of
longitudinal muscles, body-wall. x. c. nerve cord from ventral ganglion. p. p. penial papilla.

53. Section through the male-pore showing junction of the two penis sacs with the prostate. ps. the two
branches of the penis sac with the penis sheaths and penial sete. pr. prostate. c¢. m. circular muscles.
l.m. longitudinal muscles. 4 male-pore. atr. atrium, or junction between the penis sacs and the prostate.

54, Section through the muscular part of the prostate. gl. inner glandular layer. m. outer muscular layer.
blw. blood-vessels and cuticle.

55. Section through upper end of prostate. blv. bloodyessels. c¢. v. vessels between the inner and outer layer of
cells. 7. v. inner capillary vessels, lining the inner surface of the prostate.

56. Section through prostate at the junction of the muscular and glandular part. m. pr. muscular prostate; other
letters same as in Fig. 54.

57A. Section through intestine in somite xv. ch. 1. chloragogic layer. J. m./. longitudinal muscular layer. v.l.
vascular layer. b/v. blood-vessel of the epithelium. ep. epithelial layer.

57B. Section through intestine, somite xiv.

57C. Section through intestine, somite xvi.

58. Cross-section through the clitellum, showing the relative size of the various layers.

ARGILOPHILUS MARMORATUS ORNATUS, Figs. 59-130.

59. A nephridium from one of the somites posterior to the clitellum. ~*0, inner orifice or nephridio-stome engaged
in the muscular dissepiment d. 1, narrow duct connecting the funnel with the fold. p. Jf. posterior
fold. a. /. anterior fold. uw. upper part of the spongy duct, which at 15 stands in direct connection with
the narrow tube 1. J. lower part of the spongy duct gradually increasing in size towards 5, a. upper main
canal. b. lower main canal, both the upper and the lower being different ends of the same canal. 0. Culfs
point of recurrence. p.c. peritoneal cell covering or support of the nephridium. x. p. nephridio-pore.
c. t. capillary tubules of the spongy duct. 6/. blood-yessels. cap. capillary blood-vessels surrounding the
canals ~@and b. 1 to 17 mark places from which the detail drawings are taken. The nephridium is longer in
proportion to its thickness than what is shown in the figure. If the part between the two parallel lines at
18 was increased to about one-third or one-fourth the entire length of the nephridium the proportion would
be correct. If the whole nephridium had been drawn the figure would haye been too long for the plate.

60. Nephridio-stome, front view.

61. Nephridio-stome, side view.

62. Nephridio-stome, end view.

* Letters and figures in full-face type refer to those in rings on the plates.

60 CALIFORNIA ACADEMY OF SCIENCES.

63. Inner part of nephridium more highly magnified. The numerals indicate the same places as in Fig. 59. The
letters are the same as in the above figure.

64. Part of the same more highly magnified in order to show the capillaries and their connection with the inner
duct of the tube. ¢./. tube leading to nephridio-stome. a. upper canal. b. lowercanal. j. /. inner lumen
of the duct which connects at c. with the capillaries. c. ¢. capillary tubes, longitudinal view. c.¢.¢. capil-
lary tubes seen in cross-section. sp.c. spongy canal or tube directly connecting with the duct 1 and con-
taining the capillaries. 0. c. outer canal leading to the nephridio-pore. c¢. br. the narrow bridge connecting
the two main folds. p.c. point where ciliation begins. bl. v. blood-vessels, connectmg with capillary
blood-vessels surrounding the two inner canals @ and; only a part of the blood-vessels are drawn. This
figure was carefully drawn with camera lucida, though, on account of the minuteness and numbers of the
ductules c. fe. and c. ¢., it was impossible to delineate all, almost one-fifth having been left out in places where
they crowded each other.

65. View of the spongy duct between 3 and 4 showing the inner canals a and } and the connecting ductules;
letters indicate the same as in previous figures.

66. The same spongy duct near 4, end view, and partly cross section.

67. The posterior fold as seen at 5, the crosses + and X indicate the respective point corresponding on fig. 59,
in order to show the direction of the tube, which is contrary to the one shown on fig. 59. sp. ¢. the spongy
tube with the upper and lower ducts, connected by ductules c. ¢. bl. bloodvessels. cap. capillary blood-
vessels, some of which surround the inner canals, others the outer fold. At the place marked with a x the
ductules begin to connect the upper spongy tube with the lower main canal instead of with the lower part
of the spongy tube, as at c. ¢.

68. A somewhat larger figure of the last taken at the end x, where the lower part of the spongy tube has almost
vanished and become bereft of its tubules c.¢., capillary tubes entering canal b. c. chambers on the central
canal in the spongy tube, which send off tubules on the other side of the two main canals, and which also
enter the canal. 6.1. lumen of the spongy tube. In this and the previous figure will be seen how the tubules
of the spongy tube shown in figs. 65 and 66 have been found forming a more regular lumen /, which in the
next figure will be seen to emerge into or change to a regular canal.

69. Part of the upper fold at 7, showing the tubules of the spongy tube to cease, the tube itself to become more
regular, fusing itself in canalb. The arrows show the direction of the fluid or excretions from the nepridio-
stome to the pore.

70. Diagramatic section of the main fold at the place where the capillaries of the spongy mass have ceased, and at
the very point of the outer band marked fig. 59 0. c.f. The spongy mass surrounding the canals is now void
of tubules.

71. View of the anterior fold at 10, showing the inner ciliated canal—formerly the lumen of the spongy tube—
and the two canals a and 4, which are enclosed by a common glandular mass, a continuation of the spongy
tube. The lower fan-shaped lines are possibly openings or valves in the tube, connecting with the glandular
mass. As will be seen, the glandular mass is not continuous, but seemingly separated by lumens—/—which
may prove to be canals connecting with the tube at the slits above referred to.

72. A part of same fold between 9 and 10. The lumen is here void of cilia, and the glandulous mass is smaller

73. ‘The main fold as seen at the narrow place 9, 7. lumen void of cilia. «. one of the canals a or b seen on the
other side of the limen. 6/. y. bloodyessels surrounding the lumen and its glandular walls.

744. Cross section of the posterior main fold taken at 8, before the main central lumen in the spongy duct was
formed.

74B. Cross section of the posterior duct at 7, showing the blind ending of the tubules.

75. A longitudinal section of the fold at 5, showing only one of the central canals.

76. The same, the under side of the lumen being in focus.

77. A longitudinal section near 3; here also one of the canals has been cut away.

78. Peritoneal cells of the nephridial supporting covering from p. c. fig. 59.

79. The same, end view. p.c. masses of perigastric cells.

80. One of the perigastric cells more highly magnified.

81. One of the spermathecs, exterior view.

$2. Cross section of a spermatheca. a. upper sac-like part. b. lower muscular part. m. muscular layer of the
sac-like part. /. m. longitudinal muscular layer of the body-wall. c. m. circular muscular layer of the
spermatheca. gl. ep. glandular epithelium. 7. inner cavity of the spermatheca. sp. b. sperm balls. /. s.
lacunary system, or chambers for the storage of the spermatozoa. s. m. secreted mass with streaks of sperm-
atozoa. spz. spermatozoa. ¢.m. transverse muscular layer of the body-wall. spth. p. spermathecal-pore.

83. A longitudinal section of the spermatheca more highly magnified, showing the lacunary system and its position
to the epithelial cells. Letters as in the last figure.

54. Cross section of the muscular part of the spermatheca, showing the arrangement of the storage chambers oc-
cupied by sperm balls. Letters as in fig. 83.

CALIFORNIA EUDRILIDA, 61

55. One of the sperm balls more highly magnified. Of these never more than one occupies the same chamber.

SGA. A somewhat diagramatic view of the anterior somite, composed from several sections, in order to show the lo-
cation of the organs, their relative size, ete. e/. clitellum. br. brain. ce. lower commissure of yentral ganglion.
phx. pharynx. sl. gl. salivary glands. w. cesophagus. gz. gizzard. ¢.i. tubular intestine. 8.7. saccnlated
intestine, ph. cross section of the anteriot nephridia in somites ii, iii, iv. The other nephridia are not
shown, sph. spermathecw. sps. sperm-sacs. 1. sps. Jobed sperm-sacs in xii. ov. ovary. ovd. oviduct.
c. r. ciliated rosettes. sp. d. spermduct; the connection with the ciliated rosettes not shown. 4 male
pore. pr. prostate. p. s. sacs with penial sets opening in the same pore, but more ventrally than the pros-
tate. ¢. the place where the spermduct enters the prostate.

86B. The body opened from above and the interior organs exposed to view. A partially diagramatic view. The
dotted lines indicate the organs coyered by others, and which could not be seen except by removal of overly-
ing organs. Letters indicate as in the preceding figure.

STA. Diagramatic longitudinal section of gizzard.

87B. Diagramatic transverse section of gizzard, to show its compressed shape.

8S. A part of the epithelium and muscular layer of the gizzard seen in longitudinal section. ep. epidermis. ¢prt.
epithelium. 7. m. longitudinal muscles.

89. A transverse section of the gizzard. (Letters as in fig. 88.)

90. A transverse section of the alimentary canal adjoining the gizzard posteriorly. ¢. m. transverse muscles.
. m. longitudinal muscles. ep. epithelium cells. chy. supposed chylus chambers. c.c¢. chloragogie cells.
gl. glands. 61. bloodvessels.

91. The same in longitudinal section. (Letters as in fig. 90.)

92. Surface view of the same, showing the arrangement of the chylus chambers. (Camera drawing.)

93A. Section of a part of the dorsal vessel in somite xvi. m. muscular layer. b/. blood clot. c.¢. chloragogic
cells.

93B. Section through the alimentary canal in somite xiy.

944. One of the testes.

94B. Right testis in cross section.

940. Left testis in the same somite xi in cross section.

95. Ovary.

96. Section of the body in somite x, throuch testes.

97. Section of the body in somite x through the sperm funnels, very close to the septum.

98. Section through somite xi.

99. Section through somite xii. sps. a. sperm sac in x. sps. ai. sperm sac in xi, sps. xiv. sperm sac in xii.
m. longitudinal museles connecting various organs with the body-wall. 7. intestine. d.v. dorsal vessel.
v.v. ventral vessel. h. hearts. ¢. testes. v. gl. yentral ganglion. s, septum.

100. Part of the sperm-sac. coc. parasitic coccidie.

101. One of the sperm-sacs separated.

102. Oviduct a and b, front view. ‘©. side view of an abnormal oviduct with two, instead of one, upper lips. x». /.

upper lip. s. septum. m. muscular band.

103A. Cross section of the body-wall through the male-pore, showing sections of prostate and penial setw. pr. gl.
sections of glandular part of prostate. pr. m. muscular part of prostate. p. s. penial set and their sacs.
v. vascular layer of the body-wall. 7. m. longitudinal layer of muscles. ¢. m. transverse layer of muscles.
e. gl. glandular layer of clitellum. h. hypodermis. g. ganglion, ». p. ventral part of the worm.

103B. Detail of former, nuclei in the muscular part of the prostate.

104. One of the ciliated funnels freed from the attached sperm-sacs.

105. Diagramatic section through prostate, showing relative size of spermducts where they join the prostate.

106. Diagramatic representation of the prostate, showing its natural position when the body-wall is spread ont.
pr. glandular prostate. spd. spermduct. p. 8. penis-sacs. m. pr. muscular prostate. v. 7. vascular
layer. v. ventral side.

107. Detail of the aboye, to show junction of spermduct and prostate.

108. Cross section of prostate. 61. v. blood vessels. o. gi. outer glands. or prostate proper. 7. gl. inner gland
or atrium.

109. Detail of the former, showing the way the outer glands open into the atrium between the inner cell layer.

110, 111, 112, 118. Suecessive sections through prostate and spermduct, illustrating the junction of the two. In
the first figure the spermducts have not yet entered the muscular part of the prostate. In the three follow-
ing figures this junction is progressing, and finally almost accomplished. 7. m. /. inner epithelium of the
muscular part of the prostate. pr. glandular prostate. spd. spermduct. 6/7. v. blood yssels. m. 1.
muscular layer of the spermduct. In 110 the spermduet has been sectioned twice, as it is here conyoluted.

114. Cross section through the body-wall in somite xix. ». /. vascular layer. 7. m. longitudinal layer of muscles.
¢. m, transverse muscles. h. hypodermis,

62 CALIFORNIA ACADEMY OF SCIENCES.

115. Cross section through clitellam. gl. glandular cells with fine secretions. c. gl. glands from the hypodermis
projecting into the former layer. /. v. blood vessels.

116. Detail of the former, illustrating the hypodermal glands, etc.

117. Cross section through body-wall, somite xii. Letters same as above, illustrating the arrangement of the
longitudinal muscles. spd. spermduct.

118. Cross section through body-wall somite xvii, illustrating the transverse muscles.

119. Somite xiv laid open, illustrating the diagonal muscular band. ord. oviduct. s. septum. xii Xiv. ovp. ovi-
pore. J. 2.3. 4. sete. m. 1.2. 3. diagonal muscular band, beginning and ending on the body-wall.
m. 4. muscular band connecting the oviduct and septum with the body-wall. 2. /. nephridio-funnel.
is. g. intersegmental groove. v. v. ventral blood-vessel. h. ventral part of the hearts. /. v. lateral vessel
spreading on the body-wall and supplying the nephridium. v. g. ventral ganglion. £

120A. Cross section through one of the ventral papille of Argilophilus marmoratus papillifer. 4. c. tubular cells
butting on the central glandular masses. c. gl. supporting and protecting cells, with coarse grainy con-
tents. hk. common supporting cells of the hypodermal layer. /. m. longitudinal muscles. ¢. m. trans-
verse muscles. v.J. inner vascular layer. nf. nerve ganglions, connecting the glandular masses with the
yentral nerve ganglion. o. glandular papille with central nerve ganglia.

120B. The corresponding organ, or ventral papilla in Argilophilus marmoratus ornatus; letters indicate the same.

121A. Detail of the last figure showing the most lateral of the glandular masses. o. glandular masses with a

central nerve ganglion.

121B. Detail of the same figure showing the most centrally located glands.

122. One of the common sete.

123A. One of the penial sete.

123B. The tips of the former more highly magnified.

124. A part of the body-wall viewed from the inner side. A semi-diagramatic figure illustrating the arrangement
of the sete and the nephridio-pores over a certain space. /, 2, 3, 4, the various rows of sete. np. p.
nephridio-pores, indicated by round circles. Two of the nephridia are drawn in order to show their position,
as well as the relative size of the ducts.

125, 126, 127, 128. The anterior part of the body seen from the ventral side. Various specimens to show the
yariation in position and number of the ventral papillw. spth. spermathecal-pores. ovd. oviduets.
t male-pores. The ovals indicate the ventral papille.

129A. The clitellar and some of the anterior somites, viewed from the ventral side, showing the yarious pores and
the genital zone. Letters indicate the same as in the preceding figures. p.s. penial sete protruding
from the male-pores.

129B. The twenty-three anterior somites of a specimen killed in alcohol gradually increased in strength and hardened
in strong alcohol, showing the spermathecal, 9 and 4 pores, the ventral papillae, ete.

1290. The most anterior somites seen from above.

130. The two and three anterior somites of two different specimens, showing the inverted buccalic sac, the prosto-
minum, etc. a. and c. ventral; c. d. and f. dorsal; e. lateral view.

ARGILOPHILUS MARMORATUS PAPILLIFER, Figs. 131-136.

131. The anterior somites viewed from below showing the single row of ventral papillew, diminishing in size an-
teriorly. The outlines at the side represent the shape of the papille as found in the figured specimen.

132A. A medium size specimen preserved in strong alcohol. The specimen was first killed in an increasing solu-
tion of corrosive sublimate.

132B. A large specimen of the former figured while alive.

133. Section of the intestine and dorsal vessel in somite ix, in order to show the typhlosolar vessel. ¢.v. typhlo-
solar vessel in the intestine. d.v. dorsal longitudinal vessel. ch. c. chloragogie cells of the intestine.
e.p. epithelial folds of the inner lining, void of chylus glands. c.v. the chylus region of the inner
epethelial lining.

. The typhlosolar region of the intestine, detail from the last figure more highly magnified. ¢. v. typhlosolar
lining. 7. m. longitudinal muscles of the intestine. c¢. m. cireular muscular layer of the same. Other
letters as in the last figure.

an
_

135. Cross section of the intestine and dorsal vessel in somite xxiv illustrating the typhlosole. ss. 7. saceulated
intestine. /. s. blood-sinuses. ty. typhlosole.

136. Detail of the former, showing the typhlosole proper, greatly magnified. d.v. dorsal vessel. /y. typhlosole.
bl. v. blood sinuses in the intestine. /. m. longitudinal muscles. ec. m. and e. p. cirenlar muscles and
epithelial lining of the intestine.

MEM, CAL. ACAD, Il. PLATE XI

ASTRAAITION AREY oP.

_.DELTANIA ELHGANS Fie’s. 1 70 11.

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275

GUSTAV

-MEM, CAL. ACAD, Il,. PLATE XVI

SUTE.BAITTOAS RET SP.

DELTANIA ELEGANS FIe’s...49 & 50.

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PACIFIC COAST OLIGOCHATA.
i

PH@NICODRILUS TASTE; PONTODRILUS MICHELSENI; ECLIPIDRILUS FRIGIDUS.
BY GUSTAV EISEN.
PHCENICODRILUS nov. gen.

Small slender terrrestrial oligocheete closely allied to Oecnerodrilus from which
genus they differ only in the absence of a prostate. In Ocnerodrilus the organ
known as a prostate, or by some as atrium, is a prominent and important characteristic
also shared with such genera as Kerria and Gordiodrilus. These genera which form
a natural group have been chiefly characterized by one or more prostates, opening
either with or independently of the spermducts. Our present form resembles
Oecnerodrilus in all principal characteristics which distinguish that genus from Ker-
ria and Gordiodrilus, with the exception that it has not even a trace of a prostate.
There are also some other minor characteristics as will be seen from the description,
but with the knowledge of only one species it is yet impossible to know if these are
genus or species characteristics. For the present the following genus diagnosis may
suffice:

Small, terrestrial oligochetz inhabiting damp soil.

Clitellum imperfect; comprises the oviduct and the male pore.

Spermathecz with rudimentary diverticula at the inner free end—in ix. The
spermathecal pore between viii and ix.

No differentiated penial sete. The common setie sigmoid, 8 in each somite in
4 couples.

Nephridia paired, those posterior of the clitellum surrounded by large peri-
toneal cells.

Alimentary canal without gizzard and typhlosole, but with one pair of large
diverticula in ix, connecting with the tubular intestine in the posterior part of the
somite. Four pair of septal glands in y, yi, vil and viii.

No subneural vessel. Dorsal and ventral vessel connected by hearts in x and
xi. Lateral vessel projected anteriorly from each of the diverticula. Blood yellowish-
red.

Testes two pair, in x and ix. Large sperm sacs in ix, x, xi, xil. Ovaries in
xiii. Oviduct in xiv. Two pair of ciliated rosettes in x and xi. Sperm ducts not
fused, open in the male pore in somite xvii. No prostate.

64 CALIFORNIA ACADEMY OF SCIENCES.

Pheenicodrilus taste n. sp.

The size is that of a very large Oenerodrilus though even fully matured speci-
mens varied greatly as to length. My largest specimens, which first had been slowly
killed by dropping solution of corrosive sublimate in the water dish and then extended
before being hardened with alcohol, reached 2? inches by 1} line in thickness at the
clitellum. Average-sized specimens were considerably over 2inches long. This refers
to the mountain specimens collected in the Sierra El Taste; the lowland specimens
from Pescadero were much smaller not reaching the 2-inch mark. The body is
slightly tapering towards the tail end. The somites are well set, those of the clitellum
are hardly distinct. The prostomium is not long, but broad, dovetailing the peristo-
mium about one-half. From here on the somites gradually, though slightly, increase
in size until somite x which is a little the largest, xi, xii and xiii are smaller. The
somites posterior to the clitellum are slightly larger than somite x, except the last few
posterior ones.

Clitellum comprises somites xiv to viii. Vertically it ends at a line drawn
halfway between sete 2 and 3, slightly receding in somite xiv. Strictly speaking the
clitellum does not enclose the male pore, as the pore is situated more ventrally than the
thick clitellar layer, and between that and the pore there is no connecting ridge or
papilla.

An accessory copulatory swelling is seen around the outer couple of sete in
somite xiv (fig. 26, c. ¢.), the body-wall here being raised like a small mound, with
the set slightly outside of its center, from which the cells are arranged as radii in a
circle.

The male pore is surrounded only by a very small ridge or papilla, not high
enough to be seen with a magnifying glass sufficiently strong to reveal the elevated
papillee of the oviducts. But the whole zone around the male pore is often considerably
elevated, turned inwards or towards the median line of the body and rounded
forming a longitudinal groove.

Exterior pores. The spermathecal pore is situated in the intersegmental groove
between somites vili and ix, in front of and slightly outside of seta 2, the inner angle
of the pore being in line with that seta, while the body of the papilla is situated more
dorsally. The oyipore is situated close by, in front of, but not outside of seta 2. The
male pore is situated in xvii exactly in a line with sete 1 and 2 according to the lon-
gitudinal muscular fibres, but as the body-wall is slightly contracted in this somite the
pore appears as if situated slightly more ventrally; the exact location is, however, the
place left vacant by the absence of the ventral couple of sete. This couple (sete 1
and 2) are never, at least not in adult specimens, found developed in this somite,
though the tips of the young reserve sete are sometimes seen in their sac close to the
pore.

Nephropores open in line with seta 2, and are situated in the anterior one-
third of the space between the sete and the anterior septum. The pores are large,
round and easily distinguished.

PACIFIC COAST OLIGOCH MTA. 65

Body-wall. The various layers offer nothing of great interest. The longitud-
inal muscular layer is slightly thicker at the dorsal part, and so is the part situated
between the 3d and 4th sete. ‘The muscles in this layer do not show any feathery
arrangement around a central fiber, but are irregularly distributed. There are no
dorsal pores. The body-wall in somites xi and xii is thinner than the anterior somites.

Areciform muscles. The inner surface of the body-wall in somites xyii and
xviii, and partially also in xvi, are covered with numerous arciform muscles which
stretch transversely or diagonally across the body cavity, connecting the region run-
ning through the outer couple of setee with that of the male pore and the inner setze.
The number of muscular bands in these somites is very great and they vary in length
and thickness. There is also a slight variation in different specimens, the main fasci-
cles being, however, always in the main the same. These muscles are best viewed
when seen from above, the body wall being spread open and the sacculated intestine
removed. As will be seen from the drawing, which is a careful representation of the
principal muscular bands, most of the bands begin or are attached to the body-wall on
a line running through sete 1 and 2 and ending in a line running through sete
3 and 4. But there are some which begin more ventrally and end more dorsally
than either couple of setze. In somite xviii most of the fascicles run in right angle
to the median line, but the most posterior one as well as one or two more run diag-
onally backwards. In somite xvii there is one large group of fascicles beginning
around the male pore and running transversely sideways, while another group of fas-
cicles run diagonally forward connecting the male region with the anterior part of
the somite. There are also longer fascicles anterior of the male pore which run much
further sideways than the outer sete. In somite xvi there are but few fascicles,
much smaller and shorter. The male pore and spermducts are so entirely covered
up by these muscular fascicles that they can only be seen with the greatest difficulty
when viewed on the spread body-wall. In the figure (fig. 7), only the principal mus-
cles of one side are drawn. The shaded bands which are seen crossing the median
line are slight ridges in the body-wall, connecting the principal muscle fascicles. The
objects of these muscles are of course to elevate and depress the male zone. Similar
muscles were first described by Benham in Moniligaster, and later by myself in Argi-
lophilus. They no doubt exist in most species of Oligochzeta but are of great interest
and yalue as species characteristics.

The septa begin between somites vy and vi, and gradually increase in size
toward somite ix. The most anterior septum, however, is not unusually thick. The
following three septa are very thick, that between viii and ix being the thickest.

Alimentary canal. The buccal cavity is very large and occupies somites i and
ii. Pharynx begins in ili or posterior part of ii and extends to the posterior part of
iii. It is only developed superiorly and merely attached to the cesophagus in the pos-
terior part of ii. Qisophagus consists of one in the beginning narrow and compara-
tively even duct, which in iy rises upwards and then becomes considerably sacculated

66 CALIFORNIA ACADEMY OF SCIENCES.

in somites v, vi, vii and viii, again to assume a more tubular form in ix. In the pos-
terior part of this somite it is joined by the two diverticula. The tubular intestine
proper occupies the twosomites x and xi. In xii begins the sacculated intestine. The
diverticula of the cesophagus are very long, narrow and slender, more so than in any
species of Oenerodrilus which I have seen. These exterior features offer nothing
peculiar. The lateral blood vessels issue, as usual, from the anterior points of the
diverticula.

These diverticula of the cesophagus originate in the posterior part of the
somite, and not in the anterior part as in Kerria. The structure of these diverticula
corresponds with that of the same organ in Oenerodrilus. Only in that genus the
rule appears to be that the interior of the diverticulum consists of one single chamber
encroached on by a few parallel ridges. The diverticulum of Phanicodrilus taste is
in the central part four to eight chambered, divided off longitudinally, presenting the
same appearance as an orange when cut through crosswise (figs. 14-16). The number
of chambers varies, as in one specimen I found the right hand diverticulum to have
four chambers, while the left one had five chambers. In sections near to either end
these chambers fuse further and finally only two and one chambers are left.

But this fusion is uneqnal at the two respective ends. In the end nearest the
junction with the alimentary canal there is only one chamber found, and a little
further forward there are two chambers, the number increasing until generally eight
chambers are found in the center. From there on no increase is made, but the echam-
bers decrease in width, and at the anterior end suddenly fuse into one which is small
and narrow, not thicker than a blood vessel. The longitudinal blood vesssels are very
much the same as in Ocnerodrilus, but generally more in number.

The number of blood vessels in each diverticulum varies with the place where
located in the diverticulum. The section nearest to the posterior septum of the ali-
mentary canal shows the two laterals from the dorsal vessel entering the diverticula.
To begin with, they are seen on the outside of the cellular mass of the diverticulum,
while in succeeding sections they appear like a few biood vessels scattered on the out-
side of the epithelial cells. In succeeding sections these vessels are seen to increase
in number until in the center of the diverticulum they number about one hundred.

The blood supply for these organs come from branches of the dorsal vessel and
not from collective vessels from the alimentary canal.

Salivary glands are as usual found attached to the muscles of the pharynx, but
they are smaller and less numerous than in Ocnerodrilus, the pharynx being less
lobed and more compact than in that genus. These glands open through ducts, which
follow muscular strands into the pharyngeal cavity in a similar way as will be more
in detail described in Pontodrilus. In fact it is probable that all the suprapharyngeal
glands in the respective genera of Lumbricids open similarly and without any great
variation as to detail (fig. 18). The narrow ducts from the gland penetrate the pha-
ryngeal epithelium and form at its outer edge small ovoid pockets for temporarily
storing a small amount of the salivary secretion. These ducts end with the pharynx,

PACIFIC COAST OLIGOCH MTA. 67

the cesophagal epithelium neither being furnished with ducts nor storage pockets (fig.
18).

The suprapharyngeal glands are posteriorly connected with the septal glands,
not only with the nearest pair, but with all the pairs in the respective somites. In
Ocnerodrilus Beddurdi I called the attention to this fact, but I was not able to point
out the connection between all the glands, which connection, however, I do not doubt
really exists in all genera of this family, and probably in most other Oligochieta.

Septal glands(fig.2). As regards these glands our present species offers no great
peculiarities different from species of Ocnerodrilus generally. There are four pair which
surround the esophagus in the usual way in somites y, vi, vil and vill. ‘The glands are
considerably lobed (fig. 1-2), and decrease in size posteriorly. That is, the pair in y
is by far the largest, the one in vi is smaller and so on, the one in viii being much the
smallest. This gradual decrease in size posteriorly, though the most common one in
this class of Oligocheta, does, however, not always exist in all species. I have one
species yet undescribed from Guatemala in which all the glands are of the same size.
In our present species the glands are distinctly paired, but they lie so close together that
that they appear in each somite as one single gland surrounding the intestine. In
sections the glands are seen to be abundantly sealed with blood sinsues or larger
vessels.

These septal glands (fig. 2), connect one and all with the suprapharyngeal
gland, being, so to say, superposed on several main longitudinal muscular bands con-
necting the “hemmed glands with the body-wall in somite ix. Wide and narrow
ducts follow these muscles, causing the secretions of the septal glands not to empty in
the alimentary canal in the respective somites in the glands, but in the pharyngeal
cavity as shown in fig. 18.

Fig. 2 is, as far as outlines are concerned, a correct representation of these
glands from a section lateral to the cesophagus. Most details, however, are not filled
out. The glands on the upper side are those above the cesophagus, the lower row
again snase: below the esophagus, both opening on the upper side of the pharynx.

The sete occur in couples of two, as usual, the distance between sete 1 and 1
being about the same as the distance between 2 and 2. The shape is the usual sigmoid
one found in Oenerodrilus, and the size is rather large. The free points are slightly
corrugated. The sete occur in all somites after the first, but there are never any deyel-
oped setze where should be the inner couples in xvii. Very small undeveloped tips
may sometimes be seen close to and lateral to the male pore enclosed in the reserve
bag, but even they are not always present.

The blood vessels agree in all respects with those of Oecnerodrilus. There are
two pair of hearts, one each in x and xi and one pair of connecting vessels in ix.

Nephridia are found in all the somites except the first few anterior ones. In
somites iii to vy the nephridia are very small and dwarfed, but from there
on posteriorly they increase in size. Those in front of the clitellum are not fur-

68 CALIFORNIA ACADEMY OF SCIENCES.

nished with a glandular covering of peritoneal cells; those in the clitellum show a few
of those cells, while the nephridia posterior to the clitellum show a highly developed
envelop of peritoneal cells, similarly as is the case in some species of Ocnerodrilus.
In our present form the difference between the anterior and postclitellar nephridia is
very marked, the latter ones being prominently visible through the body-wall both in
alcoholic specimens as well as in alive ones. As regards the form of the nephridia,
it agrees in general with that of the various species of Ocnerodrilus, some of which I
have re-examined. The windings of the canals as well as the general arrangement
of the ducts is much the same in the genera which I have examined more in detail,
such as Oenerodrilus, Kerria, Deltania, Argilophilus, Pontodrilus. Especially is this
the ease with Oecnerodrilus and Kerria, the nephridia of which have been misunder-
stood, in several particulars, especially so as regards the windings of the canals and
the presence of blood vessels.

In Oenerodrilus as well as in Pheenicodrilus the nephridium consists of two
distinct parts, A, the folds, with the winding canals, and 8, the peritoneal covering,
with more or less numerous blood vessels. The peritoneal covering again is also diyided
in two parts, one upper almost free, and one lower surrounding or at least adjoining
the canals, about which more later on.

In Pheenicodrilus the nephrostome leads to the narrow duct which connects
with the folds of the main nephridial body. The folds of these canals are placed on
the outside of or rather on the upper edge of the peritoneal covering. The narrow
duct when it enters the fold is very narrow, in fact conspicuously so. In the neck of
the nephridium it coils itself several times around the part of the wide duct that is
enclosed in the neck. Retaining its narrow size it enters the anterior fold, in which
it is the most anterior and exterior canal, but nowhere does it appear to ramify as in
Pontodrilus, Deltania, Argilophilus, ete. It retains its narrow size all through the
windings (fig. 12), but increases in size in the posterior fold, in which the three canals
are of equal size.

In the apex of the spur, which as usual is thicker than the fold, the continua-
tion of the narrow duct connects with the very wide canal which later on forms the
bridge. After passing the bridge this wide canal becomes much narrower but still
continues as thicker than the other canals until it enters the posterior fold. It is also
less coiled than any of the other canals, in fact it is most conspicuous by being very
straight—it always occupies the under and inner side of the folds. Even in the
“windings,” which, in this species, occupy a very large part of the folds, this canal is
straight, while the two other canalsare coiled and bent. We thus find in this genus all
the principal parts of the nephridium of the much larger Argilophilus, and it may
be safely stated in all highly developed Oligocheeta the structure of the nephridia
are in the main the same. As far as Iam aware we may distinguish the following
divisions of a perfect nephridium:

PACIFIC COAST OLIGOCH MTA, 69

a. Nephrostomal part.
Nephrostome, or funnel.
Neck of nephrostome, consisting of glandular cells.
Narrow, or nephrostomal duct.
b. Main glandular part.
Neck of glandular lobe, connected with the nephrostomal duct.
Anterior fold.
The windings, or the coiled part of the two folds.
Posterior fold, in which the three canals are of the same size.
Spur, with four canals.
Bridge, with only one canal.
c. Efferent part.
Wide terminal-duct or outlet. The latter is frequently furnished with a
ceelomic bladder.
Nephropore.

When the nephridium is spread out and mounted in glycerine it is seen that
the canals are of different transparency. Thus the wide canal coming from the bridge,
together with its continuation in the spur, is much darker than the two other narrow
canals, which are conspicuous through their brillianey. These two white canals meet
in the very apex of the spur, while the two darker canals meet a little further in.
The greater obscurity of the wider canal is partly due to ciliation.

The nephrostome of Phwnicodrilus taste is furnished with a large neck, almost
as large as the rosette, containing several very large cells with conspicuous nuclei.
The nephrostome is about as Jong as the narrow tube.

In size the nephrostome ranges with the very largest, it being conspicuous
and readily dissected.

The marginal cells vary in number between eight and fourteen. There is a
larger central nucleus as usual. The glandular neck of the nephrostome contains
two of these nuclei much larger than any of those found in the rosette, though it also
contains several smaller ones. When the body of the annelid is laid open and viewed
from above, the nephrostomes are seen to lie on their side as in fig. 9, the side of
the thick rim of the rosette being uppermost. In order to see the rosette flattened
out it is necessary to dissect out the nephrostome, which operation in this species is
not particularly difficult, as the muscular tissues easily separate.

The peritoneal covering of the posterior nephridia separates itself in two dis-
tinct masses, one dorsal and one ventral, the latter being somewhat the smallest. At
the point where they join the peritoneal covering has narrowed down to a narrow
band connecting the two main parts.

The upper peritoneal mass does not surround or contain any of the canals or
ducts, but appears to be merely an appendage, a gland, as it were, engrafted on the
lower part. The canals and tubes are entirely confined to this part, as is also the case
in the nephridia of Ocnerodrilus and Kerria.

70 CALIFORNIA ACADEMY OF SCIENCES.

The peritoneal sac consists of very large cells with small, sharply defined and
very round nuclei. When these cells are empty they are very transparent and
their walls are very plain. There are a few blood vessels on the peritoneal sac, and
few of these cover also the folds.

Nephridia of Kerria (fig. 23). Having recollected Kerria McDonaldi in the
pond near Miraflores in the Cape Region of Baja California, the only place where it is
found to date, I have taken the opportunity to re-examine the nephridia in order to
ascertain their resemblance to those of our present form as well as Oenerodrilus. As
a consequence I am able to correct some errors and to add several details. The gen-
eral structure is the same as in the Oenerodrilus and Pheenicodrilus and the windings
of the canals the same as in the Argilophilus, ete. The two folds make a large
rounded loop upwards, and join posteriorly in a very long spur partly free from
peritoneal covering. The bridge connecting the junction of the spur and the
posterior fold with the junction of the anterior fold, wide duct and narrow duct,
is wide and ciliated, in fact it is the widest part of the canal. Posteriorly the
canal of the bridge projects into the spur forming the widest of the four canals in
this part. At the apex of the spur two and two of these canals are seen to join as
usual, forming two loops, one outside of and above the other. The inner and lower
one of these is the bend of the ciliated canal from the bridge, but it does not ap-
pear to be ciliated at this point.

The posterior fold contains as usual three canals, and so does the anterior fold.
The rounded stretch where the two folds meet is more irregular in outline, and con-
tains more windings than any other part of the fold, though not as many as in Ponto-
drilus. The nephrostome is connected with the main body by a slender and nar-
row tube, the connection being a little in front of the one between the wide duct and
the main body. This wide duct is almost straight with only a slight curve away from
the nephridium. It becomes slightly wider towards the nephropore, just before
reaching which it turns sharply upon itself.

Another point of considerable interest is the presence of numerous blood
capillaries on the nephridia, especially in the peritoneal sac, which they permi-
ate. The blood has its origin from two vessels, one from the branch from the ven-
tral main vessel and one from the branch from the dorsal main vessels, the two con-
necting by capillaries on the nephridial folds. Until now it has been supposed that
Ocnerodrilus and Kerria did not possess blood vessels on their nephridia, but this is evi-
dently an error as far at least as some species are concerned. Ordinarily these vessels
are not visible and not distinet from the peritoneal cell-walls but a staining with
orange G. will bring them out at once.

The generative organs, with one or two exceptions, resemble those in Ocnero-
drilus in form and general arrangement, and if it were not for the regular absence of
a prostate our species would be considered as a true Oenerodrilus. The Spermatheca
is very large and resembles in general outline that of Ocnerodrilus Beddardi. In
species of Ocnerodrilus the spermatheca always stands up and is pressed close to the

PACIFIC COAST OLIGOCH MTA. 71

anterior septum. In Phwnicodrilus taste, however, the spermatheca lies always flat
pressed against the ventral side of the body-wall, and is of sufficiently large size to reach
as far backwards as to the posterior septum between ix and x, which makes it about
equal in length to the diverticula of the cesophagus (fig. 1, spth.) The lower part
of the spermatheea is as usual more muscular than the free end, which in this species
is more or less, though always shallowly, lobed, showing a large number of incipient
diverticula irregularly formed and arranged. The spermatozoa are found principally
in these warty diverticula.

Testes are very long and situated as usual in x and xi. The posterior one at
least, and propably both pairs, connect with the sperm-sacs in the same somite.

Sperm-sacs are arranged as in some species of Oenerodrilus. They are all
paired. The anterior pair in ix is attached to the posterior septum of that somite.
This pair is very much lobed, the lobes being more in number and in shape more
round than in any species of Ocnerodrilus, the pair resembling two bunches of large
grapes, completely filling the whole available space in the somite, especially above the
diverticula and the cesophagus. The sperm-saes in x and xi are less or hardly lobed,
connecting with the the testes below, the latter, being long, slender and not branched,
reach across the somite and joining the sperm-saes in the posterior part near the sep-
tum. The sperm-sacs in xii are connected directly with those in xi, but otherwise
attached to the anterior septum of somite xii. This pair of sperm-sacs are lobed but
not as much so as those in ix.

There are two pair of ciliated rosettes in x and xi, and two pair of spermducts
as usual leading from them. The spermducets join, as is generally the case, forming a
single strand which runs close to, but a trifle more dorsally, than setze 2, until somite
xvii is reached. In this somite each spermduct enters a small muscular atrium deyoid
of prostate, and entirely confined to the longitudinal layer of the body-wall. As soon
as entering the body-wall and this muscular chamber the two lumens of the spermduct,
which until now had been separate, fuse together and enter the muscular chamber as
one single duct.

As atrium I consider a small muscular chamber entirely confined to the body-
wall, in which the spermducts open. This chamber is, however, devoid of any
glandular cell prolongation such as we are accustomed to find in Oenerodrilus, and
ordinarily it ends where the spermducts enter, which is at the upper end of the layer
of the body-wall. The atrium itself consists of an inner layer of epithelial cells,
which at the very pore are much larger and furnished with larger nuclei, but which
gradually decrease in size as they approach the place where the spermducts enter.
This layer is a direct continuation of the hypodermal layer of the body-wall. An
outside layer again consists of fine muscular fibres with smaller nuclei directly con-
tinued from the transverse muscular layer of the body-wall. We see thus that this
short chamber might in reality be nothing but the remnants of a degenerated atrium,
or rather remnants of the lower muscular part of a degenerated prostate, which gland-
ular prolongation has disappeared. That this is the case I judge from the structure

Memorrs, Vou. II, 4. February, 1895.

a

~J
bo

CALIFORNIA ACADEMY OF SCIENCES.

found in one specimen differing from any other which I investigated. Out of six
specimens which I sectioned off five agreed in all particulars as regards the absence
of a prostrate, as generally understood in Ocnerodrilus, neither did seven specimens
which I dissected show any trace of such a prostate. One specimen which I sec-
tioned, however, showed an abnormal prolongation of the muscular chamber, in every
particular resembling the lower muscular part or the atrium proper of the prostate as
characteristic of Ocnerodrilus. It ascended inwardly in the cavity of the body as
high as to sete 3 and 4, ended here blindly without any differential glandular part
or prostate proper, as is always found in Ocnerodrilus. Such “returns” to original
characters and ancestors must, of course, be expected, and are the more interesting
when encountered. We might on the other hand consider Phwnicodrilus taste as
standing on a lower grade than Oenerodrilus, the prostate not having yet appeared.
But against such a view speaks the fact that her organs are as highly developed as
in Oenerodrilus, which would hardly be expected of a lower form, in which we would
naturally look for a lower degree of development in several organs at the same time.
A degeneration of a certain organ, however, such as the prostate, would not necessitate
a similar degeneration of several other organs at the same time.

The absence of a prostate in Pheenicodrilus is of considerable interest, and I
think it clearly demonstrates that the absence or presence of this organ cannot be laid
at the foundation of families. Such absence of the prostate in an Ocnerodrilide is not
unexpected. A perusal of the various species of Oenerodrilus shows us how these
species may be arranged in a series according to the size of the prostate, the list being
headed by Ocnerodrilus occidentalis, with a very extended prostate, while at the other
extremity we find Ocnerodrilus Hendriei with the most diminutive prostate, not extend
ing outside of the somite. There is thus only a step further to Phcenicodrilus where
this organ is entirely absent. That this fact will have some influence upon our views
of the classification of Oligocheeta is evident, and several genera or even families
which hitherto have been considered far apart solely on account of the presence or
absence of a glandular prostate, must now be brought closer together.

Ovary and oviduct occupy the same somites as in Ocnerodrilus and offer no
characteristics of interest.

Habitat. 1 found this species in the Sierra el Taste, in the Cape region of Baja
California, some 40 or 50 miles north of Cabo San Lueas, at an altitude of 4,000 feet.
Later on I found specimens in great number in a garden at Pescadero on the Pacific
coast, on the western slope of the same sierra, but at an altitude of only a few feet above
the ocean. The water used for irrigation was taken from a creek coming from the
sierra.

The species lives in damp soil and oecur in great numbers, not mixed with
any other form as far as my experience goes. ‘The distribution of the species is most
interesting as on the eastern side of the Sierra in the valley of San José, [never found
any other Ocnerodrilid than Ocnerodrilus Beddardi. The question if the Sierra really
absolutely divides the habitats of the two only forms of this family found in the Cape
Region, further explorations are necessary to decide.

PACIFIC COAST OLIGOCH MTA.

~I
co

PONTODRILUS Grube.

The genus Pontodrilus, together with Photodrilus, is distinguished from all re-
lated genera by the absence of nephridia in the twelve anterior somites. In the
majority of species the nephridia commence in somite xiii, but in two species they
commence respectively in xiv and xv. A common character to all the species appears
to be the very great thickening of the septa between somite v and xiii.

I have so far on the Pacific Coast only found one species of the genus Ponto-
drilus. Examinations were made from alcoholic specimens, I having no opportunity
to examine them when [ found them in their native habitat—Mexico.

Pontodrilus Michaelseni n. sp.

This species differs from all other species, except P. Maronis,* which have been
referred to this genus in possessing a glandular crop occupying somites xiv, xv and
xvi, as well as in other minor details. The habitat of the species is the very narrow
moist line between high tide and dry soil on the shores of the Gulf of California
around Guaymas, Mexico. The soil in which it occurs is very sandy and thoroughly
soaked or moistened with the strongly saline water of the gulf. It occurred here in
large numbers, but unfortunately at my visit most of the specimens were immature,
only two possessing clitellum in the end of November, 1895. I dedicate this species
to Dr. W. Michaelsen of Hamburg, whose labors in our common field are among the
the most thorough and best.

EXTERIOR CHARACTERS.

The body of this species reaches a length‘of 3} inches with a width at the
clitellum of less than } of an inch, but the majority of specimens are somewhat
smaller. The above measurement refers to specimens slowly killed and then hard-
ened in alcohol. The body is tapering towards the tail end, the latter however being
slightly swollen (fig. 24).

The prostomium encroaches on somite i, dividing it about one-half. Somite i
is slightly larger than any of the following somites (fig. 25).

The clitellum commences in xiii and in full-grown specimens includes part of
xix. It is incomplete in a peculiar manner. In xiii to xvii inclusive it is only
developed on the dorsal side of the body. But in xviii and in part of xix the clitellum
is only developed on the ventral side of the body, though this fact cannot be ascer-
tained from exterior inspection. Viewed from the underside the clitellum appears to
be on a line drawn through seta one. Between xvii and xviii the clitellar swelling
recedes slightly, again to widen out in xviii, and here joining to a pair of ventral
cushions, between which and the clitellum proper are situated the male pores.

Another pair of swellings are noticed around the spermathecal pores, covering

“I received M. Perrier’s memoir on Pontodrilus only after this paper was partly in print.

74 CALIFORNIA ACADEMY OF SCIENCES.

External pores. There are no dorsal pores. The spermathecal pores are
found between somites vii/viii and viii/ix in front of seta 2, each one situated ona
slightly elevated cushion. The ovipore in xiy in front of seta 1. The male pores are
in xviii in front of and in line with seta 2 (fig. 266). The nephropores are in front
of seta 2.

Setw. The setse commence in somite ii, eight in each segment and in couples.
The distance between 3 and 4 is only slightly larger than that between 1 and 2. The
distance between 1 and 1 is nearly twice as large as that between 1 and 2, and the
distance between 2 and 3 is a little smaller than that between 1 and 1 (fig. 27).

Color of body pale flesh, rather transparent and marbled very much like
Deltania. Clitellum yellowish.

INTERNAL ANATOMY.

Body-wall. The body-wall appears to me to be of unusual thinness, throughout
the length of the body. The dorsal side is slightly thinner than the ventral side, at
least anterior to the clitellum (fig. 29). Dorsally the longitudinal muscular layer is
of about the same thickness as the transverse layer while on the ventral side the
longitudinal muscular layer is about twice as thick as the tranverse muscular layer.
This refers to the anterior somites. To this there is however an exception in somites
viii and ix where on the ventral side in the vicinity of the spermathecal pores the
transverse layer is thicker than the longitudinal layer. The transverse layer tapers
down towards the spermathecal pores, but this thickening is found only in the imme-
diate vicinity of the spermatheca. In the clitellar somites the relative development of
the muscular layer is very different. Here the inner or longitudinal muscular layer
is enormously thickened laterally in somites xvii and xviii or in the vicinity of the
male pores (figs. 37, 38, 40, 41), while in the anterior part of the clitellum the longi-
tudinal layer is only thickened ventrally, between the inner couples of sete it here
being at least twice as wide as it is dorsally (fig. 39).

Clitellum offers many points of interest. It has already been stated that this
organ is incomplete, that is, not simultaneously developed on the dorsal and ventral
sides. A section through an immature specimen shows (fig. 38) that the clitellar
glandular layer is developed only between the seta, that is, from seta 4 ventrally to
seta 4, while dorsally there is no trace of such cells. As regards the nature of these
cells it is to be remarked that they are unusually small or rather thin compared to the
larger and thicker cells of the dorsal part of the clitellum in the anterior somites of
that organ. These latter cells offer nothing in particular of interest, resembling those
of other genera of the family as far as I can make out. Unfortunately most of the
specimens were immature and only two possessed clitellum, but these two had unavoid-
ably not been treated, and had contracted to such very great extent that the finer
structure of the clitellum had been hopelessly lost. From eross and longitudinal see-
tions made it was, however, evident that the clitellar glandular cells, which constitute the
clitellum, do not extend ventrally further than to sets 1, thus leaving the ventral space

PACIFIC COAST OLIGOCHATA. iD

between sete 1 and 1 with the usual layers less the glandular cells (fig. 41, 5. ¢/.)
marking the points where the clitellum ceases. We have thus before us a species
with a dorsal and lateral clitellum in some somites, and with a ventral and lateral in
one somite.

In all of the clitellar somites the inner or longitudinal muscular layer possesses
an enormous development laterally, especially so in the region of the prostate, where
it reaches a width four times that at the dorsal and ventral part. The reason of this
enormous thickening of this layer is readily understood when we view one of the
sections of this region. The part of the body-wall, on which rests the prostate, is
attached by numerous arciform muscular bundles, arranged in a fan-shaped manner,
to the dorsal and lateral sides of the body. These muscles are more numerous and
stronger than any I have seen described in other species, making it possible for this
region of the body to contract in a most characteristic manner.

When the body is opened and flattened out these muscles are seen to spread out
from the male pore laterally, connecting the center of somite xviii with the two
nearest somites xvii and xix as I have endeavored to show in fig. 42. As may be
judged from the figure the muscles are arranged in regular fascicles of which we may
count twenty-two to twenty-four as being more prominent and overlapping the other—
a smaller number of fascicles situated below. The wider periphery of the attachment
is situated on the body-wall above the prostate, but the lower and narrower part of
the attachment of the fan is situated on a peculiar swelling which I have designated
as the copulatory cushion (fig. 42, ¢. ¢.; fig. 41, c. ¢.; fig. 38, ¢c. c.; figs. 44, 45),
apparently a thickening of the longitudinal muscular layer, transversed by innumer-
able other muscles in every direction, all connected by what looks like connective
tissue. The size of this copulatory cushion is very great; not only do these swellings
project considerably outward but inwardly they encroach to a great extent on the
internal cavity, especially so in mature specimens. This peculiar organ is entirely
muscular, there being no sign of any outwardly opening papillae. The prostate pore
is situated between this cushion and the projection caused by the increased thickness
of the longitudinal layer and the contraction of the fan-shaped muscles. In fig. 43
the section, a longitudinal one, is somewhat oblique, having followed the large muscular
strands which surround the outlet duct of the prostate as well as the muscles of the
copulatory cushion. Fig. 44 represents a more yertical, longitudinal section, more
interior to the large fan-muscles, which do not show in the section except their inner
attachments (a. m.) The large arciform strands vary considerably in size, those
nearest the body-wall being the smallest, those nearest the intestine the largest.
When a cross-section through somite xviii is viewed it will be seen that the smaller
strands are filled between with connective tissue (fig. 41, ¢. ¢.), while a concentric
transverse muscular strand is crossing them near the inner angle of the prostate pore.
Other muscles connect the glandular prostate with the body-wall (figs. 37, 40 m and
42 sp.m.) The large strand sp. m in fig. 42, connects the bend of the prostate, where
enters the sperm duet, with the ventral part of the body-wall under the ventral nerye-
cord,

76 CALIFORNIA ACADEMY OF SCIENCES.

Septa. The first very pronounced septum is found between somites iv and y.
3eddard has already remarked that the septa in Pontodrilus hesperidum are unusu-
ally thickened, some of them being thicker even than the body-wall at the point
where they are attached to it. The septa in our present form are almost gradually
increasing in thickness towards the one between xi and xii, which septum is the
thickest, being almost as thick as the ventral body-wall. The septum dividing xii
and xiii, while thickened, is much thinner, and those bounding the somites down to
xix are almost normal in thickness, that is, equal those posterior to the clitellum. As
a rule the attachments of the septa correspond with the intersegmental furrows. The
septa bounding vii/viii and viii/ix are, however, exceptions as far as the place of their
ventral attachment is concerned. These two septa are here affixed to the body-wall
half-way between the sete of the anterior somites, respectively viiand viii. This makes
it appear as if the spermathece opened in the centre of the somite, when in reality
they open as usual in the intersegmental groove.

Alimentary canal. The alimentary canal takes the shape of a long, narrow
duct, singularly straight and without any prominent characteristics until it reaches
somite xiv, in which somite commences a kind of gizzard of peculiar construction
(fig. 29, giz.) Though this organ resembles a gizzard in outward form it is in reality
no gizzard at all, but rather a glandular modification of the alimentary wall. With
a gizzard we must of course mean an enlargement of the alimentary canal in which
the muscular part has reached an enormous development in order to grind the food
properly. In the organ referred to in our present species the muscular layers are on
the contrary not increased in size, the thickening of the wall being caused exclusively
by a new layer composed of glandular cells, which has been interposed between the
transverse muscles and the inner epithelium, thus forming a glandular crop between
the cesophagus and the tubular intestine. This organ occupies three somites, Xiv, xv,
xvi, or very much the same place as is so frequently the location for gizzard in other
Oligochete. If we view a longitudinal section of the body through this crop (fig. 47)
we find it to be more or less tapering towards either end. The large longitudinal
blood yessel lies almost immediately on the top of the outer or chloragogic cells, in
places penetrating them with connecting vessels which supply the underlining sinus
with blood (fig. 47, d. v. ¢.).

The chloragogic layer of cells vary considerably in size. Sometimes there is
more than one row of cells, one projecting above the other. The nuclei are oyal and
situated at the place where the cells become narrower. The longitudinal muscular
layer is narrow, about two strands thick, immediately superposed the transverse layer,
which consists only of one single thickness of strands (fig. 48 to 54 ¢. m.) The case gen-
erally observed in gizzards is that this layer is composed of a great number of strands
more or less regularly arranged around a central plate. Below this transverse layer,
commences the very thick layer of glandular cells, about 12 cells wide in centre.
In the upper part of this layer are seen numerous blood laeunes, which in

ro

places join the muscular layers (fig. 49 to 53 0/.), at other times are more or less

~I

PACIFIC COAST OLIGOCHATA,

perfectly enclosed in the glandular layer, forming generally a row of vascular
lacunes near its outer margins (fig. 48 4/. s.) apparently without touching the mus-
cular layer, or doing this only at certain places. Figs. 49 to 53 represent cross-section
from different parts of the crop, from the posterior part, near the boundary of somite
xvii (fig. 49), to the anterior part of xiv (fig. 52), illustrating the variations in thick-
ness of the glandular layer. In the posterior part this layer is very thin (fie. 49 ql.)
consisting only of a few cells, from one to three cells wide. In figs. 50 to 52 the
glandular layer is seen to have increased in thickness and so has the inner epi-
thelial cells. Fig. 53 is a portion of the glandular layer, in which the inner lumen
is more plainly represented. The outer part of this glandular layer is divided up in
lobes by the numerous blood sinuses, and in each such lobe there is a wider or nar-
rower, generally, branched lumen, which, however, I have not been able to follow
down to the epithelial cells.

The sacculated intestine commences at the posterior end of the crop, and
offers nothing of great interest or characteristic.

There is no thyphlosole, but the intestine is otherwise yery rich in blood
lacunes.

Pharyngeal or Salivary Gland. Pharynx which occupies somites ii and iii is
only developed dorsally. It is superposed by a large mass of glands and muscles, as is
usual in a large number of Oligochete. In outline this glandular mass is remarkably
even, especially so at its posterior end. In a longitudinal section we see customarily
three lobes (fig. 29), supported by long strands of muscles, running back to the pos-
terior boundaries of somites yi, vil and viii.

On the ventral side there are seen three of those muscular strands, similarly
running back to vi, vii and viii, indicating that there is a row of similar strands corres-
ponding with somites ii, iii and iv. In somite vy there is a pair of smaller glands of
similar nature attached to muscles which connect with the larger strands of the main
gland (fig. 29, s.s. g/.)

A cross-section of this glandular mass (fig. 30), shows us that the glands are situated
principally on the periphery, supported by muscles (ms.), while the inner and posterior
part is principally taken up by strands and ducts. A division of the massin three more or
less distinct layers is discernible, probably corresponding to somites ii, iiliand iv. These
glands communicate directly by means of ducts with the epithelium of the pharynx.
In fig. 29 these ducts are roughly represented as dark violet. In the cross-section
(fig. 30), the darkest blotches are intended to represent the ducts, while the lighter
colored violet ones are the glands. In fig. 31 a lobe of the longitudinal section is seen
in a larger magnification, and in fig. 32 a smaller lobe, yet more highly magnified.
The glandular cells are rather of varying size, and arranged around the margin of
glandular sack, leaving the inner space open. The cell cytoplasm is massed in places,
leaving in other places larger or smaller, generally roundish, vacuoles (fig. 35, va.)

In figs. 33 and 34 I have endeavored to show the relative arrangement of the
muscular strands, the glandular ducts and the glandular cells. As will be seen these

bt |

'a) CALIFORNIA ACADEMY OF SCIENCES.

ducts are in places entirely closed in by muscles, while the glands themselves are
only supported by them.

The ducts lead directly to the pharyngeal epithelium; arrived here they branch
out sending numerous discharge-tubes between the epithelial cells (fig. 56 g/. dt.), dis-

. = . & . = . nr 4

charging the salivary mucous in the pharyngeal cavity. These ductules are fre-
quently, though not generally, branched while in the epithelial layer. Each ductule
is furnished at the distal end with a small storage chamber (564) of oblong form and
considerably smaller than the nucleus of the epithelial cells.

Septal glands. As hasalready been stated there are five pair of very small glands,
which are pricipally attached to the connecting vessels in somites v to ix, and sit-
uated on the ventral side of the cesophagus. These glands do not hang on closely to
the septum, but are apparently suspended from it only by a few tiny mesenteric tis-
sues and by a muscle or two. In longitudinal section they appear as suspend entirely
between the two septa (fig. 29), while in cross-section they are seen to be affixed to
the connecting vessels and from them project laterally, the point of affixion being
close to the ventral vessels (fig. 70 g/.) The general outline of these glands resemble
the so-called liver cells attached to the connecting and other vessels in some Lumbri-
cides, but the structure is similar to the salivary glands. The gland in v resembles
exactly the structure in the salivary glands which open in the pharynx, it being
transversed by blood capillaries, infested with the same parasites, supported by mus-
cles, and finally is only sparcely surrounded by floating, globular, coelomic cells. The
other gland in vi to ix are all surrounded by a thick coating of these floating ccelomie
cells. These glands stain in the same way as the salivary glands, their secretions being
stained deep violet with heematoxylon-orange, while the ecelomic cells stain pale yellow.
A fine and very thin duct runs backwards and upwards from the far upper end of
each gland towards the alimentary canal to its junction with the septum, but I
have some doubt about it emptying into the intestine, and it is much more
probable that in Pontodrilus, as well as in Phoenicodrilus and Oecnerodrilus, these
septal glands empty into the pharynx. None of my sections, however, show
this to be the case. Certain it is that in Pontodrilus the various septal glands
are not as closely connected with the salivary glands as in the just mentioned
genera, in which the respective glands are actually not only suspended from the same
longitudinal muscular band, but along and resting on the latter run also the collective
duets of the glands. Among the salivary and septal glands are seen numerous ir-
regular, generally oval or oblong bodies full of nuclei. These are the terminal pock-
ets of the capilaries, generally termed blood glands.

Blood glands (fig. 734, t.). Td. Perrier was the first to describe blood glands
in Pontodrilus, but he found them exclusively in the blood vessels or at the end of
the capilaries investing the nephridia. In our present species, P. Michaelseni, I have
found these glands only in the capillaries of the salivary and septal glands. They
here occur in very large numbers, especially in the former, being massed at
or near the posterior edge of the gland in varying numbers. Some specimens con-

PACIFIC COAST OLIGOCH MTA, 79

tain comparatively few blood glands, others three or four times as many. They
are of all sizes and shapes, as Perrier has shown. Some contain only one single
nucleus, then frequently surrounded by a blood clot; others again contain a very great
pumber of nuclei, which are then situated in a sac-like pocket at the end of the blood
‘apillary. In some of the larger blood vessels in the salivary gland the blood gland
takes the form of a “hertzkérper.’ The smaller ones situated on the capillaries
may be named terminal blood glands, while those situated inside the larger vessels
may be designated interior blood glands. The structure of the two are at least in this
species very similar,

In fig. 73, a. to s., [ have endeavored to illustrate the structure of these blood
glands from sections. In g.a large blood vessel with a blood clot, at the base of which
is an inner blood gland. On one side of the blood vessel is a part of a salivary gland
with brown secretions. In a. a small terminal blood gland is shown, and in 0. and e.
some of a greater development. The nuclei are not always surrounded by a distinct
cell membrane, in fact in almost every gland are found some nuclei with distinct cell
membranes while others lie loose in the granular serum. The exterior line in all
the figures represents the wall of the blood vessel, and the difference between the
terminal and inner blood glands consist in reality only in the absence or presence of
blood surrounding the glands. As far as the granular protoplasm concerns it is always
differentiated in two parts. The one at the distal extremity is more evenly dif-
fused and finer grained than the one next the capillary, which again is coarser, streaky
and which, besides, stains differently or at least more intensely than the other. Many
of the glands contain larger or smaller bodies (p., ¢. and 0.) equally of round form
and lighter in color than the cytoplasm, but sometimes they are very opaque, stain-
ing deeply as at r., the two classes probably being of entirely different character.
The former resembles a pale nucleus, while the latter opaque bodies appear only to be
secreted matter. The paler ones may possibly be parasitic protozoa.

The blood glands described by Claparede, Lankester and others in Lumbricus,
ete., are probably of a similar construction, and judging from the figure given by
Michaelsen of the “ hertzkérper” in Enchytreeus, we may conclude that it, too, is
identical with the blood gland in Pontodrilus.

Spermathece (figs. 30, 55, 56, 57, 58). There are two pair of spermathecz
found in somites viii and ix, the exterior pores being as usual in the intersegmental
grooves between vii/viii and viii/ix in line with sete 2. Hach spermatheca possesses
a tubular diverticulum, the junction of the two being in the body-wall. The position
of the diverticulum is always ventral to the spermathece proper. This is cylindrical,
quite narrow, with a larger globular chamber at the free inner end, in which the wall
is much thinner than in the cylindrical part. At the junction with the body-wall
is a much larger swelling, the lower and more strongly muscular part of the main
cylinder being greatly widened, presenting a muscular cushion partly projecting above
the body-wall, partly again being immerged in it. The spermatozoa are principally
massed in the inner globular chamber, though they are seen also in the diverticulum.

80 CALIFORNIA ACADEMY OF SCIENCES.

The main spermatheca is strongly muscular, especially at its lower end where
the muscular layer is much thicker than the inner epithelial cells. These latter are
large with oval nuclei. Outside of these cells and between them and the muscular
layer are seen a row of interstitial nuclei of much smaller size than the nuclei of the
epithelial cells (fig. 56 intes. n.) The structure of the diverticulum is somewhat
different. Interiorly we find large epithelial cells with large round nuclei. Outside
of them is a single row of interstitial nuclei. Surrounding them we find a circular
muscular layer varying in thickness (fig. 57, c. m.) and with few, small nuclei.
Exterior to this layer are seen numerous blood vessels, and outside of them a two or
three cells thick layer of glandular cells (g/. ¢.)

When the body of the worm is viewed from the interior, with the alimentary
canal removed, the spermatheca as well as their diverticula are seen to extend back-
ward, parallel with the ventral ganglion. The diverticulum is always much shorter
and narrower than the spermatheca proper.

Testes and Ovaries. The former organs consist of two pair of minute narrow-
lobed bodies (fig. 29, ¢. 59, 60), the lobes being all in one plane, parallel to the body-
wall. One pair are in x and one in xi as usual.

The ovaries consist of one pair of flat bodies with wavy margin and wide and
shallow lobes, distributed in both a horizontal and vertical plane (figs. 61 and 62).
As usual the ovaries are in xiii.

The oviduct is placed as usual with the funnel in xiiiand the pore in xiv. The
funnel part is very thick, fig. 63 drawn from a longitudinal section.

Ciliated rosettes, Spermducts and Prostates. There are as usual two pair of
ciliated rosettes in x and xi opposite the testes. The funnels are very thick and not
much crimpled or hardly crimpled with one flare on either side. The epithelial nuclei
are quite long, and their cells are superposed a thick layer of very distinct blood
vessels (fig. 65, cr. 66 and 67). The spermducts unite to a single duct which passes
immediately outside of the second setee. The duct is unusually narrow, the narrowest
I have seen in any species of this size. The spermduct enters the glandular part of
the prostate just above the intersegmental groove separating xvili/xvii (fig. 42, spd.;
68, spd.)

The prostate is tubular, very large, bent upon itself once. It starts from the,
male pore, which is situated in the center of xviii, forwards, running parallel with
ventral nerve cord. When it reaches xvii it turns backward, its apex being in the
center of xviii. The prostate consists as usual of two distinet parts, connected in the
center of xvii. The advancing part is strongly muscular, the returning part again is
glandular. The prostate is cylindrical, the two halves being almost equal in thickness.
The part which penetrates the muscular body-wall is several times thinner than the
other part. The muscular part consists of two layers, the inner one consisting of a
row of epithelial cells with oblong nuclei. The outer layer, which is very thick, con-
sists entirely of circular muscles with a few small nuclei.

PACIFIC COAST OLIGOCH ATA, $1

The glandular part of the prostate which commences at the anterior bend of
the organ consists of two or possibly of three layers. The inner lining consists of only
one layer of epithelial cells with ovoid nuclei. These long cells appear to be sur-
rounded by a narrow zone of fibrous or perhaps muscular tissue with few nuclei.
3ut by far the greatest part of the prostate consists of fibrous tissue with numerous
small roundish nuclei, with here and there a cell being visible, and with a few cells
of a glandular appearance (fig. 68), especially toward tie circumference. In cross-
section the anterior part of this prostate is triangular in outline, while the muscular
part is always circular, and in size always thicker. The very narrow part of the
prostate which penetrates the body-wall is strongly muscular of the same general
structure as the free muscular part. The general structure of the prostate appears
similar to the one of Pontodrilus hesperidum as described by Beddard. He has
pointed out the absence of a regular layer of glandular cells in the prostate of that
species, and it is possible that this construction of this organ which thus ap-
proaches that of Ocnerodrilide, is not a species but a generic character, if it does not
prove to be of even greater value.

Vascular system. There are a dorsal and ventral vessel, but no subneural, nor
any subintestinal vessel, and no thyphlosole. The two main vessels are connected
with hearts in x, xi, xii, xiii, the most posterior one of which is found in somite xiii,
immediately in front of the sacculated intestine. ‘This heart is the largest, the others
gradually decrease in size forward. The posterior part of these vessels are entirely
free of brown cells. The ventral vessel is forked in somite ix in two parallel
lateral vessels, there being no central vessel left. These two branches are always of
unequal size, both being situated immediately under the cesophagus (fig. 64 7. ».) In
the somites anterior to x, these branches of the ventral are connected by laterals with
the dorsal vessel. In one specimen the ventral fork commenced in xii (fig. 64).

Between the dorsal vessel and the ventral forks there are connecting vessels,
one pair in each of the somites y, vi, vii, vili and ix. To the ventral parts of these
connecting vessels are attached oblong glands, which again are surrounded by a coat-
ing of globular brown cells. These glands do not extend clear to the dorsal vessel,
but end laterally before reaching it. The nature of the glandular cells appear the
same as those of the pharynx, staining in exactly the same way. The cells of these
septal glands are more numerous in the anterior somites, gradually diminishing
posteriorly, while the opposite is the case with the free round cells which are more
numerous in the posterior glands (figs. 29 g/. and 71 g/.)

Nephridia (figs. 71 and 72). These organs commence in somite xiii, or in the
same somite as the ovaries. The first two anterior nephridia are furnished with a
smaller covering of peritoneal cells, but already in xvi do the nephridia attain their
full size, as in all posterior somites.

The nephridia are built upon the same general principles as those of Argilo-
philus, Deltania, Oenerodrilus and Pheenicodrilus, as well as of Lumbricus as shown
by Benham. We find here the corresponding ducts, canals, lobes, ete., and a gen-

82 CALIFORNIA ACADEMY OF SCIENCES.

eral description of them will be superfluous. The spur is directed backwards and the
two folds are directed upwards in about right angle to the spur and narrow duct as
well. The spur and the folds rest on the large lobes of peritoneal cells, one of which
is posterior and one anterior. The posterior one surrounds the spur, upon the ante-
rior one, which is the smallest, rests the two folds. The narrow duct is not unusually
narrow, while the wide duct or outlet duct is very narrow, not any wider than the
narrow duct. The neck of the anterior fold is, where it connects with the nar-
row duct, very wide, enlarged, irregular and sigmoid, gradually increasing in size to
the anterior fold. Where the two folds join, the fold is always very coiled. The tube
forming the bridge is not any wider than the clear canals, but it is less clear or trans-
parent, just as in Lumbricus. The canal leading from this bridge into the anterior
fold, is straighter, darker, and slightly wider than the two bright tubes which are
much coiled and situated more anteriorly and superiorly to the straighter canal. This
coiling ceases as soon as the big bend and windings are passed and the posterior fold
is reached.

The nephrostome is large. The marginal cells in the rosette are only slightly
decreasing in size toward the extremities or centripetal marginals. There is a large
centripetal protuberance surrounding the inner opening of the duct, as in Lumbricus.
as described by Benham, but the centrifugal cells are less regular and more scattered,
The centrifugal cells are never hidden by the centripetals as in Lumbricus, and the
whole centripetal protuberance is most prominent seen in whatever direction. ‘The
outlet duct enters the fold much closer to the narrow or nephrostomal duct than is
usual in Oligocheeta, in fact it connects with the free neck of the anterior fold, close
behind the septum.

The relationship of Pontodrilus Michaelseni to the other species of the genus
is not as clear as we might wish. Beddard’s description and notes in his paper,
“V. Some new or little known Oligocheta,” are the only comparative remarks yet
made on the few worms which are grouped under this genus, an arrangement which
must be considered as entirely preliminary. The only very characteristic features
which connect the six species of the genus is the commencement of the nephridia
posterior to somite xii, and the opening of the spermduct into the prostate, absence
of typhlosole, grape-like sperm-sacs, and no penial sete. None of these species
have been sufficiently described, an unavoidable fault attendant all species im-
mersed in alcohol without previous careful preparation and evacuation. In the
following table I have endeavored to compile the characters of the various species as
far as I can make out fromthe descriptions, no specimens for comparison being in my
possession. I include here, as suggested by Beddard, the genus Photodrilus Giard.
I have had no access to Grube’s description of P. littoralis, and have therefore
excluded it from this table.

PACIFIC COAST OLIGOCH UTA, 83
SPECIES OF PONTODRILUS AND PHOTODRILUS.
P. Michaelseni| P. arene P. hesperidum P. insularis P. Marionis | P.. phosphoreus
n. sp. | Fr. Miller & Bedd. Rosa. Perrier. Dugés.
= ais Michaelsen.
Nephridia commence yp re re
492: SOMO ara cleooe xi. xii. xili ? xill. XV Xiv
Nephropores in line
With SUC sree) tetaists |= ? ? 4. De 2:
Gizeardadeataeepaitee Rudimentary. | Absent. Absent. Rudimentary.) Rudimentary? Absent.
esophageal swellings..| None. None. None. None. None. Four in x-xiil.
Glandular crop....... Large in xiv, | None. None. None. Present. None.
XV, Xvi. | |
Septal glands.......-- In vy to ix. ? | None. None. Present. In y-ix.
|
Penial sete ........-.-| Absent. Absent ? Absent. Absent. Absent. In xviliand xii
Common set, ........ Plain, regular.| Ornamented, | Plain, regular.) Plain,irregular Plain, regular. Plain, regular.
regular. | behind.
|
. . | . . | . . . . . . .
Spermatheca.......... Two pair in| Two pair in| Two pair in| Two pair in| Two pair in One pair in ix;
viii and ix;|] viii and ix;| viii and ix;| viii and ix;| viii and ix; one diverticle.
| one diverti-| one diverti- | one diverti-| no diyerti- | one diverti-
| cle. cle. cle. cle. cle.
Sperm-sacs.......---- Two pair,| Two pair, | Twopair,rac-| Two pair, | Two pairin xi Two pair in xi
grape-like, grape-like,| emose,in xi| grape-like, | and xii. and xii.
in xiand xii.|. in xiand xii.) and xii. in xiand xii. |
| |
Glitetin eras ieco a xiii-xvili, xiii-xvii, eed |? xiii to xvii xili-xyli ?
incomplete. incomplete. | complete (?).
|
Prostomium . ........| Encroaches on! Encroaches }| ? Does not en- | Encroaches on Does not en-
somite 1. on somite 1. | croach on! _ somite l. croach on so-
| | somite 1. mite 1.
; , my ee] |
Ventral papille....... In xvii and xix One central,
xix—xx, and one
| ditto, xx—xi.
abun. pen bas ats Guaymas, Desterro, Jamaica. Aru Island. Marseilles. France.
Mexico. Brasil.

In the above table I have principally followed Beddard, but I cannot agree
with his opinion as regards the identity of Grube’s P. /ittora/is, with that ot P. Marionis
of Perrier. As Perrier remarks, the ventral median papillee described by him in P.
Marionis are not found in P. /ittoralis, in which species these papillee were paired.
Until Grube’s species has been found and redeseribed we must therefore accept it as
a separate one. It is of course not by any means impossible that one or more species
of Pontodrilus are to be found in the same locality.

In reference to the gizzard I have given it as rudimentary in two species. This
refers only to an anterior gizzard close behind the pharynx, in which the muscular
layer of the cesophagus is simply thickened as shown in fig. 46 Aands. Beddard says
that P. /ittoralis Grube has also a rudimentary gizzard, but I think that he judges by
M. Perrier’s description of what I call a glandular crop in the somites posterior to the
testes. It may not be impossible that the muscular thickening of the cesophagus
really exists in all the species more or less prominently, as it is easily overlooked.

$4 CALIFORNIA ACADEMY OF SCIENCES.

That the glandular crop really exists in P. Murionis is evident from Perrier’s drawing
(fig. 22, pl. xvi), though I cannot accept the epithelial nature of the cells. Perrier
says nothing of the place from which this drawing is taken, but I suspect that it was
from a section of the intestine between the tubular and sacculated parts, similar as in
P. Michaelseni.

LITERATURE.

Bepparp, Fr. E. Abstract on Some Investigations into the Structure of the Oligochwta. Ann. Mag. Nat. Hist.
Jan., 1891, p. 96.

Bepparkpb, Fr. E. Some New or Little Known Oligocheta. Proceedings of the Royal Physical Society, Edinburgh,
June 13, 1893.

Bepparp, Fr. E. On the Anatomy of Oecnerodrilus. Proc. Royal Society of Edinburgh, vol. xxxvi, p. 563.

Micuartsen, W. Terricolen der Berliner Zoologischen Sammlung II, page 14.

Benuam, W. B. An Attempt to Classify Earthworms. Quarterly Journal of Microscopical Science, vol. xxxi,
part il, page 243, ete.

Rosa, DANrIeLE. Sui Generi Pontodrilus, Microscolex, e Photodrilus. Boll. Mus. Torino, vol. iii.

Rosa, DanieLe. Die Exotischen Terricolen des K. K. Naturhist. Wien. Annal. K. K. Nat. Hofmus. Wien, Bd. vi,
p- 387.

Etsen, G. Anatomical Studies on New Species of Ocnerodrilus. Proc. Cal. Acad. Sci., ser. 2, vol. iii.

Etsen, G. Anatomical Structure of Two Species of Kerria. Proc. Cal. Acad. Sci., ser. 2, vol. iii.

Perrikr, EpMonp. Etudes sur l’Organisation des Lombriciens Terrestres. IV Pontodrilus. Archives Zoologie
Experimental, vol. ix, pages 175 to 248, pl. xiv. to xviil.

Eclipidrilus frigidus Eisen.

Through the kindness of a friend traveling in Sierra Nevada I have received a
small number of specimens of this interesting oligocheeta, but unfortunately all the
specimens were in a poor state of preservation and much macerated. However
I was enabled to make several very nearly continuous series of sections and thus settle
several very important points in the anatomy of this worm. My former study of the
species was entirely dependent on dissection, which could not possibly reveal all the
details of this minute species, especially as regards the spermducets, the presence of
which I am now able to demonstrate. My present researches show that the species
is less erratic in its anatomy than I first supposed, while again in many respects it
differs strangely from its nearest allies, the various genera of Lumbriculide. For
the present I retain the family of Eclipidrilidee, but not on the same grounds as
formerly, and I now consider it rather as a subfamily to Lumbriculidee than one
standing isolated, however with strong leaning towards Moniligaster.

The generative organs are situated as follows:

Testes, two pair. The anterior pair attached to the anterior septum of somite
ix. The posterior pair similarly to the anterior septum in somite x.

Ovary, one pair attached to the anterior septum of somite xi.

Oviduct in xii, opening in front of the inner pair of sete.

Spermathece, one pair in ix opening posterior to the setze and near the posterior
septum.

Atrium and prostate, one pair opening in x, posterior to the inner pair of sete.
This organ, which is very long, occupying seven to eight somites, consists of three
parts, first, one anterior atrium and prostate proper, second, a thin and narrow part

PACIFIC COAST OLIGOCH ®TA. 85

connecting the former with, third, the posterior one, a storage chamber for sperm-
atozoa,

Spermducts are two pairs, exceedingly narrow, opening close together in the
posterior part of the storage chamber.

Ciliated rosettes, two pairs, the anterior pair opening in ix. The posterior pair
in x, both in front of the septum. They are exceedingly thin and flat. The posterior
wall is attached to the septum.

Sperm-sacs consist of a pair of very long continuous sacs, which cover the
generative organs, including the spermatheca, and extending from somites x to xvii
or xviii, generally several somites posterior to the caudal part of the storage chamber.

Setw are 8 in 4 pairs in each somite, commencing with the second.

The vascular system is characterized by blind forked vessels in the posterior
30 odd somites, thus bringing the genus in close relationship with Lumbriculid:e.
Another characteristic is the two loops of lateral vessels which branch out from the
main vascular trunks in somites ix and x, and which run backwards as far as the end
of the storage chamber. The anterior eight somites contain winding lateral yessels
connecting the two main dorsal and ventral trunks. Hertzkorper in the dorsal vessel
as well as in the branches in the intestine. After these preliminary references to
the main anatomical points I will enter more fully upon the anatomical and_histo-
logical structure of the various organs.

Body-wall and Chitellum. The finer structure must be left for future study.
The inner longitudinal muscular layer is considerably thicker than the transverse
layer and hypodermis together. The longitudinal strands are very thin and ribbon-
like, some being much longer than others and reaching through the width of the
layer, others being very much smaller, situated principally close to the transverse layer.
Figs. 77, 79, 99, ete.

The clitellum comprises about 6 somites, commencing in the posterior part of
ix and extending to the center or posterior part of xiv. The clitellar glandular
cells, one layer thick, are oblong, irregular, flask-like, containing very coarse, angular,
grains (fig. 79). They are separated or interspersed by large non-staining cells.
The peritoneum is rather poor in blood vessels, but the layer is very thick, in places

almost as thick as the longitudinal muscular layer (fig. 82, pr).

Septa. None of the septa are abnormally thickened. The first distinct sep-
tum is seen between somites v and vi. The septa are st ‘aight, not cup-shaped. Those
surrounding the various divisions of the prostate are much firmer than the others.
They constrict the prostate, in fact the latter appears notched at every septum
(figs. 78, 92, 94).

Alimentary canal. The alimentary canal can properly be only divided in two
parts—pharynx and intestine. The pharynx which ends in somite v is developed
latterally and dorsally, but not ventrally (figs. 83, 77). The thickened part is very
thick, consisting of the usual narrow and almost filiform cells. The ventral part is

86 CALIFORNIA ACADEMY OF SCIENCES.

very thin, thinner in fact than any other part of the alimentary canal. Of the balance
of this canal there is no distinction between cesophagus and sacculated intestine. The
gut is every where sacculated, only increasing in thickness towards the genital somites,
where it is thickest. The alimentary canal throughout its length is lined by a
columnar, ciliated epithelium, outside of which is a very thick vascular layer, with
large blood lacunes, directly connected with the dorsal and ventral vessels, which are
closely attached to, or almost imbedded in the intestine. The latter as well as the
vessels are covered with chloragogie cells, which, especially in the region of the
dorsal vessel, are very large (figs. 85 to 91), the layer being thickest close to the
strands of mesenteric tissue connecting the intestine with body-wall.

The free eclomic lateral vessels in the eight anterior somites are similarly
surrounded by a thick mass of glandular cells, arranged around muscular strands, and
which are quite distinct from the chloragogic cells and more resemble real glands.
Their reaction to stains is entirely distinct from that of the chloragogic cells of the
main vessels and of the alimentary canal, staining very deeply with ammoniated
hematoxylon (fig. 84), and showing a coarser grainy secretion, while the real chlora-
gogic cells remain much more pellucid and contain much finer grains. Cells similar
to the former are also seen attached to the ccelomie covering of the prostate (fig.
107, ete.) They also greatly resemble the glandular cells, or multicellular glands
from the pharynx of Pontodrilus and other oligocheta, possessing pharyngeal glands.

The vascular layer of the intestine is very much developed, especially on the
ventral side, where it connects with the ventral vessel, through a thick band of mesen-
teric and connective tissue. This as well as the walls of the blood vessels were so
thickly studded with a protozoa (Hemagregarina) that the structure of the layer
could not even in a single instance be properly made out.

There are no pharyngeal glands, though a few glandular cells are seen scat-
tered about between the muscular strands connnecting the pharynx with the body-
wall. But these cells resemble more chloragogen cells than true pharyngeal glands.

The ¢estes are of no unusual structure. The anterior pair, in the specimens I
opened, are smaller than the posterior pair, which were always forked, while the ante-
rior pair was undivided.

The ovary in xi isalways sigmoid of irregular shape and present the peculiarity
that seldom more than one ovum is developed at a time, this one being situated not at
the periphery or at the free end of the ovary, but in the inner angle of the sinus. The
ovum is unusually small in size and readily detached from the gonad (figs. 78, 108).
It grows large after separation, and is found in numbers in the posterior somites.

The ovary is of large size reaching far back to the posterior septum. Its lower
end is not only attached to the septum and body-wall, but also to the narrow end of
the outer sperm funnel (or ciliated rosette) fig. 96. In one of the specimens sectioned
the ovaries either extended past the oviduct through somite xii, or there was a second
pair of ovaries in xii. Beddard has similarly remarked that the ovary in Sutroa is
attached to the cells of the spermiducal funnel.

PACIFIC COAST OLIGOCH MTA. 87

Spermatheca. As stated there is one pair situated in ix, the spermathecal pore
being posterior to the sete, and in line with the inner couples. The spermatheca
consists of two distinct parts, as is usual in this group, the lower muscular part, and
an upper rounded part consisting of an unicellular layer of dice-shaped cells. This
rounded chamber was sometimes situated in the same somite, ix, as the muscular
body, but frequently it projected backwards into somite x.

The structure of this muscular part is represented in figure 82. The inner epithe-
lial layer, consists of narrow columnar cells, with rounded nuclei and striated proto-
plasm. Exterior to this layer, there is a thin one of transverse or circular muscles,
outside of which again is an epithelial stratum one or two cells deep, with slightly
oblong nuclei, the cells themselyes being irregularly dice-shaped (fig. 82).

Prostate and spermducts. The most interesting, as well as the most complicated
structure of this species, is the prostate, of which I am now able to give a fuller ac-
count, which J believe will not leave any of the points of its structure in doubt.

I have already referred to the three main divisions of the organ, the proximal
one consisting of a long cylindrical tube, containing penis, atrium and _ prostate.
Second, a very narrow tube of almost the same structure as the prostate part of the
former, connecting with a long cylindrical chamber of somewhat modified structure,
into which the two spermducts open, quite near its junction with the narrow part or
bridge. If we, however, disregard the difference in size of these various parts and
only consider the structure, we find that the whole organ may also be divided in
three parts:

First, the proximal part, which is entirely confined to somite x. This part is
upright, so to say, does not run backwards; it is also somewhat bent, forming a right
angle with the balance of the organ. This part consists of penis, and a long tubular
part which, in want of a better name, I designate as atrium (fig. 100 air. and p.).
With somite xi commences a change of structure of this organ, common both to the
wide part and to the very narrow posterior part. I will refer to it as the prostate
proper, as it contains the thick layer of regular prostate cells so common in all higher
oligocheeta where this organ occurs; it is the “two layer”’—prostate of Beddard. The
third part or storage chamber is characterized by the absence of this layer.

I will now refer to each one of these three or four parts in succession and more
in detail, beginning with the penis and atrium.

Next to the body-wall, ending at the transverse muscular layer and from there
stretching inwards, is a reversible sae—a preputium
with very large, round, compressed nuclei and striated contents. This epithelium is
surrounded by a thin muscular layer (fig. 100, pre., 101).

Into this preputium opens a penial glans (fig. 100 p. g/s., 101, etc.), consisting of
two separate covers, posteriorly attached to a collar of larger rounded or pear-shaped
cells, at the base of which are seen a number of muscular plates. This collar is
folded on itself, one part of the fold connecting with outer and one part with the
inner cover of the glans. Through the median line of these parts runs a long, very

5

consisting of epithelial cells,

Memorrs, Vou. II, 4. February, 1895.

58 CALIFORNIA ACADEMY OF SCIENCES.

narrow excretory tube with thick walls. Nearest the male pore this tube is quite
straight, or only slightly folded, but at its distal end the windings are greatly packed
or irregularly coiled (fig. 99, tube).

The part posterior to the penial collar I have designated the atrium proper,
simply beeause it does not contain any inner layer of glandular cells, and because in its
distal end it connects with the true prostate. The excretory tube continues all through
this part and is, so to say, suspended in a more or less dense mass of fibrous tissue.
Nearer the collar this mass is reduced to a few strands only, but towards the distal end
it becomes quite dense, especially so nearest the tube, while towards the periphery it is
much less dense (fig. 92, f. ¢., fibrous tissue; /. sf., fibrous strands). Fig. 100 repre-
sents a cross-section taken near the distal end; the tube is seen coiled in the center,
while fibrous tissue connects it with the walls of the atrium.

The exterior layers of the atrium are constructed yery much as the same lay-
ers of the prostate proper and the storage chamber. There are three layers which
are common to the two different parts of the prostate. Interiorly a thin layer of cir-
cular museles somewhat variable in thickness in different parts, but generally only
three or four strands thick. Exterior to this layer runs a spirally wound layer of lon-
gitudinal muscles, arranged in band-like plates, and when seen in cross-section re-
sembling a row of fringes (figs. 100, 101, 102, 108, 106, 107, 7. m). Fig. 107 shows these
plates, highly magnified, to consist of rather rectangular muscular strands. Exterior
to this layer of /. m. muscles we find everywhere a broken row of prostate glands of
very minute size, appearing to penetrate in between the muscular plates, though on
account of the macerated condition of the tissues I could not follow their tubes. These
small glands, which barely project outside of the muscles, are found everywhere from
the distal end of the storage chamber to the penial collar, wherever this muscular
layer isfound. In places they are continuous, in others scattered about, seldom more
than one row thick (fig. 101, g/s). But as we reach the region of the narrow part of
the prostate and especially the part of the storage chamber where the spermducts
enter, we find another thicker layer of prostate glands similarly scattered over the
longitudinal muscular fringe (figs. 102, 103, 106, 107, pr. gi. s, small prostate glands;
pr. gl. l., large prostate glands). In the region where the spermducts enter the stor-
age chamber this layer of large prostate cells is almost continuous and considerably
thicker than the muscular layers combined (fig. 103).

The prostate proper contains besides these jayers of muscles and exterior
prostate glands two inner layers, which resemble and propably correspond to the two
cell-layers found in the prostate of the higher oligochta, viz.: one layer of lining
epithelium (fig. 101) and one layer of glandular prostate cells, with very large nuclei,
and separated one from the other by transparent spaces, through which possibly enter
projections of the exterior prostate glands. These two characteristic layers do not
extend to the narrowest part of the prostate. Hence the muscular layer is covered
by a single row of inner lining epithelium with large, slightly oblong nuclei (fig.
102), which in the very narrowest part are about 12 to 14 in the row.

The proximal end of the storage chamber is lined by a very thick epithelium

PACIFIC COAST OLIGOCH MTA, 8Y

with very oblong nuclei (fig. 105 ep.). This epithelium is thickened only at the
anterior side, nearest the narrow tube (fig. 104), but narrows quickly both superiorly
and posteriorly. The inner transverse muscular layer is similarly thickened in this
region (fig. 105, ¢. in.).

The storage chamber contains the same general layers as the narrow bridge.
The inner lining epithelium is narrow, with very compressed nuclei, with the flat side
lying against the muscular wall (fig. 107).

The nature of the prostate differs apparently much from that of the prostates of
some members of Lumbriculidee, especially Sutroa, as lately more minutely described
by Beddard, but more resembles in structure Lumbriculus. But I am more inclined
to compare its makeup with the prostate of Moniligaster. In Eelipidrilus the pros-
tate contains the characteristics of both Limicole and Tericole, if I may yet use the
expression. The inner two-celled layer of the prostate which is characterizing most
higher earth worms, possessing a prostate, is superposed by the layers, muscular and
glandular which characterize Tubifex, Moniligaster, ete.

Spermducts. IT am now able to describe the spermducts for the first time.
They enter the storage chamber of the prostate near to the bridge and close to-
gether, but. still entirely separate (fig. 95), and when the inner surface of the
chamber is viewed from above the entrance pores are seen as two small slightly ele-
vated papillae (fig. 95 sp.) From these the spermducts, which are of very minute
size, run forward parallel to the prostate, one on either side, except alongside of the
bridge where they run close together. The outer pair leave the prostate in xi, dip
down to the ovaries in which they are partially engaged and push their ciliated
rosettes through septum x/xi, the rosettes opening in x. The inner spermducts are
similarly engaged in the gonads in x, push through septum ix/x and open their
rosettes in ix, all very close to the body-wall.

The rosettes are only one cell thick, very thin and_ flat, with the posterior sur-
face attached to the septum, the anterior lip only being free. Only the inner deeper
surface of the rosettes is ciliated.

Sperm-sacs, one on each side and continuous from the beginning of the sperm-
atheca, reach as far back or further than the posterior end of the storage chamber of
the prostate. Generally the sperm-sacs reach two or three somites further back.
Each sac is separate from the other and consists of one continuous bag contracted
somewhat at the septa. It is not racemose and does not connect with the septa as in
so many of the higher forms, but greatly resembles those of Sutroa. The sperm-sac
only covers the prostate, but does not properly enclose it, as it does not extend to the
space between the prostate and the intestine. With the latter, however, it is con-
nected by two continuous walls of connective tissue, one on the dorsal and one on
the ventral side of the intestine. In the enclosure thus formed the prostate, as well as
the upper part of the spermatheca, lies free.

In the posterior parts of the sperm-saes are always seen very large sacs of yolk
granules. In one specimen I found a mass of these yolk granules surrounding the

YO CALIFORNIA ACADEMY OF SCIENCES.

spermatozoa in the prostate storage chamber (figs. 120 to 123). The yolk sacs are
frequently so large that they fill the larger part of the cwlomic cavity pressing
the intestine close up against the wall of body.

Nephridia. The most anterior nephrostome is found in iii in front of septum
iii/iv. The most anterior nephropore is found in iy in front of the inner couple of
sete. There are nephridia (pores) in iv, v, vi, vil and viii, while somites x, xl, xi
have no nephridia. The neck of the nephrostome contains a large glandular swell-
ing, to which is attached muscles connecting with the septum. This neck is perforated
by several narrow ductules, which occasionally branch. The duct appears to con-
sist of a single tube not covered by peritoneal glands. On account of the rather mac-
erated condition of the specimens I could not investigate the nepridial structure any
closer.

Some of the specimens collected by my friend were from a new locality and
somewhat larger in size. This locality is Three Spring Meadow, on the east
side of the North Fork of Kings River, opposite the natural bridge, at an altitude of
about 8,000 or 9,000 feet, and several thousand feet above the river bottom at that
point. They were there found under some old logs lying across the meadow, and over
which the water was flowing, the worms being attached to the surface of the decayed
wood. These specimens were about 4 larger than those from the springs at Alpine
Meadows on the South Fork of Kings River several thousand feet higher up, but I
find no distinct characteristics, though the spermatheca appeared more twisted.
Among the specimens were a few of Velmatodrilus Vejdovskyi, for which I am thus
able to note a new locality.

PACIFIC COAST OLIGOCH MTA.

PIAt eX.

9]

1B.

<)

CALIFORNIA ACADEMY OF SCIENCES.

PLATE XXX.
PHCNICODRILUS TASTE.

Semi-diagrammatic view of the body in a longitudinal vertical section, showing the position and proportion of
the various organs. pha. pharynx. sl. gl. salivary glands. sp.gl. septal glands. sp.s. sperm-sacs. cl. cli-
tellum. h. hearts and dorsal vessel. @. cesophagus. spth. spermatheca. divt. diverticulum of cesophagus.
t. testes. si. sacculated intestine. @Q. ovipore. ov. ovary. ¢ male pore.

View of the interior organs, the body-wall being laid open from above. Onthe right hand side the sperm-
sacs have been removed in order to show the underlying organs. The letters indicate the same as in figure
one. The shaded part of somites xiv to xviii indicate the clitellum proper. ac. accessory copulatory papillz
around the outer sets in somite xiv.

Longitudinal section through some of the anterior somites, from a section lateral to the cesophagus, illustrating

the connection between the septal and supra cesophagal glands and their ducts, which all open into the upper
wall of the pharynx. s. septum. bd. w. body-wall. s/.gl. salivary or supra pharyngeal glands. ss. gl. sep-
tal glands.

Spermatheca in outline.

The spermduct and male pore viewed from above or from the interior of the body. Both are covered by the
longitudinal muscles, as well as by numerous arciform muscles.

Mie Gy Acasa ll. Pare Jy

OBE, aon oclecent ae i; 7, j Wa

SSS =

3.

) Dieta rarinanriee Aca aul, RY,
‘9 HIENICODRILUS LASTE LLU ¢.
.

ee

Vi

PACIFIC COAST OLIGOCH ETA.

Ma SOLOS

94 CALIFORNIA ACADEMY OF SCIENCES.

PLATE XXXTI.
PH@NICODRILUS TASTE.

5. A transverse section of the body-wall in somite xvii, through the male pore showing the degenerated atrium
and the wider upper part where enters the spermduct. These latter have just been fused and enter the pore
as one single duct. cl. clitellar cells. Ay. hypodermal layer. cl. collar of epithelium of the atrial chamber.
ac. m. arciform muscles. (f. sp. fused spermduct just entering the atrial chamber. J. m. longitudinal mus-
cular layer of body-wall. é. m. transverse muscular layer of body-wall. 4 male pore.

6A. The section nearest anterior to the former, showing the spermducts just as they enter the atrial chamber.
The spermducts are not yet fused. Figures indicate the same as in the preceding figure. sp. spermducts just
before being fused together.

6B. Section nearest anterior to the former showing the two spermducts, ete.

7. The three somites xvi, xvii and xviii viewed from the interior, the alimentary canal and nuerve-cord having been
removed, as well as nephridia.

8. One of the sete magnified, the same relative proportion as that used in my paper on Ocnerodrilus.

9A. One of the nephrostomes seen in front views. When in proper position the face of the rosette stands
parallel to the septum. +. rosette proper. . neck with very large glandular cells. . d. narrow duct.
s. septum. f. fold of the main nephridial body.

9B. Nephrostome seen from the side, the position when the body is opened and spread out. Letters indicate the
same as in the last figure.

10. One of the muscles connecting septal glands with body-wall, in cross-section. s. septum. m. muscular
fascicle.

11. One of the posterior nephridia seen under a low magnifying power in order to show the peritoneal covering
and the general outline of the canals in the lower peritoneal lobe. a. /. anterior fold of canals. p./. posterior
fold of canals. spu. the partially free spur. s. septum. st. nephrostome. xp. nephropore. w.t. the
wide tube leading to the nephropore. br. bridge connecting the spur with one of the folds. wu. /. upper
lobe of peritoneal cells which does not contain any canals. c./. connecting lobe. /. l. lower lobe. The two
peritoneal lobes vary greatly in shape and hardly two are found exactly alike, though the |general form is the
same in all,

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Pyemconnius TASTE. HGS. «

PACIFIC COAST OLIGOCH UTA,

PLATE, Xooxtl.

96

15.

16A.

16B.

160.

16D.

161K.
16F.

CALIFORNIA ACADEMY OF SCIENCES,

PLATE XXXII.
PH@NICODRILUS TASTE.
One of the posterior nephridia more highly magnified, to show the course of the canals.

Pheenicodrilus taste, natural size, the worm having first been killed gradually with increasing strength of
alcohol. Largest specimen.

Section of the intestine at the point of connection with the diverticula, in order to show the arrangement
of the blood vessels and the subdivisions of the diverticula. This figure is taken at the junction and shows
the diverticulum as one chambered, and with a large blood vessel penetrating the diverticulum from the
outside. This blood vessel comes from the dorsal vessel. This and the following five figures are outline
drawings from slides, but the details of less importance are not filled in. 6/. small vessels in the vascular
layer of the alimentary canal. dv. diverticulum. Ol. v. large blood vessel penetrating the diverticulum
from the exterior. ep. epithelial lining of the intestine. m./. muscular layer of the intestine.

Section closely following, or anterior to the former, showing the diverticulum as one chambered and with a
division of the blood vessels into smaller ones. Letters indicate the same as in the former figure.

A section of one of the diverticula a little anterior to the former, the organ is yet one chambered, and a large
blood vessel is seen on the left side or the side nearest the intestine. J. bl. v. very large vessel. b/. v. small
vessels in the wall of the diverticulum.

A section anterior to the former, showing the beginning of the subdivision of the diverticulum.
A section next anterior to the former, showing the diverticulum two chambered.

A section near the center of the diverticulum, showing six interior chambered and a multitude of blood
vessels.

A part of diverticulum more highly magnitied, showing nuclei and striated cycloplasm.

Section of alimentary canal.

PACIFIC COAST OLIGOCH MTA.

PUATE | SOCkIII.

98

20.

28.

CALIFORNIA ACADEMY OF SCIENCES.

PLATE XXXII.
PH@NICODRILUS TASTE.

Longitudinal section of the alimentary canal (esophagus), in vi. ep. inner lining epithelium. g/. glandular
cells. 61. blood sinus. m. the two muscular layers. pt. peritoneal epithelium.

Longitudinal section of the pharynx, showing the glandular ducts and ductules penetrating the pharyngeal
epithelium. cl. ciliated surface. n. ep. nuclei of epithelium. dus. ductules. duct. large duct from the
pharyngeal glands above the cesophagus. . ep. casophageal epithelium joining the pharyngeal ciliated
epithelium.

Section of the body through one of the clitellar somites, a diagrammatic view, to show the relative size of the
different layers of the body-wall. d.v. dorsal vessel. s.i. saculated intestine. m. /. the muscular layers.
cl. c. clitellar cells. sp. d. spermducts. v.v and 7. c. ventral vessel and nerve cord.

A diagrammatic section of the body in somite ix showing the relative size, etc., of the diverticula and the
layers of the body-wall. d.v. dorsal vessel. sp. s. sperm-sacs in somites ix and x. div. diverticula of the
intestine. v.v. ventral vessel. mn. c. nerve cord.

Section of body-wall of a non-clitellar somite. g.c. unicellular glands. s. c. supporting cells. ¢. m. trans-
verse muscular layer. /. m. longitudinal muscular layer. pr. peritoneum.

KERRIA McDONALDI.

Nephridium of Kerria McDonaldi from one of the somites closely posterior to the clitellum. pr. s. peritoneal
sac with nuclei. The posterior end of this sac is not furnished with blood capillaries. c. b/. capillary blood
vessels, spreading principally through the peritoneal sac. a./. anterior lobe. p./. posterior lobe. nephr. st.
nephrostome. nephr. pr. nephropore. w.d. wide duct outlet. m. d. narrow dyet. nec. neck of the
nephridium. 67. bridge. spr. spur. s. septum.

PONTODRILUS MICHAELSENT.

A specimen, natural size, having been slowly killed and extended. One of the largest specimens, the majority
being only one-half as large.

The three anterior somites seen from the side. pr. prostomium. per. peristomium.

The anterior part of the worm more highly magnified, ventral view. cl. clitellum. ¢ male pore. spth. p.
spermathecal pores. ov. p. ovipore. c. pr. m. cushion where ends the fan-shaped fascicle of prostate or arci-
form muscles. c.c¢. exterior copulatory cushion.

Part of the body-wall iaid open and spread out to show the relative distances of the sete, the numerals
indicating the number of the seta. The ventral nerve cord is indicated.

One of the setee highly magnified.

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100 CALIFORNIA ACADEMY OF SCIENCES.

PLATE XXXIV.
PONTODRILUS MICHAELSENI.

29. A somewhat diagrammatic view of the anterior somites seen in longitudinal section. br. brain. mo. mouth.
ph.c. pharyngeal cavity. pha. pharynx. m. muscular strand connecting pharynx with the body-wall. s/. gl. sal-
ivary or pharyngeal glands opening by ducts in the pharynx, the ducts being supported by muscular strands
(stained yellow). ss. g/. smaller gland; connected with blood vessel. g/. glands connected with the lateral vessels.
@. esophagus. d.v. dorsalblood vessel. s.septa. spth. spermathecm, and their diverticula. The lower end
of the spermatheca is greatly enlarged, forming propulsory bursa. spth. p. spermathecal pore. ¢. testes.
ov. ovaries. ovd. oviduct. c. 7. ciliated rosettes. A. hearts. sp. s. sperm-sacs. These sacs are principally
yentralto the esophagus. nephr. nephridia, the most anterior one is foundin somite xiii. c/. clitellum. gl.cr.
glandular crop in shape resembling a gizzard. s. 7. sacculated intestine. pr. prostate. ¢ the muscular duct
of the prostate ent through. c.c. copulatory cushion. The dotted line indicates the outlines of the duct as
they appear in other sections.

30. Section through the posterior part of the pharyngeal glands, showing the arrangement of the glands, ducts,
muscles and blood vessels, all in vertical section. «@. cesophagus. ep. inner epithelial lining. m. @. cireular
cesophageal muscles. m. muscular strands. pha. gl. pharyngeal gland. d.v. dorsal vessel.

In this figure the glands are stained violet, the muscles appear as yellow and the duets are dark violet. The
blood vessels are black (Corosive sublimate, abs. alcohol, orange G. amm. hematoxylon. Thus in xylol.
This and the following five figures are all drawn from sections treated in the same way.)

31. A longitudinal section of a lobe, showing the general arrangement of the glands and ducts. duc. ducts from
the gland, leading anteriorly to the pharynx. gl. c. glandular cells. O/. gl. blood glands. m. supporting
muscular strands. b/. v. blood vessels.

32. A small glandular lobe more highly magnified. The letters indicate the same as in the preceding figure. The
nuclei are stained yellowish with orange G. The secretion of the cells is precipitated? and stained dark violet
with amm. hematoxylon.

33. <A part of a gland, with supporting muscles.

34. Two sections of muscular strands, showing the connection with the ducts in which the secretion has been
stained deep blue.

35. One of the glandular cells with nucleus stained yellow. ‘The vacuoles (va.) are white, the protoplasm grayish,
while the secretions are stained deep bluish-violet.

36A. The proximal end of two glandular ducts where they penetrate the epithelial lining of the pharynx, near its
dorsal side, showing the ductules or end ducts leading between the epithelial cells. 6/. blood vessels. m. ph.
supporting muscles for the ducts connecting with pharynx. ¢. m. transversal muscles cut across. gl. d.
glandular ducts leading from the gland at the distal end beyond the pharynx. These ducts are the largest
and collective ones, having received the secretion from many smaller ducts. ep. epithelial cells lining the
pharyngeal cavity. g/l. dt. glandular ductules passing between the former into the pharyngeal cavity. s. gl.
two of the smallest of the salivary glands, a few of which are found at the periphery of the pharynx.

36B, A longitudinal section of one of the small septal glands, one pair of which is found in y, vi, vii, viii, ix.
c. bl. v. connecting blood vessel between the ventral vessel c. bl. vi and the dorsal vessel. This vessel is
cut in two places, the one marked c. b/. vii being nearest to and just below the esophagus (@. v.) the ventral
outline of which only is figured diagrammatically. s. septum, outline of, between yi/vii. g/. small glands, a
few of which surround the somewhat larger septal gland. ss. gl. the largest septal gland. w. c. wandering or
perigastric corpuscles. d. ¢. discharge ducts from glands, probably connecting with the pharynx.

37. Cross-section of an immature specimen, through the intersegmental groove between somites xyii/xvili, just
anterior to the copulatory cushions. In this section no clitellar cells haye yet developed on the ventral side
of the body. hy. hypodermis. ¢. m. transverse muscular layer. /. m. longitudinal muscular layer. neph.
nephridium. s. a. septum between xyii/xviii. m. muscles connecting the glandular part of the sperma-
theca with the body-wall. g/. pr. glandular part of the prostate. spd. the sperm-duct just entering the
glandular prostate. m. pr. muscular part of the prostate. v. ne. ventral nerve cord. s. 7. sacculated intes-
tine.

38. Cross-section of the body of an immature specimen, through somite xviii, the section passing through the
copulatory cushions and through a part in which the clitellar cells on the ventral side of the body are already
developed. a. m. arciform muscles connecting the copulatory region with the dorsal part of the body-wall.
cl. c. clitellar cells developed in this somite only on the ventral side, neyer extending to the dorsal side even
in fully mature specimens. Other letters indicate the same as in the preceding figure.

39. Cross-section through somite xiii close to the anterior septum. The clitellar cells are here fully developed.
s. 8. part of septum between xii/xiii. e/. ¢. clitellar cells, only developed dorsally. v. ¢/. ventral termini of
the clitellar cells. ov. ovaries. spd, spermduct. ¢. i. tubular intestine. dv. dorsal vessel. fA. hearts.

7 Ie ey eo

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Parle =e _ A P c
PACIFIC COAST OLIGOCHATA,

102

40.

41.

42.

43.

44.

45.

CALIFORNIA ACADEMY OF SCIENCES.

PLATE XXXYV.
PONTODRILUS MICHAELSENI.

Cross-section through somite xvii of a mature specimen showing the development of the clitellar cells on the
dorsal and lateral sides of the clitellum. vv. cl. ventral termini of the clitellar cells. d. v. dorsal vessel.
s. i. sacculated intestine. neph. nephridium. gl. pr. glandular part of prostate. pr. m. muscular part of
prostate. m. muscles connecting the prostate with the body-wall. ¢. m. transverse muscles. 1. m. longi-
tudinal muscles. cl. clitellar cells. v. cl. ventral termini of the clitellar region.

Cross-section through a strongly contracted, fully matured specimen, through somite xviii, showing the copula-
tory cushions as fully developed, as wellas the clitellar cells on the ventral side. Unfortunately the only fully
developed specimen at my disposal had been badly contracted, hence the cells appear smaller than they would
have done in properly prepared specimens. c.c. copulatory cushions. a. m. arciform or fan-shaped muscu-
lar strands, which contract the copulatory region. i. ¢. m. interior transverse muscular strand, ending
respectively in the copulatory cushions and in the projecting ventral lobes of the body. Other letters as in
the last four figures.

The body-wall viewed from the inner side of somites xvii, xviii and xix, showing the genital region of the left
side. pr. m. prostate muscles connecting this organ with the body-wall. a. m. arciform muscles connecting
the copulatory cushion with the upper body-wall. c.c. copulatory cushion. sp. m. muscles connecting the
body-wall with the prostate at the junction with the spermduct. sp.d. spermduct. 1 and 2s. first and
second seta. v.c. ventral nervecord. gl. pr. glandular prostate. m. pr. muscular prostate. s. m. smaller
muscles, fan-like arranged, confined entirely to the copulatory cushion. sp. d. p. spermiducal pore overlaid
by muscles. s. septa.

A somewhat diagonal longitudinal section through the body-wall and copulatory cushion in somite xviii. The
section is parallel with the muscular fan of arciform muscles, passing through the copulatory cushion near
but not through the body-wall. Illustrating the arrangement of the muscular strands around the prostate in
the vicinity of the male pore. The letters indicate the same as in the preceding figure.

A longitudinal section through the copulatory cushion and the body-wall, the section being almost vertical.
pr. prostate, just leaving the arciform muscles, and immediately before bending down towards the prostate
pore. m. muscles connecting the prostate with the body-wall. «. m. the entrance of the arciform muscles
into the copulatory cushion.

A horizontal section of the body through somite xviii and the copulatory cushions, showing their size and
position, etc.

Pe Py N/ fre 9 f a
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ores

PACIFIC COAST OLIGOCHAHTA, 105

AY SOOOAe

104

46A.
465.
460.
47.

48.

CALIFORNIA ACADEMY OF SCIENCES.

PLATE XXXVI.
PONTODRILUS MICHAELSENT.
Section through the rudimentary gizzard or thickening of the intestinal wall in somite v.
One of the circular muscular strands of the former figure.
Section of the alimentary wall adjoining posteriorly to the rudimentary gizzard in somite vi.

A longitudinal section through the intestine showing the glandular-crop in somites xiv, xv, xvi, and the be-
ginning of the sacculated intestine in xvii. d.v. dorsal vessel. d.v.c. connection between the dorsal vessel
and the vascular system of the crop. gl. cr. glandular crop. /. s. blood sinus in the outer layer of the
glandular layer. gl. c. glandular cells constituting the bulk of the crop. clo. chloragogen cells. m. 1. the
two muscularlayers. ¢.7. the end of the tubular intestine or cesophagus.

A section of the former more highly magnified, showing the glandular cells which constitute the crop and the
branched lumens which are especially pronounced in the outer part of the layer. /.m., ¢. m. the two mus-
cular layers, which here are not more developed than in any other part of the intestine.

Cross-section through the glandular crop in the posterior part of the organ in somite xvi. In this section only
few of the glandular cells are found.

A section anterior to the former, but inthe same somite. Here many more of the glandular cells are seen. In
these two sections the chloragogen layer is thick and the cells crowded, similarly the ridges of the epithelial
cells ave high and separated by deep grooves.

A similar cross-section through the crop in somite xv. Both the chloragogen cells and the epithelial cells are
lower.

Another cross-section showing the entrance of a blood vessel from the dorsal vessel into the vascular system

of the glandular crop.

A part of the glandular cells of the crop more highly magnified showing the structure of the cells and the
branched lumens between them. g/l. c. glandular cells. /. lumen. 6/. blood sinus. ep. inner epithelial
cells.

An oblique section through the surface of the crop showing the two muscular layers, a blood sinus and a few
chloragogie cells cut through. The glandular cells immediately below are not delineated.

68

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PACIFIC COAST OLIGOCH MTA, 105

Pielke SOOCiE

106

Or
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oy
~1

CALIFORNIA ACADEMY OF SCIENCES.

PLATE XXXVII.
PONTODRILUS MICHAELSENT.

A spermatheca with its diverticulum seen from interior of the body, when the body-wall is spread out. spth. p.
spermathecal pore, d. diverticulum. spz. inner sac-like enlargement where the spermatozoa are massed.

A cross-section of the body-wall through the spermatheca near its pore. v. Jl. thin vascular layer or perito-
neum. /. m. longitudinal muscles. ¢. m. transverse muscles. hy. hypodermis. spz. spermatozoa. int. n.
interstitial nuclei. en. epithelial nuclei and cells. c. m. cireular muscles of the spermatheca. div. diver-
ticulum.

A cross-section of the diverticulum of the spermatheca. im. inner nuclei of the epithelium. inés. x. inter-
stitial nuclei. c.m. circular muscles. 5]. yascular layer. gl. c. exterior glandular layer consisting of a
double or triple row of cells.

A longitudinal section of the body-wall, somites vii, viii and ix made through the spermathecal pore. spih. p.
spermathecal pore. spth. the lower, enlarged part of the spermatheca, which forms a strongly muscular
papilla. dv. the junction with the diverticulum. sef. seta. 7. m. longitudinal muscles. ¢. m. transverse
muscles. a.s. anterior septum of somite viii. p.s. posterior septum of somite viii.

One of the testes.

A longitudinal section of one of the testes.
An ovary.

Longitudinal section of an ovary.

Longitudinal section of the oviducal funnel. s. septum.

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PACIFIC COAST OLIGOCH MTA. 107

PLATE XxXkV IIL

108 CALIFORNIA ACADEMY OF SCIENCES.

64.

69.

PLATE XXXVIII.
PONTODRILUS MICHAELSENI.

Section through the body in somite xi, showing the sperm-sacs and their relative position. s. anterior septum.
d.v. dorsal yessel. h. hearts. @. wsophagus. v.v. the branched ventral vessel. sp.s. the racemose
sperm-sacs situated much ventrally of the intestine. x. c. ventral nerve cord. J. m. and ¢. m. longitudinal
and transverse muscles of the body.

A cross-section of the same region more magnified. s. x/xi. septum between x/xi. s. xi/xii. septum between
xi/xil. sp.s. one of the sperm-sacs. spz. spermatozoa. m.c. mother cells. c. r. ciliated rosette. m. m.
muscles connecting the septum below the sperm-sac with the cesophagus.

A more longitudinal section of the ciliated rosette, showing the muscles and the blood vessels in the thick
funnel.

Another section through the ciliated rosette, showing the thick layer of blood vessels below the ciliated
epithelium.

A cross-section through the glandular part of the prostate, uear the entrance of the spermduct. ep. epithelial
cells with round nuclei. c. m. circular muscles. c. glandular cells. m. muscles connecting with the body-
wall. sp. d. spermduct.

Cross-section through the muscular part of the prostate.

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PACIFIC COAST OLIGOCHATA. 109

PEALE. XOOCx

110

70.

71A.

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to

73.

CALIFORNIA ACADEMY OF SCIENCES.

PLATE XXXIX.
PONTODRILUS MICHAELSENT.

Cross-section of the body in somite xi, showing the relative thickness of the septa and body-wall, the septa
being very much cupped, showing the cesophagus, etc., of the somite ix, as well as the septum and ciliated
rosettes of x. v.v.x. the ventral vessel in x where it connects with the hearts. v.v.ix. the two forms of
the ventral vessel. gl. septal gland surrounding the connecting vessels in somite ix. There are five pair of
these glands, one each in vy, vi, vii, viii, ix.

One of the nephridia isolated. p./. posterior lobe of the peritoneal sac of the nephridium. a./. anterior
lobe of the same. w. windings of the canals where the two folds meet. p./. and a. /. posterior and anterior
folds of the canal. xn. p. nephropore. x. st. nephrostome. spr. spur with four canals. br. bridge.
bl. v. blood vessels on the nephridium.

Nephrostome isolated. m. marginal cells. w.d. wide duct. nec. neck in which the narrow duct is seen to
be branched.

An enlarged and somewhat diagrammatic drawing of the course of the ducts, etc., of the nephridium. The
letters indicate the same as in the preceding figure. The canals have been represented as further apart
than they are in reality, otherwise their course could not have been clearly delineated. When the nephridium
is viewed from above mounted in glycerine, the outlines of the canals are only dimly discernible, being greatly
obscured by peritoneal cells and blood vessels, neither of which have been delineated. The shape of the
neck varies tosome extent, in some specimens being much wider than in others.

Various forms and sizes of blood-glands from the supra pharyngeal and septal salivary glands. Drawn from
paratine sections, hardened in Formaline, and stained: a. orange G. alcohol, Ehrlich’s hematoxylon amm.
4. rose aniline in hydrochloric alcohol, Bismark brown, and Ehrlich’s hematoxylon. The latter combination
gives by far the finest results, clearly differentiating the blood from the gland-secretion, this being very
imperfectly done by the orange G.

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CALIFORNIA ACADEMY OF SCIENCES.

PEALE xe: "
PONTODRILUS MICHAELSENT.

Cross-section through the body-wall and through the nephropore and wide duct. The latter is seen to enter
the glandular neck of the nephridium (nec), its inner duct having been cut twice, these cross-sections
being always of unequal width. The section a passes through the distal windings of the ducts, the letters
in the duct indicating that they belong to the same duct. c. c. ciliated canal, always wider than the other.
The canal c. is a branch or forking of one of the other ducts. At d. about an inch of the drawing 1s left out
in order to economize room.

Longitudinal section of the nephridium near the windings showing the forkings of one of the canals. c. ¢. is
the ciliated larger canal. 6. is the body of the nephridium, in which are seen the branched canals and lacunes,
as well as the branching blood vessels which are held back.

Part of the neck in cross-section, a larger magnification of a section similar to a 74, showing the branching of
the narrow duct x. d. The wide duct w. d. ¢. is cut twice; the part marked w. d. is the part of the wide duct
as it leaves the neck for the exterior pore of the body-wall.

ECLIPIDRILUS FRIGIDUS.

A diagrammatic view of the anterior somites of the body, composed from several longitudinal sections. Only
part of the vascular system is shown, and none of the nephridia, the principal object being to
show the arrangement of the reproductive organs and the shape of the alimentary canal. Sperm-sacs not
shown, br. brain. bd. body-wall. pha. pharynx. c.v. connecting vessels. d.v. dorsal vessel. v. v.
ventral vessel. cl. clitellum. v.c. ventral nerve cord. ¢. first and second pair of testes. spth. spermatheca.

ov. ovary. ovd. oviduct. c. 7. inner and outer ciliated rosette. cl. clitellar layer of glands. sps.
spermatozoa. 1. testes. chlo. chloragoyven cells. 4 male pore. jp. penis. atr. & pr. prostate and

atrium. pr. m. muscular layer of prostate. pr. g. glandular part of prostate. ar. atrium. sir. storage
chamber for protozoa. spd. spermduct. int. intestine. br. bridge.

PACIFIC COAST OLIGOCH ATA,

Ricks, cle

114 CALIFORNIA ACADEMY OF SCIENCES.

PLATE XLI.
ECLIPIDRILUS FRIGIDUS.

78. A diagrammatic figure of the region of the reproductive organs, including somites ix to xvii, showing the arrange-
ments of the reproductive organs. The body-wall is represented as being cut open dersally and spread out.
br, the bridge or narrow part of the prostate connecting the wide and lower part with the storage chamber of
the spermatozoa, str. storage chamber of spermatozoa. The prostate and ciliated rosettes are somewhat
stretched, as otherwise they could not be seen both at the same time.

79. Part of the clitellum in transyerse section, showing the clitellar glands, the longitudinal and transverse muscu-
lar layers.

80A. One of the blood glands or herzkérper.
80B. A single gland cell of the former.
81. A transverse section of the dorsal vessel, showing the blood gland occupying the yentral part of the vessel.

82A. Cross-section of the lower part of the spermatheca, at the spermathecal pore. bd. body-wall. ep. inner lining
epithelium. m. muscular transverse layer. ex. exterior layer. /. m. longitudinal muscular layer of the
body-wall. ¢. m. transverse muscular layer of the body-wall.

82B. Cross-section of the distal end of the spermatheca.

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ECLIPIDRILUS FRIGIDUS -FiG5.. 75.10

PACIFIC COAST OLIGOCHMTA,

PLATE XTi

116 CALIFORNIA ACADEMY OF SCIENCES.

PLATE XLII.

ECLIPIDRILUS FRIGIDUS.

$3-91. Represent somewhat diagrammatic views of transverse section of the body, respectively through somites 111,

90.

93.

94.

96.

iy, ix, x, X, xi, xi, xiv andxy. Of somites x and xi sections from the anterior and posterior parts are shown.
The following letters are common and indicate the same in each one of the figures: v. v. ventral vessel.
d.v. dorsal vessel. int. intestine. /. m. longitudinal muscular layer. 7%. m. transverse muscular layer.
v.c. ventral nerye cord. m. muscles and connective tissue uniting the intestine and body-wall. sp. s.
sperm-sacs. c. v. connecting vessels, cut in various places. pha. pharynx, shown to be developed only on
the upper side of the alimentary canal.

The intestine is here seen only to be connected with two superior muscular fascicles with the body-wall. The
connecting vessels between the ventral and dorsal vessels are covered with glandular cells, the contents of
which stains deeply with hematoxylon, and are of a different nature from the chloragogic cells.

Section through the first clitellar somite ix. cl. clitellar layer. spth. spermatheca. spz. spermatoza.

Section through somite x. In this specimen both the spermatheca project into somite x. The blood vessels
which are seen in the sperm-sacs and surrounding the spermatheca are the anterior part of the anterior pair
of lateral vessels, which project no further than here.

Section through the posterior part of x cutting through the atrium and penis. The ciliated rosettes were not
seen in this section, only some parts of the spermduct. The blood vessels seen surrounding the penis and
atrium are the anterior parts of the posterior pair of lateral longitudinal vessels.

Section through somite xi, close behind the anterior septum, showing the middle part of the outer ciliated
rosette cut through. The other rosettes having been torn were not seen on the section. The spermducts
are seen as a small circle, one on either side of the prostate.

Section through the posterior part of somite xi. The outer spermducts are seen as situated further down,
approaching each other, the approach being yet closer in the next section.

Section through somite xiv. The prostate being bent zigzag is seen as cut in three different places. At athe
part anterior to the very narrow part or bridge is seen cut. At b this very narrow part and at c the posterior
part or storage chamber where the spermducts are seen entering the chamber.

Section through somite xy. The storage chamber is cut close through its anterior end and is seen surrounded
by a thick layer of prostate glands. ‘

Cross-section of the atrium below its junction with the prostate. g/. small prostate glands penetrating the outer
muscular layer. m.s. muscular layer. ‘tube tube winding in the inner layer of fibrous tissue. mn. nuclei
of lining layer, boundaries of the cells not distinct probably on account of maceration. A few nuclei only are
seen in the fibrous mass, which is much thicker in the center surrounding the tube.

Cross-section of the prostate proper in somite xii. g/. s. small surface glands penetrating between the longi-
tudinal muscular layer. m.s. longitudinal muscular layer composed of fascicles or plates of strands. ¢. m.
transverse muscular layer. gl. c. thick glandular layer of one row of cells. /. ep. lining ep. of cells. spz.
spermatozoa.

Cross-section of the narrow part or bridge of the prostate. The layers are the same as in the preceding figure,
but of different relative thicknesses. Letters as in preceding. sp. sperm ducts. out. m. outer layer of
muscles, running in different directions.

Cross-section of the storage chamber in somite xv, posterior to the entrance of the spermducts, but very close
to it. sp. d. spermducts. gl. s./. large surface or prostate glands. gl. s.s. small surface or prostate glands.
m.s. longitudinal layer of muscles. ¢. m. transversemuscles. cep. lining epithelium with large nuclei.

A somewhat oblique section of the posterior part of the bridge, near its junction with the storage chamber,
but anterior to the entrance of the spermducts, showing the relative size of the layers and the increased
epithelium in the anterior part. br. posterior part of bridge, near its widening. ep. lining epithelium. pr.
gl. prostate glands. Jl. m., 4. m. longitudinal and transyerse muscular layers, both cut obliquely. Both the
transyerse muscles and the epithelium are thickened towards one end.

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101.

CALIFORNIA ACADEMY OF SCIENCES.

PLATE Xb Ete

ECLIPIDRILUS FRIGIDUS.
Cross-section of the neck of a nephridium.
Section of the long duct.
Nephrostome and part of duct.
Section of the lower part of the male organ, slightly oblique to the longitudinal axis of the penis, but parallel
to the atrium. air. atrium. (/f.¢. fibrous tissue. in. inmer lining of the atrium with long oval nuclei.
1. ms. longitudinal muscular layer of atrium. ¢. ms. transverse muscular layer. gl. small glands scattered
over the surface of atrium and prostate. spd.spermduct. sep. part of the septum between x/xi. sph2t. cir-
cular muscles or sphinxter. g/l. c. large glandular cells composing the collar. m. p. muscular plates, to which
the longitudinal muscles of the atrium are joined. tube. penis tube. ps. penis sheath. pre. preputium.
bd. body-wall. i

Section of inner penis tube of the former figure.

The lower end of the male organ, atrium and prostate in somite x, xi, somewhat diagrammatically figured, in
order to show the different layers, penis, ete. pr. prostate proper, projecting somewhat into x. s. septum.
1. pr. terminus of prostate. pr. m. muscular layer surrounding the prostate. This layer contains numer-
ous glands. pr. gl. glandular layer of long flask-like cells. ep. inner lining epithelium. tube tube running
from prostate through penis and atrium—a penis lumen, with thick walls connected with the atrial wall by
numerous strands, which on the upper part form a thick layer surrounding the tube, but lower down becomes
thin, at least consisting of only few strands. col. collar surrounding posterior part of penis. p. penis,
projective part. pre. preputium, eversible. bd. body-wall. /f. ¢. fibrous tissue.

A somewhat larger view of the lower end of the penis. Both this and the former drawing are made from
dissections mounted in gum-thus and yiewed from above. Letters indicate the same as in fig. 92.

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102.

1038.

104.
105.

106.

CALIFORNIA ACADEMY OF SCIENCES.

PEATE 2S ry:
ECLIPIDRILUS FRIGIDUS.

The central part of the dissected prostate and male organ, as seen when mounted in gum-thus. This figure
represents the narrow part of the prostate or tube, connecting the lower or glandular part with the upper or
storage part. The two spermducts are marked much more strongly than they appear and are more spread
or separated. The drawing represents views at two different foci. Zeiss C. Eyep. 3, t. 165. a. the posterior
part or storage chamber in which the spermducts open at j. sp. The anterior end of this chamber is lined
by a very thick epithelium, which however rapidly diminishes in thickness backwards. 6. the very narrow
part connecting the posterior storage chamber with the anterior prostate. c. the anterior prostate. out.
outer layers of the storage chamber and prostate, surface views. Ju. lumen of storage chamber. dum. lumen
of prostate. ep. very thick epithelium at the anterior end of the storage chamber. j. sp. junction of sperm-
ducts and storage chamber. /.v. lateral longitudinal blood vessels. se. septum between xiv/xv. spt. sep-
tum between xv/xvi. spd. spermducts. 67. narrow part of prostate or bridge. g/l. glandular layer of
prostate. ept. lining-epithelium of prostate,

The anterior end of the storage chamber, dissected, mounted in gum-thus. The focus is set on the inner
concave surface, showing the junction of spermducts and bridge with the storage chamber proper. sp. the
pores where enter the two spermducts. outer. coats of muscles and small prostate glands of the storage
chamber. br. bridge or narrow part of the prostate.

Longitudinal section of the body-wall, showing posterior ciliated rosette, etc., behind the male pore.

The inner end of the former more highly magnified. The letters indicate the same as in the last figure.
Only some of the larger prostate glands are figured. Zeiss C. Eyep. 3.

A part of the former more highly magnified. Zeiss Hom. Im. 1-12, Eyep. 3-155. The letters indicate the same
as in tig. 104. This section is more parallel with the longitudinal axis of the organ. The cell walls of the
epithelial cells were not distinct, probably on account of maceration.

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22 CALIFORNIA ACADEMY OF SCIENCES.

PLATE XLV.
ECLIPIDRILUS FRIGIDUS.

107. A transverse section, the storage chamber more highly magnified. Zeiss Hom. Im. 1-12 Eyp. 3. 155. pr. gl.1.
large prostate glands. pr. gl.s. small prostate glands. /. m. longitudinal muscles, arranged in strands.
t. m. transyerse muscles. n. nuclei of inner lining epithelium.

108A. A dissected ovary.
108B. <A ripe ovum.
109A. A dissected oviduct.

109B. Longitudinal section of the body-wall, at the ovipore, showing the latter to be situated in the intersegmental
grooye.

110 to 124. Various and successive stages of development of spermatozoa.
110. A resting spermatogonium (spermatospore) from the testes.

111. A spermatogonium from the outer edge of the testes ready to fall offinto the sperm-sac. The chromosomes of
the nucleus have begun to develop into winding rods.

112. Nucleus from a spermatogonium from the sperm-sac.

113. Spermatogemme (spermatosphere or spermpolyblast). The spermatocytes (spermatoblasts) are surrounding
and attached to a central, non-nucleated cytophore (or spermblastophore).

' 114. The same shortly before division of the spermatocytes.

117. A spermatogemme in the next last stage of development. The spermatocytes have through division reached
their final number. The cell divisions are indistinct. The nuclei are globular with scattered chromosomes.

118. A further developed or next last stage of thespermatogemme. The nuclei have assumed an ovoid shape, stain-

ing very dark with saffranin.

119. A part of a spermatogemme, last stage. The nuclei have again diminished in size and become perfectly round,
previous to growing out into spermatozoa.

120. Part of spermatogemme in which the nuclei of 119 have begun to grow out into spermatozoa.

121. A fully developed spermatogemme with grown spermatozoa attached to the cytophore.

22. The same in a state of dissolution, the spermatozoa detaching themselves.

123. Spermatozoa fully developed, the nuclear end being only slightly thicker than the other part of the body.

124-125. Sections of fully developed spermatogemmes, showing spermatozoa in cross-section, they having gradually
diminished in diameter since the nuclei were first concentrated (fig. 119), but having proportionately grown
in length.

126. Nuclei from 113, 117, 118, show their relative size.
127-129. Yolk sac with yolk cells.

127. The whole sac in section. Zeiss C.

128. Yolk cells without granulation.

129. Further developed yolk cells with granulation.

KERRIA McDONALDI.

130A. Longitudinal section of the alimentary canal of Kerria Mc Donaldi in the somite next posterior to the male
aperture, showing unicellular glands alternating with ciliated epithelial cells. g/. unicellular glands.

130B. A more highly magnified part of the former, showing one of the unicellular glands, surrounded by common
epithelial cells, also rudimentary glands. ;.gl. rudimentary glands. gil. large gland. jp. its pore. ep.
epithelial cells. bl. v. blood-lacune. /. m. longitudinal muscles. ¢. m. transverse muscles.

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PACIFIC COAST OLIGOCHETA.
IM,

BENHAMIA, ACANTHODRILUS, ALEODRILUS, SPARGANOPHILUS, DELTANTA,

PHG@NICODRILUS.
BY GUSTAV EISEN, PH. D.

The following genera and species are treated of in this paper:
Benhamia nana n. sp.
Benhamia mexicana Rosa.
Benhamia Bolavi Michaelsen.
Benhamia palmicola n. subsp.
Benhamia papillata n. sp.
Benhamia rugosa n. Sp.
Benhamia octonephra Rosa.
Benhamia Godeffroyi Michaelsen.
Benhamia malayana Horst.
Benhamia floresiana Horst.
Benhamia Annz Horst.
Acanthodrilus tamajusi n. sp.
Acanthodrilus Vasliti n. sp.
Aleodrilus Keyesi n. sp.
Sparganophilus Benhami n. sp.
Sparganophilus Smithi n. sp.
Sparganophilus sonomz pn. subsp.
Sparganophilus Eiseni Smith.
Sparganophilus guatemalensis n. subsp.
Sparganophilus carneus n. sp.
Sparganophilus tamesis Benham.
Deltania Troyeri var. crassa n. var.
Deltania Troyeri var. lagunz n. var.
Pheenicodrilus taste Eisen,

Pheenicodrilus tepicensis n. sp.
Memoirs, Vou. II, 5. December I4, 1895.

124 CALIFORNIA ACADEMY OF SCIENCES.

BENHAMIA Michaelsen.

The genus Benhamia was for some time considered a typical African genus,
and when later on a few extra-African species were found it was supposed that these
were recent emigrants, which had become more or less cosmopolitan. Of American
species two are known from Mexico and Venezuela, viz.: 6. Bolavi and B. mex-
icana, and one, 6. octonephra Rosa, has been described from Paraguay since this
paper was presented for publication. This latter species appears very nearly related
to L. rugosa, described below, but differs through the absence of penial sete.

The discovery of species of Benhamia at Miraflores, in the Cape Region of
Baja California, in a locality to which plants of any kind have rarely if ever been
introduced directly from foreign countries, would indicate that this genus has possessed
representatives on American soil for ages past, and that we really must consider these
American species as truly endemic. A final answer to this question of habitat must
be deferred to a future time, when more researches will have been made as regards
the distribution of these and other Benhamia species, as it is probable many more will
be found on this continent. A difficulty, which besets us from the beginning, is, that
so few species have been properly delineated, all descriptions having been made chiefly
with a view to distinguish the species from others already known, while with proper
delineations of the various organs, we would in all probability be able to make a satis-
factory comparison between species new and old. I haye received much aid from
Dr. W. Michaelsen, of Hamburg, who has described more Benhamia species than any
other investigator, and who has written extensively upon this genus. He has kindly
placed at my disposal several species of African Benhamia, as well as of B. Bolavi, for
comparison with forms found by me. This has enabled me to point out several im-
portant differences between 6. Bolavi and B. palmicola, which are sufficient to dis-
tinguish these as subspecies from each other, as well as from others previously known.
At the end of the descriptions of the various new species I append a table of charac-
teristics, etc., between species which may be confounded with our present ones, either
on account of similarity of some characters or because of their geographical distribu-
tion in the Malay archipelago, or in America.

As we now understand the distribution of this genus, the species are divided as
follows: America, 7 species; Malay archipelago, 5 species; Africa, 25 species; West
Indies, 2 species.

It may, however, be remarked that 6. rugosa described below is of uncertain
habitat having only been found in a hot-house, to which it had been imported from
unknown country.

Considering our yarious new species of Benhamia this genus may be charac-
terized as below:

Benhamia Michaelsen.

Acanthodrilid oligocheta. Setze strictly paired, ventral and lateral. Cli-
tellum generally incomplete, but in some species complete in some somites. Two
gizzards in succeeding somites. Calciferous glands generally three pairs, but some-
times only two pairs, very distinctly set off from the tubular intestine. Nephridia in

PACIFIC COAST OLIGOCH MTA. 125

respective species either diffuse or micronephridia arranged in three, four or more
rows on either side, some species showing several gradations. Spermatheca two pairs,
contracted at center with one or more small diverticula, Penial sete nearly always
present. Generally small, tropical worms.

-

DEFINITIONS OF AMERICAN SPECIES OF BENHAMIA.

Benhamia nana n. sp.

Derinition. Size
pigmented. First dorsal pore between iii/iv. Prostomium very swollen and overlapping.

Length 20 to 30 mm.; number of somites 110; skin strongly

Clitellum complete in central somites. Penial setie, the largest with 12 or more bristles.
Common sete, distance between the dorsal couples about equal to that between the ventral
couples. Oviducts, two pores, open in front of 1 and 2. Prostate pores not on papille.
Pharynx situated very far forward. Gizzards in viii and ix. Caleiferous diverticula,
two pairs in xv and xvi. Sperm-sacs in x and xi. Spermducts thicker at the base in
voi, wou and «xvi. Nephridia in three rows on either side, the posterior with calomic
mantle. Spermatheca, basal part with one diverticulum, apical part warty and irreg-
ular. Typhlosole large, begins in xviii. Color brownish red. Habitat, San Blas,
Meico, at sea level.
Benhamia mexicana Rosa.

Derrrinition. Length 30 mm.; number of somites 120. Skin not pigmented.
First dorsal pore between iii/iv. Prostomium divides the first somite completely. Cli-
tellum complete in xiii to wxi. Penial sete not ornamented (?); common sete, 3 and 4
further apart than Land 2. Oviducts, two separate pores in front of 1and 2, on small
papille. Gizzards in vii and ix. Calciferous gland 3 pairs in xv, avi and xvii. Ne-
phridia in 3 rows on either side. Spermatheca with a single short diverticulum. Alco-
holic specimen colorless. Habitat, Durango, Mexico, at 2500 meters altitude.

Benhamia Bolavi Michaelsen.

Derinition. Length 40 to GO mm.; number of somites 97. First dorsal pore
between v/a. Clitellum incomplete in all somites, xiii to xx. Penial sete, the largest
with five to eight notches, the smaller spoonlike and slightly forked; common sete couples
equidistant. Oviducts open in one single pore on a median papilla in wiv. Gizzard two
mv. Caleiferous gland three pairs in xv, xvi and xvii. Sperm-sacs one pair in wi.
Nephridia in three rows on either side. Spermatheca, the basal part with a single small
diverticulum. Saculated intestine begins in xxi. Color dingy flesh. Habitat, Venezuela
(and Hamburg).

Benhamia palmicola n. subsp.

Derinition. Length 50 to 60 mm.; number of somites 90. Skin not pigmented.
First dorsal pore between iv/v. Olitellum incomplete in all somites. Penial sete, largest
with four notches, the smaller one spoonlike and forked; common sete about equal dis-
tance between the dorsal and ventral as between the ventral couples. Oviducts open in one
single pore on a median papilla. Gizzards both in viii. Caleiferous gland 3 pairs in

126 CALIFORNIA ACADEMY OF SCIENCES.

xv, aviand xvii. Sperm-sacs in xiand xii. Spermducts of even width. Nephidia
in 3 distinct rows on either side. Spermatheca basal division globular, equal in size and
shape to the apical part, single small diverticulum. A typhlosole. Color reddish flesh.
Habitat, Baja California, Cape Region, 1000 feet; Tepic, Mexico, 4000 feet.

Benhamia papillata n. sp.

Derrnition. Size 50 to 70 mm. by 2 mm. Number of somites 125. First
dorsal pore v/vi. Prostomium divides somite I more than one-half, and is much swollen.
Clitellum complete, viii-wx. No ventral sete in xviii, but sete 3 and 4 present. Prostate
pores on large papilla, the four papille being close together in the sunk clitellar pit.
Oviducal pores on a large median papilla, but the pores are separate and on a line drawn
hetween the sete. Penial sete, largest slightly hooked with 8 indistinct notches, smallest
with a hair-like sigmoid tip. Spermathece, the apical-sac not globular, basal part
with a swell diverticulum. Sacculated intestine commences in xix. Typhlosole in xx-xiv.
Oaleiferous diverticula 3 pairs in avi, xvii, voi; the anterior one is much smaller.
Sperm-sacs in x and wi. Nepridia a consists of three distinct lobes, each one with a
calomic mantle. Color pale flesh, no pigment. Habitat, Tepic, Mexico, 4000 feet.

Benhamia octonephra Rosa.

Derrition. Compiled after Rosa. Length 20 to 40 mm.; number of somites
85 to 95. First dorsal pore between » and vi. Clitellum incomplete, in aiii to xx. Pe-
nial sete, the larger with six groups of two blunt tubercles; the smaller seta like a scalpel.
Common sete, couples equidistant. Oviducts open into a single median pore on a slight
swelling. Calciferous diverticula three pairs in xv, xviand xvii. Sperm-sacs in xi and
vit. Nephridia micronephric in four rows on either side. Spermatheca, basal part
pear-haped with a globular diverticulum. Habitat, Paraguay.

Benhamia rugosa n. sp.

Derrnition. Length 30 mm.; number of somites 118. Skin slightly pigmented
and corrugated. Clitellum incomplete, in wiii to wx. Penial sete, largest with five
notches; smaller seta forked with prongs of equal size. Common sete pointed and ven-
tral. Oviducts open jointly in a median pore. Gizzard in vii and viii. Calciferous di-
verticula in xv, xvi and xvii. Nephridia micronephric in four rows on either side.
Typhlosole present. Spermatheca much flatiened. Color reddish flesh. Habitat, Cal-
ifornia, in hothouse (imported).

Benhamia Godeffroyi Michaelsen.

Derinition. Length 90 mm. by 4 mm.; number af somites 174. Skin pig-
mented anteriorly. Clitellum incomplete (xiii) xiv to wx. Penral sete slender with nu-
merous irregular notches; muny set in each sac. The curved fossa between the pros-
tate pores with the convex side toward the median line. The anterior prostate largest.
Caleiferous diverticula three pairs. Nephridia plectonephric. Spermatheca without diver-
ticulum, but with warty protuberance. Color, anterior pale reddish, posterior part gray.
Habitat, Hayti (?).

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To face page 126.

Table A.

COMPARATIVE TABLE OF ALL KNOWN EXTRA-AFRICAN SPECIES OF THE GENUS BENHAMIA. ~

Habitat.

Number somites .......--

First dorsal pore.....----

Prostomium

Clitellum

Penial set@.........- +++

Common sete

Oviducal pore. ...-.---++

Prostate pores

Pharynx

Gizzards

Calciferous diverticula . ..

IS BPI) -BOC8 se olntnniseietatalais

Spermducts

Wephridiq: «cle seule

Spermatheca

nana 0. sp.
San Blas, Territory of Te-

Strictly tropical form.

ping.

Complete in the central
somites, xili-xx.

The largest with 12 or
more strong bristles.

Distance between the dor-

to that between the yen-
tral couples.

Two pores, open in front
of set 1 and 2,

Not on papille.

1 Situated yery far forward.

In yiii and ix,
Two pairs in xv and xvi.

In x and xi.

Thicker in xviii, xvii and
xvi.

In three rows on each side
of the median line. The
posterior ones with cwlo-
mic cell- mantle. The
nephridia in each somite
of about the same size.
The nephridia on each
side about equidistant,
but the dorsal nephridia
are much further apart
from the dorsal median
line than from the lateral
nephridia. Nephridia a
open in front of 3 and 4.

Basal part with one plain
diverticulum. Apical
part warty and irregular.

20 to 30 mm.

|
110. Skin strongly pig- |
mented in anterior so-
mites.
ill-iy.

Very swollen and overlap- |

sal couples about equal |

pic, Mexico, at sea level. |

mexicana Rosa.

| Durango, Mexico, 2565
metres above the sea level.
Temperate form.

| 30 mm.

120. Skin not pigmented.

iii-iv.

Divides the buccal somite
completely (7)

Complete, xiii-xxi.

Not ornamented (Rosa).
According to Ude, faintly
ornamented with 6 thick-
enings.

Setm 3 and 4 further apart
than 1 and 2.

Two pores in front of
sete» 1 and 2 on small
papilla, which touch with
their bases.

Not on papille.

In viii and ix.

Three pairs in xy, xviand
xvii.

In x and xi.

Not enlarged ( ?)

In three rows on either
side, probably all of the
same size. Nephridia a
correspond to sete 1 and
2. The nephridia on each
side are equidistant, and
the dorsal nephridia c ave
as equidistant from the
dorsal median line as
from thelateral nephridia
b. With eelomic mantle.

The basal part wide, grad-
ually clanging into the
apical part. Single short
diverticulum. :

Bolavi Michaelsen.
Hamburg, at Bergedorf,
imported, and Venezuela
(Ude).
40 to 60 mm.

97.

y-vi.

Does not divide buccal
segment.

Incomplete in all
meuts, xili-xx.

seg-

The larger seta with five
(or eight B) notches on
apex; the smaller is spoon-
like and slightly forked,
with no denticulation.

Those on posterior seg-
ments larger. The couples
equidistant.

One single pore on a large
median papilla in xiv.

Never elevated.

Not forward.

Two in somite vii.

Three pairs in xy, xviand
xvil.

One pair only in xi,

In three rows on either
side; the anterior ones
have no ceelomic mantle
like those posterior to
the male pore. The two
lateral nephridia of equal
size; the ventral nephrid-
ium is much narrower,
but furnished with two
separate ccelomic man-
tles, the inner of which is
smallest. The nephridia
are more separated than
in B. palmicola.

Basal part with a single
small diverticulum.
Apical division not glob-
ular.

palmicola n. subsp.

Baja California, Cape Re-
gion and Tepic, Mexico,
1000 and 4000 feet.

50 mm.

90. Not pigmented.

iy-v.

Not very pointed.

Incomplete in all somites,
xili-xx.

The larger seta with four
notches, the smaller one
spoon-like at apex and
forked.

About equal distance be-
tween the dorsal and yen-
tral as between the yen-
tral couples.

One pore on a central
media papilla.

Not on papille.

Not forward.

Two; both in viii.

Three pairs in xv, xvi and
xvii.

In xi and xii.

Not thicker in any so-
mites.

Arranged in three distinct
rows on either side of the
median line. Nephrid-
ium a@ consists of two
unequal parts, the most
yentral of which is the
smallest. Nephridium a
is much larger than the
other nephridia. The

dorsal nephridia far apart.

Nephridia a open in front
of 1 and 2.

Basal and apical divisions
globular, equal in size.
Single small divertien-
lum. 4

papillata 0. sp.
Tepic, Mexico, 4000 feet.

50 to 70 mm.

125 (largest specimen).

v-yi.

Divides partly buecal
segment; swollen.

Complete, but ventrally
thin, xiii-xx.

Larger with eight faint
notches; smaller with
a hair-like sigmoid
point.

Inner couples wanting
in xviii, equidistant.

Two clasely approxi-
mated pores on a me-
dian papilla.

Each on a large papilla.

Not forward.

Two.

Three pairs in xy, xvi
and xvii.

In x and xi.

In three rows, the pos-
terior ones with cwlo-
mic mantle. Nephrid-
ium a consists of three
lobes, each with a sep-
arate coelomic mantle.
The inner lobe is much
the smallest. The ne-
phridia are closer, over-

lapping.

Apical diyision not glob-
ular; dorsal part with
a single diverticulum.

octonephra Rosa.

Paraguay.

20 to 40 mm.

85 to 95.

xiii to xx, incomplete be-
tween the ventral sets.

The larger curved, strong-
ly curved at apex, and
with 6 groups of 2 blunt
tubercles, the smaller
expanded at apex like
a scalpel.

Strictly paired, all ven-
tral; couples equi-
distant.

A single pore on median
line in xiy, on a slight
swelling.

In line with ventral set.

Three pairs in xy, xvi
and xvii.

In xi and xii, two pairs.

Arranged in four series
on either side of the
median line; the yen-
tral one corresponds to
the dorsal seta.

Basal part pear-shaped;
apical part sac-like,
basal part with a glob-
ular diverticulum di-
rected forward.

malayana Horst,

Malay Archipelago.

20 to 30 mm.

95. Skin without pig-
mentation.

y-vi or vi-vii.

Continued into the bue-
cal segment.

Complete in the anterior,
incomplete in the pos-
terior somites, xiii to xx.

The larger seta with 4
broad bristles; the
smaller spoon - like,
but not forked.

Distance between dorsal
and ventral couples
about equal to that be-
tween ventral couples.

Two distinct pores.

The depression connect-
ing the prostate pores
concaye, with the con-
vex part turned toward
the median line, An
elevated area around
the depression.

The posterior nephridia
arranged in 3 groups
of delicate tubuli on
either side, the inter-
nal ones situated near
the dorsal seta. y

Basal part longest and
as broad as the apical
part, with a narrow di-
yerticulum.

floresiana Horst.

Malay Archipelago.

35 to 40 mm.

125. Skin without pig-
mentation.

vi-vii.

Divides somite I about
one-half.

Incomplete, xiii to xxi.

Anterior part much un-
dulated, the smaller
seta less so; three small
bristles at apex of
largest seta.

Distance between a dor-
sal and ventral couple
about three-fourths of
tliat between the two
ventral couples.

Open in an oval gland-
ular area, pores open
separately on line with
sete 1 and 2.

Not elevated, situated
at the corners of a
square area.

Arranged in four longi-

tudinal series on each
side of the median
line.

Basal part the longest,
narrow; single pear-
shaped diverticulum.

Anne Horst.

Java.

30 mm.

85. Skin withont pig-
ment.

iv-v.

Divides partly the buc-

cal segment.

Incomplete in xiii to xxi.

Undulated, with 12
(about?) slight dentic-
ulations or
The form of smaller
seta doubtful.

The four couples are
ventral and equidis-
tant.

Unknown.

Depressions connecting
pores lunate, with the
convex side turned to-
wards median line.
Area oval, not square;
a ridge borders the
grooves.

Two in vi and yii.

Xv, xvi and xvii.

In ix-xii.

A network of delicate
tubules anteriorly
spread over the body
area, except for a nar-
row dorsal and ventral
median line, Posteri-
orly arranged in four
rows on either side of
median line, and fur-
nished with c@lomic
mantle.

Basal part longest and
very narrow, with a
narrow diverticulum
situated near the con-
traction.

notches. |

TUGOKU Li. Sp.

| San Francisco, Califor-
nia, imported.

30 mm.

118. Skin slightly pig-
mented and anteriorly
corrugated.

v-vi,

Divides buccal somite
about one-half.

Incomplete in xiii to xx.

Largest seta with five
notches; smaller seta
forked, with prongs of
equal size.

Strictly ventral.

pore in xiy.

Anterior prostate pores
elevated on papille.

Not forward.
Tn vii and viii.

Three pairs in xv, xvi
and xvii.

Arranged in four rows
on either side of me-
dian line. Posterior
nephridia with ccelo-

if
| Open jointlyina median ....

mic mantles. Neph-
ridia of nearly the same
size. Neph. d and d@
about equidistant as
a and a. The most
ventral part of nephrid-
ium a@ not covered by

_ mantle. Clitellar ne-
phridia much larger
than those anterior of
clitelum.

Apical part smallest.
Both parts much flat=
tened. Very small di-
verticulum.

jo
*
2

Godeffroyi Michaeleen.
Hayti (or New Zealand”)

90 mm. by 4 mm.

174. Anterior part of
body pigmented.

Divides buceal segment
somewhat less than 2.

Incomplete (xiii), xiv-xix

Slender, numerous irreg-
ular notches, giving
knotty appearance, but
no teeth or bristles.
Many set# in each sac.

In 4 couples, all ventral.

The curved fosse between
the pores, with the con-
vex side turned towards
the ventral median line.
The anterior prostate
the largest.

Plectorephric condition,
but not known in detail,
except that they cover
the whole of the body-
wall except a narrow
strip dorsally and ven-
trally. Stronger in the
clitellar somite, where
they form a thick coy-
ering.

Spermath: with |
distinct diverticula, but

the smaller basal part

i]
~

PACIFIC COAST OLIGOCH MTA. 1

DETAILED DESCRIPTION OF NEW SPECIES.
Benhamia nana pn. sp.
Figs. 1-42.

Habitat. San Blas, territory of Tepic, Mexico. I found a small number of
specimens, of which only two possessed a perfectly developed clitellum, October, 1894.
A real tropical form.

Color. Deep reddish-brown and very opaque.

EXTERIOR CHARACTERS,

Size small, 20 to 380 mm.

Somites 110.

Clitellum comprises somites xiil to xx, being complete in the central somites,
but incomplete in xiii, xvii, xviii and xix.

First dorsal pore ii-iv.

Spermathecal pores, one pair vii-vill, and one pair between viii-ix, rather close
together in a ventral grove.

Oviducal pores open separately in xiy in front of sets 1 and 2.

Prostate pores, one pair in xvii, one pair in xix, open with penial sete.

Penial sete, two in each pocket, the largest of which is furnished with 12 or
more bristles.

Common sete, distance between the dorsal couples equal to that of the ventral
couples.

INTERNAL CHARACTERS.

Buccal cavity with a dorsal pocket.

Suprapharyngeal glands with pharynx opening far forwards.

Gizzards in viii and ix.

Caleiferous glands two pairs, one in xiv, one in xy.

Septal glands very minute in ix, x, Xi.

Sacculated intestine commences in Xiv.

Typhlosole very strongly developed in xviii to xxii.

Sperm-sacs two pairs in x, X1.

Ciliated rosettes in x, xi. The lower part of the spermducts in xvi, xvii, xviii
is thickened and twice the diameter of the anterior part.

Prostates are straight and tubular, each one confined to one or two somites.

Spermathece in vii-vili, yili-ix. The apical part much smaller than the
basal part, the latter with a small diverticulum placed high up near the constriction.

Nephridia arranged in three rows on either side of the median line. Pair a
open in front of the outer couple of sete, pair b and ¢ open laterally and dorsally as
regards the sete. The posterior nephridia furnished with a large celomie cellular
mantle. The nephridia in each somite of about equal size. The nephridia on each
side about equidistant, but the dorsal nephridia ¢ are much further apart from the
median line than from nephridia 0.

=
bo
[0 <]

CALIFORNIA ACADEMY OF SCIENCES.

DETAILED DESCRIPTION.

Body-wall. "The prostomium with bueeal cavity is strongly eversible, forming
a bladder-like apex to the body, as is so frequent in oligochzeta. Somite i is very
narrow, especially laterally, and may readily, when viewed from the exterior, be taken
for part of the prostomium.

The body-wall contains the usual layers of which the muscular fibers show the
same bipinnate arrangement as in Lumbricus, ete. This arrangement is less regular
and pronounced in the anterior somites, but quite plain in the genital and clitellar
ones, especially so on the ventral side, immediately below the nerve-cord. _

Sense organs of the epidermis. All the species of Benhamia described here
possess a continuous row of sense organs, in the equatorial plane of each somite, be-
tween the setse. Outside of this equatorial circle I haye not found them anywhere
in the epidermis; there, however, they are very plain and prominent, appearing under
low power and in longitudinal sections, as a large pellucid spot in the center of each
somite. The organ consists of two distinct kinds of cells, a double line of large lunate
cells, surrounding a row of sense cells, several layers thick. These lunate cells are
generally three or more thick in the row, evidently modifications of the common
goblet cells of the epidermis. They do not stain with the ordinary aniline colors, or
only so with difficulty, and generally remain transparent and white. These lunate
cells run continuously around the somite, and enclose between them bunches of sense
cells, which may now and then be seen to penetrate the cuticle (fig. 20). Somewhat
similar sense organs have been known in Lumbricus, etc., for considerable time, but
have lately been described more in detail by Richard Hesse and F. E. Langdon.
The sense organs of Benhamia differ from those of Lumbricus agricola (probably a
collective name used for some East American Allolobophora) in two prominent
points. Presence of the large lunate cells in Benhamia, which are not seen in
Lumbricus. The continuous and broad circle of these organs in Benhamia, while in
Lumbricus they appear to be much further apart. Unfortunately my Benhamias
were collected at a time when no special preparation for nerve structure was feasible,
and this made it impossible to work out the details as minutely as desirable. The
work had already been finished when Langdon’s beautiful paper reached me.
Fig. 20 represents a section of the body-wall in somite iv. Letters org. signify
the sense organ.

3esides these epidermal sense organs I find in all the Benhamias, observed by
me, a large zone in the buccal cavity characterized by almost cubical transparent
cells arranged in one single row deep, just as the epithelial cells in the pharynx, but of
the same nature as the transparent cells in the sense organs of the epidermis. This zone
is nearly always folded against itself like a sac, and is of considerable extent, as long
or longer even than the pharynx. In certain places apparently scattered about, but
principally near the opening of this sae, I find clusters of sense cells of the same
structure as those of the epidermis. They are all narrow, do not reach below
in the ccelomie cavity, but end in this direction in line with the pellucid cells and
connect at the end with nerve fibres. The free ends penetrate above in what ap-

PACIFIC COAST OLIGOCH ATA. 129

pears as a veritable cuticle, though I can find none above the pellucid cells. In
fact the whole structure of these organs is very much the same as the sense organs of
the epidermis of the body-wall. In connection with them I may point out the simi-
larity of structure of these organs with those described below in Acanthodrilus Vasliti,
in the vicinity of the prostates; and with those in the tubercula pubertatis of Sparga-
nophilus Smithi and tepicensis, in all of which I find the same sense cells.

Tt remains to add that this pouch-like area of sense cells and glandular cells has
previously been observed by both Michaelsen and Horst and designated as a diver-
ticulum of the buccal cavity, but the true nature of the pouch as a sense-area has, I
believe, not been previously recognized. 6. Bolavi, malayana and probably most, if
not all other, Benhamias possess this organ of the buceal cavity.

Septa. These are generally very thin, only two being thicker than the rest,
viz.: Xil-xili and xili-xiy. These septa do not strictly correspond with the inter-
segmental grooves, but are affixed much further back in the somites.

Pharyngeal and septal glands. The usual supra-pharyngeal glands are present.
They are evidently unicellular and arranged ribbon-like as will be directly described
below. The septal glands are found in ix, x and xi, are very narrow and only one cell
thick in the row as the former. These glandular masses are much thicker in B.
palmicola, but not any longer.

Intestine. In all the specimens which I could examine I found the pharyngeal
parts strongly everted and protruded to such an extent that it formed the lip or front
margin of the body, taking the usual place of prostomium proper. I cannot ascribe
this entirely toa simple protrusion of the pharynx, but believe that this part is actu-
ally situated much more forward than in other species, as I found it to be the case in
every specimen observed. ‘The pharynx was found actually on the outside of the body
(figs. 15, 16, 17, 18). The real pharynx, of course, is the zone in which open the
supra-pharyngeal glands. The wall of this part is in our present species hardly
thicker than those of the buccal cavity and the cesophagus, but it contains the usual
arrangement of narrow epithelial cells, between which penetrate the fine ducts from
the supra-pharyngeal glands. The ends or discharge pockets of these glands are
almost globular or rounded flask-like (fig. 18). The supra-pharyngeal glands are
arranged as ribbons, running singly along muscular strands. They differ from similar
glands in Pontodrilus, Pheenicodrilus, Ocnerodrilus ete., by their arrangement in
single rows, and here and there the duct of a single glandular cell may be followed
clear to the discharge pocket (fig. 16). These glands appear to consist of a single
cell with a long duct, just as the corresponding glands in Euchytr:eus, described by
Hesse. But to draw the conclusion from this fact, that all the pharyngeal and septal
glands are unicellular is, I think, premature. In Pontodrilus, at least, there
may be seen plainly numerous nuclei on the gland duets, which, of course, indi-
cates that we here have a fusion of several cells. The pharyngeal glands in that
genus do not show this ribbon-like ar rangement as in Benhamia. I could, however,
see that some of the smaller glands nearest the pharynx consisted of only one cell,
but the majority, and all the large glands, consisted of several cells, the respective

130 CALIFORNIA ACADEMY OF SCIENCES.

duets of which finally united into one. In Benhamia I could see no such union, and
the single ducts could be followed with great facility to the outlets. Fig. 17 repre-
sents some of these cells with three narrow ducts and secreted matter.

(Hsophagus, following pharyngeal division, is very long. The upper part, im-
mediately below the pharyngeal gland, is very thin-walled, consisting of only one
strand each of transverse and longitudinal muscles, and lined by a very narrow epi-
thelium (fig. 19). Posteriorly the walls of the cesophagus thicken considerably. The
two distinet gizzards are in viii and ix, as usually connected by a very thin wall of
the same general nature as cesophagus (fig. 7). The muscles of the gizzards are
columnar but not bipinnately arranged, the ribbons running parallel with the short
diameter of the body (figs. 7 and 21).

The tubular intestine extends through somites x to xiv, being of irregular out-
line. Sacculated intestine commences in xy, and is furnished with a typhlosole in
somites xviii to xxiii, or thereabout.

Epithelial cells of the alimentary canal surround glands of various forms. In
the epithelial lining of the gizzard we find club-like glands (fig. 21 g/.) consisting
each of one (seldom of more) large cell with round nucleus, a narrow duct reaching
between the epithelial cells, and ending witha large chamber, in very much the same
way as the pharyngeal and septal glands.

In the narrow thin walled part between the two gizzards I find a few clusters
of glands (?) similar to those I have described in Argilophilus. At the bottom of the
cluster we find a glandular cell, upon which are butting the peculiar lunate cells,
which again surround a lumen, which is much wider than in the corresponding organ
in Argilophilus (fig. 22 g/, and c.) The lining of the sacculated intestine and the
typhlosole are composed of three distinct kinds of cells, two of which are glandular
(figs. 24 and 25.) The common epithelial cells offer nothing of particular interest.
The glandular cells are of two distinet kinds (fig. 25). One kind is the one most
common in oligocheeta (fig. 25 g/.) and its enclosed granules are much smaller. The
other kind is probably identical with the T-shaped cells described by Benham in
Eminiodrilus, though the T-shaped form is not quite so prominent. The granulation
is coarse and highly refractive and the distal part of the cell stains intensely, especially
with methyl green. The other or first mentioned glands remain at the same time un-
affected by this stain. These dark staining cells are much less numerous than the
other kind, and are scattered about in a rather regular way. Fig. 24 4 represents
one of the lobes of the typhlosole, showing the absence of glands at the apex as well
as the general distribution of the dark staining cells.

Calciferous diverticula. I found in the two specimens dissected and sectioned
ouly two pairs of calciferous glands in somites xv and xvi, but [ am unable to say if
this will be found constant, as all other species of Benhamia possess three pairs
of calciferous pockets, which Beddard claims are characteristic of this genus.
Michaelsen’s observation that the two diverticula are of different nature is confirmed
here, as in the posterior pair no lime crystals were found, either in this or in the other
species described below. Also the histological structure of the posterior and anterior

PACIFIC COAST OLIGOCH TA. 131

diverticula is distinct. The nuclei of the anterior pair are round while those of the
posterior pair are more oval, and the cells of the latter pair are not separated by dis-
tinct walls. They contain also numerous vacuoles or bladder-like bodies, some of
which deatch themselves from the main body of the organ.

Figures 27 and 28 represent the anterior pair, 29 and 30 the posterior pair of
calciferous diverticula seen respectively under high and low power. The posterior
pairs only were found covered by chloragogen cells. These differences in structure I
also found in the ealciferous diverticula of the following species: Benhamia palmi-
cola and rugosa, and they probably hold good with all the species of Benhamia.

Typhlosole exists in somites xvil or xviii to xxii, and is greatly developed as
regards length, width and depth (figs. 7 and 14). It is widest and deepest in somite
xix, tapering or diminishing both anteriorly and posteriorly. In the following species
the typhlosole is much smaller.

Spermathece. The two pairs as usual in vii—vili and yiii-ix. The form of the
spermatheca and its diverticula may be best understood from the flgures 54 and 39,
The pores come rather close together in a general groove below the ventral ganglion.
There is a muscular and a glandular part, and the spermatozoa were fonnd as usual
principally in the diverticula.

Spermducts and rosettes. The only peculiarity of the spermducts is that they
are thickened in somites xviii, xvii and xvi. In xy the duct narrows, and continues
forward, with about half the thickness it possesses in the above posterior somites. The
thickening is due to an increase of a circular muscular layer (figs. 7, 11, 12, 15, 56,
307). The rosettes are in x, xi, as usual.

Prostates. The usual two pairs are in xvii and xix. The lower part is long,
slender and muscular, while the upper thicker part is glandular. This glandular part
consists of only one layer of cells, covered by a thin epithelium. The glandular cells
are of different length, as shown in fig. 32. The muscular part consists mainly of a
thick layer of circular muscles (fig. 386), lined by a very thin layer of interior epi-
thelial cells (fig. 38). ;

Penial sete. A sae with two penial sete open jointly with each prostate. At
least one of the sete is sculptured as shown in the figures 31 and 35, with a number
of bristles, 12 or more. The smaller seta was broken in all the specimens, and I
could not ascertain if it was sculptured or not.

Nephridia. Benhamia nana belongs to a group in which the nephridia are
arranged in three rows on either side of the median line. There is not a diffuse
nephridic condition, but each nephridium is well developed, and upon the same
principle as the mega-nephridia of Acanthodrilus. Nephridia a. >. are ventral and
in front of setee 5 and 4, while the other nephridium c., is lateral and partly dorsal.
The posterior nephridial ducts, from xvii to xviii, are superposed each on a large oval
sac of coelomic cells, while those in front of the male-pore are not furnished with any
coelomic mantle. Of the anterior nephridia those in the genital somites are some-
what smaller than those in the vicinity of the pharynx, the increase in size forward

being gradual. Blood capillaries are found in great numbers on all the nephridia, but
Memorrs, Vou. II, 5. December 14, 1895.

132 CALIFORNIA ACADEMY OF SCIENCES.

especially so on the posterior ones. The general structure of the nephridial canals
and their windings resembles those of fully developed mega-nephridia (figs. 39, 40,
41), but the smallness of these organs prevented me from definitely ascertaining the
relative size and structure of all the canals. The canal leading to the outlet duct is
strongly ciliated. Fig. 41, which represents the inner part of one of the anterior
nephridia is held somewhat diagrammatic, as the respective thicknesses of the canals
could not be properly delineated. I believe the most anterior nephridia are found in
iii, in which somite they are very large, crowding each other.

Benhamia palmicola n. subsp.
Figs. 48-55.

Habitat. Miraflores, in the Cape Region of Baja California, some 40 miles
north of San José del Cabo; also around Tepic, territory of Tepic, on the Pacific
Coast of Mexico. The altitude of Miraflores is about 1,000 feet, while that of the
City of Tepic is about 4,000. Probably the species does not descend to the hot low-
lands of the coast belt, at least no specimens have been found either at San José del
Cabo or at San Blas.

Color. The specimens are pink flesh, much less opaque than 6. nana, no pig-
ment cells.

General Remarks. For the present I arrange this as a subspecies under
B. Bolavi, from which it differs in several points of considerable importance. These
are the different form of spermathece, the situation of the first dorsal pore, the differ-
ent sculpture of the largest penial seta, the location of the gizzard, the number of
sperm-saes, ete.

EXTERIOR CHARACTERS.

Size 50 to 60 mm.

Somites 110.

Clitellum incomplete in all somites xili—xx.

Dorsal pores, most anterior one between iy—y.

Spermathecal pores in line with sete 1 and 2, between vii/vili and vilii/ix.

Oviducal pore. One central median oyiducal pore in xiv.

Fenial sete, two in each sac. The longer seta has apex curved and furnished
with four blunt, bristle-like crenulations. The smaller seta has the point slightly
forked, one prong being longer, anda thin membrane extended between the two forks.

Common sete. About equal distance between the dorsal and ventral couples
as between the ventral couples.

INTERNAL CHARACTERS.

Buccal cavity with a dorsal pocket of sense organs and glandular cells.
Pharynx not situated far forward.

Gizzards both apparently in viii.

Calciferous diverticula, 3 pairs in xv, xvi and xyiii.

Septal glands, small in ix and x.

PACIFIC COAST OLIGOCHATA. Le

eo
we

Typhlosole small,

Sperm-sacs in xi and xii.

Spermducts not thickened in the somites near the pores.

Prostates are larger than in B. nana, but mostly confined to one somite each,

Spermatheee in yii-vili and vili-ix. The two divisions are of equal size, and
globular; small tubercular diverticle pointing forwards.

Nephridia in three distinct rows on either side of the median line. Nephri-
dium @ nearest the ventral ganglion consists of two unequal parts, the most ventral
of which is the smallest. These two parts together are much larger than either of
the other two nephridia. The dorsal nephridia are far apart.

DETAILED DESCRIPTION.

Body-wall and sense organs. We find the same zone of sense organs in the
body-walis as in B. nana. In B. palmicola this zone is found in all the somites,
while in B. nana I found none in somite i.

The sense-organ zone with pellucid cubie cells is in this species larger than in
B. nana, and stands out most prominently on the upper side of the buccal cavity.

Septal glands. As has already been stated small septal glands are found in
somites ix and x or posterior to the gizzard. These glandular masses are quite small,
are situated in the anterior part of the somite, close to the intestine, but hardly pro-
jecting above the latter. I cannot find that they are in anyway connected with the
pharyngeal system of glands.

Intestine, etc. The portion of the pharynx is not far forward. The region of
its epithelial columnar cells is thick and the discharge chambers of the pharyngeal
glands are tubular, instead of globular as in b. nana. The pharyngeal glandular
mass is less lobed than in B. nana.

The region between somite iand pharynx is much longer in B. palmicola than
in nana. The specimen sectioned longitudinally showed the two gizzards as both sit-
uated in vill, and there was no septum separating them. If this character is constant
is undecided as I did not wish to sacrifice another specimen. The tubular intestine
is long and cylindrical and furnished with the regular three pairs of calciferous
diverticula in somites xy, xvi, xvii. No lime crystals in the posterior pair. The
sacculated intestine commences in xviii or xix. The typhlosole is smaller than in
B. nana, but owing to sand I could not definitely ascertain its location.

Spermathece. The four spermathece are of the same form and size. The
contraction at the center is deep, dividing the organ in two equal, globular sacs, each
sac being confined to one somite; that is, the apical part is situated entirely in the
somite posterior to the pore. The basal part carries a short narrow cylindrical and
tubular diverticulum pointing forwards. The equality in size and the globular form
of the two parts of the spermatheca appears very constant and characteristic of the
species.

Sperm-sacs are sac-like, not racemose. They are found in xi and xii, while
in B. nana they are placed in x and xi.

134 CALIFORNIA ACADEMY OF SCIENCES.

Spermducts are considerably thickened, but the lower part is not any thicker
than the upper part, as in B. nana.

Gonads are affixed high up on the septum and not at the junction with septum
and body-wall.

Prostates. These bodies are larger than in B. nana, but nevertheless generally
confined to one somite each. The penial sete are, as usual, of unequal size. The
larger seta is furnished with four notches, the smaller is spoon-like and forked at the
apex.

Nephridia. These organs are arranged in three distinct rows on either side of
the median line, but the nephridia in each somite are of unequal size. The nephri-
dium nearest the ventral ganglion or nephridium a consists of two, more or less,
separate parts, evidently a tendency to diffusion or an imperfect centralization. The
most ventral part is the smallest and the most distal the largest. The ducts run con-
tinuously between these parts, but the ecelomic cells are grouped in such a way that
the bridges between the two parts are quite narrow. The nephrostome of nephridium
a was always plain and readily seen, but I never succeeded in finding the nephro-
stomes of bande. Still, these nephridia appear perfectly formed on the meganephric
principle, and I could never see any connection by canals between @ and /, and } and
c, though sometimes the ccelomie cell masses extended more or less continuously over
and between the respective nephridia a, } and ce.

Nephridia @ open in front of setee 1 and 2, while in B. nana they open in front
of 5and 4. ‘The most anterior nephridium possessing a coelomic covering I found in
xxi. Through the courtesy of Dr. Michaelsen, I have received specimens of J.
Bolavi for comparison. The nephridium of Bb. palmicola resemble that of 6. Bolavi,
but is much larger, and the respective nephridia cover each other slightly, while in
the two specimens of 4. Bolavi which I dissected the respective nephridia were sep-
arated by considerable distance; the latter nephridia are also smaller. I had at first
intended to assign these species of Benhamia possessing several nephridia of a perfect
form under a separate subgenus, when my attention was called to the fact, by Dr.
Michaelsen, that B. Stuh/manni sometimes possessed a similar arrangement of nephri-
dia as those in B. Bolavi, ete. As the nephridia of B. Stuhimanni are generally
plectonephrie or diffuse, it became at once evident that this distinction could not be
used as a generic character of value, and that it really is impossible to draw any dis-
tinct line between a plectonephric and a micronephric condition. It is, however, en-
tirely incorrect to characterize these nephridia as a mass of tubules, etc., as wherever
they are separated one from the other, as in Bolavi, palmicola, nana, rugosa and
probably great many other species, each micronephridium is perfect in itself, and built
on the same general principle as the meganephridia of the other terricole. I would
therefore propose to. make a distinction between plectouephidia, or really diffuse
nephridia, and micronephridia, or nephridia of small size, but perfect, or built on the
meganephric plan. Such a distinction may be useful in descriptions, even if they are
not morphologically distinct.

PACIFIC COAST OLIGOCHEHTA. 135

Benhamia papillata n. sp.
Figs. 43 a, B, Cc, D, E; 52.

Habitat. 'Tepic, Territory of Tepic, 4000 feet; Mexico.

Tam unable, on account of want of time, to add any detailed description to the
short definition of this species given above. When I described Benhamia palmicola
I possessed only a single specimen of B. papillata,and I supposed that the differences
in the structure of some of the organs, as well as the presence of the prostate papille,
were due exclusively to individual variation. But while this paper was being printed
I became possessed of about twenty specimens from the same locality, all of which
resemble each other in all the points referred to in the description, and I therefore
do not hesitate now to assign to them a specific name, especially as I find that this
species is much more distinct than I at first suspected. While it in many respects
resembles L. Golavi, it differs in other points which must be considered of specifie im-
portance. b. papillata differs thus from B. Bolavi in possessing four exterior tubercles,
one for each prostate pore. The smaller penial seta in B. Bolavi is flat and somewhat
forked, while in B. papillata the smallar seta is furnished with a sigmoid tip of ex-
ceeding thinnesss. The clitellum in B. Bolavi and B. palmicola is incomplete, but
in BL. papillata it is complete even on the ventral side, showing several rows of glan-
dular cells, but the width of the layer of clitellar cells is much narrower in the central
part. Clitellum is ventrally complete only in somites xiv, xv and xvi.

The following are the other points of interest as regards the character of this
species: It is larger than B. palmicola. The four papille, each one of which
carries a prostate pore, are very distinct and prominent, and they stand close together
in the sunken genital pit of the clitellum. The median central papilla, on which open
the ovipores, is elevated and oval. The two ovipores are situated in the center of the
papilla, but entirely separate. They are in the very center of the somite and ina
line drawn between setee 1 and 1. In B. palmicola the ovipores join, and the common
pore is situated somewhat in front of a line drawn between sete 1 and 1. Longitu-
dinal section of a specimen shows that the septum separating somites xiii/xiy is very
much cupped, and the oviducal funnel is situated exactly above the central papilla in
xiv and dips down straight to the ovipore. The anterior septe are hardly thicker
than the posterior ones, but they are all very much cupped. Sacculated intestine
commences in xix. There is a strong superior typhlosole in xx to xiv. Of the three
pairs of calciferous diverticula the anterior one in xy is very much, or about four
times, smaller than each of the posterior ones. The glandular part of the prostate is
folded and quite thick, but confined to one somite.

Penial sete. ‘The penial sete are the most distinet character of this species,
besides the tubercles of the prostate pores. In general shape the two setee are much
more slender than those of B. palmicola, and several times narrower at the apex. They
are so thin that it required an oil-im. 1/12 to show their structure sufficiently to enable
me to sketch it. The larger seta is slightly curved, gradually tapering from the root
to the apex. The apex is much less curved than that of B. Bolavi or B. palmicola,
and furnished with seven or eight shallow but still distinet notches. But the smailer

136 CALIFORNIA ACADEMY OF SCIENCES.

seta differs entirely from that of the B. Bolavi group. Instead of being more or less
distinctly forked and wider at the apex, as in those species, it is drawn out to an ex-
ceedingly thin hair-like point, which is sigmoid (fig. 43 ¢). In shape the seta is almost
straight, with the two ends slightly curving.

The papille of the genital region are most characteristic. Each papilla forms
the exterior apex of a prostate. With a low power lens these papille appear round,
but under a higher power they appear rosette-like, and consist of two or three divisions,
one on top of the other. The ventral part of segments xvii, xvili and xix, carrying
the genital pores, is much contracted, and the papillee appear as ina bunch. The
distance between the male or spermaducal pore and the prostate papilla is Just about
the same as the base of the papilla.

In a section the structure of the oviducal median papilla appears to greatly
resemble that of the tubercula pubertatis of Sparganophilus. IT find there the same
relatively large number of tall, narrow cells, radiating from the ovipore, but I cannot
discern any sense cells.

The spermathecz differ in shape from those of B. palmicola. ‘The apical sac
is not globular, but pointed and narrow, as represented in fig. 52 G and H.

The nephridia also differ some from those of B.palmicola. In this species the
inner nephridium, or a, consists of two parts, each one with a ccelomic mantle separate
from the other, but in b. papillata nephridium a consists of three distinct parts, each
one with a coelomic mantle. This, of course, refers only to the posterior nephridia;
the anterior ones are not covered by ccelomic mantle.

Sperm-sacs in x and xi, not racemose. In other respects, as far as I can judge
by a hasty examination, this form resembles B. palmicola.

Benhamia rugosa n. sp.
Figs. 56-63.

Habitat. Native habitat unknown. The eight specimens in my possession
were found in the orchid house in the Golden Gate Park of San Francisco, California,
under pots. Two were adult, two imperfectly developed, the others immature.
July, 1895.

Color reddish flesh.

EXTERIOR CHARACTERS.

Size, 30 mm. by 14 m.

Segments, 118. Cephalic lobe very long, pointed.

Clitellum incomplete.

Dorsal pore, most anterior y—vi.

Oviducal pore, one single in the median line in xiy.

Prostate pores, the anterior ones elevated on tubercles.

Penial sete. Largest seta with five notches, smallest seta forked with prongs
of equal size.

Common sete. All ventral in four couples; the inner sete are present in xviii.

PACIFIC COAST OLIGOCH ETA. 137

INTERIOR CHARACTERS.

Gizzards, two in vii and yiil.

Calciferous diverticula. Three pairs in xy, xvi and xvii.

Sacculated intestine commences in xviii.

Typhlosole present.

Prostates slender, confined each to one somite.

Spermathece. The apical part a trifle smaller than the basal. Both parts
much flattened. A very small diverticulum pointing forwards.

Nephridia in four rows on either side of the median line. The posterior ne-
phridia with celomic glandular mantle. Nephridia in each somite of about equal
size. Nephridia d and d about equidistant as a and a. The most ventral part of
nephridium @ not covered by the mantle. The clitellar nephridia larger than those
anterior to clitellum.

DETAILED DESCRIPTION.

Size. The specimens had been slowly killed and extended before hardening.
All the specimens were curved backwards, thus with the dorsal line on the inner
margin of the crescent, causing the male pores to be situated at the greatest bend on
the outer margin of the crescent. Most terricole curve the opposite way.

Shape of segments. The first ten somites are very nearly of the same diameter
in the direction from head to tail. Somites xi and xii are slightly narrower. The
clitellum which comprises somites xiii to xx shows plainly the intersegmental grooves.
The posterior somites are of a somewhat shorter diameter than xi and xii. All the
anterior somites including the most posterior ones of the clitellum are sculptured by
longitudinal furrows running in the diameter from head to tail. This corrugation is
seen all around the body, and gives the anterior part of the worm a very marked
appearance. The posterior somites show a fainter corrugation. All somites are
3-ringed, except those of the clitellum.

Prostomium is long and pointed and slightly curved upwards (figs. 57 and 58)
and is a trifle longer than somite ii. Inall the specimens but one, somite i, was nearly
entirely retracted in the buccal cavity, and when viewed from the exterior only a
short portion of its dorsal part could be seen (fig. 58) and the prostomium appeared
as if projecting from somite ii. One specimen had somite i extended as figured
in fig. 59. Dr. Michaelsen has remarked a somewhat similar retraction of somite i in
B. kafuruensis. Spermathecal pores as usual, vii—vili and viii-—ix.

Clitellum is narrower than the surrounding somites. It is strongly corrugated,
especially at the anterior and posterior margins. The prostate pores are, as usual, in
xvii and xix. The exterior part of the genital region is very characteristic. In
young specimens the two prostate pores are connected by a deep furrow in the
center of which is seen the slight depression for the male pore. In the two fully
adult specimens, however, the two anterior prostate pores were each situated on a
very large globular papilla, surrounded by a deep fossa; while no such papilla and
fossa characterized the posterior pair of prostate pores. In fact, these posterior pores

138 CALIFORNIA ACADEMY OF SCIENCES.

were not visible from the exterior and could only be ascertained by dissection and
transparent light. Viewed from the exterior the clitellum appears incomplete.

Penial sete. The longer seta is hooked at the apex and furnished with five
notches on the inner side, very much as in B. Bolavi. The shorter seta is less curved
and about # as long as the larger sete. It is not hooked and the free apex is slightly
forked, each prong being of the same length. There is no flare or wing between the
two prongs (figs. 60, 61 and 62).

Oviducts. Both open together in a central pore in the median line in xiv,
very close to the groove between xiii-xiy. In the largest of the two adult specimens
the ovipore was situated abnormally between xv and xvi.

Intestine and glands. A strong supra-pharyngeal gland as usual. Gizzard in
vii and viii, separated by a thin wall, as in other species. Tubular intestine extends
from gizzard to xvii, and is in xy, xviand xvii furaished with the three pairs of
ealciferous diverticula, of which the two anterior ones are of different construction
histologically just as in other species. The sacculated intestine commences in xviii.
There is a typhlosole.

Spermathece. As usual, in vii and viii. Those in vil were slightly smaller,
and entirely confined to their somite, wh'le those in viii projected their distal part
through the septum into ix. The distal part is only a little smaller than the other
part. The spermatheca is much flattened. Seen from below it appears as in fig. 52 m,
while seen from the side it looks as in fig. 52 2. The lower part is furnished with
a very small diverticulum, pointing forwards. The duct leading to the pore is very
short, almost not set off from the lower sac.

Oviducts are slender, tubular, with a small and narrow funnel.

Nephridia. ‘Those posterior to xix are furnished with a coelomic cell mantle
of oval form, even and regular, and of the same size in the various nephridia. Pos-
terior to xix we find four rows of nephridia on each side of the median line, the rows
being reguiar and parallel. The most ventral nephridium or a is situated between
the setze couples 1 and 2 and 3 and 4. The other nephridia 6, ¢ and d are dorsal and
lateral. The two dorsal nephridia d and d are about as equidistant as nephridia a and
a, and much closer together than the dorsal nephridia of B. nana, palmicola and
Bolavi, the three other species which I have examined. The more ventral part com-
prising the outlet duct, ete., of nephridium @ is longer than the corresponding parts
of the other nephridia, and not covered by the ccelomic mantle. The clitellar nephri-
dia are larger and their ducts thicker than the other anterior nephridia. The
nephridia anterior to clitellum are very small, except those in somites iv and v, where,
as usual, the nephridial ducts are long. Each nephridium appears perfect, and built
on the meganephric plan (fig. 63).

Benhamia rugosa is readily recognized and characterized by the large papille
on which open the two anterior prostate pores, by the four rows of nephridia, by the
forked smaller penial setee, by the corrugation of the anterior somites, inclusive of
clitellum, by the pointed prostomium, by the flattened spermathéce.

Since this was written Rosa has described a new American Benhamia from

PACIFIC COAST OLIGOCH UTA. 139

Paraguay, viz.: B. octonephra, which appears to resemble my B. rugosa in several
points, especially in having four rows of nephridia on either side of the ventral median
line, or eight parallel rows in all. But it also differs in several others, sufficiently to
be arranged under separate species. BL. octonephra has about 90 somites, 2. PUGOSA
about 118. . rugosa possess penial setse, which, as far as I can judge from Rosa’s
description are different from those of L. octonephra.

Unfortunately, the characteristics of Benhamia species must be founded on
most minute characters, which cannot always be expressed in words, as long as so com-
paratively few species are known. oo few of these characters have been figured,
and many have been entirely overlooked. The penial sete, which are of the utmost
importance as species characters, should be in all instances figured. They vary much
less than do the spermathec, the shape of which is only approximately constant.

ACANTHODRILUS Perrier.

Out of thirty-five species of this genus, which have been described to date and
recognized by Beddard, none have been found as far north as Central America or
Mexico. The two new additions to the genus which I am able to describe below will
therefore prove of considerable interest on account of their habitat so far north, while
one of the species shows a combination of characters rarely met with in this genus.

DEFINITIONS OF SPECIES.

Acanthodrilus tamajusi n. sp.

Derimition. Length 15 cm., width 1 em. in thickest part in front of clitellum;
number of somites about 218. Clitellum aiii to vx. First dorsal pore posterior to clitel-
lum. Penial setw ornamented, one seta only in each sac. Common sete paired. Pros-
tate pores each ona round papilla. Gizzard inv. Caleiferous diverticula 3 pairs in
vii, nit and ix. Saculated intestine commences in xiv. Typhlosole present. Sperma-
theca large sac-like, with a few small warty diverticula. Sperm-sacs not racemose, in ix,
v, wi, vit. Prostates four, much folded, opening with penial sete. Nephridia large,
meganephridia, not alternate. Color reddish, partly pigmented. Habitat, Guatemala,
lowlands.

Acanthodrilus Vasliti n. sp.

Derimition. Length 6 em. by 2 mm. in the region of somite viii; number of
somites 92. Prostomium distinct, divides somite 1 about 4. Clitellum unknown. First
dorsal pore vii/viii; first large dorsal pore ix/x. Penial sete absent; common seta
paired, 1 and 2 not present in xvii and xix. Prostate pores not on papilla; each pore a
duplex one. Ovipores in front of sete 1 and 2. Spermathecal pores in front of
sete 1 and 2, and between somites vii—-viii, viitiix. Spermiducal pores in xviii in line
with sete land 2. Septa v to wii slightly thickened. Gizzard in v. Sacculated intes-
tine in xv. No caleiferous diverticula. Peritoneum covered with a single layer of very
large glandular cells. Very large suprapharyngeal gland, and a very small subpharyn-

geal gland. Spermatheca long tubular in viii and ix, no diverticula. Prostates 8 in
Memorrs, Vou. II, 5. December 16, 1895.

140 CALIFORNIA ACADEMY OF SCIENCES.

aumber, 2 and 2 opening close together in the prostate pore. Typhlosole. Nephridia large,
meganephridia, not alternate, opening in front of sete 3 and 4. Color milky white, no
pigment. Habitat, Tepic, Mexico, 4000 feet altitude.

Acanthodrilus tamajusi n. sp.
Figs. 87-96.

Habitat. Of this species I possess one adult and half a dozen young speci-
mens, principally from Tamaju on the rio Polochic in Guatemala. The largest in-
dividual measures about 150 mm., but even this one did not have the elitellum per-
fectly developed, but was otherwise seemingly mature. One of the smaller specimens
measuring only 70 mm. in length possessed a more perfect clitellum than the larger
specimen. These specimens were found by myself several years ago in June and
July on top of the ground after rain. It is probably a common species in tropical
Guatemala. On account of imperfect preservation of the specimens, some histological
details are wanting.

Color. Deep violet-brown and iridescent when alive. Skin pigmented strongly
on the tail, slightly on anterior part, but only on the upper side in the two mature
specimens. A large immature specimen possessed no pigment, and was of a grayish
color, but there may be some doubt as to its belonging to this species.

Size. Largest specimen 15 em. by about 1 em. in front of clitellum. The
specimen had not been extended and killed before being placed in alcohol and must
therefore have been considerably longer.

Number of somites 218 about, those in front of clitellum much larger than
those posterior to it.

First dorsal pore in the pigmented specimens only posterior to clitellum. In
the unpigmented between vil and xiii.

Prostomium doubtful (injured), but appears to incompletely or not at all divide
somite i.

Clitellum complete, xiii to $xx.

Penial setw. A sac with one penial seta at each prostate. This seta has
numerous short rows of teeth, decreasing in number towards the apex. The seta is
quite short, blunt at one end and broad, tapering gradually toward the apex which is
slightly recurved. The whole seta is irregularly sickle-shaped. The teeth are on
the concave side. One seta possessed four notches with short teeth also on the con-
vex side posterior to the other, or about where the thickest part begins (figs. 93 A, B,C).

Common sete strictly paired on ventral side of body. The distances between
the couples about the same. The ventral couples of sete are present near to the
spermiducal or male pores. They are the ordinary form but slightly larger than the
common sete.

Oviducal pores in xiy situated somewhat nearer the median line than to the
inner couples of setee, and the distance between the pores is a trifle more than the
distance between setxe 1 and 2 in the same somite.

Prostate pores situated each on a small papilla.

PACIFIC COAST OLIGOCH ATA. 14]

Gizzard is in v.

Calciferous diverticula, three pairs, one each in vii, viii and ix, all of the same
structure.

Sacculated intestine commences in xix and furnished with a typhlosole.

Spermathece in viii and ix with a few warty diverticula, one of which is
larger and furnished with two swellings. Spermathecal pores in front of sete 1 and
2. The spermathece are large, filling the space between the septa on the ventral side
of the body. No spermathecal copulary sete.

Sperm-sacs sac-like, not racemose, occupying the whole space left in ix—xii.

Prostates much folded, opening with penial setz; each prostate confined to
one somite each, the free distal end pointing backwards, and situated close behind the
male pore.

Nephridia, strictly meganephridia, not alternating.

DETAILED DESCRIPTION.

Each prostate pore is situated on a round globular papilla, between which and
the male pore runs a copulatory fossa. In xvi and xx there is in each a copulatory
ridge, running parallel with the intersegmental grooves and somewhat longer than the
space between sete 2 and 2 (fig. 91). There is a shorter ridge in xviii extending
between the male pores.

The prostomium and somites i and ii are transversely suleated, most so somites
iand ii. Also somite ili shows this sulcation in a smaller degree. The anterior nine
somites are more or less distinctly segmented in three parts, that is, they show a central
ridge on which are situated the sete. The following posterior somites show a seg-
mentation in five parts. In the clitellium the segmentation is clear, as far as regards
the intersegmental grooves, but the segmentation of the somites is not very distinct.

The supra-pharyngeal glands forma very large body, superposing the pharynx.
It is rounded posteriorly, and consists of four larger and five smaller lobes, this when
viewed in a longitudinal section just outside the median line (fig. 96).

The calciferous diverticula show all the same structure.

The spermathece are large, globular masses, as seen in figs. 94 and 95, with
warty diverticula, one of which is much larger. The spermathece overlap each other
and are bunched in a solid mass between the septa on the ventral side of the body.

Testes are in two pairs, quite or very small when viewed in sections—in x and
xi, in front of the ciliated rosettes, on the posterior face of the anterior septa.

The ciliated rosettes or sperm-funnels are in xi and xii, and are very large,
crimped and furnished with a wide muscular duct. The funnels in xii lie much
higher up than the one in xi, much closer to the intestine. The posterior part of the
funnel is very thick, and so is the duct. The ducts join and open as usual in xviii,
The rosettes lie free, but are connected by connective tissue with the anterior septa.
They thus point backwards.

The body-wall is very thick, and the longitudinal muscles are not bipinnately
arranged. The septa between vy, vi and so on until xi, xii, are much thickened, and
thicker than the other, though the one between xi, xii is not as thick as the anterior

142 CALIFORNIA ACADEMY OF SCIENCES.

ones. The gizzard is connected by several powerful muscular strands with the body-
wall in xi.

There is a very large mass of pepto-nephridial tubes situated immediately above
the posterior part of the pharyngeal gland. Longitudinal sections show the anterior
nephridial canals to be greatly folded and very narrow. ‘There are very large masses
of free ecelomie cells situated in the somites containing the sperm-sacs and the funnels
of the spermducts.

Acanthodrilus Vasliti n. sp.
Figs. 148-154.

General Remarks. his species is one of those abnormal forms which occur
in almost every large genus, and whose organization and characteristics are not readily
accounted for. It is also the most northern of any Acanthodrilus found so far, though
undoubtedly true Acanthodrili will be found much further north. Acanthodrilus
Vasliti differs from any other Acanthodrilus in possessing eight prostates or spermi-
ducal glands, arranged in four pairs, two and two prostates opening together in each
one of the four prostate pores in somites xviii and xx. Also in one other respect does
this species show an interesting characteristic. The peritoneum lining the septa and
body-wall is covered with enormous glandular cells, very much resembling those
forming the nephridial mantle in many species. The duplication of the prostates is
also found in Aerria Mc Donaldi, while abnormal development of peritoneal cells re-
mind us of certain coelomic organs found in Perichzeta and some species of Acantho-
drilus (Beddard, page 29, Monograph of Oligotheeta). Four pairs of prostates have
been described by Ude in Geodrilus singularis, but details are wanting.

Of the specimens in my possession one was sectioned longitudinally, two were
dissected and afterwards sectioned vertically. None of the specimens were fully
adult; the various generative organs were developed, but there was no trace of
clitellum.

Habitat. epic, Territory of Tepic, Mexico, at 4000 feet altitude, in the
moist ground immediately under decaying logs, in the shade of a stone-fence, about
one mile north of the city. October, 1894, Eisen and Vaslit, col.

Color. Milky white, like an Enchytreeus, without trace of pigment.

Size. Length 6 em. by 2 mm. in the region of somite vill. Slightly tapering
towards the tail, the end of which is thickened.

Number of somites 92 in the largest specimen, all of about the same size, ex-
cept the last few caudal ones, which diminish in width towards the most posterior
somite. All are smooth, the anterior ones with a faint trace of trisegmentation.

Dorsal pores. The most anterior one that is distinct is seen between ix—x.
Between yii/vili and viii/ix there is respectively a much smaller but still distinet pore.
The most anterior pore is thus between yii/yiil.

Prostomium is distinet, dividing somite i about $. The anterior somites are
more distinctly set than the others. A long narrow groove begins on the ventral
median line between somites xyi and xyiii, and extends backwards about 20 to 25

PACIFIC COAST OLIGOCHMTA. 143

somites, then changes into a narrow keel. This in the largest specimen, all haying
been slowly killed and straightened out.

Clitellum unknown, all specimens being immature.

Genital pores. Two pairs of spermathecal pores between ix—vili and yili-vii, in
front of the respective septa, the spermatheci thus being in ix and viii. The pores
are in line with sete 1 and 2, but are not prominent. Oviducal pores are in front of
setee 1 and 2 between xii/xiv.

Prostate pores in two pairs in the center of somites xvii and xix, in line with
sete 1 and 2. But each pore is really a duplex of two joint pores, which are only
separated at the very epidermis by a thin wall. Thus the two prostates of each
couple run parallel through the body-wall, each one opening separately, but the pores
being so closely joined that they appear almost as one. No sete of any kind near these
pores (figs. 151 and 152).

Spermiducal pores are seen in line with sete 1 and 2, close posterior to the
sete, only visible by strong transmitted light. The genital region in my immature
specimens did not show any particular structure with papille, ete. The median fossa
already referred to causes these pores to be situated on a slight ridge, which was not
marked off laterally. Close to each prostate pore on their ventral side is a small
tubercula pubertatis, only visible in sections.

Penial setw absent.

Common sete. Strictly paired ventral and lateral; 5 and 4 being situated
slightly ventral, below the horizontal line. Sete 1 and 2 are missing in somites xvii
and xix. In xviii the ventral sete 1 and 2 are present, but do not differ from the
other sete, which are all plain and sigmoid.

Nephropores in front of sete 3 and 4.

INTERNAL CHARACTERS.

Body-wall offers no prominent characteristics, except that the peritoneal layer
is enormously developed in all somites posterior to xiii. Peritoneum is strongly vas-
cular, but in addition to the usual small peritoneal cells we meet with a thick layer
consisting of a single row of tall peritoneal cells of varying but fairly even height.

Peritoneal cells are in places as high as the other layers of the body-wall com-
bined, while in other places they are shorter. Similar peritoneal cells cover also the
septa, principally those posterior to xiii. The center of the septa bear as a rule the
tullest cells (figs. 149 and 153). These cells show a round nucleus and a granulation
which resembles that of chloragogen cells, but which does not stain deeply as does
the one of the latter cells.

The hearts in x, xi and xii, as well as the septal glands in vii and viii, are also
surrounded by similar cells. The dorsal blood vessel again is covered by regular
chloragogen cells.

Septa. Those separating respectively somites v to xii are slightly thickened
and very strongly cupped, but not covered by any peritoneal cells. The septa sep-
arating xii to xiv are less cupped, not thickened, but all covered by a few peritoneal

144 CALIFORNIA ACADEMY OF SCIENCES.

cells. The septa posterior to somite xiv again are not much cupped, but each one is
lined on each side by a single layer of very thick tall peritoneal cells (fig. 153).

Alimentary canal. Pharynz is furnished with a large upper chamber and is
apparently only developed superiorly, although it possesses a small subpharyngeal
gland close to the ventral nerve cord.

(Hsophagus rises diagonally upwards and joins the single gizzard situated in
v. The tubular-intestine extends to xiy. The sacculated-intestine commences in xv.
There is a typhlosole in the dorsal wall of the intestine in xvi to xix. The typhlo-
sole presents a network of fibres resembling in a general way the structure of a
sponge. This typhlosole does, however, not descend into the canal, but partakes
more of the nature of a wide continuous blood-sinus.

Salivary glands. The suprapharyngeal glands form a mass with five distinet
lobes, of which the posterior one, as usual, is the largest and the anterior one the
smallest (fig. 149). There is also a subpharyngeal gland very low but rather long
(fig. 149). A thin but wide septal gland is found in vi, posterior to the gizzard,
while smaller septal glands, which are principally developed ventrally are found in
vu and yi.

Spermathece consist of two pairs of long and narrow organs in vili and ix
opening in the intersegmental grooves between vi and yiii and viii and ix. In my
specimens they were probably rather undeveloped and did not show any trace of
diverticula either externally or in the wall.

Testes are in x and xi and ovaries in xiii. The oviduct opens between the
setee and septum. Spermiducal funnels or rosettes are very small, thick and compact,
and situated in x and xi. The spermducts run separately backwards between the
longitudinal layer and peritoneum and open jointly on the center of xviii as usual.

Prostates are in four pairs as has been already stated, two and two opening in
each pore in line with sete 1 and 2, these sets, however, not being present in these
somites (fig. 150). The prostates showed no glandular part, the whole being muscu-
lar (fig. 154). They were very thin, tubular, the two prostates in each twin couple
running entirely parallel and close together along the septa, as far as the line of sete
Sand 4, A large part of this distance the muscular part is surrounded by regular
peritoneal cells, not by the large glandular ones. Each prostate remains separate
from the other and even their external pores, though situated close to each other, are
not strictly joined, though they are surrounded by a common thicker lip (figs. 150
and 151). I have already referred to the duplication of prostates in one species of
Kerria, otherwise it has not been found with certainty in Acanthodrilidee. Some of the
earliest described Acanthodrilides were, however, supposed to have a prostate and
spermducts open jointly, and it does not seem unreasonable to suppose that in some
case at least a duplicate prostate existed, and one was mistaken‘for a spermduct. This
would be quite easily done in specimens poorly preserved, especially if the prostates
should be entirely muscular as in Acanthodrilus Vasliti, in which species they are
also very narrow and thin and not really wider than the spermducts of many forms.
Ude, in describing Geodrilus singularis from Danyille, Illinois, mentions as one of the

PACIFIC COAST OLIGOCH ATA, 145

generic characteristics four pairs of prostate glands, but does not describe nor figure
these in detail. The only way I can understand the presence of four pairs of pros-
tates opening into four pores, is that two and two prostates open together. But on
the following page (70) we are told that there are two pairs of prostates in segments
18 to 22, which open in segments 17 and 19. The figure shows us four prostate pores.
If the first statement is not a misprint, we would in @eodrilus singularis have an
analogy similar to what we find in Acanthodrilus Vasliti. But Beddard states in his
large monograph on Oligochieta that @eodrilus singularis is probably identical with
Diplocardia communis. This could, of course, not be the ease if the prostates were
duplicated in the former.

Tubercula pubertatis. Although no large and prominent elevation in the
genital region is found, I have, however, satisfied myself that a tubereula pubertatis
is really present even in my undeveloped specimens. Adjoining the prostate pore is
a small tubercle, consisting of some tall supporting cells surrounding a bundle of sense
cells, the latter characterized by the usual long, oval nuclei. These nuclei are
situated much deeper than they are in the sense cells of the epidermis or in the buceal
cavity, and they are also narrower than those. Otherwise this organ shows a very
great resemblance to those in Benhamia, except for the absence of the glandular
refractive cells. At the base of the sense cells in the tuberele I find numerous smaller
round nuclei, the relationship of which my sections do not fully explain. Numerous
nerve fibrils are seen to connect with the sense cells. Although no clear glandular
cells are seen around the sense-organ in the tubercle, it may be possible that some
may develop later at the same time as the clitellar cells, none of my specimens pos-
sessing any.

Vascular system. The last heart is in xi. The dorsal vessel is single; no
subneural vessel. A large blood sinus in xvi to xix inclusive.

Nephridia are strictly paired, opening in front of sete 3and 4. Those posterior
to somite xiv are surrounded by large peritoneal cells, while those anterior to xv con-
sisted of the usual narrow ducts, free of any ccelomic peritoneal cell mantle. Numer-
ous blood vessels cover the nephridii. The most anterior nephridium is found in iii.
No pepto-nephridium. The anterior nephridia gradually diminish in size towards the
anterior part of the body.

ALEODRILUS 7. gen.
Figs. 66 to 86.

Derinition. Acanthodrilide. Paired meganephridia, not alternating. Pros-
tate pores, two pairs; one pair in wx and one pair in xxii. Svermiducal pair of pores
inva. No caleiferous glands; no penial sete. Two pairs of spermathece in viii and
ix. Olitellum wiii-ax. Two gizzards.

General remarks. I possess only one single specimen of this interesting oli-
gocheeta, collected at Ensenada de Todos Santos in the northern part of Baja Califor-
nia. My time for searching was very limited and the season unfavorable, and this
will account for the want of specimens. The single specimen was found in the dry

146 CALIFORNIA ACADEMY OF SCIENCES.

bed of the river south of town at a water-hole in the otherwise dry creek bottom
sand. It occurred here with Deltania and Limnodrilus. The sand was merely moist
on account of overlying rubbish and sacks. The species is undoubtedly a native one
and the only Acanthodrilid found on the coast so far north in the open ground, and
and on this account even of geographical interest. The intestine of the worm was
gorged with the coarse white sand of the river bed. The anterior part of the worm
was cut lengthwise, one-half dissected and the other half sectioned crosswise. The
want of specimens made a full investigation impossible, though I believe none of the
important points remains in doubt. I have no reason to believe the species is scarce,
though probably it issharing the fate with all native worms, that of being displaced by
European importations. The species is dedicated to Professor W.S. Keyes, my com-
panion in many travels in tropical Mexico.

Affinities. It is interesting to note that Aleodrilus shows considerable affinity
to the only other North American genus of this family, viz.: Diplocardia. It re-
sembled this genus by having two gizzards, no calciferous glands, meganephridia, no
sacs with penial sete. It resembles Benhamia in having two gizzards, ete., but it
differs from these two, as well as from all other Acanthodrilide, by the far backward
position of the prostate and spermiducal pores, these being in xx, xxi and xxii re-
spectively, while all other genera of this family have these respective pores in xvii,
xviii and xix. The genital male pores are thus in Aleodrilus pushed three somites
further back. Considering these and some minor characters I believe I am_ justified
in placing this worm in a new genus.

Aleodrilus Keyesi n. sp.

Derrinition. Length 7 em., by 5 mm. wide; number of somites 80. First dor-
sal pore vitirix, Clitellum complete in anterior, incomplete in the posterior somites,

brititxx. No penial sacs and sete. Common sete paired, those of the inner couple

closer than those of the outer couple. Spermathecal pores between vit/oiii and viti/ix.
Gizzards inv and vi. No caleiferous glands. Nephridia not covered by a cwlomic man-
tle. Nephropores outside of sete 4. Hearts in x, wi, avi. Sperm-sacs racemose in x,
vi, vii. Testes ina, vi. Color pale flesh, no pigment. Habitat, Northern Baja Cal-
ifornia, at Ensenada de Todos Santos.

EXTERNAL CHARACTERS.

Color is very pale, mottled and marbled, showing clearly the intestines and
blood vessels. When collected this worm resembled in delicacy of color and trans-
parency Deltania elegans and I supposed it to be this species. Spermathecal pores are
separate in front of setee 1 and 2 between vii/viii and vili/ix. Setw are ventral and
lateral, 8 in each somite approached in couples. The sete of the inner couple is
closer than those of the outer couple. The distance between the couples is about twice
as large as the distance between sete 1 and 2, and one and one-half us wide as the dis-
tance between sete 3 and 4. No penial sete in special sacs. All setee are sigmoid
without sculpture. The anterior five somites are two-ringed, that is with a single
groove in the equatorial region, while all the following somites are four-ringed, or

PACIFIC COAST OLIGOCH®TA, 147

with two parallel grooves in the equatorial region. The posterior somites are very
wide and four-ringed, but the segmentation is irregular,
Ovipore a little interior to seta 1; nephropores outside of seta 4.

INTERIOR CHARACTERS.

Body-wall. Thereisa special zone of sense organs in the anterior somites as in
Benhamia, but they are more scattered than in this genus. The longitudinal muscles
run irregularly, with no trace of bipinnate arrangement. The strands are rather
narrow. The longitudinal layer is very narrow, especially in the clitellum. The
transverse layer consists of only 3 to 4 strands. The hypodermis offers nothing of
special interest.

The longitudinal layer in the clitellum is greatly diminished in thickness, most
so in the lateral and dorsal region of the clitellum. Anteriorly in somites ii and iii,
the longitudinal strands leave the body-wall and spread out fan-shaped to the inner
wall of the prostomium, forming retractor muscles for the upper and lower lips.

Arciform muscles. In somites xx, xxi and xxii we find on each side of the
ventral nerve-chord several oblique or aciform muscles, running from the region of
the copulatory grooves to the region above the lateral sete, thus serving to depress
and relax the two grooves. These muscles are confined to a single row, and do not
show a complex arrangement, as is so frequently shown in oligocheeta.

Septa. The septa between somites vii and xiv are much thicker than the others,
especially thickened are those separating somites from vii to x. Those between x and
xiii are less thick than the anterior ones. The thickest septa, vil/vili, vili/ix, ix/x, are
much thicker than the body-wall. The most anterior thick septum is the one which
posteriorly bounds the gizzard (fig. 74).

The septum next anterior to this, the one which separates the two gizzards,
presents the peculiarity of not being attached to the body-wall, between yi and
vii, but it extends forward parallel with the intestine and passes in front of the brain
on the upper side, while the ventral side is attached to the cesophagus below the
pharynx. It forms thus a sac, as in various species of Benhamia. Anterior to this
septum I find no trace of others.

Alimentary canal. The pharynx is well developed and superposed by a very
large glandular mass, which consists of about six layers of lobes, attached to muscular
strands, as usual. The most posterior mass is the thickest. The discha rge pockets of
these glands into the pharynx are much thicker than any I have seen in other species,
but are otherwise not of any characteristic construction.

Septal glands are situated far back in somites vii to xi. They are of the same
nature as those which discharge in the pharynx, but I have good reasons to believe that
the glands in this species discharge in the tubular intestine. I have been able to
follow the discharge duct as far as to the muscular layers of the intestine, which would
hardly have been the case if the ducts had continued forwards into the pharynx, as
do those of the forward septal glands in many genera. A peculiarity of these glands

is that they are especially developed on the ventral side of the intestine (fig. 75) and
Memoirs, Vou. II, 5. January 6, 1896.

148 CALIFORNIA ACADEMY OF SCIENCES.

are distinctly paired. On the median line below the intestine the ends of each
glandular mass meet and join into one duct, the one I have just referred to. The free
ends of the glandular masses are attached by mesenteric tissue to the ventral or sub-
intestinal blood trunk.

A much smaller tubular gland of the same nature runs between the dorsal
vessel and the intestine; discharge duct unobserved. A yet smaller gland is seen
above the subventral vessel in the same somite as the former; its discharge duct could
not be followed.

These glands stain exactly as the common supra-pharyngeal glands and septal
glands, but they show no similarity as regards reagents with the chloragogice cells of
the intestine and blood vessels, in corresponding places in the somites posterior to x1.
The septal glands appear to be of about equal size, a close-examination being im-
possible, from want of sufficient material.

Other glands are found in the epithelial walls of the intestine, arranged in
clusters, like the cloves ina garlic. They are scattered about at short intervals among
the epithelial cells, and appear of the same nature as those I have described in Ar-
gilophilus, but they are not as numerous as in that species. They do not stain freely,
but stand out bright and pellucid among the darker staining cells.

Typhilosole not present.

Gizzards are connected by a very thin wall of the same nature as the cesophagus.
As far as I can make out the gizzard must be in v and vi, at least the posterior gizzard
is bound by the septum separating viand vil. The circular muscular layer is about
30 strands wide, and is at the widest place about four times thicker than the epithelial
layer and cuticle together. The whole width of the gizzard wall is little more than
twice that of the body-wall in that somite. The longitudinal muscular layer of the
gizzard is only one single strand thick, and the thickness of this strand is less than
any one strand of the circular layer of the gizzard. The epithelial layer is com-
paratively thick, about one-fifth of the whole gizzard. It contains the same peculiar
glands as I figured in Benhamia.

Spermathece. The absence of diverticula is interesting in as much as most
species of related genera possess them. There are, however, some warty elevations.
The muscular or basal duet is very long, slender and tubular, several times longer
than the upper ovoid sae (fig. 74). The muscular part offers no peculiarities of
structure. The spermathecze occupy each only one somite. They stand upright fol-
lowing close to the anterior surface of the septum.

Testes are greatly lobed and are situated high up on the septum, just as are the
ovaries. :

Spermducts and ciliated rosettes. The spermducts are separated, but enclosed
in a common muscular sheath until somite xxi is reached. Between xx and xvi the
two ducts fuse into one lumen, which opens out into the center of somite xxi. The
double lumen runs along the circular muscular layer forward until somite xiii/xu,
when the respective ducts rise upwards following the septum to the ovary and testes.
From here on forward each duct is thicker and muscular, and instead of following the

PACIFIC COAST OLIGOCH ATA. 149

body-wall passes straight through the ccelomic cavity to the anterior septa, which they
pierce immediately at the base of the ovaries and testes (fig. 78). A double lumen
is seen only after it passes posteriorly to the ovary. The rosettes are not large,
but thick with a wide base; the latter is furnished, where it passes through the sep-
tum, with several small sac-like glands, each with a distinct lumen, which I followed
through the muscular layer of the spermduct and which I suppose empties into the
neck of the rosette (fig. 79). The rosettes are not enclosed in the sperm-sacs.

Sperm-sacs. The three pairs of sperm-sacs are racemose, but not exceedingly
so. There are four or five large lobes of globular shape seen in every section sus-
pended from the septum and on the under side of the intestine, though somewhat
projecting above it. The two anterior sperm-sacs in x and xi are smaller than the
posterior one in xii, the one in x being the smallest of the three pairs. Those in
x and xi are suspended from the anterior septum separating x/xi and xi/xii. Of the
position of the posterior sperm-sacs I am uncertain, but it appears suspended from
the posterior septum, the one separating xii/xiii, as far as I can judge from a eross-
section.

Spermatogonia. In the two anterior sperm-sacs the spermatogonia offered
nothing peculiar as regards the development of the spermatozoa. There is in each
sperm-sac a large central zone of peculiar cells, staining differently. They re situ-
ated very close together, and possessed in my preparation rather shrunken nuclei
(figs. 63 and 75). This zone exists in all the various sperm-saes in the anterior as
well as in the posterior ones. The development of the spermatozoa in the two an-
terior pairs appeared entirely normal, the spermatogonia possessing the same form as
in other species, the large nuclei standing out freely like beads above the wall. But
in the posterior sperm-sacs the spermatogonia, one and all, looked quite differently.
They were here of many varying sizes, some small, some enormously large, and the
nuclei, instead of standing out from the wall of the spermatogonium, were always
bunched in the center, or strung across it as a central band (75). In other spermat-
ogonia the nuclei were arranged as ina ring along the cell walls, but without pushing
out. When the nuclei were in the center there appeared always a row of large and
small vacuoles along the cell-wall. I find two sizes of nuclei, the smaller being al-
ways less in number, and about four times smaller than the large nuclei. When
counter-stained with orange, the larger nuclei give quickly up part of their heema-
toxylon, the smaller ones giving it up slowly and not at all. The eytoplasma of the
spermatogonia presented a very strong polarity as regards its position, it being always
massed towards the cell-wall nearest the central germinative area (fig. 63). A some-
what similar-process of developing spermatogonia have been described by Vernon in
the sperm cells of Bombyx. The central germinative cell in Bombyx appears to
correspond with the central area or cell agglomeration in Aleodrilus, but I haye seen
no such budding out of the germ cells as figured by Vernon, but this may depend
on the insufficient material at my command.

A large number of similar spermatogonia were found in very large numbers
in nearly all the anterior somites, either loose in the ccelomic cavity, or in a

150 CALIFORNIA ACADEMY OF SCIENCES.

peculiar tissue which fills much of the ccelomie cavity. This tissue consists of large,
more or less connected cells, very much like the ccelomic glandular cells surrounding
so many nephridia. These cell masses are more or less diffuse, and appear not dis-
tinctly connected with any of the interior organs of the body. Among these cells
are found scattered about large masses of spermatogonia, all in about the same stage
of development. There are also numerous small free ccelomic cells, such as found in
all earthworms.

Nephridia. There is one pair of mega-nephridia in each somite. The
uephridium is very large, projecting above the lateral line of the body. The nephro-
pores—at least some of them—are seen outside of or more dorsal than sete 4. The
nephrostome again is as usually seen very near sete 1. There are no glandular
ceelomic mantles on any of the nephridia. The anterior fold is much wider then the
posterior one. The windings are very deep and twisted, and the spur generally sigmoid.
The outlet duct, which is tapering towards the pore, is much darker than the other
ducts. Its prolongation into the anterior fold forms there the central wider canal,
which is also darker than the other two. The canal in the bridge continues forwards
and upwards through the anterior fold, and is there the posterior one of the three
canals.

Vascular system. The ventral and dorsal longitudinal vessels are both single.
There are connecting vessels in somites vii to xii. In x, xi and xii, these vessels are
very large, and take the form of so-called hearts. Each heart consists of four or five
links, increasing in thickness upwards. Between each two links there isa thick
circular valye. Similar valves are seen also in the dorsal vessel at the junction with the
hearts. At the base of some of the valves are seen two rows of very large cells, the
nuclei of which are about 3 to 4 times larger than the nuclei in the valve cells (fig.
83). All the valves in the hearts point downwards or ventrally. There are no
glandular cells in the vessels, such, for instance, as are found in Pontodrilus, ete.

Posterior to the hearts we find long, tubular connecting lateral vessels, between
the dorsal and ventral trunks, in the anterior part of each somite. These vessels, one
pair in each somite (fig. 86), are of even thickness throughout, but with two short
knob-like diverticula, one above the other, about equal distances from each other and
from the longitudinal vessels. Both the dorsal vessel, as well as the laterals, are
thickly covered with coelomic chloragogen cells of a yellowish opaque color. These
cells do not cover the ventral longitudinal vessel. These lateral vessels contain no
valyes.

Each valve consists of several circular rings, each containing a number of
muscular strands enclosed within a common membrane. In longitudinal sections of
the vessel most of the nuclei lie parallel with each other, but the outside ones run as
the periphery in a circle. The smaller valves consist each of only one such lobe (fig.
73a), while the large ones are composed of several (fig. 73)).

PACIFIC COAST OLIGOCHRTA. 151

SPARGANOPHILUS Benham.

Drrrnrrion. Aquatic oligochata. Tight sete in four couples. Prostomium
not marked off from peristomium, but Surnished with a superior pit. Few dorsal pores.
Clitellum very large, from eight to twelve somites. No penial sete. Spermathecw in
bu, vii and ix, from two to eight in each somite. Sperm-sacs in xi and xii, racemose.
Speemduct always subepidermal. Male pore xvii/ix, or anterior part of xix. Pros-
tates generally present in three or four pairs situated several somites posterior to the
spermiducal pores. No gizzards, no caleiferous diverticula, no typhlosole. Four pairs
of hearts. Two pairs of lateral, longitudinal integumental vessels extending forward
From somite wiv, not connecting anteriorly with the gut-wall and median trunks.
Nephridia meganephric, commence in somite wii or wiii.

Principal species characters are derived from the position of the sete; number
and shape of spermathece in each somite; lobulation of the sperm-sacs; course of the
spermducts; shape and position of tubercula pubertatis, whether dorsal or ventral, to
the male pore or spermduct; extent of clitellum; presence of a subpharyngeal integ-
umental gland in iii; length of worm and number of somites, ete.

GENERAL REMARKS.

The first species belonging to this genus was described by Benham from the
river Thames in England. Later, Frank Smith described another species from North
America and it became doubtful, as first suggested by Benham, if the genus was
originally an American or European one. The very restricted locality or habitat of
the European species would indicate its probable importation from some other country,
and when the American species was found it became almost certain that we had to
search for the original home of this genus on the American continent. Already
When Benham’s paper was published [ had in my possession specimens of this genus
from Guatemala, California and the Central North American States, and there
remained no doubt in my mind as to the native habitat of the genus. As it now stands
we have seven species sufficiently well defined to be recognized and one more of which
no detailed description can be had, but of which we know enough to be able to
recognize it should it again be observed. Of these eight species then seven are
American and one European, and everything points to the probability that the latter
one is a lately imported species to England most likely from this country, as Ben-
ham originally supposed.

The species described here below are not. of equal value as species as might
be naturally inferred. Four of the species are well defined, which principally
is due to better preserved and abundant material for study. These species are: Spar-
ganophilus tamesis, Biseni, Benhamiand Smithi. Two are less well-known, due entirely
to want of sufficient well preserved material. These species are Sparganophilus car-
neus and guatemalensis, both of which may prove only varities of Sp. Benhami. The
remaining species Sparganuphilus sonome may prove a variety of Sparganophilus
Smithi, but just as such all the more interesting.

A point of unusual interest in this genus is the presence of prostates, or, as

152 CALIFORNIA ACADEMY OF SCIENCES.

Beddard calls them, spermiducal glands. These glands are, in Sparganophilus, situ-
ated much further from the male or spermiducal pores than in any other genus,
apparently quite independent of the pores. They are also subject to considerable
variation, in some species being three, in others four; in one species none. Of the
same nature I consider the forward parietal pair of glands in somite iii of Spargano-
philus Eiseni, and it seems not unlikely that originally this genus possessed many
more pairs of spermiducal glands, perhaps one in eyery somite. This location of the
spermiducal glands favors greatly Beddard’s view that these glands were originally
independent of the spermduct, and according to this view these glands are in
Sparganophilus the most primitive of any. There is also much difference as to the
development of these glands in the various species. Thus in Sp. Benhami the gland
consists of two distinct parts and is both glandular and muscular, while in Sp. Smithi
the muscular part is degenerate or undeveloped, the whole gland being very dimin-
utive: Another point showing the primitive arrangement of the spermiducal glands
is the absence of any copulatory sete, the common sete in their yicinity being
unmodified.

There is no doubt that a large number of species of this genus will soon be
found on the American continent, especially as specimens appear numerous and widely
distributed. In Guatemala I found them everywhere in springs and lakes, and a
hasty examination of live specimens satisfied me that there were among them about
three species, characterized by the position of the hearts, whether in viii—xi, ix—xi, or
x-xill. Unfortunately my Guatemala collection was destroyed and only few speci-
mens in poor condition remain.

In this connection I will also call attention to another species of Sparganophilus
not described below, but found by me several vears ago in the small lake known as
Laguna Puerea, situated near the ocean at San Francisco, California, or in the same
lake as I now find Sparganophilus Smithi. This species which [ have sinee been
unable to re-collect, although I have made repeated excursions to the place and dug at
the identical spot, offered much of interest and was undoubtedly a different form
from any now described below. It was much shorter and thicker, about 5 em. long
by 4 mm. thick, or about half as long and twice as thick as our smallest species now
known. The hearts were, according to a fieldnote, situated in vii, vili, ix, x. The
longitudinal lateral vessels began in xiv, but most interesting of all, the species
possessed five distinct and large eve spots on the first somite and prostomium, an
occurrence not recorded in any other of the higher earthworms.

The specimens were very rare—I could only find two or three in an hour; they
occurred in the clay soil near the shore at shallow depth.

In connection with this it is most interesting to note that at that time I found
no trace of Sparganophilus Smithi, which now occurs in the same lake in countless
numbers, a worm many times longerand much thinner. It is impossible that I could
be mistaken in regard to my former observations. My explanation is this: Formerly
the Laguna Puerea was much lower, the bottom soil near the surface was clayey, while
now the lake is several feet higher, the bottom soil near the surface is sandy. A

PACIFIC COAST OLIGOCHAHTA, eR:

number of vegetable gardens have of late years been established at the headwaters
along the ereek furnishing the lake, and it is probable that Sparganophilus Smithi
has been introduced with watercress, ete., from some other locality within the last few,
say ten years, while the eye-bearing species may yet be living in the bottom soil.

KEY TO THE SPECIES OF SPARGANOPHILUS.

I, Sete 3 and 4 ventral; no prostates, spermducts dorsal to tubereula pubertatis.
SPARGANOPHILUS TAMESIS Benham.
If. Set 3 and 4 dorsal, prostates present, situated several somites posterior to the male pores.
A, A pair of ventrally situated parietal glands in somite iii. Spermducts and spermiducal pores dorsal to
tubercula pubertatis. SPARGANOPHILUS Eisent Smith.
B. No ventral parietal gland in any somite.
x. Spermiducal or male pore ventrally situated to the tubercula pubertatis.
a. 8 spermathece in each of somites vii, viii, ix. 2 ditto in vi. SPARGANOPHILUS SMITHI n. sp.
b. 4 spermathecw in each of somites vii, viii, ix. 2 ditto in vi. SPARGANOPHILUS SONOMZ ni. subsp.
xx. Spermiducal pore dorsally situated to tubercula pubertatis.

¢. Spermathecs trowel-shaped, outline crenulated. SPARGANOPHILUS BENHAMI n. sp.

d. Spermathece trowel-like, but outline smooth. SPARGANOPHILUS GUATEMALENSIS 0. subsp.
xxx, Spermiducal pore unknown, but probably dorsal to tubercula pubertatis

e. Spermathece club-like, apical end globular, smooth. SPARGANOPHILUS CARNEUS 0. sp.

Sparganophilus tamesis Benham.

Derinition. Length 7to 10cm. Clitellum 1-2 av—1-2 xav. Tubercula puber-
tatis xvii to xxii ventral to spermiducal pore. Setw 3 and 4 ventral. Spermiducal pore
between voi and winx dorsal to tubercula pubertatis. Sperm-sacs in vi and xii minutely
lobulate. Spermathece one pair in each of somites vii, viii, ix, ventral. No prostates.
Anterior nephridium in xiii. Septal glands in v and vi. No subpharyngeal integu-
mental gland. Blood capilaries on nephridia are numerous. Continuous blood sinus in
the sacculated intestine. Hearts in viii, ix, x, xi, not Jilling the cwlom. Color pinkish
with violet iridescence. Habitat, England, Thames. Description is compiled after
Benham’s paper only, as I have not seen any specimens.

Sparganophilus Eiseni Smith.

Derinition. Length 18 to 20 cm.; width 26 em. Clitellum dorsally 1-4 xv—
wun, ventrally 1-2 aiv—xvavi. Tubercula_pubertatis 1-2 vvii-1-2 xxii, ventral spermathecal
pores. Sete 5 and 4 dorsal. Spermiducal pore in anterior part of xix, dorsal to tubercula
pubertatis. Sperm-sacs minutely lobulate. Spermathece, one pair each in vir, viii and
ww, all dorsal. Prostates 4 pairs in wxaiti-avi; muscular duct present. Anterior
nephridia commence in wiii. Septal glands in iv, v, about the same size, while the one in vi
is very small. A subpharyngeal parietal pair of glands present, opening in front of
setw 1 and 2 in somite tii. Blood capillaries on nephridia are many. Continuous blood
sinus in sacculated intestine present. Hearts in viii, ix, x, wi small. Color violet flesh.
Habitat, Itlinois, North America.

Prof. Frank Smith has kindly sent me for comparison a full set of sections of
this species, also several mature specimens in alcohol and formalin. As, however,
Prof. Smith writes me that he intends to further work out the details of the anatomy
and histology of this species, I have not considered myself at liberty to more than

154 CALIFORNIA ACADEMY OF SCIENCES.

touch upon a few essential points of importance in further characterizing this species
from its other American allies. The species is nearest related to Sp. Benhami and its
subspecies. The main points of difference and similarity may be found in the com-
parative table of the species of Sparganophilus. Sp. Benhami is a much more slender
form than Sp. Fiseni, especially as regards the body posterior to clitellum. The ven-
tral side of the genital regions in the two species differ. Thus in Sp. Benhami the
ventral depressed area between the tubercula pubertatis and the oviducal pores is
circular or oval, while in Sp. Hiseni it is contracted in the middle, the shape of a sand
time-glass. The presence of a small parietal gland in somite ii in Sp. Hiseni
distinguishes this species from all others known so far. This gland is paired, extends
considerably into the ccelomic cavity, is racemose and opens to the exterior in front
sete 1 and 2, and projects its free apex back through septum into iv. Another
character is the small size of the septal glands in yi, the corresponding pair in
Sp. Benhami being larger.

Sparganophilus Smithi n. sp.

Derinition. Length 20 em. by 3 1-2 mm. wide; number of somites 185. Dor-
sal pores, the most anterior one between i/ii, the most posterior one between vii/xiii. Clhi-
tellum dorsally 1-2 vvi-wxviit, ventrally via—vxv. Tubercula pubertatis xia—vaeori, dorsal
to spermiducal pores. Sete 3 and 4 dorsal. Spermiducal pore between xviii/xix, ventral
of tubercula pubertatis. Sperm-sacs in wi and ati. Spermathece 4 pairs in each of
somites vii, viti, ix, and 1 pair in somite vi, slender, dorsal and lateral. Prostates 3 pairs
in xvaii, waiit/naiv. No muscular duct. Anterior nephridium xvi. Septal glands in
iv, v, vi and vii about equal in size. No subpharyngeal parietal gland. Blood capillaries
on nephridia numerous. Blood sinus in sacculated intestine not continuous. Hearts in
vin, iv, xv, vi, small. Color violet flesh, somewhat iridescent. Habitat, San Francisco,
California.

Sparganophilus sonome n. subsp.

Deriition. Length 20 em. by 3 1-2 mm. wide; number of somites about 200.
Dorsal pores. Clitellum dorsally xvii—wavin, ventrally xvi-waix. Tubercula pubertatis
vix—vavii, dorsal to spermiducal pore. Sete .32 and 4 dorsal. Spermiducal pores
between xviii/vix, ventral of tubercula pubertatis. Sperm-sacs in xt and xii. Sperma-
thece, two pairs in each of vii, viii, iv; one pair in vi, not slender. Prostates 3 pairs
in wxit, vxiv, vxv, no muscular duct. Anterior nephridium in avi. Septal glands in
iv, v, vi, vii, about equal in size. No subpharyngeal parietal gland. Blood capillaries
on nephridia numerous. Blood sinus in saccuiated intestine not continuous. Hearts
in vii, iv, x, wi, small. Color violet flesh, less iridescent than Sp. Smithi. Habitat,
Sebastopol, Sonoma County, California.

Sparganophilus Benhami n. sp.

Derinition. Length 17 em. by 2 1-2 mm. in the posterior part of the clitellum;
namber of somites about 250. No dorsal pores. Clitellum, dorsally xv—wxiv; ventrally
xvii-vavi. Tubercula pubertatis xviii-vxii, ventral to spermiducal pores. Sete 3 and 4

é

§ fa ee Sas
ON: <A Ba
| 7

Sete Sang

Clitellum.|

Tubercula FE >

a \

Spermathe,
~

— Sperm-sacs

| Prostates .

a! Anterior i
ce

’ Capillaries
body-wall

-
r
x
+
= Re:
a
=

Table B.

Habitat

Length and width...

Sete 8 and 4

COLT Pyare critic crc BOP

Tubercula pubertatis. .

Spermiducal pore..

Spermathece .

Sperm-sacs .

Prostates

Anterior nephridia
commence in....

Septal glands in .

Parietal celomic
gland iniii....

Blood sinus in the
sacculated intestine... .

FL EG LB) Ss on cites
Capillaries on the ne-
phridia and in the

body-wall

England, Thames.

7 to 10 cm.
Ventral.

4xv-hxxv.

xvil—xxii, ventral to

spermiducal pore.

Between xyiii/xix, dor-
sal to tubercula puber-
tatis; so are the sperm- |
duets.

Slender; one pairin each
of somites vii, viii, ix; |
ventral in line with |
3 and4.

In xi and xii, minutely
lobulate.

Not present.

xiii.

iv, v, vi; the posterior
are the largest.

Absent.

Continuous.

viii, ix, x, xi; not filling
the ecelomie cavity.

Very numerous.

Eiseni Smith.

| Illinois River, U.S. A.

18 to 20 cm. by 2 mm.
Dorsal,

Dorsally {xv-xxv;
ventrally 4xiy—xxvi.

dxvii-}xxii, ventral to
the spermiducal pore
and to the spermducts.

On anterior part of xix,
dorsal to the tubercula
pubertatis; so are the
spermducts.

One pair in each of so-
mites vii, viii, ix, dor-
sal in line with 3 and
4; slender.

In xi and xii, minutely
lobulate.

Four pairs in xxiii-xxyi.
xiii.

iv, v, vi. Those in iv
and y about equal size;
the one in yi is very
diminutive.

One pair of glands open
in front of ventral sete
in iii; distal end ex-
tending into iy.

Continuous.

In viii, ix, x, xi not fill-
ing the celom,

Very numerous.

|
|
|
|
|
|
|
|

COMPARATIVE CHART OF SPECIES OF SPARGONOPHILUS.

To face page 154.

Benhami un. sp.

Tepic, Mexico; 4,000.

17 em. by 24 mm.
Dorsal.

Dorsally xv-4}xxv;
ventrally xvii-xxvi.

xvili-xxii, ventral to
spermiducal pore and
to spermduct.

On anterior part of xix,
dorsal to tubercula pu-
bertatis.

Robust, trowel-like, with
wavy outline; one pair
in each of somites vii,
viii, ix; dorsal.

xi, xii, less minutely
lobulate.

Four pairs, one each in
xxiii-xxvi; a long mus-
cular basal duct.

xii. The first two pairs
smaller; the first large
pair in xiv.

iy, v, vi. About equal
in size.

None.

Continuous.

viii, ix, x, xi, not filling
the celom.

Very few.

guatemalensis n. subsp.

Guatemala.

10 cm. by 2 mm.
Dorsal.

xvi-xxvi.

carneus n, subsp.

Mississippi River, Iowa.

9 cm. by 2 mm.
Dorsal.

Dorsally xv-xxiv;
ventrally ?

Parallel ridges, xviii-xxii

Robust, trowel-shaped,
with smooth outline,
in vii, vili, ix.

Four pairs in xxiv, xxv,
XXvi, xxvii.

viii, ix, x, xi, not filling |
the celom.

Dorsal, one pair in each
of somites vii, viii, ix.
Apical end globular,
not trowel-like.

xi and xii, lobulate, but
not minutely so, as in
Sp. Hiseni.

| a .
Present; four pairs in

xxiii, xxiv and prob-
ably in xxy and xxvi.

iv, v, vi; the one in yi
the smallest.

None.

viii, ix, x, xi, completely
filling the celom.

Numerous and large; on
the body-wall very nu-
merous in the six an-
terior somites.

|
|
|
|
|
|
|
|
|
|
|
|

Smithi n. sp.

Laguna Puerca, San
Francisco, California.

20 cm. by 34 mm.(or less)
Dorsal.

Dorsally 4xvi- xxviii;
yentrally xix-xxy.

xix-xxvii, dorsal to

spermiducal pore.

Between xyiii/xix, ven-
tral to tubereula pu-
bertatis.

Slender; four pairs (8)
in each of somites vii,
viii, ix, dorsal and lat-
eral; one pair only in
vi dorsal.

xi and xii, more lobulate
than in Benhami, but
less so than in Eiseni.

Three pairs, one each in
somites xii, xiii, xiv;
no muscular duct.

xvi.

In iv, v, vi, vii; about
equal in size.

None.

Not continuous.
viii, ix, x, xi, quite small,
not filling the celom.

Numerous, giving the
nephridia a pink color.

sonome n. subsp.

Sonoma Valley, California

25 to 30 cm. by 3} mm.
Dorsal.

Dorsally xvii-xxviii;
ventrally xvi-xxix.

xix-xvill, dorsal to
spermiducal pore and
to spermduct.

Between xviii/xix, ven-
tral to tubercula puber-
tatis.

One pair in yi; two pairs
(4) in each of somites
Vii, vill, ix; not slender.

xi and xii, more lobulate
than Benhami, but less
than Hiseni.

Three pairs in xxii, xiii,
xly.

xvi.

iv, v, vi, vii, about equal
in size.

None.

Not continuous.

Viii, ix, x, xi, quite small,
not filling the ceelom.

Very numerous.

PACIFIC COAST OLIGOCH #®TA. 155

are dorsal. Spermiducal pore on anterior part of wix. Sperm-sacs in vi and xii, not
minutely lobulate. Spermathecw one pair in each of vii, vivi and ix are dorsal. Sperm-
iducal glands or prostates 4 pairs in waiii-wavi, with a long muscular duct. Most
anterior nephridium in vit. Septal glands in iv, v, vi of about equal size. A pair of
subpharyngeal parietal glands not present. Blood capillaries on nephridia few. A
continuous blood sinus in sacculated intestine. Blood glands many. Hearts in viii, ix,
xv, xt. Color violet, strongly iridescent. Habitat, Tepic, Mexico, 4000 feet.

Sparganophilus guatemalensis n. subsp.

Derinition. Length 10 cm. by 1 1-2 mm. wide; number of somites 260. Dor-
sal pores none. Clitellum xvi-vani. Tubercula pubertatis xviii-vxii, parallel ridges.
Sete 3 and 4 dorsal. Spermiducal pores unknown. Prostates four pairs in wriv, xxv,
wevi, wav. Spermathece one puir each in vii, viii, ix; wide free end, wide, flat, outline
smooth. Hearts very strong. Color deep bluish violet iridescence. Habitat, Guatemala
City.

Sparganophilus carneus n. subsp.

Derinition. Length 9 em. by 2 mm. wide; number of somites 160. Clitellum
dorsally 1-2xv—xxv. Tubercula pubertatis unknown. Sete 3 and 4 dorsal. Spermidu-
cal pores unknown. Sperm-sacs in xi and «ii, lobulate, but less minutely than in Sp.
Hisem. Spermathece one pair each in vii, viii, ix dorsal; free end globular. Prostates
unknown. Septal glands in iv, v, vi, the one in vi the smallest. No subpharyngea]
integumental glands. Blood capillaries enormously large and many in the anterior six
somites. Hearts very large in viii, ix, x, vi. Color reddish flesh, much darker when in
alcohol than Sp. Benhami. Habitat, Mississippi River, near Clayton, Lowa.

After these preliminary definitions I will now describe in detail the various
species of Sparganophilus examined by me.

Sparganophilus Smithi n. sp.
Figs 120-122, 124, 129-39.

Habitat. This species is very abundant in a small lake or pond known as
Laguna Puerea and situated between Lake Merced and Golden Gate Park at San
Francisco. Adult form in June to October, more frequent in the latter month. It
occurs at or near, above and below the water’s edge, the earth being thrown up in
abundance and in heaps. I have already referred to nonpresence of this species
some ten years ago in the above locality.

EXTERIOR CHARACTERS.

Color. A brownish flesh with violet reflex, but much less so than Sp. Benham
or Sp. guatemalensis. In alcohol this species becomes much paler than either of the
two just referred to, almost pure white, looses its irridescence entirely. In formalin
the color is pale, the irridescence very faint, strongest on the clitellum, and hardly
perceptible on any other part of the body.

Length varies some, but generally reaches in fully grown and largest speci-
Memorrs, Vot. II, 5. January 6, 1896.

156 CALIFORNIA ACADEMY OF SCIENCES.

mens 20 em., while the width is about 54 mm. posterior to clitellum. This and the

following species are the longest forms so far known of the genus. The worms are
not remarkably lively when out of the ground.

Sade. The couple 3 and 4 are distinctly dorsal, and it may be of interest to
note that in all species found in America the dorsal portion of these setee appear
characteristic, while in Sp. tamesis the setee 3 and 4 are ventral or sublateral. I can
find no character in the form of the sete in the various species. They are allslightly
sigmoid, slightly hooked and devoid of ornamentation. There are no penial sete and
no modified sete in any part of the genital region. The sete of Sp. tamesis as figured
by Benham are more hooked than those of my new forms.

Prostomium and pygidium. I cannot find any good characters in the form of
these part in the respective species. The small pit in the prostomium of Sp. tamesis,
deseribed by Benham, is found in all the species. The pygidium yaries greatly.
While some specimens have the anal orifice elevated and dorsal others have it central.
The shape and size of the last somite varies also to such an extent that no species
character can be derived from it. I will then in the following not refer to these part
in any of the species described below.

Clitellum. In all the species of this genus the clitellum is very large, oecupy-
ing from 8 to 15 somites. In Sp. Smithi it is located as follows: dorsally, $xvi-}xxviii;
ventrally, xix—xxy. Continuous all around the body. In all preserved specimens
the body is bent towards the ventral side just at the clitellum, and special pains must
be taken if a straight specimen is desired.

Tubercula pubertatis consists of a very elevated, continuous ridge, which is
broken or depressed at the intersegmental grooves. It extends through somites
xix—xxvil and is situated dorsally to the spermiducal duct and its pore. The ridge is
further generally concave on the dorsal and convex on the ventral side. The ventral
area between the two tubercula pubertatis ridges is thus much wider than in any of
the other three new species described here. Anteriorly the tubercula pubertatis
ridge is continued forward in a kind of semicircle which ends at the groove between
somites ix/x. But these anterior ridges are much lower than the tubercula pubertatis
proper, but nevertheless very sharply defined and distinct.

Spermiducal pore is situated between xix/xx, laterally to the sete 1 and 2 and
about three times as far from 2 as 2is from 1. But the pore is ventral to the tuber-
cula pubertatis ridge. This is an important characteristic and only shared with Sp.
sonore. In Sp. tamesis, Kiseni and Benhami, the spermiducal pores are dorsal to the
tubercula pubertatis ridge.

Oviducal pores are plain in front of sete 1 and 2 on somite xiy.

Prostate pores externally not visible, but situated in front of setee 1 and 2.

Somites. All the anterior somites are three-ringed, except i, ii, ili, which are
smooth.

Dorsal pores. The most anterior pore is situated between i/ii, and the most
posterior one between xii/xiil.

PACIFIC COAST OLIGOCH MTA. 157

INTERIOR CHARACTERS.

_ Septa. Thickest septum is found between somites xi/xii. Anterior septa are
gradually diminishing in thickness forward, and similarly the septa posterior to xi/xii
are diminishing in thickness backwards. The most posterior thickened septum, the
one between xiv/xv, is hardly thicker than the one next posterior. The central
part of the thickened septa is very much thicker than the part near the body-wall.
(fig. 127t.m.) The septa correspond more to the intersegmental grooves than do those
in Sp. Benhami. When the body is viewed in cross-section it will be seen that the
septa are principally attached to the body-wall at four points nearly half way between
the sete (fig. 131).

Suprapharyngeal and septal glands. There are dorsal septal glands in iy, vy,
vi, vil, about equal in size, and ventral septal glands in y, vi, vil, also of about equal
size. ‘The latter are much larger than in Sparganophilus Benhami, and reach above
the center of the body (fig. 120). In addition to these glands I find an accessory
septal gland attached to the central anterior surface of septum 1x/x, just above the
intestine and extending downwards. This gland is closely attached at every point to
the septum, and projects only slightly into the ecelomie cavity. In height this gland is
not much thicker than the central part of the septum (fig. 152). Its free surface is very
smooth and even. I could follow ducts with precipitated secretions running along the
septum downwards towards the intestine, but its connection with the latter, if any, I
could not ascertain. Beddard suggests that as the nephridia only commence in the
thirteenth segments the septal glands described by Benham in Sp. tamesis are homo-
logous with the nephridia. In all species of Sparganophilus investigated by me I find
no such mucous glands as in Pontoscolex, only regular salivary glands, which open
into the pharynx, and of the same structure as the supra-pharyngeal glands generally.
These salivary glands in Sp. tamesis occur exactly in the same somites as in all other
Sparganophilus species, viz.: iv, y and vi.

Body-wall offers no other characteristics, except that it is throughout of almost
the same thickness, and not thinner along the dorsal parts of some of the central
somites, as in Sp. Benhami.

Chtellum is continuous all around the body, but it is much thicker dorsally
than ventrally. This thicker dorsal portion commences with the tubercula pubertatis,
gradually increasing in thickness dorsally towards the median line, where it is four

‘times thicker than at the ventral median line between the tubercula pubertatis. This

refers only to the clitellar cells in the central clitellar somites. The long clitellar
cells are confined to the latero-dorsal part, while the ventral part contains only
the short clitellar cells, the point where the former cease being the tubercula
pubertatis. This is the case in all the species examined by me.

Tubercula pubertatis. It has already been stated that they form a continuous
ridge, only broken in the anterior two somites xvii and xviii. Their structure differs
considerably from that of Sparganophilus Benhami. We find the tubercle cells less
differentiated (fig. 138), and more resembling the short clitellar cells, of which they
are only a variety. Another characteristic is the presence of a large blood vessel in

158 CALIFORNIA ACADEMY OF SCIENCES.

the center of the tubercle. The most interesting feature, however, consists in the
presence of three or four arciform muscular bands which connect the opposite sides
of the tubercular projection, and which, of course, serve to further push out the tip
of the tubercle, or to relax it, as the occasion may demand. These bands are entirely
confined to the clitellar cell layer, and penetrate to the very cuticle, to which at least
some of the muscles are attached, while others seem attached to the cells themselves.
Besides these epidermal arciform muscles, there are numerous colomic arciform
muscles, which also penetrate the glandular layers and serve to connect the two sides
of the clitellam on either side of the tubercula pubertatis (fig. 188). While the
arciform muscles of the epidermal layer have been described by Cerfontaine in Lum-
bricus, they are by no means known from many species. Their presence varies con-
siderably in different species of Sparganophilus, but they are especially numerous
and strong in this one.

(Hsophagus and intestine. As regards the general shape of the alimentary canal
I can see no marked difference in the various species so far examined. Again as to
structure I find two points worthy of mention. The chloragogen cells which are
everywhere covering the intestine are much larger and more numerous in this species
than in Sp. Lenhami, covering as they do both tubular and sacculated intestine.
Another point is the absence of a continuous blood-sinus. A continuous blood-sinus
in the sacculated intestine has been described by Benham in Sp. tamesis, by Smith in
Sp. Hiseni and is also found in Sp. Benhami, as will be recorded further on. In this
species, Sp. Smithi, the sinus is not quite continuous, the respective blood lacunes are
quite close and in places run together, but they are nowhere continuous in the same
way as in the species referred to above. The sacculated intestine commences in xiii.

Spermathece (fig. 150). The most characteristic feature of Sparganophilus
Smithi is the occurrence of numerous spermathece in at least three somites, while in
one somite there is found only one pair. Sevenadult specimens from Laguna Puerea
were opened and agreed in the following arrangement and number of spermathecz:

Somite yi: 2 spermathece, in front of or slightly dorsal to setee 3 and 4.

Somites vil, vill, ix: 8 spermathec in each, in front of, slightly dorsal and
lateral of sete 3 and 4.

The location of these spermathece is not strictly constant asin some specimens,
as well as insome somites of the same specimen, the spermathecx were shoved a little
dorsally or ventrally of sete 3 and 4. One specimen possessed three spermathece
on one side and four on the other side in the same somite, but all the other specimens
possessed eight spermathece in each of the somites, except in vi where inyariably
only one pair was found,

These spermathece are very large (fig. 125), tall and slender, and viewed
in cross-sections of their body they are seen to extend from one end of the ccelom to
the opposite side, touching both body-walls. They are generally directed forward
and crowding each other; they occupy nearly all of the available room in the somite.
I believe there is no other case known in oligocheta where so many and so large
spermathece are known to occupy the same somite. The size of the respective sperm-

PACIFIC COAST OLIGOCH UTA, 159

athece varied some, but the general shape appeared quite constant within certain
limits. The figures appended represent the spermathece taken from one specimen.
There is a narrower muscular basal tube and a more swollen non-muscular chamber
as usual, the structure of which offers no characteristics.

Sperm-sacs. Of the two pairs of sperm-sacs the anterior one in somite xi is
much the narrowest in the direction of the median longitudinal diameter of the body.
It is principally ventral in location, but extends upwards and joins the one from the
opposite site. It is closely surrounded by a sperm reservoir without enclosing mem-
brane. The sperm-sacs in xii are much broader and longer, but are generally only
dorsal and do not enclose the intestine in any of the specimens sectioned. This rela-
tive size of the two pairs of sperm-sacs appeared constant. Somites ix and x are
transformed into two very extended sperm reservoirs.

Ovaries in the various species are long and flat, not lobed, and in both longitu-
dinal and transverse section offer to view a single undivided surface.

Oviducts are very broad and furnished each with an ovisae consisting of an in-
vagination of the septum, which in all my specimens were filled with large gregarine,
rarely containing any ova.

Ciliated rosettes are similar to those of Sp. benhami in size. The spermduets run
ventrally to as the tubercula pubertatis, about four times as far laterally and dorsally
from seta 2, as 2 is distant from seta 1. The spermiducal pore is similarly situated
ventrally to the tubercula pubertatis, This characteristic is in all the species only
shared by Sparganophilus sonome. All other species are characterized by having the
spermduets run dorsally as to the tubereula pubertatis. The spermiducal pore is simi-
larly in Sp. Smithi situated ventrally to the tubercula pubertatis, and is found in the
intersegmental groove between somites xix/xx.

Prostates or spermiducal glands (Beddard). There are three pairs of spermi-
ducal glands, one pair each in somites xxii, xxiii, xxiv. One specimen possessed only
one gland in xxiv, and there may possibly be found some variation in number when
more specimens have been investigated. In all cases these glands were much smaller
than in Sparganophilus Benhami, but otherwise almost similar in structure, except for
the entire absence of the basal muscular duct. The glandular part was never large
enough to be folded on itself lengthwise, but its tube was much twisted in the direction
of its long diameter, often to such extent that cross-sections always showed the tube
as three or four circular openings, surrounded by a wall two cells thick. The museu-
lar basal part is absent, and the lower tube, where it enters the muscular layers, being
very short, is surrounded by a thin layer of connective tissue. The whole organ in
mature specimens projects only slightly above the body-wall, and the glandular part
is in either diameter not any thicker than the body-wall.

Nephridia are in this species covered and perforated by numerous blood
capillaries, to a much greater number than in Sparganophilus Benhami, but similar to
what is described in Sp. tamesis and Sp. Hiseni. Otherwise the nephridia of the re-
spective species appear to be of the same general size and structure.

Vascular system. The hearts begin in xi, and extend forwards to viii. In xii

160 CALIFORNIA ACADEMY OF SCIENCES.

and xiii there are no connecting vessels. In xiv to xvii we find the dorsal vessel en-
larged and folded on itself in a zigzag manner. Both hearts and dorsal vessels are
thickly covered with chloragogen cells. I have already pointed out the absence of
a continuous blood sinus in any of the various parts of the intestine. All the various
organs of the body are thickly covered with blood capillaries, especially so the neph-
ridia, spermathec, the clitellum, in which strong capillaries separate every two or
three rows of the large clitellar cells. In the center of the tubercula pubertatis is
always seen a dense mass of larger and smaller capillaries.

A characteristic of the vascular system is also the great scarcity of blood
glands, these being very numerous in Sp. Benhami.

Sparganophilus sonome n. subsp.
Figs. 123, 126.

Habitat. Creeks and springs around Sebastopol, Sonoma County, California.
Adult specimens very numerous in April.

General Remarks. I consider Sparganophilus sonome as probably a sub-
species under Sp. Smithi, which it resembles in most points, two of which are of
the greatest importance, viz.: The position of the spermduct and spermiducal pore,
and the duplication of the spermathece in several somites. But as long as I found
the number of spermathec constant, and some other minor points of difference, I
thought it best to describe this form more carefully, leaving the question of species or
subspecies as a matter of choice, and for future consideration and study.

EXTERNAL CHARACTERS.

Tubercula pubertatis are the only external organs which differ from Sp. Smithi.
While in Sp. Smithithey form a continuous ridge on either side, they are in Sp. sonoma
broken up in numerous tubercles, generally two or three on each somite (fig. 125).
I believe there is another characteristic worthy of mentioning. The single puberty
groove is in the center of the tubercle, while in Sp. Smithi there are two grooves, one
at the base on either side of the tubercle. But I freely confess that I have not sec-
tioned up a sufficient number of specimens in order to know if these points are really
constant, or if not rather they are subject to considerable variation. My supply of
mature specimens of Sp. sonome was very limited, but in all I found the tubercula
pubertatis broken into a succession of little knobs, but all together forming a ridge
in outline like that of Sp. Smithi.

INTERIOR CHARACTERS.

Spermathece. While in all the specimens of Sp. Smithi I found eight sperma-
thece in each of somites vii, viii, ix, the subspecies possess them as follows: Somite
vi: one pair with the pore in front of seta 4. Somites vii, viil, ix: two pairs (4) in
each, with pores in front of setee 4, and between 3 and 4. Each pair consists of two
separate spermathecie, one of which opens in the intersegmental groove in front of
seta 4, while the other opens similarly in front of a line drawn 4 the distance between
sete 3 and 4, the 4 being towards 4 and the } towards 3. Both pairs are, therefore,

PACIFIC COAST OLIGOCH TA. 161

dorsal. The direction of the spermathecz is in the diameter of head to tail. They
lie closely pressed to the body-wall. As regards size, the most anterior pair is the
smallest, and the most posterior the largest.

Sparganophilus Benhami n. sp.
Figs. 97-119.

Habitat.—In the mud of springs, October, November, 1894. Tepic City, Ter-
ritory of Tepic, Mexico, 4000 feet altitude. Of some fifty specimens collected only
three were well iceeloned as regards clitellum and tubercula pubertatis.

EXTERIOR CHARACTERS.

Color, reddish-yiolet, yery strongly iridescent. The violet tint is very pro-
nounced, much more so than in any other species that I have examined except Sp.
guatemalensis, where the bluish-violet tint is very deep and intense.

Body. ‘This isa long and slender species tapering towards the tail end. The
cephalic lobe is superiorly not distinct from somite i, but there is a small pit on the
boundary between the two. Somites i, ii, iii are less prominent. Somites iy to ix are
strongly convex, but x to xv are much less so. The clitellar intersegmental grooves
are only distinct between the ventral sete.

Dorsal pores. The most anterior dorsal pore is seen between i/ii; the most
posterior one between v/vi. There is a central pit in the dorsal side of somites i, ij, iii.

Chitellum, as in other species; the extent of the clitellum is different on the
dorsal and ventral sides. This I think can best be expressed thus: dorsally, }xiv—
xxiv; ventrally, xvii—xxvi. The clitellum is continuous, but much less developed
ventrally between the tubercula pubertatis than dorsally. This is also characteristic
of the other species I have examined. Viewing the clitellum exteriorly, it appears
as discontinued between the tubercula pubertatis ridges.

Tubercula pubertatis begin in xviii and end in xxii. Each one consists of a
straight ridge on the top of which runs a groove parallel to the ridge. Posterior ‘to
the tubercula pubertatis proper the eel part of the clitellum is concaye deepening
posteriorly and ending in a deeper pit in the posterior part of xxvi. Anteriorly the
tubereula pubertatis Ree continue forwards almost straight and outside of the ventral
sete 1 and 2 to the center of xii, where they end in the region of the ventral sete. ‘These
two almost parallel ridges are very thin and sharp, several times narrower than the
tubereula pubertatis proper. They only appear in fully adult specimens, and are very
characteristic. I believe that the underside of the clitellum will furnish good species
characteristics if carefully noted. But in order to bring out this region properly the
specimens must be slowly killed and then as rapidly as Hens be passed through the
alcohols into that of 96%. Formalin specimens do not show the region as well as
alcoholic specimens.

Male pore or spermiducal pore is situated on the anterior } of xx, just lateral
to the tubercula pubertatis. The pore is not prominent, but may ii readily seen.

Prostate pores are not prominent, but in adult specimens they may be seen in
front of sete 1 and 2 in somites xxiii, xxiy, xxv, xvi.

162 CALIFORNIA ACADEMY OF SCIENCES.

Ovipores are separate between somites xili/xiy, or slightly on the anterior part
of xiv.
Nephropores prominent in front of sete 1 and 2 in all somites posterior to xii.

INTERIOR CHARACTERS.

Body-wall. In somites x to xiv the body-wall is much thinner on the dorsal
side of the body than on the ventral side. As Benham does not mention this peculi-
arity in Sp. tamesis, and as I have not observed it in Sp. Smithi, I take it for granted
that this character is peculiar to this species. The clitellum is complete, but much
thinner on the ventral side between the sete, as shown by figure 119, which repre-
sents the ventral body-wall in somite xix. It will be seen that the longitudinal muscular
layer is several times thicker than the transverse layer. The strands are separated
by irregular layers of granulated tissue of various thicknesses. The circular layer
again consists of only one row of fascicles, each fascicle being tubular, and inclosing
numerous irregularly distributed muscular fibers, strongly striate. When viewed in
cross-section it becomes apparent that the strands of the longitudinal muscles are of
varying thickness, the thicker ones being situated nearest the coelomic cavity, from
there gradually decrease in size towards the circular muscles.

Clitellar cells. The clitellar cells are especially interesting on account of their
relationship to the cells of the tubercula pubertatis, which will be described later on.
Figs, 107, 108 represent a longitudinal section of the body-wall in somites xxiii and xxiy.
The muscular layers are much thinner, the clitellar cells are only developed in the
vicinity of the prostates. Below the clitellar cells are seen long tubular cells, some-
what like those of the tubercula pubertatis, and between them are seen, now and then
(fig. 107 d.ch.), the deeply staining discharge chambers of the clitellar cells. The uni-
cellular glands, which are here very irregular, are entirely confined to the zone
surrounding the intersegmental grooves. In fig. 108 is seen a more magnified part
of, the region close to the prostate. Only one unicellular gland, though of unusual
size, is seen. It isastray one, as theyare rare in this particular zone. The structure
of the clitellum of Sparganophilus has been well described by Benham in his paper
on Sparganophilus tamesis, and I can only add a few points of interest. For studying
the clitellar and tubereula pubertatis cells I have used principally the iron-lack
Heidenhain process, but by adding a light tint of Ehrlich-Biondi to the ammonio-
ferric-alum solution the most perfect differentiation may be had of the various clitellar
elements. The clitellum in Sparganophilus contains the following various varieties
of cells:

1. Regular epidermal cells, secreting the cuticle, furnished with oval nuclei.
Even on the fully developed clitellum of Sparganophilus Benhami they occur all
around, both on dorsal and ventral sides. Their inner ends are drawn out into one or
more threads, some of which extend to the innermost margin of the epidermis
(fig. 103, 1).

2. Unicellular goblet glands, entirely confined within the outer layer of the
foregoing cells. They are very irregular as to size. These cells are the only ele-

PACIFIC COAST OLIGOCH &TA, 163

ments in the clitellum which stain deeply with hematoxylin. These cells offer
nothing characteristic of the species. They are the last to be affected by tropwolum
and orange, their mucin remaining intensely blue. Their nuclei are compressed and
placed in the posterior part of the cell in the triangular mass and extremity, staining
black in the iron-lack (fig. 103, 2).

3. Short club cells. These are of very uniform size, their free ends forming a
marked line all around the elitellum interior to the goblet cells. The free end of
this cell is club-shaped and generally regular in outline. The heights of the indivi-
dual cells are quite even, and, as these cells stain darker than the following kind,
they form a marked contrast to all others. The granulation is coarse and regular, but
does not stain readily with hematoxylin. It takes, however, deeply the red aniline
stains. On the dorsal side of the clitellum these cells reach inward about 4 of the
whole width of the epithelium, while on the ventral side, they constitute the only
so-called clitellar cells. The nuclei of these cells are oval and always prominent
(fig. 105, 3).

4, The pre-eminently clitellar cells consist of several, 3 or 4, different lengths.
This class of cells are in many genera, such as Lumbricus, etc., massed in columns,
often very regular, and separated by septa. In Sparganophilus these cells are not
arranged in columns, but distributed almost regularly all through the epidermis, of
which they occupy about 3 of the whole mass. At certain intervals, however, may
be seen a thin triangular septa of connective tissue with nuclei, projecting down-
wards or outwards from the muscular layers, dividing this part of the epidermis in
numerous irregular pyramidal masses, which latter are homologous with the columns
in Lumbricus, ete. The nuclei of these clitellar cells are shrivelled up and situated far
back in the triangular apex where the cellular plasma is especially agglomerated.
There is no such granular secretion in this class as in the former, and they stain much
less intensely than No. 3. These cells occur only dorsally to the tubercula pubertatis,
and are entirely absent from the ventral side between the tubercula pubertatis.

5. Here and there we find intermediate varieties of oval and club-shaped
cells longer and wider than the epidermal supporting cells and furnished with large
round nuclei.

6. Connective tissue cells with large round nuclei, especially prominent
between the circular muscular layer and the tall clitellar cells. At intervals this
tissue forms triangular projections outwards, separating bunches of cells four or five
wide and of different sizes. This tissue is generally accompanied by numerous
capillaries.

Tubercula pubertatis. These organs are very prominent, and situated on a
nearly continuous ridge occupying somites xviii-xxii. Benham has described this
structure in detail in his species Sp. tamesis, and called attention to the fact that it is
composed of long narrow cells, which are modified clitellar cells. In Sparganophilus
Benhami these cells are of two kinds, as far as regards the form.

a Some which are very narrow, longer than the others, and narrowest at the

free end. All of this class are grouped in such a way that they radiate outwards
Memoirs, Vou. II, 5. January 7, 1896.

164 CALIFORNIA ACADEMY OF SCIENCES.

towards and around a small concave pit or groove in the outer edge of the epidermis
(fig. 105).

}. Interior to these taller cells are shorter and thicker cells, indicated asd in the
above figure.

Exterior to these, the regular and characteristic tubercula pubertatis cells
proper are seen numerous unicellular glands, the same as the goblet cells of the
epidermis.

In addition to these glandular cells, I find at certain intervals a few sense cells
opening out into the narrow pit or groove just referred to. They are everywhere very
few in number, tall, narrow, with now and then the tip projecting through the cuticle
into the shallow pit (fig. 105s.¢./.) There is not a continuous row of these cells, but
here and there are found bunches of half a dozen cells, opening close together. They
do not seem strictly parallel, but bulge and diverge in such a way that in sections,
which show the common cells parallel, rarely more than one single sense cell is in
view to any great part of its length. This, together with the small number of these
cells, is undoubtedly the reason why they are frequently overlooked.

In connection with these sense organ cells, I have re-examined the ventral
papillee found in Argilophilus marmoratus, and I find that these structures are really
nothing but tubercula pubertatis nature, or at least sense organs furnished with sense
cells, a description of which will be deferred to a future paper. As a general con-
clusion, I may state that the tubercula pubertatis are really sense organs, furnished
with sense cells of the same nature as those found in other parts of the epidermis, and
described by me in Benhamia, and by Vejdovsky in Rhynchelmis, by Cerfontaine
and Langdon in Lumbricus, by Hesse in Lumbricus and Allolobophora, ete.

Septa. The septa are not all of the same thickness, neither is each individual
septum of the same thickness throughout. The first distinct septum is found between
iv/y. It is of regulation thickness, or as thick as septum x/xi and those following
posteriorly. The anterior septa increase in thickness forwards and backwards in such
a way that septa vi/vil, vii/viii are of about equal thickness, while those in front and
behind these are thinner in proportion as they are more distant. The anterior five
septa are much thicker in their central area, and thin out towards the periphery and
-body-wall, but even then, at the thinnest part, they are about four times thicker than
the posterior septa. The six anterior septa do not strictly correspond with the seg-
ments, but are attached to the body-wall about one-fifth the distance forward from the
posterior intersegmental groove.

Suprapharyngeal and septal glands. The glandular mass superposed on the
pharynx and opening into it, is prominent, but situated far back, and with its lobes
pointing forward, or in the same direction as the septal glands. The opposite is gen-
erally the case, and has been so in all other species examined by me.

Longitudinal sections show that the regular septal glands are present in three
somites. The most anterior pair is in iv, immediately behind the suprapharyngeal
glands, and not separated from the latter by any septum. The other two pairs are in
yand vi. In longitudinal sections the suprapharyngeal glands are seen to be com-

PACIFIC COAST OLIGOCH MTA. 165

posed of two distinct lobes of about equal size, one situated closely posterior to the
other. This is also the structure of the septal glands in iv and y, but the one in vi is
solid, consisting of only one lobe, when viewed in longitudinal section. The main
lobes are situated on the same longitudinal muscular band, with their discharge ducts
running forwards into the pharyngeal cavity. In each of the three somites there are
two distinct glands on either side of the median line. One larger supraintestinal
consisting of three parts extending laterally, and one smaller subintestinal. These
latter ones also open into the pharynx, but into its ventral side, the pharynx
thus being partially developed even on the ventral side. The discharge tubes and
chambers are very large, and the latter oceupy more than one-half the width of the
pharyngeal wall. They stand very close together, and are all of about the same height
and form (fig. 112). The dorsal wall of the pharynx is much thicker and denser
than the ventral one, and the discharge chambers stand closer and are of more uni-
form size, more tubular. The discharge chambers on the ventral side are thicker,
more pear-shaped and much fewer in number. Fig. 112 represents the lining of
the dorsal wall of the pharynx; fig. 115 the ventral wall of the same, and fig. 114
the wall next posterior to the main dorsal section.

Hsophagus and intestine. The description given by Benham of the histology
of the intestine of Sp. tamesis may in the main points be applied to this species.
The cesophagus is cylindrical with parallel sides and slightly nipped by the septa. It
is sparsely covered by chloragogen cells, which latter are more numerous on the
intestine. The cilie of the inner epithelium are much longer in the esophagus,
and so are the epithelial cells themselves. Seen in longitudinal section (fig. 110)
we find that the transverse muscles surrounding the cesophagus are more numerous and
present in several rows, while in the intestine they are few and far between, and
arranged only in one row. There is thus a distant approach to a gizzard in the
esophagus. The longitudinal muscular layer is reduced everywhere in the intestine
to a single strand, but in the cesophogus it is double, sending out strands to the septa
and to the mesenteri¢ sac.

Blood sinus. Both Benham and Smith have shown the existence of a conti-
nuous blood sinus in the intestine of the species described by them. Inthe wsophagus
of our present species we find only a vascular network and confluent smaller lacunes,
but in the intestine proper we meet with a continuous blood sinus all around the
epithelial cells. The radiated appearance of the blood in the sinus, as well as in the
other vessels, must as Benham suggested be due to the erystalization of heematine. In
Sp. Benhami and Sp. Smithi these crystals are so many and so heavy that they invari-
ably destroyed the edge of the microtome knife and made sectioning most difficult.
But the crystalization presents some peculiarities, and in places appears as if there
was a mass of radiating fibers always from the side nearest the center of the body
radiating towards the periphery, but never the contrary. In Sp. Benhami these crystals
are much more slender than in Sp. Smithi. In the latter species their nature is not
to be doubted. They are also found in the hearts, but more rarely in the main longi-
tudinal vessels.

166 CALIFORNIA ACADEMY OF SCIENCES.

In the intestine we find between the inner ends of the epithelial cells a layer
of connective tissue, strengthening as it were that side of the perienteric sinus (fig.
111). Both cesophagus and intestine are enclosed by a thin mesenteric sac, which
is nipped by the septa (fig. 111, mes.) and pressed close to the chloragogen cells.

Spermathece. These are much broader than those of Sp. Smithi or Sp. tamesis,
but resemble more those of Sp. Hiseni and Sp. gquatemalensis. Their position in
somites vil, vill, ix, is in the anterior part of the somite, opening in the intersegmental
groove and pointing backwards. ‘Their pores are in line with the dorsal couple of
sete, or 3and 4. The muscular part is tubular, smaller, and consists of two layers,
one inner of epithelial cells, one outer thicker, of circular muscles. The inner epi-
thelial layer, which is a direct continuation of the epidermis of the body-wall, consists
of very tall, narrow, columnar cells, while the thick muscular coating is a direct con-
tinuation of the circular muscular layer of the body-wall. The free end of the
spermatheca is wavy and warty in outline, and consists of much shorter epithelial
cells, simply covered by the peritoneum. The spermatheca is very broad and very
flat (fig. 118). .

Sperm-sacs. There are two pairs of lobulate sperm-saes in xiand xii projecting
from the anterior septum. They are situated principally dorsally, and resemble those
of Sp. tamesis, but are much less lobulate than those, and very much less lobulate
than the sperm-sucs of Sp. Hiseni, Judging from sections of the species sent me by
Prof. Frank Smith for comparison.

Large masses of free spermatogonia and spermatozoa are seen in front of the
ciliated rosettes.

Ciliated rosettes (fig. 109) are large and very regularly folded. Each rosette
sends out a long tubular lip into the sperm-saes, in way which I have figured in fig.
119¢e. Ina cross-sectioned specimen I found this lip far back in the posterior part
of xu. The lower convolute lip of the funnel is almost absolutely regular, and similar
in each of the four rosettes. The spermducts appear to resemble those of Sp. tamesis
and Sp. Hiseni. The spermidueal pore is situated just outside of the tubercula puber-
tatis in the anterior part of xx, Just as in Sp. Hiseni.

Prostates. Smith is the first to describe the prostates in Sparganophilus. In
Sp. tamesis they appear not to be present, as Benham does not mention them. In Sp.
Benhami there are four pairs opening in somites xxiii, xxiv, xxv and xxyi, in front
of sete 1 and 2. The prostates are constructed on the same principle as the prostates
in Acanthodrilidee and Cryptodrilidee, and consist of two parts; one basal and
muscular, one apical and glandular. The glandular part is tubular, straight or folded,
of considerable length, but confined to one somite. The glandular part contains an
inner epithelium, and surrounding it club-like glandular cells of varying lengths,
giving to the surface of the prostate a wavy and irregular appearance. The prostates
in this species difter from those of Sp. Smithi by having a muscular duct or basal
part. ‘This latter does not exist in Sp. Smithi, the glandular part in the latter species
being immediately attached to the body-wall.

Nephridia. ‘There is undoubtedly some difference in the location of the most.

PACIFIC COAST OLIGOCHMTA. 167

anterior nephridia in the various species. In Sp. Benhami the most anterior pair is
in xii, but they are smaller than those in xiii. In xy we find the first very large
nephridium covered with a thick coelomic cell mantle. This mantle covers the ducts;
sometimes we also find a mass of similar cells attached to the nephridium as a rounded,
nearly separate mass, only connected with the nephridium by means of a very narrow
part. Owing to the great opaqueness of the nephridia, [ have not been able to make
out the run of all the canals, and can illustrate only a little more than the outline of
the organ. It appears, however, that some of the canals are doubled. The outlet
duet is very heavy, wide, and its walls are thick. It runs far up into the windings.
There is a bridge, and I can distinguish all the various principal parts described by
me in the nephridium of Pontodrilus. There are comparatively few blood vessels on
the nephridia, and in this respect the species differs from Sp. Hiseni and Sp. Smithi,
in which the nephridia are thickly covered and penetrated by capillaries, causing
them to be of a deep pink color. As regards size, the nephridia are as high as the
diameter of the coelom. The nephropores are in front of sete 1 and 2, and very
wide.

Vascular system. Benham’s description of the vascular system of Spargano-
philus is so complete that I can add but little; the various species seem to agree to a
very great degree. Sp. Benhami is distinguished by a scarcity of capillaries on the
various organs of the body, such as clitellum, nephridia, etc., while in Sp. Simithi
and Sp. Hiseni capillaries are so abundant that they, for instance, almost obscure the
nephridial surface. The crystalization of the blood has already been described. It
is found in all the vessels of the body, but especially in the mesenteric blood sinus.
I have found swimming free in the blood two distinct cell elements, some of which
are very large-—possibly leucocytes, while others are extremely minute, more round
and dense—possibly erythrocytes. But as their description requires more time and
study, I will defer it to a separate paper.

The walls of the blood vessels present a banded appearance, caused by par-
allel bands of thicker tissue, furnished with large circular nuclei, arranged in rather
regular rows.

A characteristic feature of the capillaries of this species are the numerous
blood glands, similar to those I have described in Pontodrilus and Argilophilus. They
are especially numerous in the nephridia and in the septal glands. They also contain
a large number of nuclei.

Sparganophilus guatemalensis n. subsp.

Habitat. Guatemala. While in this Central American State several years
ago I found a great number of specimens pertaining, as I believe, to at least three
distinct species of Sparganophilus. ‘These, as well as my other oligoch:etological col-
lections made there, were mostly destroyed by accident, few specimens being saved.
These are now not in good condition for description, and this must account for the
imperfect data I am able to furnish for all species now described from Guatemala. I
found Sparganophili in that country in the most varied localities —City of Guatemala
at Los Bafios, Los Arcos, Laguna Amatitlan, Coban, Panzos Ysabal, Duejias, ete.
The present form is from Los Banos and from Coban.

168 CALIFORNIA ACADEMY OF SCIENCES.

GENERAL REMARKS.

I consider this a subspecies of Sparganophilus Benhami, at least until a closer
investigation of better material may reveal other characters, if any there are. Spar-
ganophilus guatemalensis is one of the smaller species about 10 cm. long by 2 mm.
wide; the specimens were considerably stretched. Color is deep flesh, with an intense
deep, dark violet lustre, much darker than in any other species and well preserved in
alcohol, in which the specimens appear blackish violet.

Setw are lateral and dorsal.

Clitellum extends from xvi-xxvi, but cannot be well defined on the ventral
side. The tubercula pubertatis are in the shape of two parallel ridges extending from
Xviii to xxii or specimens from Patal and Coban, which I take to belong to the same
species as those from Los Baiios in the City of Guatemala.

INTERNAL CHARACTERS.

Of these I could only distinguish a few. The spermathece are in three pairs,
one.each in vii, vili, ix, opening in the anterior intersegmental groove. The apical
end in the fully developed spermatheca is very wide, flattened like a mason’s trowel.
Several of the spermathecz possessed a slit or opening in the apical end which com-
municated with a longer or shorter sac, continuous with the exterior lining of the rest
of the spermatheca. ~ It is possible that this is only a result of maceration, as I have
seen nothing like it in other species of Sparganophilus, and it is not probable that we
here have an analogy with the spermathecs of Enchytreeus or Sutroa, where this
organ communicates with the intestine.

Prostates are found in four pairs and situated in somites xxiv—xxvil or one
somite further back than in Sp. Benhami. This in specimens from Sapote. If this
character holds good it is one of considerable importance.

Hearts were in vill, ix, xX, xi strongly developed, but not filling the ccelom.
Some of the Guatemala Sparganophili possessed only three pair of hearts in somites
ix, X, xi, with the ventral vessel branching in xy/xvi. Those were specimens from
Los Arcos. Others again from Amatitlan possessed the hearts in x, xi, xii, xiii, with
the ventral vessel divided in xv/xvi, while those from Los Banos, Guatemala City, had
the ventral vessel branched in xiv/xv. I believe those from Amatitlan and Los
Arcos belong to different species, but, as all the specimens are lost, I can only eall
attention to the differences and to the importance of further investigations.

Sparganophilus carneus n. subsp.

Habitat. Mississippi river near Clayton, Iowa, in soil at the water’s edge,
under pieces of boards and lumber. Numerous specimens at end of August, 1890,
only few of which were adult.

General remarks. It is probable that this is only a northern form of Sparga-
nophilus Benhami, from which species it differs principally in the form of the sperm-
athece, and by a much lighter color. The specimens at my command were not in proper
condition and the shape and position of the tubereula pubertatis could not be made out.

PACIFIC COAST OLIGOCH ATA, 169

My object is merely to call attention to this form which after all may be more distinct
as more of its structure is known. In regards to the form of the spermatheecs it
differs much more from Sp. Benhami than does the other subspecies Sp. guutemalensis.
It is also much smaller.

CHARACTERISTICS,

Color is much lighter, more flesh and less violet than that of Sp. Benhami and
Sp. guatemalensis, and the irridescence more faint. The species is smaller than
Sp. Benhami, being about 9:em., about one-half the size of that species.

Clitellum in the most developed specimen occupied xy—xxiy ventrally, dorsally
it could not be properly defined. Prostates present with certainty in xxiii, xxiv, but
owing to the somites being torn in xxy, xxvi I could not with certainty ascertain their
presence there.

Sete are similar to those in Sp. Benhami. The anterior xxii somites have
much larger sete than the other, except the most posterior ten. The sets increase
in size from somite ii to somite xy, then diminish backwards to xxiii, in which somite
they are smaller yet, this size continuing to the tail, where at about ten somites from
the anus the sete again begin to increase. There is no parietal gland in somite
iii as in Sp. Hiseni, and this is the chief reason why I class this species under Sp.
Benhami. The spermathece are in three pairs and open in front of 3 and 4 dorsally.
Their shape resembles much more those of Sp. Hiseni than they do those of Sp.
Benhami. The apical end is globular and very much larger than the basal part.
The spermathecz are not trowel-like. I give some outline drawings of the sperm-
athecee taken from one and the same individual.

Sperm-sacs are very large in xi, xii, the one in xxii projecting into xxiii filling
the somite also. The sperm-sacs are coarsely lobulate, much less so than in Sp. Kiseni.

The capillaries resulting from the lateral integumental vessels are very large
in the anterior six somites. The hearts also are very large and seem to fill all available
space on the ecelomic cavity in vill, ix, x, Xi.

Deltania Troyeri Eisen, n. var. crassa.
Figs. 142, 143.

Derinition. Size 45 mm. by 2 1-2mm. Septal glands in v and vi, about twice
as thick as the septum. Spermathece each with two diverticles, which have the apical
ends globular and inflated. Prostates thick, with the glandular part about 1 1-2 times
longer than the thin muscular duct. Habitat, Baja California.

In other respects like the type, judging from two dissected specimens.

Habitat. Of this variety I possess only three specimens, from Ensenada de
Todos Santos, in the northern part of Baja California, on the Pacific Coast side. They
were taken in the same locality as A/eodrilus Keyesi, in the pure sand in the creek
bottom, in a spot which had been kept moist by the accumulation of some sacks and
rubbish. The specimens became strongly contracted, on account of the necessity of
immediately placing them in alcohol.

170 CALIFORNIA ACADEMY OF SCIENCES.

EXTERNAL CHARACTERS.

The variety resembles the main form in most respects, but is much larger,
especially thicker. The contracted specimens were as long as the longest of the main
form from San Francisco, when fully extended, and twice as thick, and must, when
alive, have measured 45 mm. by 25 mm. Other external characters agreed exactly
with those of Deltania Troyeri type from San Francisco, California. The deltoid
arrangement of the ventral sete in the genital region was identical, which satisties me
that we in this arrangement have most important species characters between the
species of this genus.

INTERNAL CHARACTERS.

The variety differs from the type in two particular points.

The septal glands are present only in two somites, y and vi, and are exceedingly
thin, or about one-fourth as thick as those of D. Troyert type. This refers to those
superior to the intestine. Those below the intestine I could not discern, as no sections
were made. The glands were lamellie-like, and only about twice as thick as the
septum.

The spermathece were thicker than those of the type, with more inflated diver-
ticula.

The prostate was thick, and its glandular part about 15 times longer than the
very thin muscular duct. Two or three fully developed penial sete with each
prostate.

Deltania Troyeri Eisen, n. var. lagune.
Figs. 144-147.

Derinition. Size 32 mm. by 1 1-2 mm. No copulatory papiile. Sete a and
b next posterior to clitellum, reach their final distance in the fifth somite posterior to clitel-
lum. Spermathece in ix twice as large as those of the type, contracted at center, apical
part swollen. Prostates more slender than in the type. Large ovisacs. Habitat, Baja
California, Sierra Laguna.

DETAILED DESCRIPTION.

Habitat. The exact locality in the Cape Region of Baja California and Sierra
Laguna is a camping-place called La Joya. There, where the trail crosses the creek
the water had backed up behind some fallen logs causing a miniature mudflat covered
with leaves and sediment. The specimens were found in the rotten wood and under
the decayed leaves in the mud. Altitude about 5500 feet.

EXTERIOR CHARACTERS.

The average specimen is larger than the type from San Francisco, the largest
specimen of the latter being smaller than the average mature specimens of the
variety /agune. The two round copulatory palpille figured by me from D. 7royeri
are not seen in the variety. The sacs of penial set in several dissected specimens
possessed each two penial sete fully developed, one haying three, but of which one

PACIFIC COAST OLIGOCH®TA. 171

was undeveloped. One of the sete is a trifle more curved than the other. The
point of each penial seta shows a slight and indistinct ornamentation (fig. 144).
Since my first description of D. 7royeri I have found some more mature specimens at
the old locality in San Francisco. One of these specimens possessed two penial setie
in each sae, both slightly sigmoid, with trace of ornamentation at the free apex. In
the variety /agune the ornamentation is more distinct.

Common setw resemble the type, but sete a and} next posterior to clitellum
reach their proper and final distance from each other only in the fifth somite posterior
to clitellum, while in D. Troyeri type they reach the final distance in the third somite
posterior to clitellum.

INTERNAL CHARACTERS.

Spermathece are much larger in the variety /agune, or about twice the size of
those in D. Troyeri, with the main body distinctly contracted at the center, and with
the apical part much enlarged. The diverticula are distinctly tri-digitate and reach
up to the narrow contraction of the main sae.

Prostates or spermiducal glands are much more slender and longer than in
D. Troyer type, and generally but not always confined to one somite.

Ovisacs are two, projecting from the Septum xili/xiv into xiv, occupying the
largest part of the ccelomic cavity in the latter somite. The structure of the ovisac
is characterized by numerous trabecula forming an extensive system of round pockets
of various sizes. Where there are no pockets there are solid zones of round or slightly
oval nuclei (fig. 147) imbedded in a dense mass of tissue without distinct cell-walls.

PHCENICODRILUS Eisen.

The finding of a new species of this genus enables me to more properly define
it. The genus was originally based on the absence of a prostate at the male-pore,
there being, however, a short muscular atrium, in which opened the spermduct. The
type was Phanicodrilus taste, a species from the Cape Region of Baja California.
The new species described below under the name of Phenicodrilus tepicensis possesses
amuch more developed atrium, in shape and structure resembling a spermatheca.
There is a total absence of glandular cells, such as are found in prostates, the whole
atrium being muscular. There is besides another difference between Pheenicodrilus
and Ocnerodrilus, though of much smaller importance. In Phcenicodrilus we find a
large number of muscular fascicles divided in three paired groups in somite xvii, radiat-
ing from the male pore. In Ocnerodrilus these arciform muscles are few in number.
In Pheenicodrilus these muscles are so thick and numerous that the atrium itself ean
only be seen in sections, not by suface view of the inner side of the body-wall.

Through the possession of a muscular atrium this genus comes rather close to
Nannodrilus Beddard. This genus, of which so far only one species is known from
Africa, possesses a muscular bursa copulatrix, in which opens separately the anterior
prostate and the anterior spermduct. The great similarity between the atrium in

Pheenicodrilus and the bursa copulatrix of Nannodrilus is very striking, especially as

Memorrs, Vou. II, 5. January 28, 1896.

Lif CALIFORNIA ACADEMY OF SCIENCES.

the spermducts in Phoenicodrilus also open intothe atrium. But the question that still
remains to be settled is this. Is the atrium in Phcenicodrilus a modified bursa ecopu-
latrix, or is it simply the remains of deteriorated prostate. The absence of a penis
in Pheenicodrilus makes the latter alternative seem most probable. While in Oene-
rodrilus the reduction of the prostate consists in the loss of one layer of glandular
cells in the apical part, this reduction has proceeded further in Phoenicodrilus, the
whole glandular apical part here being wanting.

The principal differences between the genera Ocnerodrilus and Phoenicodrilus
are then as follows:

Ocnerodrilus. A prostate of varying size is always present at the male pore.
This prostate consists of two parts, one glandular, lined by a single layer of cells, and
one muscular basal part.

Phenicodrilus. No prostate, only a short or rudimentary muscular atrium, in
the lower part of which opens the spermduct.

Pheenicodrilus taste Eisen.

Derrnirion. Atrium rudimentary, muscular and bulbous, only about twice as
thick as the spermducts. No ovisac. Spermathece covered with warty excrescences.
Arciform muscles in xvii comparatively few.

This species, which I first described from the west coast of the Cape Region
of Baja California, has since been found by Messrs. Eisen and Vaslit in two widely
separated localities. It was found in great numbers at Miraflores, on the east side of
the Sierra of the Cape Region, some 30 miles north of San José del Cabo, together
with Ocnerodrilus Beddardi, Sept., 1894. It was, however, quite unexpected to find
this species also common in garden soil at San Blas, the seaport in the territory of
Tepic, several hundred miles south of the Cape Region peninsula. I can find no
differences in the anatomy, nor in the exterior of the specimens from these respective
localities. The small copulatory papilla in somite xiy, lateral to the oviduct, is present
in all. So are the numerous muscles in the copulatory region around the male pore.

Pheenicodrilus tepicensis n. sp.
Figs. 155-160.

Derinition. Atrium about twice as long as the body wall, but hidden by arciform
muscles in somite xovi. A median ovisac in aiv/xin. Spermathece entirely smooth,
without warty excrescences. Arciform muscles in xvii very numerous.

Halutat. Numerous specimens in and around the city of Tepic, territory of
Tepic, Mexico, 4000. In moist soil under logs, in gardens, ete. November, 1894.
Found together with Acanthodrilus Vasliti. Only few specimens were adult in
November.

EXTERIOR CHARACTERS.

Size 4 em. by 15 mm. Number of somites 75.

Clitellum incomplete, comprising somites xiii—xix.

PACIFIC COAST OLIGOCH MTA. 173

Sete strictly paired, sigmoid plain. The pair 1 and 2 is wanting in xvi.

Spermathecal pores in ix in front of setee 1 and 2.

Oviducal pores in xiy in front of sete 1 and 2.

Spermaducal pores in the center of xvii, where otherwise would be the sete
land 2. A small copulatory papilla around each male pore, but it is not connected
with a zone. No dorsal pores.

INTERNAL CHARACTERS.

Muscles. Numerous arciform muscles in the copulatory region, but especially
in xvii, connecting the copulatory papillse with the body-wall in line with sete 3 and 4.
These arciform muscles are arranged in three distinct groups. The central group,
which branches out fan-shaped from the inner surface of the copulatory papillie, is by
far the largest, consisting of about 10 to 12 distinct fascicles. Anterior to this is
another group of 6 to 7 fascicles, and in the posterior part of the somite we find a
swollen group of about 5 fascicles. To what extent these vary as to numbers, ete., is
uncertain, but my observations in Phanicodrilus taste are that they are quite constant,
and might be used as valuable characters in determining the species, as with a change
in the muscular strands is also connected one in the exterior copulatory zone. Between
xvii and xiii, as well as between xvii and xix we find several pair of arciform muscles
in each somite.

Septal glands are well developed. The suprapharyngeal glands are very jong
and extend far backwards. ‘The septal glands in the following somites diminish
posteriorly.

Spermathece. One pair in ix open between viii/ix in front of sete 1 and 2.
Form sac-like, very thin walls, pellucid, outline smooth, without any warty diverticula.
No distinction between a muscular and glandular part. The size is large, about as
long as the somite is wide.

Testes, ovaries and ciliated rosettes not characteristic.

Ovisac. There is a median ovisac in xili/xiy (fig. 157), consisting of a
pouching backwards of part of the septum. ‘This is the only species in the three
genera, Ocnerodrilus, Gordiodrilus and Pheenicodrilus, which possesses an ovisae.

Spermducts are of the same width throughout, without any muscular enlarge-
ment near the male pore. The two ducts run together, but their lumens are separate
until they reach the muscular atrium, where, just before they enter it, the two lumens
fuse into one (figs. 158, 159).

Muscular atrium. As has already been stated the usual prostates in Ocnerodrilus
are replaced by a pair of muscular pouches, entirely covered up by the numerous
arciform muscles. There are no glandular cells, and these atria can best be compared
to the basal muscular parts of the Oenerodrilid prostate only they are yery much
thicker, and consist entirely of muscular cells with the lumen in the body-wall lined
by columar epithelium. In the main pouch this epithelium consists of short cells with
round nuclei, around which extends a thin musenlar layer, which becomes wider at
the base. This atrium is very short and cannot readily be seen when the opened

174 CALIFORNIA ACADEMY OF SCIENCES.

body cavity is viewed from the interior surface. It is attached to the at this point
very much thickened body-wall (fig. 158) by numerous muscles and connective
tissue. The various species of Ocnerodrilus, especially those with short prostates,
require reinvestigation in order to ascertain the structure of the prostates. Should
intermediate forms be found it might become proper to merge the two genera into one.

Nephridia. ‘Those anterior to clitellum are not covered with a ccelomie cell
mantle, while those posterior to the clitellum are surrounded by a very thick one.
The structure of the nephridia appears to be similar to those of Phwnicodrilus taste,
at least in a general way.

LITERATURE REFERRED TO.

BenuAm, W. B. A new English Genus of Aquatic Oligocheta (Sparganophilus), belonging to the family of Rhino-
drilide. Quarterly Journal Micros. Science, vol. xxxiv.

Bennam, W.B. Report on an Earthworm collected for the Nat. Hist. Dept. of British Museum by Emin Pasha in
East Africa. Journal Roy. Microse. Society, February, 1891.

Bepparp, Fr. E. Two New Genera, Comprising Three New Species of Earthworms from Western Tropical Africa.
Proceedings Zoological Society, 1894, pages 379, ete.

Bepparpb, Fr. E. A Monograph of the Order of Oligocheta. Oxford, 1895.

CEeRFONTAINE, P. Recherches sur le Systeme Cutané, etce., du Lombric Terrestre. Archives de Biologie, x, 1890.

E1sen, G. California Eudrilide. Memoirs Cal. Academy of Sciences, vol. ii, No. 3.

Eisen, G. Pacific Coast Oligochwta I. Memoirs Cal. Acad. of Sciences, vol. ii, No. 4.

E1sen, G. Anatomical Studies on New Species of Ocnerodrilus. Proceedings Cal. Academy of Sciences, ser. 2,
vol. iii.

Garman, H. On the Anatomy and History of a New Earthworm (Diplocardia communis). Illinois State Labora-
tory of Nat. History, iii, 1888.

Hessz, Ricuarp. Uber the Septaldriisen der Oligocheten. Zoological Anzeiger, No. 456, 1894.

Hessr, Ricuarp. Zur Vergleichende Anatomie der Oligochwten. Zeitschrift f. Wissensch. Zoologie, 1894, pages
406.

Horst, R. Earthworms from the Malay Archipelago. Zoolog. Ergebn. e. Reise in Niederland. Ost-Indien. Dr.
Max Weber, B. ii, Leiden, 1892.

Lanopon, F. E. Sense Organs of Lumbricus. Journal of Morphology, vol. xi.

Micuar.sen, W. Oligocheten des Nat. Hist. Museum, Hamburg, iii.

MicHartsen, W. Oligochwten des Nat. Hist. Museum, Hamburg, No. iv.

MicHar.isen, W. Regenwiirmer der Thierwelt Ost-Afrikas, Bd. iv, Berlin, 1895.

Rosa, Dr. D. Annailen d. K. K. Nat. Hist. Hofmuseums. Die Exotischen Terricolen, Wien, 1891.

Rosa, Dr. D. Contributo allo Studio dei Terricoli Neotropicali. Accademia Reale delle Scienze di Torino, 1895.

Smirn, Frank. A Preliminary Account of Two New Oligocheta from Illinois. Bulletin Illinois State Laboratory
of Nat. History, vol. iv, 1895.

Une, H. Beitrage zur Kentniss auslandischer Regenwurmer. Zeitschr.f. Wissens. Zoologie, 57, 1894, page 69, ete.

Vespoysky, Fr. Organogenie der Gordiodrilen, Zeitschr. f. Wissensch. Zoologie, Bd. lyii, p. 642.

Vespovsky, Fr. System u. Morphologie der Oligochwten, Prag., 1884.

Verson, E. von. Zur Spermatogenesis, Zool. Anzeiger, xii, 1889, p. 100.

PACIFIC

COAST

OLIGOCH MTA,

REFERENCE TO LETTERS ON PLATES.

a. c. gl. anterior.caleic gland.
a. f. anterior fold of nephridium.
a. nephr. anterior nephridia.
a. spd. anterior spermduct.
atv. atrium.

bd. body-wall.

bl. blood vessel or capillaries.
bl. s. blood sinus.

br. bridge in nephridium.
buc. buccal cavity.

c. clusters of lunate cells.

ec. c. columnar cells.

c. ep. celomic epithelium,

c. gl. calciferous glands.

ch. discharge chambers of glands.
chlo. chloragogen cells.

cil. ciliated epithelium.

el. c. clitellar cells.

el. clitellum.

c. m. circular muscles.

com. commissures.

c. 7. ciliated rosettes or sperm-funnels.
c. t. connective tissue.

cu. cuticle.

d. gl. dark staining glands.

d. pr. dorsal pore.

d.v. dorsal vessel.

e. epithelial cells.

ept. epithelium.

gl. glands.

gl. c. unicellar glands.

gl. pr. glandular part of prostate.
gizz. gizzard.

h. heart.

hy. hypodermis.

i. s. gr. intersegmental groove.
l. c. large cells.

l. m. longitudinal muscles.

m. muscles.

m. atr. muscular atrium.

mes. mesenteric tissue.

m. pr. muscular prostate.

n. nephridia.

n.c. nerye cells.

ne. nerve ganglion.

nec, neck of nephridium.
neph. nephridium.

neph. p. nephropore.

es. esophagus.

ov. ovary.

ovd, oviduct.

org. sense organ.

p.c. gl. posterior calciferous diverticulum.
p.f. posterior fold, nephridium.
phx. pharynx.

p. neph. posterior nephridia.
pr. prostate.

p- spd. posterior spermduct.
pr. sto. prostomium.

ps. penial sete.

prt. peritoneum,

se. secreted matter.

s. int. sacculated intestine.
sep. gl. septal gland.

sep. septum.

s. gl. suprapharyngeal glands.

s. org. b. sense organ cells in equatoreal zone.

8. org. pr. sense organ cells in prostomium.
spe. sperm-cells.

spd. spermduct.

spgn. spermatogonia.

spr. spur of nephridium.

S. ps. sac with penial seta.

sp. 8. Sperm-sacs.

8. ph. sgl. subpharyngeal glands.
spth. spermatheca.

spth. p. spermathecal pore.

t. testes.

t. m. transverse muscles.

trb. trabecula.

th. spd. thicker part of spermduct.
tu. tubular intestine.

tub. p. tubercula pubertatis.

ty. typhlosole.

u.c. anicellular glands.

v. g. ventral ganglion.

v. v. ventral vessel.

v. sgl. yentral gland.

w. ‘windings ” of nephridial ducts.

1

6

CALIFORNIA ACADEMY OF SCIENCES.

©

PLATE XLVI.
BENHAMIA PALMICOLA.

Longitudinal section of the genital somites.

BENHAMIA NANA.
A specimen; natural size.

The yentral side of the anterior somites, showing the exterior pores, etc.

The ventral side of the most anterior somites, showing the everted pharynx and the very narrow somite i, etc.

Side view of the same somites.

‘The anterior twenty somites laid open, exposing the various organs, ete. The alimentary canal is removed

and the whole figure is held somewhat schematic. atr. thickened part of the spermducts.

Part of a posterior somite viewed from the inner side, showing the relative position of sete and nephropores.
1, 2, 3, 4 refer to the set; a, b, c to the nephridia.

now om
is ol

ne ww

Mem. CAL.Acan.IT PLare XLVI

BENHAMIA PAPILLATA Fig 0. BENHAMIA NANA Figs. 170 6.

22TH. SAUTTEN & ASY SE

PACIFIC

OLIGOCH ETA.

REFERENCE TO LETTERS ON PLATES.

a. c. gl. anterior caleic gland.
a. f. anterior fold of nephridium.
a. nephr. anterior nephridia.
a. spd. anterior spermduct.
atr. atrium.

bd. body-wall.

bl. blood vessel or capillaries.
bl. s. blood sinus.

br. bridge in nephridium.
buc. buccal cavity.

c. clusters of lunate cells.
c.c. columnar cells.

c. ep. ccelomice epithelium.

c. gl. calciferous glands.

ch. discharge chambers of glands.
chlo. chloragogen cells.

cil. ciliated epithelium.

cl. c. clitellar cells.

el. clitellum.

c. m. circular muscles.

com. commissures.

c. r. ciliated rosettes or sperm-funnels.

c. t. connective tissue.

cu. cuticle.

d. gl. dark staining glands.
d. pr. dorsal pore.

d.v. dorsal vessel.

e. epithelial cells.

ept. epithelium.

gl. glands.

gl. c. unicellar glands.

gl. pr. glandular part of prostate.
gizz. gizzard.

h. heart.

hy. hypodermis.

i. s. gr. intersegmental groove.
l. c. large cells.

l. m. longitudinal muscles.
m. muscles.

m. atr. muscular atrium.
mes. mesenteric tissue.

m. pr. muscular prostate.
n. nephridia.

n. c. nerve cells.

ne. nerye ganglion.

nec. neck of nephridium.
neph. nephridium,

neph. p. nephropore.

es, cesophagus.

ov. ovary.

ovd. oviduct.

org. sense organ.

p. c. gl. posterior calciferous diverticulum.
p. f. posterior fold, nephridium.
phx. pharynx.

p. neph. posterior nephridia.
pr. prostate.

p. spd. posterior spermduct.
pr. sto. prostomium.

ps. penial set.

prt. peritoneum.

se. secreted matter.

s. int. sacculated intestine.
sep. gl. septal gland.

sep. septum.

s. gl. suprapharyngeal glands.

s. org. b. sense organ cells in equatoreal zone.

8. ory. pr. sense organ cells in prostomium.
spe. sperm-cells.

spd. spermduct.

spgn. spermatogonia,

spr. spur of nephridium.

8. ps. sac with penial sete.

sp. 8. sperm-sacs.

s. ph. sgl. subpharyngeal glands.
spth. spermatheca.

spth. p. spermathecal pore.

t. testes.

t. m. transverse muscles.

trb. trabecula.

th. spd. thicker part of spermduct.
tu. tubular intestine.

tub. p. tubercula pubertatis.

ty. typhlosole.

Z. c. unicellular glands.

2:

. g. ventral ganglion.
v. ventral vessel.
sgl. ventral gland.
. windings ” of nephridial ducts.

esge

178

CALIFORNIA ACADEMY OF SCIENCES.

PLATE XLVII.

BENHAMIA NANA.

A somewhat diagrammatic figure representing the anterior part of a specimen sectioned lengthwise, composed
from several sections, showing the location and size, etc., of various organs. 2. beginning of the muscular
part of the spermduct.

Cross-section through the most anterior part of the body, through prostomium and buccal division, showing
the pharyngeal glands in four distinct divisions. The section is anterior to the cephalic ganglion.

Section posterior to the former, through somite ii.
Section through somite vii.

Section through somite xiv; the calciferous gland with the intestine belongs to somite xiv; the ovary to xiii.
The section passes just in front of the oviduct. a. the limit of the clitellum.

Section through somite xvi. The calciferous diverticula belong to somite xv.

Section through somite xvii and the anterior prostate pore. zx. ending of clitellar cells.

Section through somite xx, showing the typhlosole.

Longitudinal section through the most anterior part of the body, showing the projected pharyngeal division
with the gland ducts. phx. pharyngeal division. The dark line indicates the row of discharge chambers of
the ducts from the glands. A more magnified view is represented in fig. 18.

A couple of strands of the unicellular salivary septal glands with their muscles.

Some of the septal unicellular glands more highly magnified.

Section through pharynx, showing the discharge ducts and pockets from the septal and pharyngeal glands.

Lining of the esophagus below the salivary giands, showing its exceeding thinness.

Longitudinal section through the body-wall, showing the sense organ region in the equatorial line of the
somite. c.e. central lumen or organ proper, in which are seen the sense cells, together with nuclei of
supporting cells. These supporting cells are surrounded by peculiar lunate or sac-like transparent cells, also
found in the sense organ of the pharynx. x. c. nerve cells and their nuclei.

Longitudinal section through gizzard, showing glands with their discharge tubes and small terminal pockets.

Section through the alimentary canal between the two gizzards, showing glands. c. cluster of lunate cells
surrounding a lumen.

Longitudinal section of the tubular intestine from somite xvi. w.c. unicellular glands. 6/. s. blood sinus.

One of the lobes of the typhlosole, showing distribution of glands, some of which are dark staining, others
not.

Section through the intestine adjoining the typhlosole.

Two glandular cells from the former section, more magnified. The two glands belong to different types.

A pair of calciferous diverticula from xiv. The right-hand gland is much smaller, which is an exception, as
generally they are of equal size.

Detail of a section of lamella from one of the anterior calciferous diverticula.

Two cells more highly magnified.

Section through one of the posterior calciferous diverticula.

Part of the same more highly magnified. Zeiss Hom. Im. Eyp., 2.

One of the prostates with sac with penial setz.

Cross-section through glandular part of prostate.

Part of a penial sete near the apex; the latter is broken.

XLVit

1. tia
Li. PLATE

iu.

{EM CAL.ACAI

PACIFIC COAST OLIGOCHXTA,

REFERENCE TO LETTERS ON PLATES.

a. c. gl. anterior caleic gland.
a. f. anterior fold of nephridium.
a. nephr. anterior nephridia.
a. spd. anterior spermduct
atr. atrium.

bd. body-wall.

bl. blood vessel or capillaries.
bl. s. blood sinus.

br. bridge in nephridium.
buc. buceal cavity.

c. clusters of lunate cells.

c. c. columnar cells.

c. ep. ecolomic epithelium.

c. gl. calciferous glands.

ch. discharge chambers of glands.
chlo. chioragogen cells.

cil. ciliated epithelium.

el. c. clitellar cells.

el. clitellum.

c. m. circular muscles.

com. commissures.

c. r. ciliated rosettes or sperm-funnels.

c. t. connective tissue.

cu. cuticle.

d. gl. dark staining glands.
d. pr. dorsal pore.

d. v. dorsal vessel.

e. epithelial cells.

ept. epithelium.

gl. glands.

gl. c. unicellar glands.

gl. pr. glandular part of prostate.
gizz. gizzard.

h. heart.

hy. hypodermis.

i. s. gr. intersegmental groove.
1. c. large cells.

1. m. longitudinal muscles.
m. muscles.

m. atr. muscular atrium.
mes. mesenteric tissue

m. pr. muscular prostate.
n. nephridia.

n. c. nerve cells.

ne. nerve ganglion.

nec. neck of nephridium.

neph. nephridium.

neph. p. nephropore.

ws, cesophagus.

ov. ovary.

ovd. oviduct.

org. sense organ.

p.c. gl. posterior calciferous diverticulum.
p. f. posterior fold, nephridium.
phx. pharynx. :

p. neph. posterior nephridia.

pr. prostate.

p. spd. posterior spermduct.

pr. sto. prostomium.

ps. penial sete.

prt. peritoneum.

se. secreted matter.

s. int. sacculated intestine.

sep. gl. septal gland.

sep. septum.

s. gl. suprapharyngeal glands.

s. org 6. sense organ cells in equatoreal zone.
8. org. pr. sense organ cells in prostomium.
spe. sperm-cells.

spd. spermduct.

spgn. spermatogonia.

spr. spur of nephridium.

8. ps. sac with penial sete.

sp. 8. sperm-sacs.

s. ph. sgl. subpharyngeal glands.
spth. spermatheca.

spth. p. spermathecal pore.

t. testes.

t. m. transverse muscles.

trb. trabecula.

th. spd. thicker part of spermduct.
tu. tubular intestine.

tub p. tubercula pubertatis.

ty. typhlosole.

u.c. unicellar glands.

v. g. ventral ganglion.

v. v. ventral vessel.

v. sgl. ventral gland.

w. ‘‘ windings” of nephridial ducts.

180

42.

43A.
43B.
43C.
43D.
43E.

444A.
44B.
440,
45.

CALIFORNIA ACADEMY OF SCIENCES.

PLATE XLVIIIL.

BENHAMIA NANA.
Section through one of the prostate pores, showing prostate, sac with penial sete and the thickened spermduct.
x. at this point the clitellar cells cease. c. ¢. connective tissue between the two muscular layers.
Cross-section of the thickened part of the spermduct, showing the two lumens separate.

Section through the muscular part of the prostate, showing the epithelium and some of the muscles nearest
the lumen.

One of the posterior nephridia; the shaded sac-like part represents the ccelomic mantle. It is generally covered
with capillaries to such an extent that the structure of the organ is obscured. br. bridge. nec. neck of nar-
row duct. spr.spur. o./f. anterior fold. p.f. posterior fold.

One of the anterior nephridia, showing absence of ccelomic glands, also the arrangements of blood vessels.
Part of the nephridium more magnified. No capillaries are represented.
Part of celomic mantle of one of the posterior nephridia. se. secreted matter staining very deep. zn. nuclei of

the glandular cells.

BENHAMIA PAPILLATA.

A specimen, natural size, from Tepic.

The ventral zone of the clitellum.

The same more highly magnified, showing the papill# and prostate pores.

The two penial sete in one sack. Zeiss D. Eyep., 2.

The tips of the two penial sete. Zeiss 12, Eyep., 2, 1.

BENHAMIA PALMICOLA.
The ventral side of the anterior somites.
The ventral region of the clitellum.
A specimen, natural size.

Penial sete. A. apex of the shorter sete. B. apex of the longer setw. C. two setw, whole.

Mine ioe Area TT Td pep UTAITIT
MEM LAL. ACAD ll, FLAIE AuvILL

SA

Le

i i ) ( D)

GSusmav Ersun Ten, . + ; Tr 9
HENHAMIA NANA Figs. 367042 UENHAMIA PAPILLAIA ic. 43

PACIFIC COAST OLIGOCH MTA. 15]

REFERENCE TO LETTERS ON PLATES.

a. c. gl. anterior calcic gland.
a. f. anterior fold of nephridium.
a. nephr. anterior nephridia.
a. spd, anterior spermduct.
atr, atrium.

bd. body-wall.

bl. blood vessel or capillaries.
bl. s. blood sinus.

br. bridge in nephridium.
buc. buccal cavity.

c. clusters of lunate cells.

c. c. columnar cells.

c. ep. celomie epithelium.

c. gl. calciferous glands.

ch. discharge chambers of glands.
chlo. chloragogen cells.

cil. ciliated epithelium.

el. c. clitellar cells.

cl. clitellum.

c. m. circular muscles.

com. commissures.

c. r. ciliated rosettes or sperm-funnels.

c. t. connective tissue.

cu. cuticle.

d. gl. dark staining glands.
d. pr. dorsal pore.

d. v. dorsal vessel.

e. epithelial cells.

ept. epithelium.

gl. glands.

gl. c. unicellar glands.

gl. pr. glandular part of prostate.
giz. gizzard.

h. heart.

hy. hypodermis.

i. s. gr. intersegmental groove.
1. c. large cells.

l. m. longitudinal muscles.
m. muscles.

m. atr. muscular atrium.
mes. mesenteric tissue.

m. pr. muscular prostate.
n. nephridia.

n. c. nerve cells.

ne, nerye ganglion.

nec. neck of nephridium.

neph. nephridium.

neph. p. nephropore.

es. esophagus.

ov. ovary.

ovd. oviduct.

org. sense organ.

p. c. gl. posterior calciferous diverticulum.
p. f. posterior fold, nephridium.
phx. pharynx.

p. neph. posterior nephridia.

pr. prostate.

p. spd. posterior spermduct.

pr. sto. prostomium.

ps. penial seta.

prt. peritoneum.

se. secreted matter.

s. int. sacculated intestine.

sep. gl. septal gland.

sep. septum.

s. gl. suprapharyngeal glands.

s. org. b. sense organ cells in equatoreal zone.
8. org. pr. sense organ cells in prostomium.
spe. sperm-cells.

spd. spermduct.

spgn. spermatogonia.

spr. Spur of nephridium.

8. ps. Sac with penial sete.

Sp. 8. sperm-sacs.

8. ph. sgl. subpharyngeal glands.
spth. spermatheca.

spth. p. spermathecal pore.

t. testes.

t. m. transverse muscles.

trb. trabecula.

th. spd. thicker part of spermduct.
tu. tubular intestine.

tub. p. tubercula pubertatis.

ty. typhlosole.

u. c. unicellular glands.

v. g. ventral ganglion.

v. v. ventral vessel.

v. sgl. ventral gland.

‘of nephridial ducts.

’

w. “windings

182

46.

52A-

52K.

52F.

52G-

52K-

CALIFORNIA ACADEMY OF SCIENCES.

PLATE XLIX.

BENHAMIA PALMICOLA.

A somewhat diagrammatic figure, composed from several longitudinal sections. s. org. pr. sense organ zone
of the prostomium. ss. org. b. the sense organ zone of the body wall, on the equatorial line of the somite.

Some of the sense organ cells of the prostomium more highly magnified. Zeiss. 12; Hom. Im. Eyrp. 2. The
sense cells are seen in the center between the pellucid sac-like cells. Between the sense cells are seen nuclei
of supporting cells.

Part of a posterior somite laid open, showing the three nephridia on one side.

A part of a nephridial cell mantle, showing various kinds of cells, some of which appear to contain calculi.

Cephalic ganglion, ete.

The spermathecal zone seen from the inner side of the body. The large globular apical part of each sperma-
theca is situated in the somite posterior to the pore.

Outlines of the spermatheew of various extra African species of Benhamia, some of which are copied from
various sources. ‘

B-C-D. Spermathecw of Benhamia Bolavi, from specimens from Hamburg, kindly furnished by Dr. W.
Michaelsen.

Spermatheca of Benhamia malayana after Dr. Horst.
Spermatheca of Benhamia floresiana after Dr. Horst.

H. Two spermathece from the same side of a specimen of Benhamia papillata from Tepic, Mexico.
Spermatheca of Benhamia octonephra after Rosa.

L. Spermathecw of Benhamia palmicola from Miraflores, Baja California.

52M-N. A spermatheca of Benhamia rugosa seen from two different sides.

SS

SSS SSS
wT = =

——

se
‘}
yj Ii
ao — A (se iS
, os «
) foe
j } |
ed We —
“Hf 1X
7 *y ) C ) 3 52A
dl. ( De =i

Fenhama Bola

G.

Fenhanua Bolavt E | |
| IN

———__—__,>
Benhanua Mealayana Benkamia Floyestana. Benjama pafellate

\ A=

| /
tena /

SS MM.
Bendamia rugest:

=a ; >

Penhemia oclone re

2 \ S Vew

\\ \ | Oo
(0) /

h / A ‘
JE |

Lenhamen nara

Lenhamea nana

— — + Benhame palmicola

ee = '
os a -
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e ‘
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} :
2
; a
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or <
i
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’
i!
\

PACIFIC

OLIGOCH MTA. 183

REFERENCE TO LETTERS ON PLATES.

a. c. gl. anterior caleic gland.
a. f. anterior fold of nephridium.
a. nephr. anterior nephridia.
a. spd. anterior spermduct.
atr. atrium.

bd. body-wall.

bl. blood vessel or capillaries.
bl. s. blood sinus.

br. bridge in nephridium.
buc. buceal cavity.

c. clusters of lunate cells.

c. c. columnar cells.

c. ep. coelomic epithelium.

c. gl. calciferous glands.

ch. discharge chambers of glands.
chlo. chloragogen cells.

cil. ciliated epithelium.

el. c. clitellar cells.

el. clitellum.

c. m. circular muscles.

com. commissures.

c. r. ciliated rosettes or sperm-funnels.

c. t. connective tissue.

cu. cuticle.

d. gl. dark staining glands.
d. pr. dorsal pore.

d.v. dorsal vessel.

e. epithelial cells.

ept. epithelium. a

gl. glands.

gl. c. unicellar glands.

gl. pr. glandular part of prostate.
gizz. gizzard.

h. heart.

hy. hypodermis.

7, s. gr. intersegmental groove.
l. c. large cells.

1. m. longitudinal muscles.
m. muscles.

m. atr. muscular atrium.
mes. mesenteric tissue.

m. pr. muscular prostate.
n. nephridia.

n.c. nerve cells.

ne. nerve ganglion,

nec. neck of nephridium.

neph. nephridium,

neph. p. nephropore.

es. esophagus.

ov. ovary.

ovd. oviduct.

org. Sense organ.

p.c. gl. posterior calciferous diverticulum.
p. f. posterior fold, nephridium.
phx. pharynx.

p. neph. posterior nephridia.

pr. prostate.

p- spd. posterior spermduct.

pr. sto. prostomium.

ps. penial seta.

prt. peritoneum.

se. secreted matter.

s. int. sacculated intestine.

sep. gl. septal gland.

sep. septum.

s. gl. suprapharyngeal glands.

s. org. 6. sense organ cells in equatoreal zone.
8. org. pr. sense organ cells in prostomium.
spc. sperm-cells.

spd. spermduct.

spgn. spermatogonia.

spr. spur of nephridium.

8. ps. sac with penial seta.

sp. 8. sperm-sacs.

s. ph. sgl. subpharyngeal glands.
spth. spermatheca.

spth. p. spermathecal pore.

t. testes.

t. m. transverse muscles.

trb. trabecula.

th. spd. thicker part of spermduct.
tu. tubular intestine.

tub. p. tubercula pubertatis.

ty. typhlosole.

u. c. unicellular glands.

v. g. ventral ganglion.

v. v. ventral vessel.

v. sgl. ventral gland.

w. “ windings ” of nephridial ducts.

184

53.

CALIFORNIA ACADEMY OF SCIENCES.

PLATE L.
BENHAMIA PALMICOLA.

Part of the intestine and hearts dissected out.
Prostate and sac with penial seta.
BENHAMIA RUGOSA.
The ventral side of clitellum, showing the copulatory papill# on which are situated the anterior prostate pores
in xyil. The clitellum is much crenulated.
Anterior somites seen from aboye, showing prostomium, somite i, ii and ili, the first of which is much swollen.
The same seen from the side. In these two figures somite i is retracted in the buccal cavity.
Another specimen; the anterior somites, but i, is fully extended.
Larger penial set, free end.
Smaller penial setz.

A partly diagrammatic representation of the interior surface of a posterior somite, showing the nephridia.
1, 2, 3, 4, small, refer to sete; 1, 2, 3, 4, large, refer to nephridia.

ALEODRILUS KEYESI.
A specimen, natural size.

The anterior somites seen from the ventral side, showing the generative pores and the position of set.
Prostomium and somite i.
The sexual foss# and papille.

A somewhat diagrammatic view of the anterior somites, composed from several longitudinal sections.

es

PACIFIC

REFERENCE

R

.c. gl. anterior calcic gland.
. f. anterior fold of nephridium.
a. nephr. anterior nephridia.
a. spd. anterior spermduct.
atr. atrium.

bd. body-wall.

bl. blood vessel or capillaries.
bl. s. blood sinus.

br. bridge in nephridium.
buc. buccal cavity.

c. clusters of lunate cells.
c.c. columnar cells.

c. ep. ecelomic epithelium.

c. gl. calciferous glands.

ch. discharge chambers of glands.
chlo. chloragogen cells.

cil. ciliated epithelium.

cl. c. clitellar cells.

cl. clitellum.

c. m. circular muscles.

com. commissures.

a

c. r. ciliated rosettes or sperm-funnels.

c. t. connective tissue.

cu. cuticle.

d. gi. dark staining glands.
d. pr. dorsal pore.

d. v. dorsal vessel.

e. epithelial cells.

ept. epithelium.

gl. glands.

gl. c. unicellar glands.

gl. pr. glandular part of prostate.
gizz. gizzard.

h. heart.

hy. hypodermis.

i. s. gr. intersegmental groove.
1. c. large cells.

l. m. longitudinal muscles.
m. muscles.

m. atv. muscular atrium.
mes. mesenteric tissue.

m. pr. wauscular prostate.
n. nephridia.

n. c. nerve cells.

ne. nerve ganglion,

COAST OLIGOCH ATA. 185

TO LETTERS ON PLATES.

nec. neck of nephridium.

neph. nephridium.

neph. p. nephropore.

es, cesophagus.

ov. ovary.

ovd. oviduct.

org. sense organ,

p. c. gl. posterior calciferous diverticulum.
p.f. posterior fold, nephridium.
phx. pharynx.

p. neph. posterior nephridia.

pr. prostate.

p- spd. posterior spermduct.

pr. sto. prostomium.

ps. penial sete.

prt. peritoneum.

se. secreted matter.

s. int. sacculated intestine.

sep. gl. septal gland.

sep. septum.

s. gl. suprapharyngeal glands.

s. org. b. sense organ cells in equatoreal zone.
8. ory. pr. sense organ cells in prostomium.
spc. sperm-cells.

spd. spermduct.

spgn. spermatogonia.

spr. spur of nephridium.

s. ps. sac with penial sete.

sp. 8. Sperm-sacs.

s. ph. sgl. subpharyngeal glands.
spth. spermatheca.

spth. p. spermathecal pore.

t. testes.

t. m. transverse muscles.

trb. trabecula.

th. spd. thicker part of spermduct.
tu. tubular intestine.

tub. p. tubercula pubertatis.

ty. typhlosole.

u. c. unicellular glands.

v. g. ventral ganglion.

v. v. ventral vessel.

v. sgl. ventral gland.

w. “ windings ” of nephridial ducts.

80.

81.

CALIFORNIA ACADEMY OF SCIENCES.

PLATE LI.
ALEODRILUS KEYESI.

Cross-section through somite ix, showing the septal glands and the thickened septa. ss. gi. septal gland, the
prolongation of which probably connects with the intestine and the median blood vessel. «x. at this point the
ventral vessel and the septal gland have been bent and cut twice. 5

Cross-section through somite xii, showing the posterior sperm-sac. tra. trabecula. v. v. ventral vessel in two
successive somites. spc. sperm-cell germinative area of cells with narrow nuclei.

Section through apical part of spermatheca.

Section through intestine and septal gland in somite x, anterior to fig. 72, showing discharge duct of septal
gland. The ventral vessel and heart have been folded and cut three different times; similarly part of the septal
gland has been cut twice. d. discharge duct of septal gland; it probably reaches in between the epithelia)
cells of the intestine.

Section through dorsal median vessel, where it connects with two lateral vessels; the section goes only through
one of the points of connection. The lateral vessels are covered with chloragogen cell.

Section through the suspended posterior part of the spermduct in the anterior part of somite xii. The duct is
seen suspended between two mesenteric strands, one connecting with the heart, the other with the septum
close to the body wall. h.v. heart valve. gl. gland emptying in spermduct.

Section through spermduct in somite xi, the cut going through testis, which is immediately superposed on the
duct or rather the posterior part of the ciliated rosette.

Section through spermduct going through the ovary in xiii. a. spd. spermduct from anterior rosette, already
attached to the body wall. p. spd. spermduct from posterior rosette, yet suspended and passing through the
ovary. ov. ovary with full-grown ova, with germ-cells and with a germinative area where cells are in mitosis.
The nuclei in the germ-cells are oval, while those in the grown ova are round. Zeiss D. Eyp. 2.

Section through spermduct through one of the posterior somites where the two ducts have joined in the body-
wall. clit. clitellum in xiv.

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PACIFIC COAST OLIGOCH TA.

REFERENCE TO LETTERS ON PLATES.

a.c. gl. anterior caleic gland.
a. f. anterior fold of nephridium.
a. nephr. anterior nephridia.
a. spd. anterior spermduct
atr. atrium.

bd. body-wall.

bl. blood vessel or capillaries.
bl. s. blood sinus.

br. bridge in nephridium.
buc. buccal cavity.

c. clusters of lunate cells.

c. c. columnar cells.

c. ep. ccelomic epithelium.

c. gl. calciferous glands.

ch. discharge chambers of glands.
chlo. chloragogen cells.

cil. ciliated epithelium.

cl. c. clitellar cells.

el. clitellum.

c. m. circular muscles.

com. commissures.

ec. r. ciliated rosettes or sperm-funnels.

c. t. connective tissue.
cu. cuticle.
d. gl. dark staining glands.
d. pr. dorsal pore.
d. v. dorsal vessel.
e. epithelial cells.
ept. epithelium.
gl. glands.
gl. c. unicellar glands.
_ gl. pr. glandular part of prostate.
gizz. gizzard.
h. heart.
hy. hypodermis.
i. s. gr. intersegmental groove.
l. c. large cells.
l. m. longitudinal muscles.
m. muscles.
m. atr. muscular atrium.
mes. mesenteric tissue
m. pr. muscular prostate.
n. nephridia.
n. c. nerve cells.
ne. nerve ganglion. ;

nec. neck of nephridium.
neph. nephridium.

neph. p. nephropore.

ws. esophagus.

ov. ovary.

ovd. oviduct.

org. sense organ.

p.c. gl. posterior calciferous diverticulum.
p.f. posterior fold, nephridium.
phe. pharynx.

p- neph. posterior nephridia.
pr. prostate.

p. spd. posterior spermduct.
pr. sto. prostomium.

ps. penial seta.

prt. peritoneum.

se. secreted matter.

s. int. sacculated intestine.
sep. gl. septal gland.

sep. septum.

s. gl. suprapharyngeal glands.

s. org b. sense organ cells in equatoreal zone.

8. org. pr. sense organ cells in prostomium.
spe. sperm-cells.

spd. spermduct.

spgn. spermatogonia.

spr. spur of nephridium.

8. ps. sac with penial sete.

sp. 8. sperm-sacs.

s. ph. sgl. subpharyngeal glands.
spth. spermatheca.

spth. p. spermathecal pore.

t. testes.

t. m. transverse muscles.

trb. trabecula.

th. spd. thicker part of spermduct.
tu. tubular intestine.

tub p. tubercula pubertatis.

ty. typhlosole.

u. c. unicellar glands.

v. g. ventral ganglion.

v. v. ventral vessel.

v. sgl. ventral gland.

w. ‘windings ” of nephridial ducts.

CALIFORNIA ACADEMY OF SCIENCES.

PLATE LIL.

ALEODRILUS KEYESI.
Section through the epithelium of tubular intestine, showing a cluster of glands between the epithelial cells.
bl. blood lacune.
Section through one of the valves of the heart. /. c. large cells at the base of the valves.
Some spermatogonia from the sperm-sacs in xii.
Cells from the central germinative and in the sperm-sac.

One of the nephridia with an anterior lateral vessel, connecting dorsally with a branch from the ventral vessel.
d.v. diverticula of the lateral vessel. Jat. v. lateral vessel.

ACANTHODRILUS TAMAJUSI.

The largest of my specimens, natural size. This specimen is strongly contracted, and was when alive con-
siderably longer.

The anterior somites, ventral view.

The anterior somites, dorsal view.

The anterior somites, side view.

The ventral part of the genital region in the clitellum, showing genital papillw, the three transverse genital
ridges, penial set#, common seta, etc. The clitellum is not well developed, but in somite xx may be seen its

posterior termination. The other one of my specimens, which possessed a more developed clitellum, had
these papille less pronounced, though of the same form and arrangement.

Cross-section of the body-wall, showing the common sete.
The free end of the penial seta.
The spermathece, dorsal view.

One of the spermathece dissected out.

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PACIFIC COAST OLIGOCHATA. 189

REFERENCE TO LETTERS ON PLATES.

a. c. gl. anterior caleic gland.
a. f. anterior fold of nephridium.
a. nephr. anterior nephridia.
a. spd. anterior spermduet.
atr. atrium.

bd. body-wall.

bl. blood vessel or capillaries.
bl. s. blood sinus.

br. bridge in nephridium.
buc. buccal cavity.

c. elusters of lunate cells.

c. c. columnar cells.

c. ep. celomic epithelium.

c. gl. ealciferous glands.

ch. discharge chambers of glands.
chlo. chloragogen cells.

cil. ciliated epithelium.

cl. c. clitellar cells.

cl. clitellum.

c. m. circular muscles.

com. commissures.

c. r. ciliated rosettes or sperm-funnels.

c. t. connective tissue.

cu. cuticle.

d. gl. dark staining glands.
d. pr. dorsal pore.

d. v. dorsal vessel.

e. epithelial cells.

ept. epithelium.

gl. glands.

gl. c. unicellar glands.

gl. pr. glandular part of prostate.
gizz. gizzard.

h. heart.

hy. hypodermis.

i. s. gr. intersegmental groove.
l. c. large cells.

l. m. longitudinal muscles.
m. muscles.

m. atr. muscular atrium.
mes. mesenteric tissue.

m. pr. muscular prostate.
n. nephridia.

n.c. nerve cells.

ne. nerve ganglion.

nec. neck of nephridium.

neph. nephridium,

neph. p. nephropore.

ws. cesophagus.

ov. ovary.

ovd. oviduct.

org. sense organ.

p. c. gl. posterior calciferous diverticulum.
p. f. posterior fold, nephridium.
pha. pharynx.

p. neph. posterior nephridia.

pr. prostate.

p. spd. posterior spermduct.

pr. sto. prostomium.,

ps. penial sete.

prt. peritoneum.

se. secreted matter.

s. int. sacculated intestine.

sep. gl. septal gland.

sep. septum.

s. gl. suprapharyngeal glands.

s. org. b. sense organ cells in equatoreal zone.
8. org. pr. Sense organ cells in prostomium.
spe. sperm-cells.

spd. spermduct.

spgn. spermatogonia.

spr. spur of nephridium.

8, ps. sac with penial seta.

sp. 8. sperm-sacs.

s. ph. sgl. subpharyngeal glands.
spth. spermatheca.

spth. p. spermathecal pore.

t. testes.

t. m. transverse muscles.

irb. trabecula.

th. spd. thicker part of spermduct.
tu. tubular intestine.

tub. p. tubercula pubertatis.

ty. typhlosole.

u. c. unicellular glands.

v. g. ventral ganglion.

v. v. ventral vessel.

v. sgl. ventral gland.

w. ‘windings ” of nephridial ducts.

190 CALIFORNIA ACADEMY OF SCIENCES.

PLATE LIII.
ACANTHODRILUS TAMAJUSI.

96. A somewhat diagrammatic view composed from several longitudinal sections of the body. +. m. retractor mus-
cles of gizzard. c.c. masses of free ccelomic cells.

SPARGANOPHILUS BENHAMI.

97. Adult specimen, natural size.

98. The anterior somites, ventral view, including clitellum, tubercula pubertatis, etc., magnified four times.

99. Longitudinal section of the anterior somites passing through the spermathecal and prostate pores, composed
from seyeral sections. Somites xix-xxii are not figured. sep. gl. septal glands which open in the dorsal part
of the pharynx. J. sep. gl. lower septal glands which open in the ventral part of the pharynx. The sperma-
thece have been cut through diagonally and the figure does not give any idea of their real shape.

101. Section of the body-wall in yentral part of somite xii. p. peritoneum. 2. ep. nuclei of supporting cells.

102. Longitudinal section of the clitellum. 7. Supporting cells. 2. Unicellular glands. 3. Short clitellar cells,

staining deeply, and furnished with perfect nuclei. 4. Largest clitellar cells, staining less deeply and with
imperfect nuclei. Zeiss D.

103. A part of the same section, more magnified. Iron, hematoxylin, Haidenhein, Ehrlich 3 acid. Numerals indi-
cate the same as in fig. 102. d.n. degenerate nuclei of the 4 cells. x. sec. cell plasma around the nuclei
pushed far back in the apical part of the cell. x. sh. c. nuclei of the short clitellar cells surrounded by cell
plasma and mucin. ec. ¢. connective tissue. J. c. blood capillaries which are very scarce in the clitellum of
this species.

104. Section of the body wall in somite xiv.

105. Cross-section through clitellum and tubercula pubertatis. 7. Supporting cells. 2. Unicellular glands. 3.
Shorter clitellar cells staining darkly. 4 The longer clitellar cells. 5. tpc. tubercula pubertatis
cells, largest kind. A few of these are seen to contain free but shrunken nuclei. 6. tpc. tubercula puber-
tatis cells of the narrow kind, with cell plasma in the apical end, but also with shrunken nuclei. ¢. p. tuber-
cula pubertatis groove. x. /. nerve fibers. cl. c. 2. large unicellular glands, a large form of the usual uni-
cellular epidermal glands. Above these is seen a wide lumen, probably a blood vessel.

106. Longitudinal section of the body-wall, through spermatheca.

107. Longitudinal section of body-wall through prostate and pore in somite xxiy. ¢.c. tubular supporting cells.
g.c. glandular cells confined to the zone around the intersegmental grooves.

108. The same as 107, more magnified.

109. The inner whorl of the ciliated rosette.

110. Section through the esophagus. i. c. interstitial cells. g/. interstitial glands.

111. Section through intestine in somite xii. fbr. blood crystals and fibrine. w. bl. w. wall of blood sinus.

112. Section through upper pharyngeal wall, showing epithelial cells, and between them the discharge tubes of the
glands. Zeiss 12, hom. im.

113. Section through the same part, showing a three-forked duct.

114. Section through the intestinal wall, next posterior to the real pharynx, showing larger discharge tubes and
chambers.

115. Section through ventral wall of pharynx, with ducts and storage chambers of glands, and between them shorter
epithelial cells. m. and d. muscles along which run the ducts. x. d. nuclei of ducts.

116. Surface view of one of the larger blood vessels.

117. Outline of one of the nephridia.

\18. Outline of a spermatheca,

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PACIFIC

COAST OLIGOCH MTA.

REFERENCE TO LETTERS ON PLATES.

a. c. gl. anterior calcic gland.
a. f. anterior fold of nephridium.
a. nephr. anterior nephridia.
a. spd. anterior spermduct.
atr. atrium.

bd. body-wall.

bl. blood vessel or capillaries.
bl. s. blood sinus.

br. bridge in nephridium.

buc. buecal cavity.

ce. clusters of lunate cells.

c. c, columnar cells.

c. ep. ccelomic epithelium.

c. gl. calciferous glands.

ch. discharge chambers of glands.
chlo. chloragogen cells.

cil. ciliated epithelium.

cl. c. clitellar cells.

el. clitellum.

c. m. circular muscles.

com. commissures.

ce. r. Ciliated rosettes or sperm-funnels.
c. t. connective tissue.

cu. cuticle.

d. gl. dark staining glands.

d. pr. dorsal pore.

d.v. dorsal vessel.

e. epithelial cells.

ept. epithelium.

gl. glands.

gl. c. unicellar glands.

gl. pr. glandular part of prostate.
gizz. gizzard.

h. heart.

hy. hypodermis.

7. 8. gr. intersegmental groove.
l. c. large cells.

1. m. longitudinal muscles.

m. muscles. >

m. atr. muscular atrium.

mes. mesenteric tissue.

m. pr. muscular prostate.

n. nephridia.

n.c. nerve cells.

ne. nerve ganglion.

nec. neck of nephridium.
neph. nephridium.

neph. p. nephropore.

@s. esophagus.

ov. ovary.

ovd. oviduct.

org. sense organ.

p.c. gl. posterior calciferous diverticulum.
p. f. posterior fold, nephridium.
phx. pharynx.

p. neph. posterior nephridia.
pr. prostate.

p. spd. posterior spermduet.
pr. sto. prostomium.

ps. penial sete.

prt. peritoneum.

se. secreted matter.

s. int. sacculated intestine.
sep. gl. septal gland.

sep. septum.

s. gl. suprapharyngeal glands.

s. org. b. sense organ cells in equatoreal zone.

8. org. pr. sense organ cells in prostomium.
spe. sperm-cells.

spd. spermduct.

spgn. spermatogonia.

spr. spur of nephridium.

8. ps. sac with penial seta.

sp. 8. Sperm-sacs.

8. ph. sgl. subpharyngeal glands.
spth. spermatheca.

spth. p. spermathecal pore.

t. testes.

t. m. transverse muscles.

trb. trabecula.

th. spd. thicker part of spermduct.
tu. tubular intestine.

tub. p. tubercula pubertatis.

ty. typhlosole.

wu. c. unicellular glands.

v. g. ventral ganglion.

v. v. ventral vessel.

v. sgl. ventral gland.

w. ‘‘ windings ” of nephridial ducts.

191

192 CALIFORNIA ACADEMY OF SCIENCES.

' PLATE LIV.
SPARGANOPHILUS BENHAMI.

119. Five cross-sections through various parts of the body.

a. Through posterior part of pharynx in somite v. s. g/. dorsal septal glands. J. s. gl. lower septal glands.

b. Through spermathec and somites viii and ix.

c. Through somite x. ¢.c. 7. tube of ciliated rosette.

d. Through oviduct, somite xiv with sperm-sacs.
Through clitellum in somite xix, showing tubercula pubertatis.
clitellar cells. spd. spermduct running below the muscular layers.

e. cl. c. 1 long clitellar cells. cl. c. 2 short

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PACIFIC

COAST OLIGOCHATA. 193

REFERENCE TO LETTERS ON PLATES.

.¢. gl. anterior calcic gland.
. f. anterior fold of nephridium.
a. nephr. anterior nephridia.
a. spd. anterior spermduct.
atr. atrium.

bd. body-wall.

bl. blood vessel or capillaries.
bl. s. blood sinus.

br. bridge in nephridium.
buc. buccal cavity.

c. clusters of lunate cells.

c. c. columnar cells.

c. ep. celomic epithelium.

c. gl. calciferous glands.

ch. discharge chambers of glands.
chlo. chloragogen cells.

cil. ciliated epithelium.

cl. c. clitellar cells.

cl. clitellum.

c. m. cireular muscles.

com. commissures.

Re a

c. r. ciliated rosettes or sperm-funnels.

c. t. connective tissue.

cu. cuticle.

d. gl. dark staining glands.
d. pr. dorsal pore.

d.v. dorsal vessel.

e. epithelial cells.

ept. epithelium.

gl. glands.

gl. c. unicellar glands.

gl. pr. glandular part of prostate.
gizz. gizzard.

h. heart.

hy. hypodermis.

i. s. gr. intersegmental groove.
l. c. large cells.

l. m. longitudinal muscles.
m. muscles.

m. atr. muscular atrium.
mes. mesenteric tissue.

m. pr. muscular prostate.
n. nephridia.

n. c. nerve cells,

ne. nerve ganglion,

nec. neck of nephridium.

neph. nephridium.

neph. p. nephropore.

es, esophagus.

ov. Ovary.

ovd. oviduct.

org. sense organ.

p. c. gl. posterior calciferous diverticulum.
p. f. posterior fold, nephridium.
phx. pharynx.

p. neph. posterior nephridia.

pr. prostate.

p. spd. posterior spermduct.

pr. sto. prostomium.

ps. penial sete.

prt. peritoneum.

se. secreted matter.

s. int. sacculated intestine.

sep. gl. septal gland.

sep. septum.

s. gl. suprapharyngeal glands.

s. org. b. sense organ cells in equatoreal zone.
8. ory. pr. sense organ cells in prostomium.
spe. sperm-cells.

spd. spermduct.

spgn. spermatogonia.

spr. spur of nephridium.

s. ps. sac with penial seta.

sp. 8. Sperm-sacs.

s. ph. sgl. subpharyngeal glands.
spth. spermatheca.

spth. p. spermathecal pore.

t. testes.

t. m. transverse muscles.

trb. trabecula.

th. spd. thicker part of spermduct.
tu. tubular intestine.

tub. p. tubercula pubertatis.

ty. typhlosole.

u. c. unicellular glands.

v. g. ventral ganglion.

v. v. ventral vessel.

v. sgl. ventral gland. :

w. ‘‘ windings ” of nephridial ducts.

194

123.

124.

126.

127.
128.

129.

CALIFORNIA ACADEMY OF SCIENCES.

PLATE LV.

SPARGANOPHILUS SMITHI.

A large specimen, natural size, ventral view.

Anterior part of the body, side view, showing tubercula pubertatis ridges, and their anterior extension to
the center of somite xi. The dorsal and ventral termini of clitellum marked by x.

The anterior somites, ventral view. The spermiducal pore is seen about half way between set 1 and 2 aid
the tubercula pubertatis in xix/xi.

Two of the clitellar somites, side view, showing tubercula pubertatis ridge. v. ventral. d. dorsal side.

SPARGANOPHILUS SONOM2.

The same as 122, but from Sparganophilus sonome, showing the tubercula pubertatis ridge to consist of a suc-
cession of separate tubercles, two or three on every somite, with the groove very nearly in the center of each
papilla.

SPARGANOPHILUS SMITHI.

Longitudinal section of the anterior somites, composed from seyeral sections. v. sg/. ventral septal glands in
vy, vi, vii. s.v.spermtanks. s. s. sperm-sacs. f. four large folds of the dorsal vessel in somites xiy—xvili.
sp. gl. prostates.

A series of outline drawings of spermathec# from one specimen.
SPARGANOPHILUS SONOM 2.
Outlines of spermathece from a single specimen.
SPARGANOPHILUS SMITHI.
Cross-section of septum xi/xii.

Cross-section of the body; the Roman numeral indicates the somite.

Cross-section of the body; the Roman numerals indicate the somites.

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PACIFIC COAST OLIGOCH ATA.

REFERENCE TO LETTERS ON PLATES.

a. c. gl. anterior calcic gland.
a. f. anverior fold of nephridium.
a. nephr. anterior nephridia.
a. spd. anterior spermduct
atr. atrium.

bd. boey-wall.

bl. blood vessel or capillaries.
bl. s. blosd sinus.

br. bridge in nephridium.
buc. buccal cavity.

c. clusters of lunate cells.

c. c. columnar cells.

c. ep. eelomic epithelium.

c. gl. ealeiferous glands.

ch. discharge chambers of glands.
chlo. chioragogen cells.

cil. ciliated epithelium.

cl. c. clitellar cells.

cl. clitellum.

c. m. circular muscles.

com. commissures.

c. r. ciliated rosettes or sperm-funnels.

c. t. connective tissue.

cu. euticle.

d. gl. dark staining glands.
d. pr. dorsal pore.

d. v. dorsal vessel.

e. epithelial cells.

ept. epithelium.

gl. glands.

gl. c. unicellar glands.

_ gl. pr. glandular part of prostate.
giz. gizzard.

h. heart.

hy. hypodermis.

i. s. gr. intersegmental groove.
1. c. large cells.

l. m. longitudinal muscles.
m. muscles.

m. atr. muscular atrium.
mes. mesenteric tissue

m. pr. muscular prostate.
n. nephridia.

mn. c. nerve cells,

ne. nerve ganglion.

nec. neck of nephridium.

neph. nephridium.

neph. p. nephropore.

ws. esophagus.

ov. ovary.

ovd. oviduct.

org. sense organ.

p.c. gl. posterior calciferous diverticulum.
p. f. posterior fold, nephridium.
phx. pharynx.

p- neph. posterior nephridia.

pr. prostate.

p. spd. posterior spermduct.

pr. sto. prostomium.

ps. penial seta.

prt. peritoneum,

se. secreted matter.

s. int. sacculated intestine.

sep. gl. septal gland.

sep. septum.

s. gl. suprapharyngeal glands.

s. org b. sense organ cells in equatoreal zone.
8. org. pr. sense organ cells in prostomium.
spe. sperm-cells.

spd. spermduct.

spgn. spermatogonia.

spr. spur of nephridium.

s. ps. Sac with penial sete.

Sp. 8. sperm-sacs.

s. ph. sgl. subpharyngeal glands.
spth. spermatheca.

spth. p. spermathecal pore.

t. testes.

t. m. transverse muscles.

trb. trabecula.

th. spd. thicker part of spermduct.
tu. tubular intestine.

tub p. tubercula pubertatis.

ty. typhlosole.

u. c. unicellar glands.

v. g. ventral ganglion.

v. v. ventral vessel.

v. sgl. ventral gland.

w. ‘‘ windings” of nephridial ducts.

€

5

196 CALIFORNIA ACADEMY OF SCIENCES.

PLATE LVL.

SPARGANOPHILUS SMITHI.

130-136. Cross-sections through various somites. The details are indicated diagrammatically. The Roman numer-
als indicate the number of the somite or somites through which the section passes. s. m. septal muscles.
a. s. gl. accessory septal gland of septum between somites ixand x. spz. ¢. spermatozoa in spermtanks. spz. s.
spermatozoa and spermatogonia in sperm-sacs. os. ovisac. a.m. ¢. arciform muscles of tubercula puber-
tatis, connecting the opposite surfaces of the tubercula pubertatis. a. m.c. arciform muscles passing from
opposite sides of clitellum through the cwelomic cavity.

137. Section through sacculated intestine. 06]. c. blood capillary reticulum.

138. Tubercula pubertatis, Zeiss Hom. Im. ,';. af. m. arciform muscles passing through celom. af. t. arciform
muscles confined to the epidermal cells of the tubercula.

139. Section through the body-wall through one of the prostates. 7. m. retractor muscles passing to the cuticle
and the outermost epidermal cells. ct. connective tissue.

SPARGANOPHILUS CARNEUS.

140. Two spermathece from the same specimens. The free or apical end is seen to be almost globular.

SPARGANOPHILUS GUATEMALENSIS.

141, aand b. Two spermathecw. The apical end is flat and trawl-like, bearing a sac-like pouch which connects
with the interior of the spermatheca, through a small slit.

DELTANIA TROYERI var. CRASSA.
142. A prostate gland with penial sete.
143. A spermatheca.
DELTANIA TROYERI var. LAGUN.
144. One of the penial set with the tip more highly magnified.
145. Two spermathecs, both from the same specimen. 4A and B are seen from the broad side, C from the narrow
side.
146. Two prostates with penial set. A is seen from the flat side, B is seen from the broad side.

147. Section of one of the ovisacs. Zeiss 4, Hom. Im.

{40%

Beary Bese Diet,

OPARGANOPHILUS SMITH! Figs. 130 To 199

SPARGANOPHILUS CARNEUS Fig. 140.

GPARGANOPHILUS GUATEMALENSIS Fis. 141. [ELIANA TROVERI VAR. CRASSA Figs. 142 & 143 TIELTANIA TROYER! VAR. LAGUNE Figs. 144 70 147

|

PACIFIC

COAST OLIGOCH MTA. 197

REFERENCE TO LETTERS ON PLATES.

a. c. gl. anterior calcie gland.
a. f. anterior fold of nephridium.
a. nephr. anterior nephridia.
a. spd. anterior spermduet.
atr. atrium.

bd. body-wall.

b1. blood vessel or capillaries.
bl. s. blood sinus.

br. bridge in nephridium.
buc. buccal cavity.

c. clusters of lunate cells.

c. c. columnar cells.

c. ep. celomic epithelium.

c. gl. calciferous glands.

ch. discharge chambers of glands.
chlo. chloragogen cells.

cil. ciliated epithelium.

cl. c. clitellar cells.

el. clitellum.

c. m. circular muscles.

com. commissures.

c. 7. ciliated rosettes or sperm-funnels.

c. t. connective tissue.

cu. cuticle.

d. gl. dark staining glands.
d. pr. dorsal pore.

d.v. dorsal vessel.

e. epithelial cells.

ept. epithelium.

gl. glands.

gl. c. unicellar glands.

gl. pr. glandular part of prostate.
' gizz. gizzard.

h. heart.

hy. hypodermis.

i. s. gr. intersegmental groove.
1. c. large cells.

l. m. longitudinal muscles.
m. muscles.

m. atr. muscular atrium.
mes. mesenteric tissue.

m. pr. muscular prostate.
n. nephridia.

n. c. nerve cells.

ne. nerve ganglion.

nec. neck of nephridium.

neph. nephridium.

neph. p. nephropore.

@s. esophagus.

ov. ovary.

ovd. oviduct.

org. sense organ.

p.c. gl. posterior calciferous diverticulum.
p.f. posterior fold, nephridium.
phe. pharynx.

p. neph. posterior nephridia.

pr. prostate.

p. spd. posterior spermduet.

pr. sto. prostomium.

ps. penial sete.

prt. peritoneum.

se. secreted matter.

s. int. sacculated intestine.

sep. gl. septal gland.

sep. septum.

s. gl. suprapharyngeal glands.

s. org. b. sense organ cells in equatoreal zone.
8. org. pr. sense organ cells in prostomium.
spe. sperm-cells.

spd. spermduct.

spgn. spermatogonia.

spr. spur of nephridium.

Ss. ps. sac with penial sete.

sp. 8. sperm-sacs.

s. ph. sgl. subpharyngeal glands.
spth. spermatheca.

spth. p. spermathecal pore.

t. testes.

t. m. transverse muscles.

trb. trabecula.

th. spd. thicker part of spermduct.
tu. tubular intestine.

tub. p. tubercula pubertatis.

ty. typhlosole.

u. c. unicellular glands.

. g. ventral ganglion.

v. v. ventral vessel.

I~

i~

. sgl. ventral gland.
w. ‘windings ” of nephridial ducts.

.

198 CALIFORNIA ACADEMY OF SCIENCES.

PLATE LVII.
ACANTHODRILUS VASLITI.

148. The largest specimen, natural size, but not yet adult; the clitellum not developed.

149. Anterior part of a young specimen; clitellum not developed. The nephridia and some other detail not figured.
s. pl. sgl. subpharyngeal salivary gland. prc. ss. peritoneal cells forming solid masses, probably surrounding
rudimentary sperm-sacs. prc. large peritoneal cells lining the septa as well as the peritoneum. a. p. pr.
anterior pair of prostates opening close together. p. p. pr. posterior pair of prostates.

150. Part of the ventral part of the body in somites xvii, xviii, xix laid open, showing prostates and thick septa
with peritoneal cells. m. copulatory muscles. «a. pr. anterior prostate. p. pr. posterior prostate both of
the same pair.

151. Cross-section of the body-wall in somite xix, showing the prostate pore, and the two prostates opening close
together.

152. Section through the same pore, three or four sections further back, showing the tubercula spubertabe sense
organ, ete. deb. debris. sup. c. supporting cells.

153. Part of a septum, showing the very large peritoneal cells, some of which are broader, others longer and nar-
rower.

154. Cross-section of the two prostates. p. pr. posterior prostate pore. a. pr.anterior prostate pore. prt. perito-
neal cells and connective tissue connecting the prostates with the body wall.

PH@NICODRILUS TEPICENSIS.
155. Spermathecs, side and front view of one and the same.

156. Cross-section of the body in somite xvii through the male pore, showing the copulatory region with muscular
atrium and spermduct. The longitudinal muscular layer is greatly increased at the male pore. The muscu-
lar atrium is seen in cross-section, the cut passing through the whole length of that organ. This is also
seen in figures 158, 159, the latter being taken from a section posterior to the former.

157. The oviduct, ovisac and part of the ovaries, interior view of the body.

158. Section through muscular atrium in cross-section of the body. The spermducts have not yet united, nor have
they entered the atrium. The section passes through the whole length of the atrium. The arciform mus-
cles must have been torn, as they did not show in the section.

159. The same parts as seen in fig. 158, but four sections further back, the section passes through the outer layers
of the atrium. The two spermducts have united and are entering the atrium in line with the longitudinal
muscular layer.

160. Arciform muscles around one of the male pores, viewed from the interior surface of the body. Only one side
is shown.

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INDEX

WonvuwE LE

OF SCIENTIFIC NAMES.

New names in heavy-faced type.

ACANTHODRILIDA 144, 145, 146, 166
Acanthodrilus 45, 131, 139, 142
georgianus 45
tamajusi 123, 139, 140
Vasliti 123, 129, 139, 142, 144, 145, 172
Aleodrilus 145, 146, 149
Keyesi 123, 146, 169
Allolobophora 41, 128, 164
Argilophilus 21, 29, 41, 44, 45, 47, 48, 49, 53, 55, 56,
65, 68, 70, 81, 130, 148, 167
marmoratus 164
ornatus 44, 53
papillifer 44, 55

BeENHAMIA 124, 125, 128, 129, 130, 131, 134, 138, 139,
145, 146, 147, 148, 164
Annex 123
Bolavi 123, 124, 125, 129, 132, 134, 135, 136, 138
floresiana 123
Godeffroyi 123, 126
kafuruensis 137
malayana 123, 129
mexicana 123, 124, 125
nana 123, 125, 127, 131, 132, 133, 134, 138
octonephra 123, 124, 126, 139
palmicola 123, 124, 125, 129, 131, 132, 133, 134,
135, 136, 138
papillata 123, 126, 135, 136
rugosa 123, 124, 126, 131, 134, 136, 138, 139
Stuhlmanni 134
Bombyx 149

CRYPTODRILIDH 21, 166

DELTANIA 21, 22, 23, 24, 40, 42, 43, 48, 52, 68, 81, 146
Benhami 22, 32, 34, 35, 36, 37, 38, 39, 40
dubia 22, 24, 36, 40
elegans 22, 23, 24, 25, 29, 32, 33, 34, 35, 36, 37,
39, 40, 146

Poultoni 22, 24, 40

Troyeri 22, 28, 31, 32, 33, 34, 37, 38, 39, 40, 170,
171

var. crassa 123, 169
lagunee 123, 170, 171

Diploecardia 146
communis 145

EcLiPIpRILipm 84
Eclipidrilus 4, 89
frigidus 84

Eminiodrilus 130
Enchytreus 33, 49, 79, 129, 142, 168
Eudrilidw 21, 44
Eudrilus 21, 45

dubius 23, 24
GEODRILUS SINGULARIS 142, 144, 145
Gordiodrilus 63, 173

H2aMAGREGARINA 86
Heliodrilus 45
Hyperiodrilus 45

Kerria 63, 65, 68, 69, 70, 144
McDonaldi 70, 142

LimicoLz 89

Limnodrilus 146

Luimbriculids 1, 4, 84, 85, 89

Lumbriculina 1, 3

Lumbriculus 1, 6, 89

Lumbricus 27, 33, 47, 48, 79, 81, $82, 128, 158, 163, 164
agricola 128

Microcumra 32

Microscolecini 21

Microscolex 21, 22, 23, 24
algeriensis 22, 23
dubius 23, 40
modestus 22, 23
nove-zelandixe 23
Poultoni 23, 24, 40

Moniligaster 65, 84, 89
indicus 43

NANNODRILUS 171

OcNERODRILID® 81
Ocnerodrilus 5, 28, 63, 64, 66, 67, 68, 69, 70, 71, 72, 78,
$1, 129; 171, 172; 173, 174
Beddardi 30, 67, 70, 72, 172
Hendriei 72
occidentalis 5, 32, 72
Oligocheta 1, 45, 47, 65, 67, 68, 72, 82, 84, 86
Oligochetie 76, 77

PericHmta 54, 142

Pheenicodrilus 63, 68, 70, 72, 78, 81, 129, 171, 172, 173
taste 64, 66, 69, 71, 72, 123, 171, 172, 173, 174
tepicensis 123, 171, 172

200 CALIFORNIA ACADEMY OF SCIENCES
Photodrilus 21, 22, 73, 82, 83 SPARGANOPHILUS 136, 151, 152, 153, 154, 155, 157, ¢.
Phreatothrix 4 162, 163, 166, 167, 168 ; i
Plutellini 21, 56 Benhami 123, 151, 152, 153, 154, 155, 156, 187,
Plutellus 21, 45, 48, 55, 56 158, 159, 160, 161, 162, 163, 165, 166, 187,
Pontodrilini 21 168, 169
Pontodrilus 21, 22, 66, 68, 70, 73, 78, 79, 83, 86, 129, carneus 123, 151, 153, 155, 168 3
150, 167 Biseni 123, 151, 152, 153, 154, 156, 158, 159, 166,

arene 83 167, 169

hesperidum 76, 81, $3 guatemalensis 123, 151, 153, 155, 161, 166, 167,

insularis 83 168, 169 E o

littoralis 82, 83 Smithi 123, 129, 151, 152, 153, 154, 155, 156, 158,

Marionis 73, 83, 84 159, 160, 162, 165, 166, 167

Michaelseni 73, 78, 82, 83, S4 sonome 123, 151, 153, 154, 156, 159, 160

phosphoreus 83 tamesis 123, 151, 153, 156, 157, 158, 159, 162, 163,
Pontoscolex 157 165, 166

corethrurus 45 tepicensis 129

Sutroa 2, 3, 4, 5, 6, 89, 168
rostrata 2, 3
RuopopriLus 21, 23

minutus 22 2 TELMATODRILUS VEJDOVSKYI 90
noye-zelandie 22 Tericolw 89

Rhynchelmis 2, 4, 5, 6, 7, 164 Trichodrilina 1
limosella 2, 3, 4 Tubifex 89

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