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THE ANNALS

AND

MAGAZINE

OP NATURAL

HISTORY

INCLUDING

ZOOLOGY, BOTANY, and GEOLOGY.

(UEIXG A continuation OF THE ‘ANNALS ’ COMBINED WITH LOUDON AND

CHARLESWORTh’s ‘ MAGAZINE OF NATURAL HISTORY.’)

CONDUCTED BY

ALBERT C. L. G. GUNTHER, M.A., M.D., Ph.D., F.R.S.,

WILLIAM S. DALLAS, F.L.S.,

WILLIAM CARRUTHERS, F.R.S., F.L.S., F.G.S.,

AND

WILLIAM FRANCIS, Ph.D., F.L.S.

VOL. XIY.— FIFTH SERIES.

LONDON:

PRINTED AND PUBLISHED BY TAYLOR AND FRANCIS

SOLD BY LONGMANS, GREEN, AND CO. ; SIMPKIN, MARSHALL, AND CO. J

KENT AND CO.; WHITTAKER AND CO.: BAILLIERE, PARIS:

MACLACHLAN AND STEWART, EDINBURGH :

HODGES, FOSTER, AND CO., DUBLIN : AND ASHER, BERLIN.

1884.

*' Omncs res creatce sunt divinee sapientise et potentiae testes, divitise felicitatis

hurnanoe : — ex barum usu honitas Creatoris ; ex pulchritudine sapientia Domini ;

ex oeconomia, in conservationo, proportione, renovatione, potentia maje.stati.s

elucet. Earurn itaque indagatio ab bominibus sibi rclictis semper acstimata;

4 vere eruditis et sapicntibus semper exculta; male doctis et barbaris semper

inimica fuit.” — Linn^us.

“Quel que soit le principe de la vie animale, il ne faut qu’ouvrir ies yeux pour

voir qu’elle est le chef-d’oeuvre de la Toute-puissance, et le but auquel se rappor-

tent toutes ses operations.” — Bruckner, Theorie du Systhne Animal, Leyden,

1767.

The sylvan powers

Obey our summons ; from tbeir deepest dells

The Dryads come, and throw their garlands wild

And odorous branches at our feet ; the Nymphs

That press with nimble step the mountain-thyme

And purple heath-flower come not empty-handed,

But scatter round ten thousand forms minute

Of velvet moss or lichen, torn from rock

Or rifted oak or cavern deep : the Naiads too

Quit their loved native stream, from whose smooth face

They crop the lily, and each sedge and rush

That drinks the rippling tide : the frozen poles,

Where peril waits the bold adventurer’s tread.

The burning sands of Borneo and Cayenne,

All, all to us unlock their secret stores

And pay tbeir cheerful tribute.

J. Tavlor, Norwich, 1818.

ACES LIBRARY

5”0

AN

sen 5

V, 1^

CONTENTS OF VOL. XIV.

[FIFTH SERIES.]

4^

vS»

r-

NUMBER LXXIX.

Page

I. On the CryptoniscidcB. By Prof. R. Kossmann 1

II. On the Spongia coriacea of Montagu, = Zewooso/enfa coriacea,

Bk., together with a new Variety of Leucosolenia lacunosa, Bk.,

elucidating the Spicular Structure of some of the Fossil Calcispougise;

followed by Illustrations of the Pin- like Spicules on Verticillites

helvetica, De Loriol. By H. J. Cabter, F.R.S. &c. (Plate I.) .... 17

III. Some Remarks upon the Variability of Form in Luhomirskia

baicalemis, and upon the Distribution of the Baikal Sponges in general.

By Dr. W. Dybowski. With P.SS. by H. J. Carter, F.R.S. &c.

(Plate II.) 29

IV. Descriptions of three new Species of Moths from the Island

of Nias. By Arthur G. Butler, F.L.S., F.Z.S., &c 84

V. On the Systematic Position of the Pulicidm. By Dr. Karl

Krapelin. (Plate HI.) 36

VI. New Investigations on the Development of the Viviparous

Aphides. By Dr. Otto Zacharias 54

VII. Notes on the South-Russian Spongillid<B. By Dr. W. Dy-

bowski 68

VIII. On the Synonymy of some Ileterocerous Lepidoptera. By

Rudolph Rosenstock, B.A 63

The System of the Monactinellidce, by Dr. R. von Lendenfeld ; On

Orhulina universa, by M. C. Schlumberger ; On the Ascidian

Genus Rhopalea, by M. L. Roule ; On the Process , of Diges-

tion in Salpa, by Dr. Ch. S. Dolley ; On a Species of Tachina

occurring on the Tracheal System of Carabus, by M. N. Cholod-

kowsky 65 — 74

09536b

VI

CONTENTS.

Page.

XXXV. On Astylosponnidce and Anornocladina. By Karl A.

ZiTTEL 271

XXXVI. Contributions towards a General History of the Marine

Polvzoa. By the Rev. Thomas Hencks, B.A., F.R.S. (Plates

VIII. & IX.) 270

New- Bn )/.’p : — Memoirs of the Geological Survey of India. Palaeon-

tologia Indica, being Figures and Descriptions of the Organic

Remains procured during the progress of the Geological Survey

of India. Series x. Indian Tertiary and post-Tertiary Verte-

brata. Vol. II. Part 6. Siwalik and Narbada Carnivora. By

R. Lydekker, B.A., F.G.S., F.Z.S. — Report on the Zoological

Collections made in the Indo-Pacific Ocean during the Voyage

of H.M.S. ‘ Alert/ 1881-82 ‘285, 291

Proceedings of the Geological Society

294—296

On the Copulation of Dijjiugia globulosa, Duj., by Dr. Carl F.

Jickeli ; How Lycosa fabricates her Round Cocoon, by Dr. H.

C. McCook 297, 298

NUMBER LXXXHI.

XXXVII. Ophryocgstis Butschlii, a Sporozoan of a new Type. By

Aim E Schneider. (Plate X.) 301

XXXVIII. Descriptions of Palaeozoic Corals in the Collections of

theBritish Museum (Nat. Hist.). — No. II. By Robert Etheridge,

Jun., and Arthur FI. Foord, F.G.S. (Plate XI.) 314

XXXIX. Diagnoses of new Species of Blenrotomidce in the British

Museum. By Edgar A. Smith 317

XL. The Auditory and Olfactory Organs of Spiders. By Fried-

rich Dahl. (Plate XII.) 329

XLI. Description of a new Species of Microgale. By Oldfield

XLH. Notes on the Palaeozoic Bivalved Entomostraca. — No.

XVH. Some North -American Leperditics and allied Forms. By

Prof. T. Rupert Jones, F.R.S., F.G.S 339

XLHI. Description of a new Species of the Coleopterous Family

Cetoniidce from Madagascar. By Charles 0. Waterhouse 343

XLIV. On a Polythalamian from "the Salt-pools near D6va, in

Transylvania. By Dr. Eugen von Daday 349

CONTENTS.

Vll

Page

Nev) Boohs: — Vergleichende Morphologie und Biologie der Pilze,

Mycetozoen und Bacterien, Von A. de Bary. — Our Insect

Allies. By Theodore Wood 363, 366

Note on the OccuiTence of some rare Foraminifera in the Irish Sea,

by Charles Flcock ; On the Occurrence of a Process resembling

Copulation in Comatula mediterranean by Dr. C. F. .iickeli ; On

the Organization of Anchinuin by M. N. Wagner ; On the

Anatomy of the TijrogUfphi, by t)r. Alfred Naleper; On the

Luminosity of the Glow-worm (Lanipyris splendidula), by M.

Wilhelm Kaiser . 366 — 372

NUMBER LXXXIV.

XLV. Description of an Impregnated Uterus and of the Uterine

Ova of Echidna hystj'ix. By Sir Richard Owen, K.C.B., F.R.S.,

(fee. (Plate XIII.) 373

XLVI. On the Coleopterous Genus Macrotoma. By Charles O.

Waterhouse 376

XLVII. Notes on Batrachians. By G. A. Boulenger 387

XL VIII. Notes on the Palaeozoic Rival ved Entomostraca. — No.

XVIII. Some Species of the Entomididcs. By Prof. T. Rupert

Jones, F.R.S., F.G.S. (Plate XV.) 391

XLIX. On new Species of Lepidoptera recently added to the

Collection of the British Museum. By Arthur G. Butler, F.L.S.,

F. Z.S., &c 403

L. Descriptions oftwo new Moths from Madagascar. By Arthur

G. Butler, F.L.S., F.Z.S., &c 407

LI. Local Colour-varieties of Scyphomeduscs : a new Species pro-

duced in Forty Years. By R. von Lendenfeld, Ph.D 409

LII. Notes on Hawaiian Neuroptera, with Descriptions of new

Species. By the Rev. Thomas Blackburn, M.A 412

LIIT. Contributions to our Knowledge of Hydromediisa, a Genus of

South-American Freshwater Turtles. By Dr. A. Gunther, F.R.S.

(Plate XIV.) 421

LIV. Note on some East-African Antelopes supposed to be new.

By Dr. A. Gunther, F.R.S 425

LV. Description of a new Species of the Carabideous Genus

Callistomimus. By Charles 0. Waterhouse 429

LVI. Description of a new Species of Julodis (Coleoptera, Bupres-

tidee). By Charles 0. Waterhouse 429

Vlll

CONTENTS.

New Books Journal and Proceedimr.s of the Roval Society of New

South Wales for 1833. Vol. XVII. Edited by Prof. A. Liver-

siDGE, F.R.S. — Internationale Zeitsch rift fiir allgem eine Sprach-

wissenschaft. Edited by F, Techmer 430, 431

Proceedings of the Geological Society 432—437

On PalucHcella erecta, by Mr. E. Potts ; On a new Insect of the Genus

Phylloxera {Phylloxera solids, Liclit.), by M. J. Lichtenstein. 437, 439

Index

410

Plate I.

II.

III.

IV.

V.

VI.

VII.

VIII

IX.

X.

XL

XII.

XIII.

XIV.

XV.

PLATES IN VOL. XIV.

Leucosolenia lacunosa. — Pin-like spicules on Verticillites.

Lubomirskia baicalensis.

Systematic position of the Pulicidse.

New species of Walckenaera.

Lycopodites Stockii. — Lepidodendron rimosum.

New species of Alveolites and Aniplexopora.

Development of the Common Shore-Crab.

Marine Polyzoa.

Ophryocystis Biitschlii.

New Palaeozoic Corals.

Auditory and olfactory organs of Spiders.

Female organs and uterine ova of FJchidna hystrix.

Ilydroniedusa platanensis.

New species of Entoraididae.

THE ANNALS

AND

MAGAZINE OF iNATURAL HISTORY.

[FIFTH SERIES.]

“ per litora sparjgite museum,

Naiades, et ciroiim vitreos considite fontes;

Polliee virgineo teneros hic carpite flores :

Floribus et pictum, divae, replete canistrum.

At VOS, o Nymphae Craterides, ite sub undas ;

Ite, recurvato variata corallia trunco

Vellite muscosis e rupibus, et mihi conchas

Ferte, Deae pelagi, et pingui conchylia succo.”

N.PartheniiGiannettasii Eel. 1.

No. 79. JULY 1884.

I. — On the Cryptoniscidse. By Prof. R. Kossmann *.

For the completion of my investigations upon the parasitic

Isopods living upon Crustacea, the Epicaridia, the Royal

Academy of Sciences granted me last autumn a considerable

travelling stipend. The completion of my labour of many

years has been brought into the immediate future by this

liberality, for which I here with pleasure express my gratitude ;

but its publication will nevertheless occupy so much time that

a preliminary communication of the more important results of

this last journey would seem to be justified.

This journey was devoted to the investigation of the Cryp-

toniscida0, and therefore of that subdivision of the Epicaridia

which has been least studied, and which, on account of its

extensive retrogression, presents the greatest difficulties to

investigation.

In the year 1843 Rathkef, in his “ Beitrage zur Fauna

Norwegens,” described, under the name of Liriope pygmeea^

* Translated by W. S, Dallas, F.L.S., from the ‘ Sitzungsberichte der

k. preuss. Akademie der Wissenschaften,’ April 24, 1884, pp. 457-473.

t Dathke, Beitrage zur Fauna Norwegens,” in Nova Acta Acad

Leop. Carol, xx. p. 60, tab. i. figs. 8-12.

Ann. & Mag. N. Hist. Ser. 5. Vol. xiv.

1

2 Prof. R. Kossmann on the Cryptoniscidse.

a minute creature which lie had found in the cavity of the

mantle of Peltogaster^ which was discovered by him at the

same time, a cirripede parasitic upon hermit-crabs ; he

regarded it as an Amphipod which had been swallowed by

the Peltogaster,

In the same year, in an article on the sexual characters of

the Cirripedia * * * §, Goodsir published his discovery of a similar

animal which occurred in Balanus halanoides^ and which the

author regarded as the male of that cirripede, the herma-

phrodite nature of which was not then placed beyond doubt.

In the course of a dissertation upon Peltogaster^ Steenstrupf

found occasion to speak of Liriope^ and indicated in a some-

what doubtful manner that it might possibly be a Bopyrid

living upon Peltogaster.

In the same year Darwin \ corrected Goodsir’s statement,

recognizing the latter’s supposed male Balanid as a parasite,

and, indeed, as belonging to the loniens,” i. e. to the

Bopyridaa.

About the same time (text 1852, figure 1855) Dana§

described under the name of Cryptothir minutusj as the male,

a similar parasite from Creusia^ and referred it, with Liriope^

to the Tanaidag.

Lilljeborg H then (1861) came back upon this Liriope^ and

also showed that Liriope was an Isopod, and, indeed, a Bo-

pyrid. He succeeded particularly in this, that besides the

larviform and excessively minute creatures that Rathke and

Dana had found, he also discovered the mature female form

of Liriope^ and he came to the conclusion that the former

were the young males.

Buchholz^ arrived at a somewhat different conclusion in

some respects. Without knowledge of the statements of

Goodsir and Darwin, he carefully described anatomically the

animal observed by them. He regards the forms found by

himself, so far as they look like Liriope^ as older larvae, and

* Goodsir, On the Sexes, Organs of Reproduction, and Mode of Deve-

lopment of the Cirripeds,” inEdinb. New Phil. Journ. xxxv. p. 88, pis. iii.

and iv.

t Steenstrup, “ Bemarkninger om slaegterne Pachyhclella og Peltogaster

in Oversigt af Danske Vidensk. Selsk. Forhandl. 1854.

X Darwin, ‘ Monograph on the Subclass Cirripedia ’ (Ray Society),

vol. ii. p. 271.

§ Dana, ‘ Crustacea in U. S. Exploring Expedition under Commander

Wilkes,’ vol. ii. p. 801, Atlas, pi. liii. tig. G.

II Lilljeborg, ^^Liriope et Peltogaster^’’ in Nova Acta Soc. Sci. TJpsal.

ser. 3, vol. iii. p. 1, and Suppl. p. 73 (see ‘ Annals,’ ser. 3, vol. vii. p. 47).

% Buchholz, Ueber Hemioniscus halanp’ in Zeitschr. fiir wiss. Zool.

Bd. xvi. p. 303.

3

Prof. R. Kossinann on the Cryptoniscidas.

tlic male as still unknown. Nevertheless he was the first to

give us an idea of the adult parasite, as Goodsir had only

figured the head and the first four thoracie segments, but

neglected all the remaining deformed ])art of the animal.

A further enlargement of our knowledge U])on this group

was furnished in the year 1871 by a memoir of Fritz Muller’s*.

Under the name of Cryptoniscus planarioides he described a

parasite resembling Liriope^ but which displaces its host, a

Peltogaster^ from the hermit-crab ; only the roots of the Pelto-

gaster are preserved, and are apparently made use of by the

Cryptoniscus for its own nourishment.

Soon after the appearance of Miiller’s memoir, and without

any acquaintance therewith, I myself published notes t upon

some forms belonging to this group from Semper’s Philippine

collections, namely an internal parasite from Sacculina pisi-

formisj which I named Eumetor Uriopides^ an external para-

site on the abdomen of a Porcellana [Zeuxo porcellance) ;

another on the head of an Alpheus [Zeuxo alphei) ; and,

lastly, a parasite from the brood-cavity of a Bopyrus [Cahira

lernceodiscoides) , Any exact anatomical investigation of

these forms was impossible, as I had only single spirit-speci-

mens of them.

Passing over Hesse’s unscientific and useless memoirs in the

^ Annales des Sciences Naturelles,’ I come to the most recent

and important work upon this subject, namely Fraisse’s

memoir “ Hie Gattung Cryptoniscus^ Fr. Muller ” To its

contents I shall have to refer repeatedly.

There has long, as we have seen, been a general conviction

of a near relationship between the Bopyridge and Cryptonis-

cid^. When, therefore, I formed the plan of undertaking a

monographic description of the Bopyridse, I could not think

of passing over the Cryptoniscidge. Moreover, there were

points of difference between the statements of Buchholz and

Fraisse and the results of my own occasional investigations,

which made a thorough testing of the investigations of these

two meritorious observers appear indispensable.

I have therefore twice, in 1882 and 1883, thoroughly

studied Hemioniscus halani in Christiansand itself, and in the

autumn of 1883, with the aid of the Royal Academy which

* Fritz Muller, ‘‘ Bnichstiicke zur Naturgeschiclite der Bopyriden,” in

Jenaische Zeitsclirift fiir Medicin und Naturwissenschaft, Bd. vi. p. 61

Taf. iv. figs. 12-19. ’

t Kossmann, Beitrage zur Anatomie der schmarotzenden Ranken-

fiissler, Anhang,” in Arbeiten aus dem zool.-zootom. Institut der Univ.

Wurzburg, Bd. i.

X Arbeiten aus dem zool.-zootom. Inst, der Univ. Wurzburg, Bd. iv.

1*

4 Prof. II. Kossmann on the Cryptoniscidge.

lias been already gratefully mentioned, Cryptomscus paguri

at Mahon, in the island of Minorca ; at other opportunities I

have also examined single specimens of nearly all the forms

pertaining here. Upon these investigations the following

brief expositions are founded.

But before I enter upon the actual subject, a formal neces-

sity presses itself upon me. As in my monograph I must, of

course, employ a critically sifted and correctly established

nomenclature, and yet must not lay too great difficulties in

the way between the preliminary and the detailed work, I

think I ought here to give a small table relating to this

matter.

I. Parasites on Cirripedes.

a. Upon non-parasitic Cirripedes {Balanus halanoides and Creusia).

1. Cryptothih, Dana, 1852 (U. S. Expl. Exped., Crustacea, p. 801).

Synonymy : — Hemioniscus, Bucliholz, I860.

Cryptothiria (p. parte), Spence Bate & Westwood, 1868.

b. Upon parasitic Cirripedes.

a. Free in the mantle-cavity.

2. Eumetor, Kossmann, 1872 (Beitr. z. Anat. d. schmar. Rankenf ussier,

Anhang).

/3. Perforating the mantle from within.

3. Liriopsis, Max Schultze, 1859 (‘‘ Anm. zu einer Aufsatz von Fritz

Miiiler,” in Wiegmann’s Archiv, Bd. xxv. p. 310).

Synonymy: — Liriope, Rathke, 1843, nom. praeoccup. (Lesson, 1837,

Trachymedusae) .

(This form is referred by F. Muller and Fraisse to Cryp-

toniscus, by Spence Bate to Cryptothiria).

y. Attacking the peduncle from without, and displacing the cirri-

pede down to the roots.

aa. Upon Peltogasterj with short cephalon.

4. Cryptoniscus, Fritz Muller, 1870 (Jena. Zeitschr. Bd. vi. p. 61).

/3/3. Upon Sacculina, with long cephalon penetrating deeply into

the Brachyuran.

5. Zeuxo, Kossmann, 1872 (Beitrage z. Anat. d. schm. Rankenf.), nom.

praeoccLip. by Templeton, but suppressed by Dana in favour of Tanais.

II. Parasitic on Ostracoda.

6. Cyproniscus, Kossmann, gen. nov. Referred by Sars (“ Oversigt af

Norges Crustaceer,” in Christ. Vidensk. Selsk. Forhandl. 1882,

no. 18, p. 73) to Cryptothiria.

III. Parasitic on Isopoda.

7. Cabirops, Kossmann, gen. nov.

Synonym \—Cahira, Kossmann, 1872 (Beitrage &c., Anhang), nom.

prseoccup. (Treitschke, 1825 Cahera, Jodoh'sky, 1837

Cahira, Lepidoptera).

Referred by Sars (Oversigt &c. p. 74) to Cryptothiria.

Prof. R. Kossmann on the Cryptoniscidaj. 5

What from the first caused the greatest difficulty was

the ascertainment of the sexual relations. The form first

described by Rathke with eyes and natatory feet has been

interpreted sometimes as tlie male, sometimes as the larva.

Rathke himself evidently regarded his animal as adult, but

without deciding any thing as to its sex. Dana took the

corresponding form, which he found in Greusia^ for a male,

but made no remark about its age. Lilljeborg’s discovery

})roved that the animals described by Rathke and Dana were

young forms ; and it seemed to Lilljeborg impossible to regard

them as young female forms, because, although already settled

upon the host, they showed no commencement of the trans-

formation into the adherent female form. He further com-

pared them with the youngest male Bopy rides found by

Kroyer, and came to the conclusion that they were immature

males. Buchholz regarded this same form (from Balanus

halanoides) as an old larva without recognizable sex ; he

found no males, but at the same time declared that the sexually

mature animals [Gryptothir balani) found by him were deci-

dedly not hermaphrodite. Spence Bate, who had already

seen and named the young animal*, thought, in 1868 f, it

might perhaps be a male, and adds to the word immature”

a note of interrogation within brackets. He therefore doubted

as to the immaturity of the animal, but without in any way

proving that it was a male, still less a mature one.

Lastly, Fraisse J asserts with almost perfect certainty that

the copulation must take place in the stage preceding

attachment,” and accordingly describes both male dnd female

animals of this stage (representing Rathke’s Liriope)^ of

which, however, he regards only the former as sexually

mature. But his proof would not satisfy most reader^

Thus, as regards the males, the testes are scarcely indicated

in Fraisse’s figure ; their form, aperture, or even structure he

has not described at all ; the semen, which was squeezed out

by crushing the animal, is not removed beyond the reach of

doubt, on account of the mode in which it was obtained and

the statements as to the form of its elements, and indeed it is

rendered absolutely suspicious by the fact that Fraisse sup-

posed he saw it also in the body-cavity of the female attached

to the ovaries, where its presence may be pronounced to be

impossible. Fraisse found these supposed males swimming

* Spence Bate, ‘ Report of the British Association,’ 1860, p. 225.

t Spence Bate and Westwood, ‘ History of the British Sessile- eyed

C/i-iistacea,’ vol. ii. p. 267.

I Fraisse, “ Hie Gattimg Cryptoniscus," loc. cit. p. 23, Taf. xv. figs. 30

and 32.

6 Prof. R. Kossmami on the Cryptoniscidse.

freely ; but he observed exactly similar animals creeping or

swimming about close to the Sacculina^ and these he regarded

as females. He saw no genital apertures, or, indeed, genital

organs of any kind in them, but observed their conversion into

the adherent animal, which undoubtedly is of the female sex.

As, however, in this, in a stage which is already strongly

metamorphosed, the ovaries are still quite immature, the

female larva, according to Fraisse’s own opinion, can only be

quite immature. Nevertheless, although he had “ no oppor-

tunity ” of observing the copulation, and did not even himself

see the spermatophores adhering to the female,” he is of

opinion that this immature larviform female must already be

fecundated. Thus, therefore,” he writes, the fecundated

female attaches herself, while the male retains its form and

probably perishes after the act of copulation.”

Evidently all that he has seen (or not seen) is opposed to

Fraisse’s own opinion, and tends to show that the male is

indeed sexually mature and copulates in the above-mentioned

larviform stage, but that the female becoming sexually

mature at a much more advanced stage is also sought out and

impregnated by the male only in this sessile condition.

That the male is really larviform I can positively assert

from careful examination of such stages. We can very dis-

tinctly recognize the genital apertures at the base of the last

pair of pereiopoda ; we find the mature testes in the trans-

verse section, and witness the brisk movement of the sperma-

tozoids. In the Gry'ptoniscidce^ therefore^ the mature male is

larviform and still furnished with natatory feet upon the pleon.

That the female is coupled before its sexual maturity there

are no observations to show, and nothing warrants any such

supposition. But that on arriving at sexual maturity it is

sought out and copulated by the male is supported by the

observation of Buchholz, who writes with regard to Crypto-

thir (Hemioniscus) : — Nearly in every Balanus which con-

tained one of the sacceiform animals there occurred one or

more small, elongated, brownish animalcules” (here follows the

description of Bathke’s Liriope-ioxm) ; as also by Fraisse’s

own figure (Taf. xii. fig. 1), in which such a larva is shown

clinging to the metamorphosed female ; and, finally, by the fact

that, in 1872, 1 found such a male animal, as I then supposed,

in the Philippine Eumetor liriopides^ and in 1883 in a Neapo-

litan species of the same genus, three of them, and, indeed^

clinging fast to the female. The natatory power of these

males makes it very easy to understand that we do not always

meet with them with the females as among the Bopyridai ;

they probably often spontaneously quit the female, and arc

7

Prof. K. Kossmaiin on the Cryptoniscidse.

certainly still more frequently missed by the observer. In

short I accept it as proved that the female is copulated only in

the metamorphosed state.

We have therefore free-swimming larviform males, and

adherent, strongly retrograded females, which copulate with

each other. But this is not all. My investigations furnish

the most convincing indications that the two forms are only

different stages of development of the same individual ; in

other words, that among the Cryptoniscidse we have to do

with a protandrous hermaphrodism. According to the inves-

tigations of Bullar and Paul Mayer such a thing is not unpre-

cedented among Isopoda, but rather undoubtedly recognized

among Cymothoidse ; there, however, the sexual maturity of

the male occurs much later, after the biramose natatory feet

upon the pleon, characteristic of the larva, have already

become transformed into branchial feet. Protandry with

larval sexual maturity was previously entirely unknown.

Of course it has not been possible for me to trace one and

the same individual through its whole course of development ;

to observe how, as male, it performed the duties of its sex,

and how, after the metamorphosis had taken place, it was

copulated as a female, and produced eggs. I can therefore,

as already stated, only bring forward evidence in favour of

my assertions.

First of all negative : — I have never found a free-swimming

Cryptoniscid larva of the last stage that had not male-deve-

loped sexual glands. I have found and examined of Grypto-

thir halani a considerable number, and of Eumetor three — all

males. Fraisse, indeed, describes female larvge, but these

were already fixed, and not truly female, but immature,

neuter; the free-swimming examples of which he found some,

were males. It is clear that the protandry is proved, unless

we succeed in discovering females or neuters in the same stage

of development as the males, hitherto exclusively found.

A second piece of negative evidence is the following : —

While in all Bopyridae s. str., and in the Entoniscidas the

male becomes sedentary and remains with the female, it is in

all Cryptoniscidae free-swimming and exceedingly active,

and is often no longer to be met with near the fecundated

female. How should this difference between two so nearly

related groups be explicable if the male had not still another

task to fulfil elsewhere after the fecundation of the female ? and

what other can it be except that of itself growing into a female

upon another host ?

Finally, we have a piece of positive evidence in the presence

in the mature female of a gland which is to be regarded almost

8 Prof. R. Kossmann on the Cryptoniscidse.

with absolute certainty as a retrograded testis. This gland

was discovered by Buchholz* in Cryptothir balanij but is

wanting in none of the Cryptoniscidae examined by me, while

no female Bopyrid or Entoniscid possesses even the smallest

trace of it. Buchholz thought that this gland was to be re-

garded as an accessory organ of the sexual parts ; notwith-

standing repeated endeavours he could discover no efferent

duct, but he found the granulated contents to be like those of

the terminal section of the female genital duct. His figure

and description represent the organ as a thin cord inflated

in three places, situated on three sides above and outside of

the ovaries, and filled with a finely granular substance. The

inflated parts form cellular diverticula.

Now this organ extends through the last three segments of

the pereion, therefore the same in which the testis is situated

in the male. These three segments of the larva are widely

separated immediately before the last change of skin, and

afterwards enormously enlarged by colossal reception of

nutriment. At the same time, however, the triple division

continues recognizable by transverse constrictions. If there-

fore a testis no longer in function be present, what is more

natural than that this should retain its original thickness in

the three segments, but become stretched into a thin cord in

the intervals — in short, acquire the form represented by Buch-

holz ? and is not also the deficiency of the efferent duct, ascer-

tained by me, in favour of the view that we have to do with

an organ no longer in function ? Lastly, as regards the con-

tents. In my transverse sections these appear as a finely

granular, strongly refractive, and very strongly colouring

detritus, which in these three properties is absolutely similar

to the contents of the testes in the males of the Bopyridm

(provided there is no semen in them).

From all this we may regard it as almost proved to demon-

stration that the Cryptoniscidae are really protandrous herm-

aphrodites, in which the testis attains its maturity in the

final larval stage, and is then visible in the mature female as

a rudimentary organ without an efferent duct.

The question now arises, what circumstances have operated

for the production of this kind of hermaphrodism, seeing that

the Bopyridm, which are so nearly allied, are certainly not

hermaphrodites ?

We generally find hermaphrodism espeeiallyin slow-moving

or adherent animals, and to these it gives an essential advan-

tage in the struggle for existence. In tlie first place, it renders

* Buchholz, loc, cit. p. 31G, Taf. xvi. figs. 2 & 3, G.

9

Prof. R. Kossmann on the Cryptoniscidse.

self-fertilization possible in cases where a meeting of two

individuals can occur only with difficulty or not at all. Now

self-fertilization is certainly in other respects injurious ; as

intensified incestuous breeding it has an exceedingly corrupt-

ing influence upon the organization of the race. It is no doubt

for this reason that in some hermaphrodite groups of animals

the habit of mutual fecundation has been brought about. In

this certainly the above-mentioned advantage disappears ; the

contact of two individuals is, as in other cases, necessary.

On the other hand, another advantage results from it, namely,

when the comparatively obstructed chance of contact does

occur, then at least two individuals are fecundated, and

consequently there exists twice as much probability of the

preservation of the species as if the animals were not herm-

aphrodite.

But if in this way hermaphrodism with mutual fecundation

may in certain species replace that with self-fertilization wfith

favourable results, this applies only to such as move with

difficulty and not to sedentary species. In the latter, contact,

and consequently mutual fertilization, is impossible. Here

therefore we must rest satisfied with self-fertilization, if the

animal were really sedentary during its whole existence. But

this is the case in no animal proceeding from an egg j and all

animals proceed from an egg, if not in every generation,

nevertheless in generations recurring regularly after a certain

time.

From this it follow^s that even in animals which become

sessile contact may take place, always supposing that one of

the two individuals is not yet sessile. During this contact

they might fertilize each other, if both kinds of sexual organs

were already developed in both. This, however, for econo-

mical reasons is usually impossible ; for parasites, at least, it

is attachment that usually secures that quantity of nutriment

which is necessary for the egg-production ; and, on the other

hand, parasites generally require for the maintenance of their

species such a colossal fertility, that the egg-formation of

itself deforms the body and compels it to become attached.

Hence, with special exceptions, it is not well possible that

the free-moving individual should already possess ovaries ;

consequently neither a mutual fertilization nor the one-sided

one supposed by Fraisse can be accepted as taking place

between two free-swimming animals. On the other hand,

the animal may well be capable of the production of the

semen, of which no great quantity is necessary, even before

the commencement of adhesion, and thus the protandry above

described would be brought about. An advantage over the

10 Prof. R. Kossmann on the Cryptoniscidae.

simple dimorphism of the sexes is, however, also obtained.

If I assume (quite arbitrarily) that the individual occupies a

week from hatching to male maturity, and then three weeks

from hatching to female maturity, then to obtain 10 (or n)

broods in tlie case of dimorphism, 10 (or n) individuals must

escape all the dangers that threaten them each for a week,

and 10 (or n) individuals each for three weeks (40 or

weeks), while in the case of protandry only 10 (or n) indivi-

duals need to exist each for three weeks and 1 more for one

week ( = 31 or 3w+l weeks). And the advantage is even

still greater than these numbers show, as it is precisely the

first week in which the animal (in our case) swims freely

about, and consequently is much more exposed to dangers

than during the next two weeks, when it is already adherent.

Those 9 (or n — 1) weeks which in our example are saved by

protandry are, as one may easily convince one’s self, all first

weeks of life. It is the first week, that of the free-swimming

stage, of nine female individuals that is saved.

The notions as to the nature of the brood-cavity in which

the ova in the Cryptoniscidac are sheltered until hatching

have hitherto been very defective. Buchholz* found that

in Cryptothir halani the deposited eggs float to and fro . . .

apparently free in the body-cavity but in reality are en-

closed in a special, extremely delicate- walled, and perfectly

transparent vesicle.” His further statements upon this subject

do not seem to me very clear. He finds this vesicle attached

to the outer wall of the body at the spot “ at which the four

genital apertures are situated If we separate the

pedicle of the vesicle from this spot we obtain it in connexion

with the four oviducts, which remain attached to it uninjured,

and the outer extremities of which seem to pass directly into

the vesicle.” Thus, while Buchholz originally saw the ovi-

ducts open outwards, he sees them afterwards open into the

vesicle ; both observations which I can confirm as correct.

In spite of this the original four sexual apertures are said to

persist on the outer surface. Nevertheless,’"’ he says, the

presence of external sexual apertures at this spot, simultane-

ously with the opening of the oviducts into the egg-reservoir,

is difficult to understand.” In my opinion, it is not to be

understood ; and his attempted explanation, which I sliall not

reprint here, is quite unintelligible to me. The true condi-

tion of things in Cryptothir^ as I have observed with certainty,

is, that there is a sinking in of the region of the genital aper-

tures, at first in the form of a transverse groove. In this way

^ Buchliolz, loc. cit. p. 315.

Prof. R. Kossmann on the Cryptoniscidaj. 11

a brood-space with a transverse entrance-fissure is formed by

invagination, and as its inner surface is tlie former surface of

the animal in the neighbourhood of tlie oviducal aperture, the

oviducts of course no longer open outwards, but into this

newly-formed brood-space.

Somewhat more complicated, but perhaps more primitive,

is the arrangement in the other Cryptoniscidm, at least in

Cryptoniscus and Liriopsis. Fraisse* falls into an error,

which, however, his predecessor had eseaped ; he says, This

brood-eavity was previously present, for it is simply the body-

cavity,” and adds, How the ova get into it I eannot say

and, in faet, it would be hardly possible to establish a con-

ceivable hypothesis upon this view : deposition of the ova in

the body- cavity would be something unheard of among Crus-

taeea. Into this brood- or, aeeording to Fraisse, body-cavity

lead two “ respiratory ” apertures, already very fully de-

scribed by him, one of them in the neighbourhood of the mouth,

the other further back, and the two united with eaeh other by

a longitudinal groove (upon the ventral surface). Of this

groove Fraisse says (p. 13) : — When the larvas are ready for

a free existenee, a fissure bursts whieh formed between the

two respiratory apertures during the third stage, and was

previously covered and closed by a thin cuticular layer.”

Through this the larvae are set free.

The aetual course of events, as I have ascertained by the

study of numerous transversely-seetioned specimens, is as

follows. Here also, first of all, the two oviduets (whieh

Fraisse, L c. p. 9, was unable to detect) open on the ventral

outer surfaee of the body. Their originally cireular aperture

becomes elliptical and is soon drawn out before and behind

into a shallow groove, so that we may easily recognize two

such parallel longitudinal grooves in the female when not yet

quite mature. Now the wall between the two grooves begins

to sink in, and thus we obtain, instead of the two shallow

grooves, an elongated depression, the two side walls of whieh

curve into one another before and behind. These side-margins

now, however, grow towards one another until they touch

throughout nearly their whole length ; there remains conse-

quently only a hair-like slit, which passes at its posterior and

anterior ends into a rounded hole, and these two holes lead

into a cavity formed by the sinking in of the wall lying be-

tween the two genital furrows. This cavity, into which of

course the oviducts open, is the brood-cavity, but it has abso-

lutely nothing to do with the body-cavity. It becomes filled

* Fraisse, /oc. cit, p. 12.

12 Prof. R. Kossmann on the Cryptoniscidaa.

with deposited ova and thereby enlarges in proportion as the

ovaries (and at the same time also the alimentary organ)

diminish in volume.

Tliat the ova obtain the ehange of water necessary for respi-

ration, is provided for by brisk pumping movements, which,

as Fraisse correctly describes, are effected in this later stage

by the musculature at the two brood-space or respiratory

apertures ; and that the current of water sliall not wash out the

ova, by a system of villiform valves which close the apertures

eelpot fashion, as Fraisse has also shown. This whole ar-

rangement is produced, moreover, before the last moult of the

animal, and therefore, in a certain stage, the fissure of the

brood-chamber still appears, as Fraisse says, covered and

closed by a thin cuticular layer.” Subsequently, however,

the fissure still holds together, only because the two margins

are to a certain extent interlocked ; with a little effort they can

be readily forced asunder without tearing anything. But spon-

taneously the fissure certainly only opens when the brood-

cavity is overfilled, and the parent animal performs the most

violent contractions. As Fraisse correctly describes, these

often still continue when all the ova are already expelled ; and

as, at this time, the ovary and alimentary apparatus, the only

coloured organs of the animal, are completely retrograded,

the animal, which now resembles a torn and perfectly trans-

parent rag, and yet contracts violently, presents a very

remarkable appearance.

Passing over many details of less general interest I cannot

abstain from stating something with regard to the mode of

taking nourishment and the alimentary organs.

In GryiJtothir [Hemioniscus) j as Buchholz has already

shown, the cephalon and the pereion as far as the antepenulti-

mate segment remain larviform throughout life ; in accord-

ance with this the small boring and sucking apparatus,

consisting of the labrum and labium, between which two

styliform mandibles are placed, serves through life as the

organ for the inception of nourishment. This anterior divi-

sion of the body is, however, also deformed in the other

genera. The genus Zeuxo^ which 1 discovered in 1872,

lives upon parasitic Cirripedes, especially upon Saccalina, a

parasite of the Brachyura. It perforates this animal, the

pedicle of which ramifies like a root in the body of a common

crab, at the point where the pedicle enters into the body of

the crab, and draws nourishment from it, after the fashion of

a plant ; it consequently intercepts the nourishment of the

Sacculinaj and often causes it to die away altogether with

only the exception of the roots. These roots, singularly

13

Prof. ll. Kossmann on the Cryptoniscida3.

enough, remain alive, and are made use of by tlic Zeuxo.

The head of the latter, which is finally inserted deeply into

the body of the crab, although always still in a great lacuna

of the radiciform pedicle of the Sacculina, presents, besides

the buccal aperture, only four cylindrical processes, of which

one pair is usually longer than the other. By their form and

})Osition they give rise to the supposition that they are the

antenna3 of the larva which have lost their articulations. They

evidently effect the fixation of the animal. In some species

the fore part of the body, from the place where it enters into

the body of the host to the mouth, is drawn out into a long

peduncle ; in others it is rather short.

Upon another parasitic Cirripede [Peltogaster) lives the

genus Cryptomscusj F. Muller, under exactly similar vital

conditions. Its head docs not form a peduncle, but within

the aperture which it has perforated we find four pad-like

swellings surrounding the mouth, which, from analogy, we

may also regard as modified antennae. That this genus

almost always brings about the destruction of the Peltogaster

itself has been already indicated by Fritz Muller and con-

firmed by Fraisse; now and then, indeed, we find a specimen

which seems not to be seated directly upon the hermit-crab,

but has bored somewhere into the mantle of the Peltogaster ;

but if such stray examples, on the one hand, do not quite

cause the destruction of the Peltogaster ^ on the other hand

they do not seem themselves to arrive at female sexual

maturity.

It is otherwise with the genus Liriopsis^ Max Schultze

{Liriope^ Bathke), which also lives upon a Peltogaster.

This animal (the anterior and posterior ends of which have

been hitherto mistaken) does not perforate the pedicle of the

Peltogaster^ but slips into the cavity of its mantle, and per-

forates the mantle from within. Thus the anterior half of

the body is inserted into the blood-lacunge of the mantle,

while the posterior half lies free in the mantle-cavity, and

the perforation which the parasite has made causes a median

constriction of the animal. But in this case, not only the

head, but at least five segments of the middle-body are

inserted into the host; and as the aperture through which

the brood of the Liriope swarms out is formed upon these seg-

ments, the parasite, when the brood is mature, or perhaps a

little earlier, must also break through the outer wall of the

mantle. This is probably effected less by boring than because

its own growth exerts such a pressure upon the tissues before

it that the latter become atrophied and finally burst. The

parasite then remains with the abdomen in the mantle-cavity

14 Prof. R. Kossmann on the Cryptoniscidse.

and the fore part of the body outside in the open. I have

been unable any longer to detect antcnnse or buccal organs in

the stage of female maturity, and my experience was similar

in some other genera. Among these are Eumetor^ which

was likewise discovered by me in 1872, and has now been

more accurately studied, and CahirojoSj which inhabits the

brood-cavity of the Bopyridae, its nearest allies. All these

three genera, in the adult condition, have the head free, and

consequently need then no apparatus for fixation or boring.

It is true also that they can then take no more nourishment.

But this is no longer necessary to them ; Gryptoniscus and

Zeuxo also at this time take no more nourishment, although

they have the head still inserted into the blood of the host.

This follows with certainty from the retrogression of the

digestive apparatus, already ascertained by F raisse in Crypto^

niscus.

The knowledge of the nature of this digestive apparatus

has advanced by various roundabout ways ; I will here only

briefly refer to its course hitherto. Rathke* ascribed to the

Bopyridse a liver consisting of seven pairs of follicles opening

separately into the intestine ; Cornalia and Pancerif, certainly

in a different genus, describe, instead of these fourteen folli-

cles, two tubes running parallel to the intestine, their opening

into which they did not see. BuchholzJ found in Crypto-

niscus halanij intercalated between the oesophagus and the

rectum, a comparatively enormous vesicular reservoir, drawn

out posteriorly into two cseca, which, from its appearance, is

really to be regarded as the intestinal canal and a similar

condition of things in Entoniscus was described first by Fritz

Muller § and afterwards by Fraissejl, the former characterizing

the organ as the liver, the latter as rectum. This latter inter-

pretation, as a section of the intestine, is also maintained by

Fraisse for a corresponding organ in the Cryptoniscidse, which

certainly, by its exceedingly vigorous growth, soon loses all

trace of a division into parallel tubes. I have already else-

where, with regard to the Bopyridse and Entoniscidge, adopted

the opinion of those who regard this organ as a homologue of

the so-called liver of the Crustacea. I do so also uncondi-

tionally with respect to the digestive organ of the Crypto-

Ratlike, ‘De Bopyro et Nereide,’ 1837, p. 9, tab. i. fig. 7 b.

t Cornalia e Panceri, “ Osservazioni sopra im nuovo genere di cro-

stacei isopodi sedentarii, Gyye hmnchialis,'' 1858, p. 16, tab. ii. fig. 6e.

\ Bucbholz, Hemioniscus halani, loc. cit. p. 310.

§ Fritz Miiller, Entoniscus porcellance, in Archiv fiir Naturg. Bd. xxviii.

p. 11.

jl Fraisse, Entoniscus Cavoknii, 1878, p. 17

15

Prof. R. Kossmarm on the Cryptoniscidaa.

nisciclas. Both the development of the organ from a pair of

cylindrical csecal tubes opening into the anterior part of the

intestine and its histological nature prove the homology with

the so-called liver. But just as I have already said with

regard to the Bopyridas that this so-called liver does not

function exclusively as such, but evidently performs a func-

tion as a section of the intestine,” so must I also assert deci-

dedly that the lumen of this so-called liver receives the food

of the parasite, which is identical with the blood of the host,

in immense quantities ; that in this place this nutriment is

digested and absorbed, during which the organ gradually

shrivels up ; and that consequently the name of liver is by no

means physiologically applicable to the organ. But at the

same time it has been sufficiently demonstrated that the

so-called Crustacean liver is no liver at all. The name

reposes on an error called forth by the most insignificant

superficial character, namely the colour of the organ. Hoppe-

Seyler* and Krukenbergf found in the secretion of the

so-called liver of the higher Crustacea a diastatic, a peptic,

a tryptic, and a fat-decomposing enzyma. Max Weber |

believed that he found in the epithelium of the liver, besides

the true hepatic cells, a second kind of cells, which he supposed

to fill the above more pancreatic function, and he named the

organ hepatopancreas. But Hoppe-Seyler showed in the

freshwater crayfish, and Frenzel § has lately done so in many

marine Crustaceans, that no biliary constituents at all are

present in the secretion. There are no biliary acids or their

soda and potash salts, no bilifuscin or the allied pigments ;

and bilirubin was sought in vain. To this must be added

that Frenzel has also ascertained that Weber’s assertion that

there are two different kinds of epithelial cells is erroneous.

In my Epicaridia nothing of the kind can have existed. In

short this organ, wherever it has the form of a gland, is

clearly the digestive gland of the Crustaceans, a glandula

intestinalis. But in the Epicaridia, and especially in the

Cryptoniscidge, the lumen of the intestine does not suffice for

the reception of the food, and then this organ takes part in it

in a very remarkable manner. Thus from a glandula intes-

tinalis it becomes an intestinum glandulare^ a reservoir,

* Hoppe-Seyler, ‘ Physiologisclie Chemie,’ p. 276,

t Krukenberg, Vergleicb.-physiol. Beitr. zur Kenntniss der Verdaii-

ungsvorgiinge,” and “ Zur Verdauimg bei den Krebsen,” in Untersucliun-

gen aiis den physiol. Institut in Heidelberg, Bd. ii.

X M. Weber, ^‘Ueber den Ban imd die Thatigkeit der sog. Leber der

Crustaceen,” in Arch. f. mikr. Anat. Bd. xvii. p. 385. °

§ J. Frenzel, “ Ueber die Mitteldarmdriise der Crustaceen,”in Mittheil.

a. d. zool. Station zu Neapel, Bd. v. p. 50.

16 Prof. P. Kossmann on the Cryptoniscidoe.

functioning as an intestine, with a secreting and at the same

time absorbent epithelium.

The sucking stomach so characteristic of the Bopyridm and

Entoniscidg0, with the large papillae projecting into its lumen,

is deficient not only in Gryptothir^ in which, indeed, the larvi-

form fore body would afford no space for it, but also in the

other Cryptoniscidae ; in them, evidently, the intestinum glan-

dulare^ the walls of which perform lively movements, assumes

its function.

Finally, as regards the rectum. Its connexion with the

anterior part of the intestine is interrupted in the more strongly

deformed animals, and we find it only in larva© and young

females. As Buchholz, F. Muller, and Fraisse have already

shown, it is inflated into a pear-shape not far from the anus.

(In opposition to Fraisse’s statements, I find an anus even in

the older animals.) Of the bacilliform elements of the con-

tents described by Buchholz I have seen nothing any more

than Fraisse; in my sections the lumen of this part is quite

empty, as the large cells of the wall project into it like

papilla©, and leave only a small stelliform space free in each

transverse section. I cannot confirm the statements of Buch-

holz and Fraisse that pigment cells surround the rectum ; the

brown pigment deposited in the neighbourhood of the intes-

tine is extracellular, and is probably to be regarded as urinary

concretion. Whether we have to do here, as Blanc * sup-

poses, with a separation-product of the fatty body, may be

regarded in this case as doubtful, as no distinct fatty-body

elements could be demonstrated even in the youngest stages

investigated by me. This, however, is no absolute impedi-

ment. In the Bopyridm and Entoniscida© the fatty body,

which is at first very large, is reduced in proportion as the

ovary enlarges by the maturation of the ova ; and it is com-

prehensible that in the Cryptoniscida©, in which sexual

products, i. e. male products, are so very early developed, the

fatty body will also be reduced very early ; and in a certain

sense it is rendered unnecessary by the intestinum glandulare^

which, indeed, does not serve for the accumulation of already

assimilated nutritive material like the fiatty body, but never-

theless accumulates unassimilated food in enormous quantities ;

and this, as it is assimilated, goes directly to the advantage

of the ova. It may, however, be supposed that the urinary

masses are separated from a fatty body which existed during

the larval period ; they are actually present in the greatest

Blanc, Observations faites siir la Tanais Oerstediii' in Zool. Auzeiger,

1883, p. G37.

17

On the Spongia coriacca of Montagu,

quantity during the male sexual maturity, and diminish sub-

sequently in amount, not only relatively but absolutely.

Nevertheless, I only wish to indicate a possibility ; it seems

a more probable supposition that these urinary concretions

originate in the blood and are deposited in the wall-less blood-

lacuna3, the most important of which indeed run along the

intestine. The decrease of these pigment-secretions coincides

Tvdth the commencement of the sedentary mode of life, and

therefore also with the complete change of nourishment, and

may consequently be caused thereby instead of by the disap-

pearance of the fatty body.

That the rectal vesicle produces a strongly smelling sub-

stance, as Fraisse asserts, may be correct ; but I cannot confirm

it, as unfortunately (or shall I, as a zoologist, say fortunately ?)

I possess a very feeble sense of smell.

The other internal organs of the Cryptoniscidse show no

great differences from those of the Bopyridae ; what there is

to be said about them and about the details of the external

organization I reserve for my monographic publication.

II. — On the Spongia coriacea of = Leucosolenia

coriacea, Bk.^ together with a new Variety of Leucosolenia

lacunosa, Bk.^ elucidating the Spicular Structure of some

of the Fossil Calcispongice ; followed hy Illustrations of the

Pin-like Spicules on Verticillites helvetica, De Loriol, By

H. J. Carter, F.E.S. &c.

[Plate I.]

In 1871 Annals,’ vol. vii. p. 278) I gave a nomenclatural

account of Montagu’s Spongia coriacea=Grantia clathrus,

Sdt., coriacea^ Bk.,= Clathrina clathruSj Gray,

under the last name, which was subsequently changed by

Hackel into Ascetta clathrus Die Kalkschwamme,’ vol. ii.

p. 30) , and now I propose to add the result of a structural

examination, chiefly on living specimens, of this calcisponge

from this place, viz. Budleigh-Salterton, South Devon.

In limine^ however, it is necessary to clear up the confusion

that has arisen from Hackel having made a separate species

of Schmidt’s Orantia clathrus under the name of Ascetta cla-

thrus ^ with a different form of spicule from that which Schmidt

has given as characteristic of it (Spong. Adriatisch. Meeres,

Suppl. p. 24, Taf. iii. fig. 3 a), and which accords with that

Ann, & Mag, N, Hist, Ser. 5. Vol, xiv. 2

18

Mr. H. J. Carter on the

of all the other authors mentioned but Hackel, who, instead of

an equilateral triradiate with straight arms, obtusely pointed,

has given one with undulating rays, which are inflated at

the ends {op. cit. vol. ii. p. 36, Atlas, Taf. v. figs. ^d-f).

Now had Hiickel offered any explanation of this with re-

ference to Schmidt’s original announcement, one could have

understood the discrepancy ; but he neither refers to Schmidt’s

characteristic figure of the spicule of Grantia clathrus {1. c.),

nor does he use the specimens which Schmidt gave himi

1868 {op. cit. vol. ii. p. 32), but gives the result of his exami-

nation of those which he himself found in the spring of 1871

on the coast of the island of Lesina, in the Adriatic Sea, as

typical of Ascetta clathrus [ih. p. 33). Thus we have to

choose between Schmidt’s published figure of his own speci-

men in 1864 and Hackel’s of his own in 1872, in which

dilemma it is evident that the latter could not have been

Grantia clathrus^ and therefore that Hackel had no reasonable

grounds for making it so. Hence the species at Budleigh-

Salterton, being the original Spongia coriacea of Montagu,

must be viewed as Schmidt’s Grantia clathrus of 1864, and

not as Hackel’s Ascetta clathrus oi 1872; while as Gray’s

genus Clathrina^^ is founded on ClatJirina sulphurea^ —

Schmidt’s Grantia clathrus j which, again, is equal to Leuco^

solenia coriacea^ Bk., and occurs here, as elsewhere, occasion-

ally under sulphur-yellow and scarlet colours respectively, we

must adopt Bowerbank’s or Gray’s names for the whole ; and

as that of the latter is most expressive of the anastomosing

tubular structure of Spongia coriacea Mont., while it does

not indicate any particular colour or contain any other diffe-

rent structure, such as Leucosolenia hotryoides^ Bk., which is

simply branched Hie Kalkschwamme,’ Atlas, Taf. ix.

fig. 10), it is evident that Clathrina'''' is the most preferable,

as the original generic name of Montagu, viz. Spongia

must necessarily be changed by some one.

Of all the sponges growing on the rocks here that I have

seen, no species is more strikingly beautiful than Montagu’s

Spongia coriacea^ which, when fresh and extending over an

area of about four square inches, presents itself under the form

of a reticulated structure of an icy-white colour, in which the

reticulation is only just visible to the naked eye, but, when

magnified, contrasts favourably, as it veils the dark rock

beneath with the most chaste and exquisite network that could

be produced artificially. No representations of it hitherto do

it full justice in this respect, least of all those of Johnston and

Bowerbank.

Spongia coriacea 19

Tin’s network is composed of a hollow anastomosing thread

or fibre of variable diameter, whose interstices or meshes (for

a parallel illustration of this mesliwork see the body of

Leitcosolema locunosa^ var. PI. I. fig. 2, a, fj g) are

subject to infinite variety both in size and shape throughout

the specimen, which, when large, thins out from a massive

variable amount of thickness in the centre, seldom more than

2 or 3-12ths of an inch, to a single reticulated layer at the

circumference. These thickened centres are numerous in a

specimen about the size mentioned, and form several little

monticular elevations scattered over the surface, whose sum-

mits respectively end in a little short open tube wdth naked

margin, which is continuous with the neighbouring branches

of the hollow thread- work, and thus forms an osculum

or vent to this part of the sponge (see also PI. I. fig. 3),

wdiile the other ends of the thread are blind and attached to

the rock over which the sponge may be growing. So that

the hollow of the reticulated thread is homologous with the

cloacal cavity of the other forms of Calcisponges.

If we follow the development of this structure from the

youngest form that can be seen, viz. that wdiich has just come

from the embryo, it will be found to consist of a simple erect

sac, whose upper end is open and w^hose lower one is attached

to the rock on wdiich it may be growing. This is the com-

mencement of the hollow thread out of w^hich the largest

specimen is finally constructed. It may now be about l-30th

inch long by 1-1 20th inch broad, narrowing to a point at the

fixed end. Such are the measurements of the smallest forms

which are just now (April 10th) to be seen of all sizes and all

stages of development on the rocks at Straight Point”

here, where the species grows abundantly. In the next stage

the sac sends out a tubular bud, which may also descend to

the rock, and then, as the individual grows upw^ards and out-

w^ards, other similar buds are put forth which either descend

to the rocks for fixation or otherwise anastomose with each

other, until at last the reticulated structure first described is

attained ; but how the anastomosis is effected, that is, how the

loops are formed, I have not been able to perceive.

Thus the development is very simple, although the adult

form appears to be so complicated, and the reticulated struc-

ture not by any means confined to Grantia clathrus alone, but

common to a great many different species of Calcisponges, of

which the beautiful little Leucosolenia lacunosa^ Bk., of the

British shores is a stipitate form and also of different spicula-

tion j still, whatever the spiculation may be, the soft parts

2*

20

Mr. H. J. Carter on the

appear to be similar in all. Grantia clatlirus consists simply

of tlie spicular structure, tlie spongozoa (Geisselzellen) , and

the granuliferous sarcode (syncytium, Hackel).

The form of the spicules or skeletal support, which are

chiefly situated in the outer part of the wall of the hollow

thread, is of one kind only, viz. equiangular and equiradiate,

with simple straight rays, obtusely pointed, as already men-

tioned.

The spongozoa do not appear to differ in form from those

of the other Calcispongige, but instead of being arranged in

juxtaposition around the interior of globular or sacciform

cavities, with their respective cilia projecting into the in-

terior, they appear to form in juxtaposition a continuous layer

throughout the inner surface of the tubular thread after this

manner.

The granules ” of the sarcode, however, are very remark-

able from their size and dominant presence ; and although they

accompany the transparent sarcode everywhere, they appear

when in situ among the spicules and spongozoa to be loosely

grouped around a delicate nucleated cell respectively, the

“ Kern ” of Hackel.

It is now thirty-five years ago that I gave an illustrated

description of this granuliferous sarcode in Spongilla An-

nals,’ 1849, vol. iv. p. 91, pi. iv. fig. 2, a-f) in the living

state ; and as this seems to apply very nearly to that of Grantia

dathrns (so far as the dead state of the latter goes in a prepa-

ration to be presently mentioned), I will here quote the

original paragraph, viz. : —

‘‘ If a seed-like body [statoblast] which has arrived at

maturity be placed in water, a white substance will, after a

few days, be observed to have issued from its interior through

the infundibular depression on its surface, and to have glued it

to the glass ; and if this be examined with a microscope, its

circumference will be found to consist of a semitransparent

substance, the extreme edge of which is irregularly notched or

extended into digital or tentacular prolongations precisely

similar to those of the protean [amcBhcL]^ which, in progression

or in polymorphism, throws out parts of its cell in this way

(pi. iv. fig. 2, dd). In the semitransparent substance maybe

observed hyaline vesicles of diflferent sizes contracting and

dilating themselves as in the protean (fig. 2, e e), and a little

within it, the green granules so grouped together (fig.

as almost to enable the practised eye to distinguish in situ

the passing form [polymorphism] of the cells to which they

belong ; we may also see in the latter their hyaline vesicles

with their contained molecules in great commotion, and

21

Spongia coriacea of Montagu,

between the cells themselves the intercellular mucilage ”

[syncytium of lliickel] .

One cannot help observing here that, as the illustration to

the paragraph represents, the “ granules ” appear to have

been draggedoff their cells (Kerne), to become scattered in the

pseudopod ial sarcode, which thus also appears to be as homo-

geneous as that of an Amcpha. Yet it seems questionable

whether the cells from which this apparently homogeneous

sarcode has been derived do not still retain their individuality,

seeing that, in the conjugation (zygosis) of two Khizopods,

they with their granules appear to flow together as intimately

as two drops of water, that is, their individuality becomes

lost ; they put forth their pseudopods afterwards as if

thoroughly amalgamated ; and yet, after a little while, they

separate and appear to be the same in every respect as they

were before the conjugation. Or, these cells and their

accompaniments can unmake and remake themselves as the

occasion may require and with the materials that are nearest,

— so inexplicable are the phenomena manifestedby polymorphic

sarcode !

Such facts would lead us to infer that the syncytium is

composed of a congeries of polymorphic cells, which thus

simulate a homogeneous substance, just as Kostafinski, and

previously to him his teacher, A. de Bary, I think, has

stated respecting that wonderful moving fungus ^thaliumy

viz. : — That “ the contents of the spores at i' e time of germi-

nation, give rise at first either to a naked zoospore provided

with a nucleus, a contractile vacuole and long cilia [? two] ,

or to an amoeboid. These zoospores or amoebaB flowing to-

gether in masses give rise to mobile plasmodia ” (Kostafinski,

Dr. J., ^Monografia Sluzowce,’ p. 83, in Polish, 1875; ap.

Cooke, ^Myxomycetes of Great Britain,’ p. 1) ; while in my

observations on AEthalium at Bombay in 1861 these appa-

rently homogeneous masses or plasmodia evinced, during the

restless unceasing changes in form of the fungus, the power

of moving about and running together like so much water, of

constricting themselves isthmus-like almost to separation, of

flowing back together again, of spreading themselves out den-

dritically, and finally of ending in a motionless, circular, con-

vex mass, which soon became a heap of black-brown spores !

Keturning to the syncytium of Grantia clathrus one finds

the granules so much more strikingly developed relatively

here than in the other forms of Calcisponges, that one cannot

help questioning their nature and import.

Taking the granule singly, it is spherical, translucent, and

glairy, glistening from refraction of light, of a faint yellow

22

Mr. II. J. Carter on the

tinge, and varying under 1-GOOOtli of an inch in diameter,

altliough rarely attaining tills size in this state. They are,

wlien in situ^ congregated round a nucleated cell (the “ Kern ”)

which is often so indistinct here as to be very difficult to see,

owing to its delicate (? polymorphic) structure and the opaque

mass which the granules form when closely applied to it in

juxtaposition ; or they are scattered throughout the syn-

cytium in the same way as in the Foraminifera, as the

preparation,” to which I have before alluded, which was

made after Schultze’s method, described by him in his exami-

nation of Euplectella aspergillum Challenger ’ Keports,

separate copy, p. 5), plainly shows, where the granuliferous

protoplasm or syncytium can be seen in a reticulated branched

form extending across the cavity of the tubular thread, very

much like that of Grornia oviformis^ represented by Max

Schultze in his Organismus d. Polythalamien ” (1854,

tab. i. fig. 1). So that one feels inclined to infer that, ex-

cepting for its spicules and the spongozoa, the sponge "would

be very nearly allied to a Foraminifer in this respect.

Iodine does not turn them purple, nor does liquor potassee

dissolve them ; but strong nitric acid appears to destroy their

sphericity, which may be brought back again by the addition

of liquor potassge. This glairy refractive appearance gives

them the aspect of fat or albumen ; while, like the green

granules in Spongillaj they appear in the sulphur-yellow and

scarlet varieties of Grantia clathrus to be the seat of these

colours respectively, when they might be termed pigmental.”

It is possible that they grow into the larger cells of the pro-

toplasm (the Kerne ”), from which they appear to be

derived, when they may fulhl other offices ; for Lieber-

kiilin has long since shown that the Korperparenchym,” =

syncytium, can enclose and extract nourishment from Infu-

soria in the same manner as Actiriophrys soV (Muller’s

Archiv, 1857, Heft iv. p. 388). So the particles of wood

taken into the plasmodia of JEthalium indicate the same

consequence. But whatever the office of the granules may

be no one as yet has demonstrated beyond conjecture what

they are or what purpose they may subserve either in the

sponges or in the lihizopoda, — so they are still called the

granules.”

The canal-system consists of the usual inhalant and

excretory divisions : the former of minute pores which

can only be seen by the microscope on the outside of a

dried well-preserved specimen, where they are bordered by

the granuliferous sarcode or syncytium ; here, too, probably,

in the living state, composed of a congeries of distinct nu-

Spongia coriacea of Montagu, 23

cleated granxdiferous cells or bodies like the investing mem-

brane” of Spongillaj which, in combination there appeared

to me to have the power of opening and closing a pore

wherever they liked (see Ultimate Structure of Spongillaf

Annals,’ 1857, vol. xx. pp. 24, 25, pi. i. tigs. 6, 7) ; the

Wanderzellen ” or migrating, amoeboid cells of Schulze

(Zeit. f. w. Zool. 1878, Bd. xxx. S. 409 &c.) ; — and the latter

or excretory division, consisting of the general tabulation

of the reticulated thread, opening in the way and at the vents

mentioned ; homologous with the cloaca in the other forms

of the CalcispongifE, as before mentioned.

The process of reproduction by ova &c. is probably the

same as that of the other Galcispongige ; but in the hope of

determining this as well as when the elements of reproduction

begin to appear, I have gathered living specimens of Glathrina

coriacea^ Grantia compressa^ and Grantia ciliata^ var. spini-

spiculum (being together), from the “ Rocks ” here every full

moon (7. e. the springs ”), since the 11th of March last in-

clusive, at which time I could see no trace of these elements

in either of these species. They began to appear in the two

latter on the 10th of April, and were strikingly developed,

especially in Grantia compressa^ on the 12th of May, but not

advanced then beyond the w/?segmented stage. In Glathrina

coriacea no trace had then appeared ; nor is there any now on

h e 9th of June, although the fragments (taken from diffe-

rent localities) were placed in pure spirit directly they were

taken off the Hocks,” which preserv^es the collar and the

cilium of the spongozoa in their extended state. It is there-

fore plain that Glathrina coriacea^ in point of time, does not

develop its ova and spermatic cells so soon as Grantia com-

pressa and Grantia ciliata^ var. spinispiculum. My observa-

tions on the former in this respect for the other summer

months will be communicated hereafter.

As before stated, the general structure of Glathrina coriacea

= Grantia clathrus &c., that is, the reticulation formed by

the continuous anastomosing of a hollow thread-like fibre or

tube, is common to many Calcispongise, which Hackel has

divided according to their spiculations respectively, so that

they appear in several genera of the first two families of his

^‘natural system,” viz. the “ Ascones ” and “Leuconesj”

but of themselves they equally form a natural group in general

structure as distinct as it is totally different from that of the

other Calcispongiie, that is, the anastomosing reticulation.

Moreover, the little Australian calcisponge which I have de-

scribed and illustrated under the name of Leucetta clathrata

(‘Annals,’ 1883, vol. xi. pi. i. figs. 13-17) must form the

24

Mr. II. J. Carter on the

type of a division of tliis family, in which the anastomosing

thread-like fibre is solid instead of hollow — a form entirely

absent in the Kalkschwamme ” of Hiickel, but one of

much interest, as I have heretofore shown, in elucidating the

structure of some of the fossil Calcispongi^, which I hope to

still further advance by a description of the following new

variety of Leucosolenia lacunosa.

Leucosolenia lacunosa^ Bk., var. Hillierij Crtr.

(PI. I. figs. 1-5.)

Small, stipitate, erect ; body globular, obconic, rather

compressed and turned to one side ; stem cylindrical and

long, rather bent upon itself and compressed in its upper

part. Body not hollow, but composed throughout of massive

clathrous structure; stem solid (PI. I. figs. 1, 2). Colour

pale yellowish white. Consistence firm, resilient in the

head, hard and unyielding in the stem. Clathrous struc-

ture or network (fig. 2, g) consisting of a mass of reticu-

lated anastomosing thread-like tube (fig. 2,/ ) issuing from

the stem in several divisions (fig. 2, (/), and terminating

in the summit by a central dilatation into which the neigh-

bouring branches of the reticulated structure are gathered

together centripetally (fig. 3, Z>), finally opening by a single

naked aperture more or less protruded, which is the osculum

(figs. 2 h and 3 a) . Pores minute, in the wall of the tubular

structure. Stem consisting of a compact solid mass of spicules,

compressed in its upper part, which is expanded scopiformly

into the divisions ” mentioned (fig. 2, <f), which, being solid

like the stem at first, pass respectively by transition into the

tubular form which characterizes the structure of the body

(fig. 2, a), terminating below in a root-like expansion which

is fixed to the object on which the sponge may be growing

(fig. 2, e). Structure and composition of the wall of the

reticulated fibre the same as that of Glatlivina coriacea

mentioned, only the meshes of the network are elongated

vertically, which of course is followed in direction by the

branches of the tubular thread, i. e. from the stem to the

summit. Spicules of three forms, viz.: — 1. Triradiate, equi-

angular, inequiradiate, rays straight, smooth, rather obtusely

pointed, the longest, which in the largest of these spicules is

three or four times longer than either of the other two,

directed backwards (fig. 4, a) ; the rest infinitely variable in

size generally and in the unequal length of their rays, some

nearly equiradiate (fig. 4, h) ; longest ray of the larger tri-

radiates (fig. 4, a) about 48-6000tlis inch long by 2-(i000ths

inch broad at the base. 2. Linear, acerate in appearance, but

25

Spongia coriacea of Montagu,

consisting of two unequal portions divided by a slightly in-

flated node which belongs to the longest part, and is therefore

a little excentric (fig. 5, c) ; divisions smooth, but more or

less varying in thickness here and there, espeeially towards

the ends, which are obtusely pointed ; one division straight

and the other a little curved so as to form a very slight angle

with the straight one ; largest average size about 9l-6000ths

inch long by 2-6000ths inch broad (fig. 5, a). 3. Linear-

vermiculate, smooth, attenuated towards the extremities, which

are pointed ; divided like the foregoing by an excentric node

(fig. 5, c c) ; amount of vermiculation and total length very

variable, the smallest perhaps about 16-6000ths inch long by

l-6000th inch broad, but immeasurable generally, from the

amount of contortion (fig. increasing in size and de-

creasing in vermiculation so as at last to reach an interme-

diate form (fig. 5jd). The triradiates are equally present in

the body and stem ; but the linear and vermiculate spicules

are exclusively confined to the latter, where they form the

outer layer and the triradiates the axial or internal structure ;

they do not begin to appear before the stem begins to divide

into the branches leading to the head (fig. 2, o?), and then go

on increasing in number and robustness, although not in

length, down to the root-like expansion or oldest part, as is

usual in most sponges. Size of largest specimen (fig. 1)

9-12ths inch in total length, of which the body is 3i-12ths,

and the stem the rest, viz.5J-12ths inch; greatest diameter of

the body about 3-12ths inch, that of the stem close to the

body l-12th, and that towards its base l-24th inch.

Hah. Marine ; growing on hard substances.

Log. Kamsgate pier, Kamsgate.

Ohs, Independently of the interest attaching to this sponge

as a variety of Leucosolenia lacunosa^ Bk., it is still more inter-

esting as presenting a spiculation and structure which reveal

the nature of the filiform spicules ” and structure of the

fibres in some of the fossil Oalcispongias from the “ Coral

Bag” of Faringdon in Berkshire.

These spicules, although first represented by Zittel, in 1878,

in Peronella multidigitata (Abh. k. bayer. Akad. d. W. ii.

Cl. Bd. xiii. 2 Abth. Taf. xii. fig. 3), were just afterwards,

that is in the same year, more particularly described and illus-

trated under the above name, viz. filiform spicules,” by

Sollas (‘ Annals,’ 1878, vol. ii. p. 356, pi. xiv. figs. 1-5) ;

and subsequently described by myself (^ Annals,’ 1883, vol. xi.

P-22)..

Until Zittel had kindly convinced me by a microscopic

preparation of Feronella multidigitata from the Cretaceous

2G

Mr. II. J. Carter on the

of Le Mans, that Calcisponges existed in a fossilized state, I

was inclined to discredit the fact, as my actual experience of

the delicate structure and perishable nature of the spicules of

the Calcispongia3 then seemed to point out that their structure

and spiculation was such that they must inevitably go to

pieces immediately after deatli, and therefore that the proba-

bility of a Calcisponge becoming fossilized was very doubtful.

When, however, convinced of the error I fully expected

that recent specimens would be discovered which would ex-

])lain all the then anomalous structure and spiculations in the

ibssil ones ; and the first that tended chiefly towards this was

the discovery, by Dr. Hinde, that the fibre of his Verticillites

d* 0 rhignyi from the Upper Greensand of Warminster was

composed of three- and four-rayed calcisponge-spicules, which

were so far loosened by disintegration that they could be

easily extricated entire, and thus viewed under the microscope,

mounted in balsam or otherwise, indeed a simple lens is

sufficient Annals,’ 1882, vol. x. p. 192 et seq. pi. xi.).

At the same time Dr. Hinde discovered in his Sestrostomella

rugosa from the Cretaceous of Vaches Noires, near Havre

(ibid. pi. X. fig. 4, and pi. xii. fig. 12, &c.), the two-pronged

tuning-fork ’’-shaped minute spicule first represented by Dr.

Bowerbank (Mon. Brit. Spong. vol. i. p. 268, pi. x. fig. 237)

from a recent calcisponge at Freemantle, in S.W. Australia,

and subsequently by Hackel in his Leucetta pandora from

the Gulf of St. Vincent &c. in S. Australia (^ Die Kalk-

schwamme,’ Atlas, Taf. xxiii. fig. A). I was myself able

also to confirm these observations respectively in Verticillites

anastomans from the Coral Bag of Faringdon and in a speci-

men of Sestrostomella ^Yom. the Jura, kindly sent me by Zittel,

when I also published the illustrated description of the little

calcareous sponge from Freemantle, in w^hich the clathrous

structure was shown to be formed by the reticulated union of

a thread-like element similar to that of Clathrina coriacea and

Leucosolenia lacunosa^ but, as before stated, solid like the stem

of the latter, and not hollow like the tubular thread of the head,

being composed of a layer of small triradiates externally with

a much larger and different triradiate-form axially or within

(‘ Annals,’ 1883, vol. xi. p. 33, pi. i. figs. 13-15) ; and now I

have had the opportunity of describing one from the coast of

England, in which the filiform spicules,” together wdth the

solid fibre in the fossil species, also receive an explanation from

a recent species.

The specimens of this sponge, which are in spirit, were

gathered on the pier at Bamsgate by Mr. Hillier, after whom

I have designated the variety, and presented to me by Mr.

27

Spongia coriacea of Montagu.

B. W. Priest in September 1882, when I tliouglit, from tlieir

resistanee and apparent durability on being handled, that

had I been acquainted with them earlier I should never have

discredited the fact that a calcispongc could be fossilized.

Thinking, however, from their resemblance that they were

specimens of Leucosolenia lacunosa^ I put them aside under

this belief ; but lately I have had to examine them in con-

nexion with the foregoing species, viz. Glathrina coriacea.^

and then I perceived that the solidity of the stem and its

spicular composition were like the fibre of Peronella multi-

digitata^ Zittel, of 1 Scypliia perplexa^ Quenstedt (tab. 125.

fig. 63), and of Manon peziza^ also Quenstedt (t. 132. fig. 30),

respectively ; that is, that it was composed of triradiates in the

centre faced by a layer of linear and vermiform spicules, each

of which indicated by the kind of node mentioned (PI. I.

fig. 5, ccc) near the centre, which slightly projects, that it

represented the aborted state of a third ray, and thus a modi-

fication of the triradiate.

Now, when we consider that the stem, as it approaches the

body (Pi. I. fig. 2, d)j divides into a multitude of branches,

each of which, although solid in the first instance, becomes

transformed into a tube to form the tubular thread of the body

(fig. 2,/) , which by branching and anastomosing produces the

clathrous structure in which the linear and vermicular spicules

are entirely absent^ and that the linear and vermicular spicules

thus cease to appear where the transformation takes plaee^ it fol-

lows that had the branches continued solidYi\<iQ the thread of the

clathrous structure in Leucetta clathratajQxiY. {op. et he. cit.)^

they would have been identical in spicular composition and

arrangement with the fibre of the fossils mentioned, where, on

account of their contortion being perhaps more generally

greater than that in Leucosolenia lacunosa^ var. HiUieri^ the

extreme thinness of the microscopic slice cutting off the bends

above and below, seldom allows one to be seen entire. Indeed

the more contort ones in Leucosolenia Hillieri during the

boiling out in liquor potass93, from this together with the

brittleness of the material, for the most part, come out broken.

Thus the filiform spicules ” of the fossil Calcispongige seem

to be elucidated.

Pin-like Spicules (? parasitic) on Verticillites anastomans

and A. helvetica. (PI. I. figs. 6-10.)

At the conclusion of my paper on the Fossil Calcispongiae

of Faringdon (^Annals,’ 1883, vol. xi. p. 33) I had only

just time to mention Dr. Harvey B. IIoll’s discovery of pin-

like spicules in that variety of Verticillites designated “ helve-

28 On the Spongla coriacea of Montagu,

tica ” by De Loriolj so I returned to the subject {ih. vol. xii.

p. 26), when by having made and mounted microscopic

sections myself, I was enabled to give a more detailed de-

scription of the fact, and to announce that such spicules also

existed in the same position in my specimens of VerticilUtes

anastomans horn. Faringdon; but on neither occasion had I

time to illustrate this interesting discovery, which is so

likely to pass unnoticed without representations, that I have

availed myself of the present opportunity to fill up a vacant

space witli these, taken from my own preparations (PL I.

figs. 6-10).

EXPLANATION OF PLATE I.

Fig. 1. Leucosolenia lacwiosa, var. Ilillieri, n. var. Natural size.

Fig. 2. The same, mag'nihed 3 diameters, to show: — a, body, composed

throughout of massive clathrous or reticulated structure. Body

not hollow, h, vent or osculum ; e, stem ; d, scopiform expansion

of stem into reticulated structure of body j e, root-like attach-

ment; f, tubular thread-like fibre; g, meshes of clathrate

structure.

Fig. 3. The same. Summit more magnified, to show the continuity of

the vent with the reticulated tubular structure of the body.

a, vent ; b, tubulated thread or fibre ; c c, meshes or interstices

of the reticulated structure.

Fig. 4. The same, Triradiate spicules of the body and centre of the

stem respectively, a, largest form ; h, smallest ; c, dotted line,

illustrative of the inequiradiate forms.

Fig. 5. The same. Substraight linear and linear contort spicules of the

stem, a, substraight linear ; b b, linear contort ; c cc, nodes ;

d, intermediate form.

N.B. — All these spicules are drawn to the scale of l-24th to

l-6000th inch.

Illustrations of the Pin-like Spicules (^parasitic') ^c. on Verticillites.

Fig. 6. Verticillites helvetica., De Loriol. Horizontal section of the wall

of a cylinder at the inflation, magnified 4 diameters, to show

the structure of the wall and its hourglass-shaped openings or

canals, a, cavity of inflated chamber ; b, wall, apparently com-

posed of little oval and quadrangular elements, because, in some

instances, the section has passed through the hourglass-shaped

canals, and in others not. Diagrammatic.

Fig. 7. The same. Vertical section of part of the wall of a cylinder at

the inflation, viewed from the inside ; magnified on the same

scale, to show the structure of the wall and its hourglass-

shaped openings in this view, a, wall, now seen to be continuous

and not formed of separate elements, as the foregoing figure

apparently represents ; b, inner opening of the hourglass-shaped

cauals, with a dot in the centre, to represent the narrow part.

Diagrammatic.

Fig. 8. The same. Horizontal section of two of the so-called oval

elements ” with the hourglass-canal between them filled with

sand ; magnified on a scale of l-24th to l-1800th inch. «, out-

side of inflation ; b, inside ; c c, so-called oval elements,” com-

posed of homogeneous crystalline calcite, with minute fibrous

\n n Ma fj Xa I . Ihst .S.:> In/ /.'J. //. /.

LIBRARY

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UNIVERSITY OF ILLINOIS

Dr. W. Djbowski on Lubomirskia balcalcnsis. 29

ptructiire (? product of fossilization) ; hourglass-shaped

canal filled with grains of quartz-sand ; e e e, pin-like spicules

in the calcite,” arranged around the funnel-shaped openings

of the hourglass-canals outside respectively, as will he better

understood by the next figure, but here only seen in the section,

where they may be observed to slope inwards with the head ex-

ternally ; ff, row of triradiate spicules in the “ calcite ” within

the pin-like spicules. Diagrammatic, with the detail relatively

magnified.

Fig. 9. The same. Pin-like spicule, more magnified, to show its shape

and relative proportions.

Fig. The same. Vertical section of a portion of the wall at the infla-

tion, viewed from the inside, magnified to the same scale (see a

less magnified portion, fig. 7). a a, wall composed of the homo-

geneous crystalline calcite with minute fibrous structure before

mentioned ; hhh, constricted parts of the hourglass-shaped

canals, respectively filled with quartz-sand, also as before men-

tioned ; ccc, position of the triradiate spicules in the wall

around the hourglass-canals, shown by their truncate ends ; d,

part of the slice where the layer of triradiates has been

ground off, showing that eee, the cross-sections of the pin-like

spicules, are in circles, indicative of their infundibular mode

of arrangement around the external openings of the hour-

glass-shaped canals. Diagrammatic, with the detail relatively

. magnified.

III. — /Some Remarks upon the Variohility of Form in Liibo-

mirskia baicalensis, and upon the Distribution of the Baikal

Sponges in general. By Dr. W. Dybowski With P.SS.

by H. J. Carter, F.E.S. &c.

[Plate II.]

During the printing of my memoir on the sponges of Lake

Baikal f, I received from Irkutsk, from my brother Dr. Bene-

dict Dybowski, a photographic representation, prepared by

him, of Lubomirskia baicalensisj and also a communication

upon the general occurrence and distribution of sponges in

Lake Baikal. These notes possess no little scientific interest,

and may therefore serve to complete my memoir, so that I

regard it as advisable to publish them as a brief supplement

to my work above cited.

* Translated from a separate copy, communicated by Mr. Cai'ter, of the

paper published in the ^Bulletin de I’Academie des Sciences de St.

Pelersbourg/ tome xxvii. pp. 45-50.

t W. Dybowski, Studien fiber die Spongien des russischen Eeiches

mit besonderer Berficksichtigung der Spongien-Faiina des Baikal-Sees,”

in Mem. de TAcad. des Sci. de St. Petersb. ser. 7, tome xxvii. no. 6

(1880).

30 Dr. W. Djbowski on Lubomirskia baicalensis.

The variability of the sponges in regard to their mor-

phology is a generally known fact. That Lubomirskia

baicalensis follows this general rule we have a satisfactory

proof in the material before us.

The morphological variability of Lubomirskia baicalensis

has already attracted the attention of Miklucho-Maclay * ;

but, as may be concluded from his words (/. c. p. 8), he was

acquainted only with inconsiderable variations. In order to

furnish as complete a description as possible of the form and

structure of our sponge, I will here summarize all that is

already known upon the subject, and enlarge it by my own

observations. But, remarkable as are the differences in the

form of the sponge under consideration, all these morpho-

logical deviations may always, by careful investigation, be

referred to one and the same type.

The simplest and therefore typical form of our sponge is

that of an arborescent stem with cylindrical erect branches f.

The branches originate sometimes at different heights (see

Miklucho, 1. c.), but sometimes at the same level (see l)y-

bowski, 1. c.).

This simplest form as just described- is modified in various

ways, and the most important modifications are the fol-

lowing : —

I. Forms in wliicli the type is distinctly recognizable.

a. The erect cylindrical branches of the sponge are not free throughout,

but unite with one another by several transverse anastomoses of

various thickness and length (see Dybowski, 1. c. p. 12, tab. i.

fig. 1).

h. The erect cylindrical branches of the sponge stand so close to each

other or to the stem that at the surfaces of contact they coalesce

either with one another or with the main stem. By this means

there are formed very variously shaped, elongated, more or less

flattened bodies, from the top of which larger or smaller branches

originate (see PI. II. fig. 1 h).

II. Forms in which the type is almost entirely effaced.

a. Arborescent sponges.

a. The branches do not stand erect, but form a more or less acute

angle with the main stem. The individual branches are not

cylindrical, but thinner at the free extremities than at the base.

("Specimen in my collection.)

j3. The short nearly cylindrical branches are pinnately arranged ; but

* Miklucho-Maclay, Ueber eiuige Schwamme des nordlichen Stillen

Oceans und des Eismeeres,” in Mem. de I’Acad. des Sci. de St. Petersb.

ser. 7, tome xv. no. 3.

t See Dybowski, /. c. tab. i. fig. 1 ; Miklucho-Maclay, 1. c. tab. i. fig. 5 ;

and Middendorfi;, ‘Sibirische Keise,’ Bel. iv. Theil ii. Lief. 1, p. 10(35.

Dr. W. Djbowski on Lubomirskla baicalensis. 31

they oricrinate only from one side of the stem (semipinnate)

(see PI. II. fig. I 5).

h. Bush-like sponges. When numerous twigs grow forth in various

directions from a short and thick base, a bush-like form is pro-

duced (PI. II. fig. 1 «). In such a bush we observe very

differently formed twigs. Most of them are furcate and grow

together in their lower part. The size of a bush is sometimes

enormous.

With regard to the occurrence and distribution of the

sponges in Lake Baikal, my brother sends me the following

information * : —

In the south-western portion of Lake Baikal f, that is in

the whole stretch from Listwiennischnaja and Possolsk on

the one hand, to Kultuk on the other, the sponges occur

wherever the necessary conditions are present |.

As a rule the sponges occur wherever the bottom of the

lake is stony and wdiere large blocks of rock or wood are

lying about on it ; further numerous sponge-stocks are found

in those places where the steep rocky shore forms terraces

projecting into the water in steps. On the other hand, if the

bottom be sandy, muddy, or covered with small easily movable

stones, no sponges occur. At a depth of 100 metres there-

fore, where the bottom is alw^ays covered wdth fine mud, a

few small coating stocks occur only where large blocks of

stone or logs of wood project out of the mud. On the western

shore these conditions are abundantly realized, so that here

sponges are almost everywhere met -with. Close to the

shore, and in only inconsiderable depths, turf-like or cushion-

like sponge-stocks exclusively occur ; globular ones are rare,

arborescent forms are never found. The sponges usually are

firmly attached to the surface of large pieces of w^ood or rock ;

they show, however, a special inclination for rotting w^ood,

so that in certain places almost every log of wood bears

sponges. Sometimes one finds large logs which are regularly

coated with a crust of sponges.

In considerable depths, as, for example, at a depth of 3

fathoms ( = 6*3 metres), often quite close to the steep rocky

shore, bush-like sponges exclusively occur. In fine still

water, when the surface of the lake is as smooth as a

* In the absence of a monographic treatise on the Baikal sponges the

observer only distinguishes the arborescen t from the turf-like or cushion-

like stocks. * The separate species or varitjties are not specially treated,

but only the conditions of the Baikal sponges in general.

t See Dybowski, 1. c. p. 7, fig. I.

X Extract from a letter dated “ Irkutsk , 1878.”

32 Dr. W. Dybowski on Lubomlrskia baicalensis.

miiror, if we go about in a boat we can delight ourselves

with the sight of numerous beautiful colonies of sponge-

bushes growing up from the bottom of the lake. The bushy

sponges sometimes attain an enormous size. One of them

brought up by the dredge was so large that it by itself filled

the whole bag of the dredge*. Such large stocks, however,

are rare ; usually the size of the sponge-bushes does not

exceed 60 centim.

At a depth of 6-25 metres, arborescent or fruticose sponges

occur ; at greater depths only cushion- or turf-like sponges.

At a depth of 100 metres (the greatest depth at which

sponges have as yet been taken) we find only occasionally a

few small and flat stocks, and even these only when the above-

mentioned conditions are realized.

On the eastern shore the bottom of the lake is generally

covered with sand or with small easily movable stones ; there-

fore the sponges are here much scarcer. On the larger blocks

the sponges only rarely occur, because the west winds, which

are here prevalent, cause a continual succession of waves,

which is evidently injurious to the growth of the sponges.

On the rocky terraces of the shore there are a few cushion-like

sponges, but not in such abundance as on the other side, on

the western coast of the lake.

In the Angara (see Dybowski, I, c. fig. 1) the sponges occur

everywhere in great abundance from the mouth up to the

Taltzinskaja manufactory. They are exclusively turf-like or

lamelliform sponges, which adhere to large logs of wood and

blocks of stone or to smaller pieces of wood ; arborescent

forms never occur ; evidently the sponges require still water

in order to become developed in the arborescent form.

Between the Taltzinskaja manufactory and the city of

Irkutsk sponges occur much more rarely, as here large stones

and logs of wood are less numerous. Above Irkutsk the

Angara has not yet been examined for sponges.

If we briefly summarize all that is known with regard to

the distribution of the Baikal sponges, the following may be

said : —

1. Close to the shore of the lake, at a depth of 2-6 metres,

only turf-like.

2. At a depth of 6-25 metres, arborescent or fruticose.

3. At a depth of 25-100 metres turf-like stocks again occur.

At all these depths, of course, the sponges only exist under

the above-mentioned favourable conditions.

The hag holds about 40 pounds of mud.

Dr. W. Dybowski on Lubomirskia baicalensis. 33

The colour of the sponges is for tlie most part more or less

dark grass-green ; sometimes, however, they occur olive-green

or brown ; only those sponges which come from considerable

depths (60-100 metres), or have grown under stones, are

almost colourless. The sponges of an indeterminate dingy

greyish colour are probably dying or dead stocks.

The following parasites have hitherto been observed upon

the Baikal sponges — Gammarus 'parasiticus^ G. violaceus,

and G, violaceus var. virens^,

[P.S. — On the 29th of May last I received from Dr. Dy-

bowski small fragments of all the freshwater sponges from

Lake Baikal in Central Asia and the Pachabica-See at its

south-west extremity that he had deseribed and illustrated in

the ^ Memoirs of the Imp. Aead. of St. Petersburg ’ (tomes

xxvii. and xxx. nos. 6 and 10, 1880 and 1882 respectively),

together with one of his ‘‘^Bosilia (?) 8tepanowiij^ from the

neighbourhood of Kharkow in Southern Kussia, to whieh I

alluded in the ^Annals’ of April last (p. 272), by whieh I

have been enabled, through mieroseopieal examination, to

confirm all that he has stated of the forms of their several

spieules, espeeially those of the latter, of whose statoblast Dr.

Dybowski kindly sent me a sketeh on the 30th May last, to

show that it possessed a tubular extension of the chitinous

coat, accompanied by the cirrous appendages and armed with

statoblast-birotules of different lengths, like those of Mr. H.

Mills’s Carterius (olim Carterella) tuhisperma from the

Niagara Biver ; but on comparison with a mounted specimen

of the latter, I find D. Stepanowii sufficiently different to

merit a distinct appellation. During my examination of the

material above mentioned I have been struck with the ability

and accuracy of Dr. Dybowski’s observations, hence look

forward with much pleasure to his description and illustrations

of the statoblasts of his “ Dosilia (?) Stepanowii^''' which he

intends to publish on the earliest opportunity. — H. J. C.

P.P.S. — In another letter from Dr. W. Dybowski, dated

1st June, he sends me sketches of the statoblast of Mr. Potts’s

Spongilla friabilis^ Leidy, var. segregata^ which was also

found near Kharkow, in Southern Russia, thus adding another

locality to those already mentioned of this species. — H. J. C.]

* Dr, B. Dybowski, ‘^Beitrage ziir naheren Kerintniss der in dem

Baikal-See vorkoinmenden niederen Krebse aus der Griippe der Gamma-

riden,” in the Horae Soc. Entom. Boss. Beiheft zu Bd. x. p. 75, tab. x.

fig. 3, p. 76, tab. xii. fig. 5, p. 147, tab. iii. fig. 3 (St. P<5tersbourg, 1874).

Ann. & Mag. N. Hist. Ser. 5. Vol. xiv. 3

34

. ' Mr. A, G. Butler on three new Moths,

EXPLANATION OF PLATE II.

Fig. 1. Luhomirskia Pallas, sp. About one third of the natural

size. Three specimens, viz. a, 6, and c. From a photograph.

Fig. 2. The same. Skeletal spicule, lateral view and transverse section.

Magnified 650 times, a, lateral view ; h, transverse section.

(Mem. de I’Acad. Imp. d. Sc. St. Petersbourg, 7® serie, t. xxvii.

no. G, Taf. ii. fig. 5, h.)

IV. — Descriptions of three new Species of Moths from the

Island of Nias. By A. G. BuTLER, F.L.S., F.Z.S., &c.

The three following species were added last year to the

National Collection ; two of them are especially interesting,

as affording an admirable instance of protective assimilation.

Agaristidse.

Ophthalmis decipiens^ sp. n.

Allied to 0, mollis ] grey-blue, with a faint greenish tinge ;

veins black j the basal third of primaries crossed obliquely by

irregular black stripes, the last two of which form an ^-shaped

character from the subcostal nearly to the submedian vein ; an

oblique black-bordered oval marking immediately beyond the

cell ; a very irregular black band from costa to submedian

vein, where it runs inwards to join the oval marking; the

veins between this band and the external border more broadly

black than on the rest of the wing ; external border black,

gradually narrowing from costa to external angle, and den-

tated internally upon the veins ; fringe white at apex and

external angle : secondaries with the basal fourth blackish ;

a large black spot over the end of the cell, beyond which the

veins are black ; a broad and very irregular black external

border enclosing a nearly marginal series of elongated blue-

grey spots ; fringe white : body above black, spotted in front

with white, but the abdomen and sides of thorax banded with

pale bluish grey ; anus orange. Wings below nearly as above,

but the black markings much broader and the blue conse-

quently narrower ; all the wings with a large black spot over

the end of the cell : body orange ; legs striped, and base of

venter banded with black. Expanse of wings 51 millim.

An^o Ma^. Nal. Ih.^L. S. .5 . VoL. M-. It. II.

I

650/1 .

MmterrL Bros .litK.

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UWIVERSITV OF ILLINOIS

Mr. A. G. Butler on three new Moths. 35

ChalcosiidaB.

Laurion zehra^ sp. n.

Allied to L. ohliquaria of Borneo and Malacca ; primaries

black, with a > -shaped blue marking at basal third of costa,

followed by a broad, oblique, snow-white, externally blue-

edged belt from costa to external angle ; veins beyond the

belt bluish ; a blue and white subbasal dot : secondaries

reddish orange, with the base, costa, and a rather broad

external border black ; a strongly dentated greenish-blue

marginal stripe ; thorax black ; head dotted with blue and

white ; abdomen dull black, regularly banded with creamy

white. Wings below' somewdiat as above, but the primaries

w’ith one or two subbasal blue-edged w'hite spots, an oblique

lunulated bluish stripe just before the middle, apical area

greenish blue, w'itli black longitudinal stripes between the

veins ; secondaries with a pale blue basal spot ; the dentated

blue margin covering the outer half of the external border.

Pectus black, spotted with bluish white; venter broadly banded

with white. Expanse of wings 68 millim.

At once separable from L. ohliquaria by the great width and

white colour of the belt across the primaries, and the absence

of the black spot on the orange area of the secondaries.

Euschemidae.

Pancethia simulans^ sp. n.

Has the general aspect of Ophthalmis decipiens^ but is of a

paler blue-grey colour ; the wdngs are crossed by four oblique

series of black markings, the first consisting of unequal oval

spots, the third much the largest, but those on the secondaries

subconfluent ; the seeond series consists of tw’o reversed curved

lines united by cross lines at the extremities upon the costal

'margin and the first median branch ; below this is a pyriform

■spot on the internal border, follow^ed by a squamose black line

across the secondaries ; the third series consists of unequal

oval spots, the second and fifth large and double on all the

wings; lastly, a series of elongated black spots gradually

lengthening and widening towards the costa of the primaries,

where they unite so as to represent the black border on the

Ophthalmis ; body blue-grey, thorax banded with dull grey ;

abdomen with the last three segments bright ochreous. Wings

below duller than above, the black markings badly defined ;

pectus grey ; venter bright ochreous. Expanse of wings

48 millim.

A more elegantly formed species than P. georgiata and

difierently marked.

3*

36

Dr. Karl Krapelin on the Pulicidae.

V. — On the Systematic Position of the Pulicidae.

By Dr. Karl Krapelin

[Plate III.]

After my investigations on tlie buccal organs of the Diptera

and Bhynchota | had led tne to the conclusion that in the

former the true sucking-tube (not to be confounded with the

labium, which serves only as its sheath) was formed by a

dorsal and a ventral lialf-gutter (labrum and hypopharynx),

and in the latter by two double half-gutters laterally inter-

locked, it seemed natural to study also the aberrant members

of the two series in the light of this criterion, which applied

to all typical forms, in order to arrive at greater clearness

with regard to their relationships. In this respect no small

interest undoubtedly attaches to the group Pulicidag, which,

notwithstanding much difference of form, presents such a

uniformity of organization, and as to the systematic position

of which for more than a century the most different opinions

'have been expressed, without any generally acceptable and

well-established view having yet been arrived at.

The history of these opinions has already been given

pretty completely by Taschenberg in his Monograph on the

Fleas J, so that here a short recapitulation may suffice.

Linn4, as is well known, created an order Aptera for the

wingless insects, Myriopods, Spiders, &c., and in this the flea

found its place. A similar position was assigned to it by

Geoffroy, Cuvier, and Dum^ril, as also by Gervais j while, on

the other hand, the order Aptera was by many rejected as

unnatural, and the relationship of the Pulicidse with various

winged insects was asserted. Thus Kirch er referred them to

the Orthoptera, Fabricius and Illiger to the Khynchota, Kosel,

Oken, Strauss-Durckheim, Newman, Burmeister, Walker,

. Von Siebold, and others to the Diptera. Lastly, there were

also very early naturalists who would associate the flea with

none of the existing orders of insects, but postulated a distinct

order for it. The leader in this direction is De Geer. He

'was followed by Lamarck, Latreille, Kirby and Spence,

MacLeay, Leach, Dughs, Bouclffi, and Van der Hoeven, and,

* ‘Festschrift zimi 50-jahrigen Jubilaiim des Real gymnasiums des

Johanneiims,’ tiamburg, 1884. Translated by W. S. Dallas, F.L.8.

+ In part set forth in the preliminary communication “Ueber die

Mimdwerkzeuge der saugenden Insekten ” (Zool. Anz. 1882, pp. 574-70)

and in a memoir, “Zur Anatomie und Pbysiologie des Russels von

Musca ” (Zeitscbr. f. wiss. Zool. xxxix. pp. 683-719).

X Taschenberg, ‘Die Flohe ’ (Halle, 1880).

37

Dr. Karl Krapelln on the Pullci(la3,

amon^ later investigators, by Landois and Tasclienberg.

But although the last two authors especially pronounced most

decidedly in favour of the independent position of the Fleas in

the system, and although the most accepted special works

upon the Diptera exclude the Fleas as not belonging to the

series of forms in that order we find that even in the most

recent manuals of zoology the group of insects in question is

almost without exception cited as a suborder of the Diptera.

This may pass in the first place as a proof that really stringent

arguments have not yet been brought forward in favour of

either view ; but we might also derive the hesitation felt by

many zoologists to raise the rank of the Fleas (even under

otherwise sufficient grounds) from the circumstance that they

lead a parasitic existence, and by this means have possibly

undergone profound and peculiar morphological changes by

adaptation,” as is sufficiently established for other groups of

parasitic forms. In opposition to this, however, it must be

remembered that with only isolated exceptions (the females of

the Sarcopsyllidge) the Pulicidse are not stationary, but only

temporary parasites, that their whole development is completed

without parasitism, and that therefore we cannot well assume

any considerable adaptation to a parasitic mode of life. But

if this be so, if we succeed in proving that the Pulicidae possess

a series of morphological characto’s which cannot be regarded

as acquired by parasitism, we must necessarily, in judging of

their position in the system, consider the same points of view

to be prescriptive that have been generally adopted for the

establishment of orders, suborders, and families in the class of

insects.

These general points of view, however, do not offer us a

very brilliant prospect. The Linnean principium divisionisj

the form, number, and texture of the wings, having proved to

be untenable, we find on the one hand the kind of transforma-

tion and its various stages, and on the othei the structure of the

organs of the mouth, raised into the most important criteria

of the nearer or more distant relationship of the groups of

insects. But, as is always the case, when a single character

is thrown too much into the foreground, and the general

morphological relations of the two series of forms are not

allowed to be prescriptive, difficulties make their appearance

even with these apparently so thorough-going principles of

division, which considerably diminish their value. The

^ It is interesting that the well-known work on the Diptera of the

^ Fauna Austriaca ’ by Schiner certainly expresses itself decidedly enough

in the above sense, but then gives a dednition of the true Diptera, which

might very well embrace the Pulicidae.

38

Dr. Karl Krilpelin on the Palicldae.

group of the Orthoptera, which is certainly not very natural,

and their multifarious relations with the Neuroptera, the suc-

torial Apidm, the biting Mallophaga, and lastly the pupal

rest of the male Coccid^, may sufficiently establish this pro-

position. It is still worse, however, as regards general avail-

ability, with the distinctive characters of the orders generally

cited — the segmentation of the thorax and tarsi, the structure of

the wings, of the different buccal organs, antennae, &c. The

mere fact of the agreement or differenceof these organs individu-

ally cannot give us certainty as to the systematic relationship

of two series of forms, but only the examination whether the

general organization of one group, as expressed in the deve-

lopment of all morphological characters, shows or does not

b\\ow phylogenetic relations with those of another group; in

other words, whether the observed differences in the structure

of the parts may be referred equally well to a different fun-

damental plan ” in their arrangement, as to simple changes of

form and reductions, such as may be explained by altered

function. Self-evident as this proposition appears in the

light of modern zoology, the history of opinion as to the sys-

tematic position of the flea nevertheless shows very plainly

how little it has hitherto been taken into consideration by

entomologists. One important aid in such investigations

upon the true phylogenetic relationships of forms is unfortu-

nately at present still almost wholly shut out from us. I refer

to the anatomical structure of the organs. The knowledge of

this, and especially that of the generative organs, is at present

so imperfect that a detailed consideration of the internal

organization seems to be of little use in the classiflcation of

insects.

After these prefatory remarks upon the principles which are

or should be of force in the grouping of insect-forms, the

question as to the systematic position of the Pulicidse may be

])ostulated as follows : — Do they or do they not, in the totality

of their organs, show near relations of affinity with any of the

other groups of insects ? In the former case we should have

to arrange them in this group of insects ; in the latter we

must establish an independent order for them.

I naturally commence my examination with that order of

insects which, in the judgment of zoologists, has the most

right to receive the Pulicidae into it, namely the Diptera.

The series of the Diptera must decidedly be called a unitary

one ; but the two characters so often brought prominently

forward (a perfect metamorphosis and suctorial buccal organs)

do not alone establish this unity, seeing that we must also

ascribe them to the Lepidoptera, the Apidje, and the male

Dr. Karl Krapelin on the Pulicida3. 39

Coccida). Nay, even if we add the footless larvae and the

fusion of the thoraeic segments as further eriteria, we might

perfectly well unite the Bees with the Diptera. It is not the

simple fact of the suctorial buccal organs that is of importance,

but their specific structure, the position and arrangement of

the parts composing the suctorial apparatus. If we fix our

attention upon this point we at once recognize that the fly’s

proboscis is constructed upon a perfectly different fundamental

plan from that of the Apidm, that the two are not directly

phylogenetically referable to each other, but that, on the

other hand, the great variations in the buccal apparatus of the

Diptera only represent modifications of one and the same

type, distinctly demonstrable throughout. The characteristic

of the bee’s trunk consists in the development of the lower

parts of the mouth into the sucking organ, while the man-

dibles retain their original function; that of the fly’s pro-

boscis, on the contrary, in the employment of the labrum and

hypopharynx for the formation of the suckiog-tube, with

which the mandibles and maxillae associate themselves as

stjlets more or less developed as required, while at the same

time the labium in all cases has to form a protective sheath

for the comparatively delicate tube through which the fluids

ascend. This fundamental plan of the employment of the

parts of the mouth occurs, as already pointed out in the

introduction, in all the groups (except the Pulicidac) which

have hitherto been placed in the group Diptera, in the piercing

Culicidsc, Tabanidgs, and Asilidgs, the different families of

honey-suckers, and the Pupipara, which are so depressed in

position through parasitism ; nay, a bridge seems even to be

thrown over towards the rudimentary buccal organs of the

CEstridge, through the structures which occur in Cuterebra. In

figs. 1-3 (PI. III.) I have drawn transverse sections of the pro-

boscides of those groups of flies which, upon one hand or the

other, have been referred to as allied to the flea. While those

of Tahanus and Culex (figs. 1 and 3) agree not only in the

position but also in the number of the pieces composing the

proboscis, that of Meloj)hagus (fig. 2, the representative of the

Pupipara) shows a great reduction, which finds its expression

in the entire absence of the mandibles and maxillge* ; but

* The two valves embracing the proboscis of the Pupipara have

been very erroneously interpreted as maxillc©, their palpi, or even

as a bipartite epipharynx (Meinert). From the whole arrangement of

the proboscis, which is freely movable in a wide cavity of the head

extending as far as the prothoracic ring, we can here have to do only

with a conical prolongation of the head which has become paired, some-

what such as we should obtain if we imagined the slight emargination

at the apex of the frontal cone of lihingia carried down to its base. The

strongly projecting cheeks of many Conopidae might also perhaps be

regarded as analogous.

40

Dr. Karl Krapelln on the Pulicidae.

nevertheless it is easy even here to recognize the typical

position of the pieces forming the sucking-tube (dorsally the

labrum and ventrally the hypopharynx) , and the labium

which encloses these as a sheath. Further, the latter bears

at the end that enlarged portion which is so characteristic of

all Diptera, and which is probably to be interpreted as formed

by uniarticulate labial palpi.

The same unity in the Diptera appears also in the special

structure of the thorax and its appendages. That this appears

always separated from the head by a deep incision is cer-

tainly not without significance ; but it can furnish no decisive

datum for the collocation of the Diptera. Of more importance,

no doubt, is the fusion of the thoracic segments into a compact

thoracic mass, which occurs in all the forms referred to this

group. It is indeed true that in orders of insects (I refer

particularly to the Khynchota) the formation of the thorax

as regards the separation or fusion of the segments composing

it shows manifold differences, without its being necessary that

we should separate forms which are united for other reasons,

seeing that the fusion or separation of the thoracic segments

has to do essentially with a function of the mechanism of

flight, and the free segmentation of the thorax in a wingless

form may very well be explained as a correlative phenomenon

of adaptation. But the conditions are different if, on the

contrary, a wingless form exhibits complete amalgamation of

the thoracic segments. In my judgment it thereby demon-

strates most unmistakably its descent from winged insects,

and in this sense the compact structure of the thorax, with

the characteristic process of the mesothorax described as the

scutellum,” in Melophagus^ the Nycteribiidae, and the Brau-

lidse, decidedly acquires the significance of a still uneffaced

relationship with the winged groups standing next to them.

And just as on account of this character the assumption is

justified that the forms just mentioned stand in close phylo-

genetic relationship with winged insects, so does the ex-

amination of the dorsal appendages of the thorax lead to the

same conclusion. All Diptera do not possess a pair of

wdngs and a pair of halteres; but the two organs which,

because special, are certainly of such great importance in

characterizing the Diptera, disappear so gradually in the

continuous series of forms, that we may trace their progress

to the rudimentary state, as it were, step by step. An

Ornithohia pallida which, as Lipoptena cervij follows a per-

fectly different mode of life, enables us at once to understand

the case, when we see MelophaguSy which is never parasitic

upon birds, entirely destitute of wings. But as regards the

41

Dr. Karl Krapelin on the Pullclda?.

lialteres, these, notwithstanding Schiner’s assertion to tlic

contrary, are quite recognizable in the sheep-tick, while in the

Nycteribiida3 they show all gradations down to quite minute

points, so that the complete abseneeof these apparently insig-

nificant organs in the Braulidm need not give us any further

disturbance. The ventral thoracic appendages, the legs, cer-

tainly present but few differences in the group of the Diptera,

nevertheless the five tarsal joints whieh are usually present

are not always eonstant ; and further, other orders of insects

sufficiently prove how little importance attaches in general to

the number of tarsal joints and the development of the different

sections of the legs.

Tlie developmental stages of the Diptera do not show a

community of type so distinctly as the structural characters

just referred to. The larv^ are certainly throughout distin-

guished by the absence of jointed thoracic limbs, which is of

special interest in the case of those forms which live free upon

leaves by prey (many larvss of Syrphid^e) ; but with regard

to the structure of the head, the armature of jaws, and the

development of the tracheal system, there are, as is well

known, such important differences, that they have been

successfully employed for the systematic division of the order

into several suborders and sections. Nevertheless even here

intermediate grades are not wanting between the different

structural characters (witness the variable development of the

first cephalic segment) ; nay, in Brauer’s* opinion, the family

Lonchopterid£e may possibly prove to be a perfect transitional

group between the Orthorapha and Cyclorapha, so that the

multifarious forms of the larvte at least offer no veto against

the unitariness of the stem of the Diptera. The same thing can

also be said of the pupae, which indeed likewise fall under two

main types, but are so far brought together by Brauer’s inves-

tigations, that these furnish a proof that the so-called tun-

pupae” (obtected pupae) show very different grades of structure,

and in many of them the enveloping larva- skin bursts exactly

as in the ordinary moulting, and consequently is to be referred

simply to a delayed moulting at the close of the larval period.

In the latter case, moreover, if the appendages of the segments

of the body are not so closely attached to each other and to

the body as in the naked and consequently less protected and

more easily injured “ mummy-pupge,” no important objection

against the natural relationship of the two groups can be

derived from this circumstance, which evidently results from

* F. Braiier, ‘ Die Zweifliigler des Kais. Museums in Wien/ p. 9

(Vienna, 1883) ; also in the Denkschr. d. math.-naturwiss. Ivlasse d.

k.-k. Akad. d. Wiss. Bd. xlvii.

42

Dr. Karl Krapelin on the Pulicidae.

altered eondition. The mummy-pupae,” however, show

many differences among themselves with regard to the closer

or looser appression of the appendages of the body, as may

be demonstrated by a comparison of the pupae of the Asilidae,

which rest in the ground, and those of Ti'pulce which live in

the water.

Of anatomical peculiarities of the Diptera especial mention

must be made of the “ sucking-stomach,” which is always

j)resent, as also of the large thoracic salivary glands, the

efferent ducts of which, wherever the buccal organs perform

any function, unite into an unpaired closed canal, which,

running along in the cavity of the hypopharynx, opens at its

extremity. The testes are almost always two ; the Malpi-

ghian vessels almost as regularly four. As regards the

tracheal system, the constant absence of the first thoracic

stigma and the small number of abdominal stigmata are to

he noticed ; while the nervous system, as is well known,

shows all possible forms of development, from the most ex-

treme concentration to a very considerable segmentation of the

ganglionic chain.

If we turn from this brief account of the Dipterous type to

the characters of the Pulicidse, we must admit, in the first

place, that in a whole series of points of comparison an agree-

ment between the Diptera and the Fleas can be demonstrated.

Like the Diptera, the Fleas have a suctorial buccal apparatus,

a perfect metamorphosis^ and footless larvse ; as in them also

the tarsi are five-jointed, there are four Malpighian vessels,

and one pair of testes. But, as has already been indicated at

page 38, we could only ascribe decisive weight to this agree-

ment if all these characters were peculiar to the Dipterous stem

alone, and if at the same time, by more detailed comparison,

real tenable parallels could be drawn between the different

parts of the organs, as between the different stages of develop-

ment. This, however, is by no means the case. The number

of Malpighian vessels and of testes recurs in the same way in

the Rhynchota, and therefore proves no more in favour of the

relationship between the Fleas and the Diptera than the

number of the tarsal joints or the annulation of the terminal

knob of the antennae, which may be recognized in all possible

groups of insects. At the first glance more importance seems

to attach to the agreement of the two groups in the larval

state, which in fact goes so far, that Brauer* has no hesita-

tion about arranging the larva of the flea in his group of

orthoraphal eucephalous Dipterous larvae. In opposition to

Brauer, Kurze Charakteristik der Dipterenlarven,” in Verb, k.-k,

zool.-bot. Ges. in Wien, 18G9, p. 84G.

43

,Dr. Karl Krapelin on the PulicidoB.

tills, liovvevcr, we must not forget that maggot-like larvsQ also

occur in groups far removed from the Fly-type, in llyrneno-

])tera and Beetles, and therefore cannot possibly be of decisive

importance in judging of relations of alHnity ; as also, on

the other hand, that the pupa of the Fleas with its quite

separate limbs differs so much at least from the general

type of the mummy-pupje, that from this very fact it has

been attempted to set up a relationship of the Fleas to the

Hymenoptera Hence the point of the question how far

the analogous characters in Diptera and Pulicidas depend

upon true phylogenetic affinity would have to be sought

in the investigation whether the construction of the sucking-

apparatus is carried out in both cases on the same plan,

i. e. with the same employment of homologous parts. That

it is only from this discussion and from that as to the

structure of the thorax and its appendages that a real decision

of the question before us can be arrived at, may indeed be

deduced from the consideration that in these organs we find

the only characters which, on the one hand, are confined to

the order Diptera, and, on the other, may be traced through-

out their whole series of forms, and therefore must be regarded

Kar eioyjiv aS typical.

The structure of the buccal apparatus of the Pulicidse has

been very frequently discussed without the question of its

relationship to the sucking-apparatus of other groups of

insects having as yet been solved. Thus to cite only a

few : — Dugbs f thinks that the proboscis of the flea may be

placed side by side with that of the Tabanidse, but also finds

resemblances to the Hippoboscida3 and Apidee. L. Landoisj:

suggests a resemblance of the mouth-apparatus of the Puli-

cidge to the rostrum of the Hemiptera; while Taschenberg§,

again, thinks he recognizes the Dipterous type, and espe-

cially calls attention to the presence of a tongue ” as

the most characteristic part of the mouth of a fly. This ex-

traordinary diversity of opinions is principally to be ascribed

to the uncertainty of the interpretation of this very tongue ”

of Taschenberg’s. The mandibles, maxillas, and labium

have long since been recognized with certainty ; but the un-

paired piercer ” (to express myself neutrally) has been referred

to as the labrum (Westwood, Haller, Bonnet), as the hypo-

* As by Duges, in bis Recbercbes sur les characteres zoologiques

du genre Pulex, in Ann. Sci. Nat. tome xxvii. p. 157. °

t Loc. cit, p. 151.

X L. Landois, ‘^Anatomie des Hundeflobes,” in Nova Acta Acad

Leop.-Car. 1866, p. 56,

44

Dr. Karl Kriipelin on the Pulicidse.

pharynx (Gerstfeldt), as the epipharynx (Karsten), and lastly,

as already mentioned, as the tongue” (Savigny, Taschen-

berg) , and therefore all serious homologizing must have been

prevented, the more, as even the real components of the suck-

ing-tube were not made out with certainty.

In figs. 10 and 13 I give two transverse sections through

the anterior part of the Pulicid proboscis. Fig. 10 represents

a section from Palex irritans ; fig. 13 a similar section, but

nearer the base of the proboscis, from Sarcopsylla penetrans^.

The sections show at once that the structure of the siicking-

tube in the two most distant groups of the Pulicidae is quite

accordant. Tn both cases it is the mandibles [md)^ which, in

conjunction with the unpaired piercer,” form the true food-

canal ; embracing the latter above and laterally, they join

firmly together in the median line below. A glance of com-

parison at figs. 1-3 shows that this unpaired piercer ” is

hollowed into a groove on the underside exactly in the same

way as the lahrum of the Diptera, and that to begin with

there is no hypopharynx, but at the utmost perhaps an

epipharynx. But if we trace the further course of this struc-

ture by the aid of longitudinal and transverse sections it is

easily seen that its upper covering immediately after its en-

trance into the capsule of the head is in chitinous union with

the upper margin of the arch of the head, while the inferior

plate, i. e. the one which immediately forms half the sucking-

channel, passes continuously into the chitinous covering-wall

of the pharynx. Consequently we find in the organ in ques-

tion precisely the same conditions as in the labrum of the

Diptera, and there is no doubt at all that we have to do here

with a true labrum. A connexion of this with the labium

by means of a strongly chitinized, brown uniting piece, as

asserted by Duges [1. c. p. 150), really has no existence at allf,

and this removes the last possibility of regarding this struc-

ture as a tongue,” i. e. as an extension or appendage of the

labium.

The interpretation of the “ unpaired piercer,” as labrum,

being thus established beyond a doubt, the comparison of the

proboscis of the flea with that of the Diptera can present no

further difficulties. The employment of the labrum [Ir) as the

unpaired covering lamella of the food-canal is apparently the

same in both groups. But it is otherwise with the other

components of the sucking-tube. In place of the horizontally-

The material was kindly sent to me from Assumption by my honoured

cjlleague Dr. H. Tbppen.

t This chitinous piece rather forms the lever for moving the mandible,

as will be shown elsewhere.

45

Dr. Karl Kriipelin on the Pulicida3.

placed mandibles of the TabanidaB and Culicldse, whicli, as is

proved by those Diptera which do not pierce, are only secon-

darily implicated in the closure of the sucking-canal, we see

in the Pulicidm the vertically- placed mandibles, bent in

towards each other laterally, appear as integral parts of that

tube — a different inferior closure, such as exists in the hypo-

pharynx throughout the whole group of the Diptera, being

here entirely deficient. This absence of the hypopharynx,

which, as is clear from what has been said, has as its conse-

quence a totally different importance of the mandibles, and

consequently a perfectly peculiar type of sucking-tube*,

proves in like manner of importance as regards the discharge

of the salivary glands. The unpaired salivary duct in the

lumen of the hypopharynx is replaced in the Pulicidae by

paired extremely fine half-tubes (fig. 13, 5), each of which,

running along the inner side of a mandible, may be traced

from the basal part of the latter as a closed duct into the

interior of the head, and, further, as far as the thoraeic

salivary gland t*

Equally great differenees in their arrangement and physio-

logical importance may be demonstrated by a comparison of

the other constituents of the proboscis of the flea with the

homologous organs of the Diptera., A labium unpaired

throughout its whole length, and at the utmost furnished at

its apex with one-jointed terminal lobes, oecurs nowhere

among the Pulicidse, although something of the kind was

formerly aseribed to Sarcopsylla. The labium of Sarcopsylla

at least presents (as fig. 8 may show) a biarticulation of the

“ palpi,” even with an indication of further segmentation, so

that in this point also the unity of the Pulicide group appears.

This differenee of the segmentation of the labium in Diptera

and Fleas, with whieh a typical difference in the relative length

of the unpaired basal part to the paired section to be regarded

as palpi, goes hand in hand, can, however, hardly be so highly

estimated in its phylogenetic significance as the further fact

that the labium of the Diptera shows quite a different attaeh-

ment to the head, and so has quite a different physiological

value from that of the Pulicidge. In the former it generally

attaches itself by its gradually widening base to a more or

* Particular attention may liere be directed to the two peculiar lateral

lamellae of the labium, which apparently, by their elasticity, force the

upper parts of the mandibles asunder, and thus bring about a closer appo-

sition of their lower parts.

t Kraft and Landois believe that they have demonstrated an opening

of the thoi-acic salivary glands into the oesophagus not far from the region

of the neck (see Landois, 1. c. p. 18).

46 Dr. Karl Krapelin on the Pulicidas.

less developed cephalic cone, with the upper lateral parts of

which it is connected, and so is enabled from the base onwards

to form that sheath of the delicate piercing apparatus (the

two pairs of jaws as well as the labrum) which often arches

together above so as to constitute almost a closed canal. In

the Fleas, on the contrary, there is no such union of the

labium with the lateral or upper parts of the head ; it simply

articulates with a firm brown chitinous piece (fig. 9, ch)

in the median line of the lower surface of the head, and this

union, as is well known, is frequently so loose that it is

difficult to obtain Sarcopsyllw^ for example, with the labium

preserved *. Hence, in its basal part, it does not form the

sheath for the piercing-apparatus, but shows only a compara-

tively shallow groove (fig. 15), which only in the ante-

rior section of the proboscis, when the stem of the labium has

become cleft into the paired palpi, becomes developed, at least

in PuleWj into two haps, embracing the piercing-organ at the

sides (fig. 10, Ip), But to make up for the deficient protec-

tion of the basal part of the sucking-tube (and in this we have

a fundamental deviation from the type of the Diptera) the

maxilla3 hava come in, originating as two broad plates from

the whole length of the side of the head, and taking here, not

only the constituents of the piercing-apparatus, but also the

base of the labium, under their protection, as shown by fig. 15

in Pulex,

We seek in vain for analogies to all these characters among

the Diptera, and we may therefore be justified in asserting

that all the parts of the Pulicide proboscis (with the sole

exception perhaps of the labrum) differ so much in position

and employment from the homologous parts in the Diptera,

that we cannot well speak of direct phylogenetic relations

between the two types of proboscis.

We arrive at precisely similar conclusions as to the rela-

tionship of the Pulicidge and Diptera when we take into

consideration the second group of characters peculiar to the

Diptera, which appear in the structure of the thorax and its

dorsal appendages. Instead of the always freely movable

head of the Diptera, we find a broad union of it with the pro-

thorax in the Pulicidse \ instead of the compact thorax with

its scutellum, which is so characteristic even of the wingless

Pupipara, we have three sharply separated thoracic segments,

without a trace of any such dorsal mesothoracic process ; and

instead of the pair of wings and the halteres, the latter of

which are aborted only in the most extreme cases of parasitism,

* Even in recent handbooks we may find the statement that the

labium of ^arcojisylla is indistinct.

47

Dr. Karl Krapelin on the Pulicidge.

there is notlimg^ absolutely notliing, that could lead us to

conclude that the Fleas were formerly in possession of any

such organs. Even the marked tripartite condition of tlie

thorax ought h 'priori to have banished the idea of rudi-

mentary wings ; nevertheless the older authors (Kirby, Dug^s,

&c.) have fallen into the serious error of regarding separated

lateral margins of the thoracic segments as such. But these

processes of the pleuras,” as Taschenberg among others

has conclusively proved, have nothing at all to do with wing-

rudiments, and are to be regarded as characteristic structures

sui generis. When Taschenberg therefore for this reason

declares the generally- employed denomination of Apliani-

ptera,” founded upon this erroneous conception, to be inad-

missible, we can only agree with him. It is only by giving

up this name that we can seriously hope that the deeply

rooted notion of the Dipt^res sans ailes,” as Strauss-Durck-

heim called the Fleas, will be completely suppressed.

The wide gap which exists precisely in the most important

characters between the Pulicidse and the Diptera must have

been made sufficiently evident by the preceding remarks.

That it is also expressed in other systems of organs than those

hitherto considered may therefore only be briefly indicated.

The sucking- stomach, which apparently is met with in all

groups of Diptera, is entirely wanting in the Pulicidge ;

while, on the other hand, the proven triculus beset with nume-

rous chitinous spines of the latter has no analogy among the

Diptera. The sucking-mechanism of the pharynx or of the

so-called “ fulcrum of the Diptera is formed by a single

powerful pair of muscles ; in the Fleas, on the contrary (as

in the Bhynchota), a whole seiies of separate pairs of muscles

(which, however, are interpreted by Landois as flexors and

retractors of the labrurn) are present for this function. Lastly,

the presence of a stigma in the protliorax of the Fleas indi-

cates more profound differences in the tracheal system ; while

as regards the simple ocelli of the Pulicida3 and the deep

lateral pits of the head, we may find analogous phenomena

among the Ehynchota, but not among the Diptera.

After all this the Fleas cannot well remain in the order

Diptera. There remains then the investigation of the ques-

tion whether they show near relations to any of the other

groups of insects. Hymenoptera and Ortlioptera, of which

earlier authors have thought in this connexion, cannot well

come into the question in the present state of our knowledge,

as it would be opposed to all rational system to assert a rela-

* Zoc. cit, p, 21.

48

Dr. Karl Kriipelin on the Pulicidse.

tionship of the Fleas to the Hymenoptera upon the sole

accordance of the pupse, or to the Orthoptera upon the

segmentation of the thorax. The order Lepidoptera also

cannot agree in a single one of the more important characters

with the PulicidEG, and thus there remains only the group

Phynchota for serious comparison. As a matter of course,

considering the fundamental difference of development between

Pulicid^ and Khynchota, we can hardly expect to find real

intimate relations between the two groups, at least not so

close as we must postulate for forms of one and the same

order ; nevertheless I think I may indicate some points of

view which deserve to be well considered in judging of the

phylogenetic connexion between Fleas and Phynchota.

In the first place there can be no doubt that the order

Phynchota does not even approximately present a unitary

type in the same degree as tliat of the Diptera. We find

united in it animals with suctorial and masticating buccal

apparatus, with perfect, imperfect, and without metamorphosis.

The head is sometimes freely movable, sometimes attached by

'a broad surface to the prothorax. The thorax, so very uni-

formly constructed in the Diptera, shows all possible stages

of structure, from the enormous development of the sepa-

rated prothorax in Scutata and Membracina, to the compact

thorax showing scarcely an indication of segmentation of the

Pediculina, or that of many Mallophaga more or less sharply

divided into three distinct segments \ and like the thorax

itself, its dorsal appendages also present no unity of type.

With such polymorphism of almost all organs it is easily intel-

ligible that we should be able to find in this Protean group ana-

logies for a whole series of characters of the Pulicida3. Thus the

segmentation and winglessness of the thorax in the Fleas may

be without difficulty placed side by side with the similar con-

ditions among the Mallophaga, which, at the same time,

present examples of the antennary pits of the head already

mentioned. The absence of facetted eyes in Pulicidas agrees

with what occurs in Coccidse, Pediculinas, and Mallophaga,

the pupa enclosed in a cocoon unites them with the Coccidje ;

the absence of sucking-stomach and the number of the

Malpighian vessels and testes are even common to them and

to all forms of Rhynchota.

For the reasons above given, however, we must not ascribe

a serious significance to all these agreements unless the

Rhynchotan type sought for finds expression at least in the

last of the characters to be discussed, those of the buccal

apparatus, and shows near relations to the homologous organs

of the Fleas. According to the present state of our know-

49

Dr. Karl Kriipelin on the Pulicidae.

ledge it cannot well be maintained that there is a clear uni-

tariness of structure in the buccal organs of the Rhynchota,

as, at any rate among the Aptera (tlie Pediculina and Mallo-

phaga), conditions occur which depart widely from those of the

more highly organized groups. But as the arguments upon

this point are not yet closed and I have made no investigations

upon these lower forms, we must content ourselves with

examining at least the sucking- apparatus of the Hemiptera

and Cicadae in search of any agreement with the proboscis

of the Fleas that may exist. With regard to the arrangement

of the parts of the mouth in these higher groups of the Rhyn-

chota, I have already published some statements in a previous

note *, and these observations have since been confirmed and

extended by Geise f- According to these the true sucking-

tube of the proboscis is formed by the two maxi lias closing

laterally againsf each other into a double tube, while the

mandibles are placed alongside of this tube as lateral piercing-

setae. From more recent investigations I do not hesitate to

declare this view J so far erroneous that it is not the maxillm

but rather the mandibles that interlock in the median line to

form the sucking-tube (see figs. 11, 14). I am led to this

changed interpretation of the two pairs of jaws in the first

place by the fact that in transverse sections through the head

the lateral setae finally come to be the lower ones, as, indeed,

Geise correctly shows in his figs. 25 and 31. Secondly, I

think that in the Cicadas I have found distinct traces of basal

joints of the maxillae connected with the outer setae. Fig. 12

shows the lower part of the face of a large tropical Cicada.

On each side of the broad labrum (Ir) there is here an oblong

plate (pl)j which terminates almost in the middle line

beneath the labrum in a blunt hairy tubercle and a peculiar

whip-like appendage (fig. 6,/) . If this structure be prepared

out of the head, a connexion, certainly only by articulation,

with the lateral piercing setae may be easily demonstrated,

for protrusion and retraction of which not only the chitinous

sinews (fig. 6, sp and sr)j but also the corresponding muscles

(fig. 6, pm and rm) are attached to this chitinous piece. If

this interpretation of the chitinous piece occurring in all

Cicadae, I ulgorinae, &c., as the basal part of a jaw, perhaps

even with palpiform appendages, be correct, this must, of

* Zool. Anzeiger, 1882, p. 574.

t Geise, ‘ Die Mimdtlieile der Rliynclioten ’ (Bonn, 1883).

X On my part this resulted merely from what I now believe to be a

wholly mijustified homologiziiig with the buccal organs of the Lepido-

ptera, the sucking-tube of which is undoubtedly formed of the maxillge

(see also Kirbach, Zool. Anz. 1883, p. 553).

Ann. (& Mag. N. Hist. Ser. 5. Vol. xiv.

4

50

Dr. Karl Krapelin on the Pulicidse.

course, be a maxilla, and thus the composition of the sucking

tube out of the two mandibles would be finally decided. Bu

then we should at once be enabled, in one point, to carry ou

a corresponding comparison between the buccal organs of the

Pulicidai and Khynchota, inasmuch as we need only suppose

the labrum of the latter, which is indeed often enough developed

into a long, slender, stylet-like organ, to sink from above be-

tween the mandibles*, in order to arrive at conditions which

might perfectly well be placed side by side with those occur-

ring in Pulicidae (compare fig. 11 with fig. 15). It appears

further that upon the basis of my conception a connexion

might be established between the modes of discharge of the

saliva in the Pulicidae and Khynchota, if we assume that the

paired half-grooves along the inner side of the mandibles of

the Pulicidae (fig. 13, s) have coalesced in consequence of the

changed adhesion of these jaws, caused by the emergence

of the labrum, as a constituent of the sucking-tube, into an

unpaired efferent canal (%s. 11 and 14, s). The variable

part taken by the two mandibles in Hemiptera and Cicadas

(see fig. 14) in the formation of this salivary tube would

come in support of this hypothesis. Among the Khyn-

chota, as is well known, a hypopharynx is not developed as a

separate organ, or only as a rudiment (in Cicadas) , so that in

this circumstance also a parallelism between Pulicidas and

Bugs may be found.

The labium of the Khynchota consists of four consecutive

cylindrical joints furnished with a deep longitudinal groove

along the upper surface. It has been said that it is destitute of

palpi, but I think that this mode of expression is not correct. A

labium divided into four or five successive rings is in complete

contradiction to the plan of the organ deduced from the con-

sideration of the masticating mouth. But notwithstanding

Geise’s assertion to the contrary {1. c. p. 11), there is nothing

to prevent our regarding the cylindrical and often much more

voluminous basal part of the labium as the submentum

and mentum, as a direct continuation of which arise the

multiarticulate palpi fused together in the median line.

That there is really an amalgamation in the terminal

joint of the labium is rendered probable by the circum-

stance that both in the Hemiptera and in Cicadse a pretty

* Geise asserts sometliing of the kind when he represents the labrum

in Corixa and Sigara as taking- part with the constituents of the sucking-

tube (/. c. p. 53, fig. 29) ; unfortunately I must reject this assertion —

welcome as it would be to me for the homology attempted above — as

positively erroneous.

51

Dr. Karl Krapelin on the Pulicida3.

considerable notch appears at the apex*, although the side

lobes thus produced are not jointed off from the unpaired piece

in the same way as is usually the case, with the labella of the

Diptera for example. But if this conception of the structure

of the labium of the Rhynchota be correct, a comparison of it

with that of the Pulicidae presents no difficulties. A fusion

of the longitudinal fissure of the labium of Sarcopsylla (fig. 8),

for example^ nearly to the apex, would essentially realize for

us the conditions existing in Rhynchota (compare the labium

of Cicada in fig. 5). And with this apparent equivalence of

the parts an approximately similar physiological application

of them would be associated.

It has already been pointed out that the labium of the

Pulicidse has undertaken the guidance of the sucking-tube

only in its distal and not in its proximal part. But exactly

the same thing may be asserted of the labium of the Rhyn-

chota, which in the basal section of the rostrum shows an

effacement of the dorsal furrow and decidedly turns down-

wards, and thus devolves the guidance of the sucking-canal

and of the piercing setae entirely upon the labrum. In the

latter circumstance, indeed, there is an essential difference

between the proboscis of the Fleas and that of the Rhyn-

chota, as in the former the labrum, which has become one

of the constituents of the sucking-tube, cannot possibly be

employed to envelop the whole apparatus. But precisely

this different application of the labrum renders intelligible

a further fundamental difference between the two types of

proboscis, which must be found in the physiological appli-

cation of the maxilla. In the sucking-tube of the Rhyn-

chota, which, under the double guidance of the labium and

labrum, is sufficiently enveloped and protected throughout its

whole length, the maxillae might, without damage, be brought

in to complete the true piercing-apparatus ; they have become

long thin structures, destitute of palpi, flanking the sucking-

tube. In the Pulicidse, on the contrary, in which the basal

section of the sucking-tube, in consequence of the peculiar

employment of the labrum, was destitute of an envelope,

the maxillae, developed into broad plates (figs. 4 and 7 and

15, w), had this important function of protection transferred

to them. That under such a change of function the palpi

* The section across the tip of the rostrum of Notonecta (fig*. 14) shows

the labium as consisting of two perfectly separate parts. Gteise’s state-

ment that in Corixa the thu’d and fourth joints of the labium are com-

pletely cleft, depends, according to my investigations, upon an erroneous

interpretation of the conditions coming into view at the tip of the

rostrum.

4*

52

Dr. Karl Krapelin on the Pulicidse.

also came to full development and importance, can hardly

be regarded as a serious obstacle to the homology here

attempted.

The preceding indications will suffice to prove that in fact,

without any great violence to the data given, a certain parallel

may be drawn between the buccal organs of the Fleas and

those of the higher Khynchota, and that this comparison is at

least far easier to carry out than that between the Pulicidse

and the Diptera. If we bring the other agreements and diffe-

rences of the three groups in question into the account, the

result must be a phylogenetic alliance, although a distant one,

of the Fleas with the Rhynchota rather than with the Diptera.

But I repeat that the demonstrated relations certainly by no

means justify a union of the two groups. The only possi-

bility that presents itself is therefore to place the Pulicidse as

an equivalent order Siphonaptera * side by side with the two

most nearly allied orders.

The entire series of insects with suctorial mouth-organs

would consequently have to be divided in the first place into

two groups, one of which (Hymenoptera, Lepidoptera) is

characterized by having the lower parts of the mouth, maxillae,

and labium employed in the formation of a sucking-apparatus,

while in the other, on the contrary, it is almost exclusively

the upper parts (labrum and mandibles) that are implicated

in the formation of the true food-canal. This latter group

would include the three orders Diptera, Siphonaptera, and

Rhynchota, which 1 may, in conclusion, briefly characterize

as follows : —

1. Diptera. Insects with perfect metamorphosis. Head

free, with facetted eyes. Sucking-tube formed by a dorsal

and a ventral half-channel (labrum and hypopharynx), more

or less enclosed throughout its length by the labium, which is

bent up like a sheath and furnished with uniarticulate apical

palpi. Mandibles deficient or styletiform, pushing in between

the labrum and hypopharynx. Maxillae, when present, with

palpi. Salivary efferent duct an unpaired closed canal in the

interior of the hypopharynx. A sucking-stomach.” Tho-

racic segments amalgamated, usually with a pair of wings and

a pair of halteres.

2. Siphonaptera. Insects with perfect metamorphosis.

Head attached to the thorax by a wide surface, without

facetted eyes. Buccal organs suctorial. Sucking-tube formed

* As the name “ Aphaniptera ” is inadmissible for reasons already

given, and that adopted by Taschenberg, Suctoria,” has already been

employed twice, for a group of Cirripedes and for the Acinetae, I think

it best to fall back upon Latreilie’s name “ Siphonaptera.”

/Uin.d- Nai. Ili.sl S.S). Vol I'i- rUH

K-KpiPpehn del

Mintern Brea .Sc.* imp

LiBBARY

OF THE

(JNIVEeSITY OF ILLINOIS

Dr. Karl Krapelin on the Pulicidae. 5S

by a dorsal and two lateral channels (labrum and mandibles),

its anterior section only more or less enclosed laterally by the

multiarticulate terminal palpi of the labium, and at the base,

besides the latter, by the lamelliform palpigerous maxilla3.

Salivary efferent ducts paired, developed as a channel along

the inner surface of the mandibles. No ^^sucking-stomach.”

Thoracic segments free, without wings and halteres, with

pleural processes upon the last two segments.

3. Khynchota. Insects usually with imperfect metamor-

phosis. Head free or broadly united to the thorax, with or

without facetted eyes. Buccal organs usually suctorial.

Sucking-tube (in the higher groups) composed of( two lateral

half-channels (the mandibles), only in the anterior portion

enclosed by the labium and its apical multiarticulate palpi,

which are united nearly to the apex ; at the base by the

labrum. Maxillte styliform, without palpi, applied laterally

to the mandibles in the channel of the labium or the labrum.

Salivary efferent duct unpaired, formed by two half-channels

of the mandibles closing together from the sides. No

“ sucking-stomach.” Thoracic segments free or amalga-

mated. Four, two, or no wings ; no halteres.

EXPLANATION OF PLATE III.

The letters in all the figures refer to the same parts: — Ir, labrum;

m d, mandibles ; m, maxiUse ; m t, maxillary palpi ; I, labium ; I labial

palpi ; h, hypopharynx ; w, food-canal ; s, salivary duct.

Tig. 1. Transverse section through the proboscis of Tahanus, sp., anterior

third.

Fig. 2. Transverse section through the proboscis of Melophagus ovinus,

middle.

Fig. 3. Transverse section through the proboscis of Culex pipiens

middle.

Fig. 4. Maxilla of Sarcopsylla penetrans^ side view.

Fig. 5. Labium of Cicada, sp., side view.

Fig. 6. Lower part of the maxilla of Cicada sp., and its union with a

lamelliform appendage {pi) of the fore part of the head. /,

whip -like process of the plate ; pm, protrusor; rw, retractor of

the maxilla ; sp and s r, the sinews belonging to them. At x the

sinew of the protrusor articulates with a chitinous rod which is

perpendicular to the surface of the plate, and therefore does not

appear distinctly in the figure.

Fig. 'll. Maxilla of Pulex irritans.

Fig. 8. Labium of Sarcopsylla penetrans from above.

Fig. 9. Labium of Pulex irritans, side view, ch, basal chitinous piece.

Fig. 10. Transverse section through the proboscis of Pulex irritans, an-

terior third.

Fig. 11. Transverse section through the rostrum of Notonecta glauca,

basal third.

54

Dr. O. Zacharias on the Development

Fif/. 12. Front view of the head of Cicada sp. pi, plates with which

the maxillae articulate.

Fif/. 1 3. Transverse section through the proboscis of Sarcopsylla penetrans,

middle.

Fif/. 14. Transverse section through the rostrum of Notonecta glauca,

apex.

Fif/. 1/5. Transverse section through the proboscis of Pulex irritans, base.

VT. — New Investigations on the Development of the Viviparous

Aphides. By Dr. Otto Zacharias

Since the a])pearance of Metschnikoff’s ‘ Embiyologische

Studien an Insecten ’ (1866) the development of the embryo

of the viviparous Aphides has not again been made the sub-

ject of a monographic investigation. What the Russian

author established with regard to the mode of development of

the pseudo va” of Aphis Rosce and A. Pelargonii passes pretty

generally for all that is observable at present. Metschnikoff’s

description of the development of Aphides (at least in its

fundamental features) is regarded as a “ rocher de bronze,”

which presents no point of attack for an incisive criticism.

This, however, is not the case, and I will, in a memoir that

will appear very shortly, furnish the proof that Metschnikoff’s

description of the first developmental stages (as far as the

formation of the S-shaped germinal streak, and even some-

what later) by no means agrees with the facts. For the

subsequent stages I have also obtained quite different results

of investigation, which I shall venture to summarize at the

conclusion of this preliminary note.

The observation of the embryonic development of the vivi-

parous Aphides is for many reasons a difficult matter.

Besides the minuteness and delicacy of the objects with which

we have to do, there is a third condition which causes many

obstacles to the investigation, namely the clearness and strong-

refractive power of the protoplasmic contents of the egg. If

in the case of the eggs of many other insects we have to

contend with the obscurity of their yelk, it is in the Aphides

the crystal clearness of the latter which frequently acts very

prejudicially : prejudicially, inasmuch as under the circum-

stances indicated the upper half of the egg constantly acts

upon the lower half (or vice versa) j like a lens with a very

short focus, and not only enlarges but also distorts those

* Translated from the ‘ Zoologischer Anzeiger/ no. 168, May 26, 1884,

pp. 292-296,

55

of the Viviparous Aphides,

parts of the embryo which lie in a plane passing through the

middle of the egg parallel to the object-slide. This opens up

a rich source of illusions for those who come uncritically to

the investigation ; but for those who are aware of the optical

behaviour of the pseudo vitellus there arises the unconditional

necessity of correcting every surface-picture observed by the

side view corresponding to it, and, if possible, by the other

surface-picture (that directly opposite to the first). In the

former case the embryo must be turned 90°, in the latter 180°.

Without this method of rolling, already mentioned by Harting

in his well-known work on the microscope, it is not possible

to make out the earliest development of the Aphis-QvoJoryo.

Of course it is not easy to practise the method referred to,

and many a fine preparation is sacrificed by clumsy handling

of the wire which is employed in producing the rotation. In

my memoir I will describe in detail the rolling method as it

should be constantly employed in the more delicate investi-

gations in insect embryology.

I will now briefly indicate in what principal points the

results of my investigations differ from those obtained by the

distinguished Eussian naturalist.

The pseudovum possesses no chorion, but only a vitelline

membrane, Huxley’s pseudo- vitelline membrane.” This

encloses the whole contents of the egg, which, at a certain

early period (as Leuckart first remarked *), shows a distinction

between peripheral and central cells. We read also in

Huxley as follows : — They [the pseudo va] exhibit a central

darkish matter surrounded by a clear cortex.” Upon this point

the simplest observation gives clear information. The deve-

lopment of the embryo now starts from the clear cortex,” the

blastoderm^ which forms a multilamellar vesicle, and, indeed,

in this way, that at its lower pole {i, e. that turned towards

the vagina) a thickening is formed, from which the germinal

streak grows forth laterally (and near to the inner wall of the

blastoderm) in the form of a small thick tongue. The yelk

at this time contracts strongly, and places itself, as a rounded

mass, also at the inferior pole of the germinal vesicle. This

is the profile view, so to speak. If we now roll the pseud-

ovum through 90° we obtain a view en face ; and Metschni-

koff appears to have this alone in his eye when he speaks of

a germinal and a vitelline hill,” the appearance of which

characterizes the earliest embryonic stage of the viviparous

Aphides. In the surface-view our glance of course falls first

upon the broad side of the tongue, which now looks like a

* ‘ Zur Kenntniss des Geuerationswechsels und der Partheuogenese bei

den Insekten ’ (1858), p. 20.

56 Dr. O. Zacharias on the Development

and behind it rises the contracted yelk, and also

appears like a hill. This therefore explains how MetschnikofF

came to the notion of a germinal and vitelline hill. But such

a notion is not justified by the facts, and still less that of a

special genital hill from which the reproductive organs are to

originate. It is moreover quite incomprehensible that so

practised an observer as Metschnikoff even then was could

overlook the fact that the tongue of the germinal streak

growing freely into the cavity of the blastoderm immediately

shows a deep groove in its median line, becomes rounded off

on both sides throughout its whole length, and thus produces

two distinctly marked germinal pads. Metschnikoff, on the

contrary, repeatedly remarks that the embryo in its early

stage shows no trace of such pads

This negative judgment I can only explain by the fact that

Metschnikoff apparently does not practise the method of

rolling,” and therefore did not get to see the different aspects

of the embryo. In this opinion I am only strengthened by

the examination of his figures 16, 17, 18, 19, and 20 (pis.

xxviii. and xxix.), as also by the reading of the text relating

to them (pp. 444-448). Every one who comes fresh and

unprejudiced to the investigation of the development of the

Aphides will make the surprising observation that the deve-

lopmental processes which finally lead to the formation of the

S-shaped germinal streak are not performed in the plane in

which Metschnikoff places them in his figures, but in one

standing directly perpendicular to it. In my memoir I shall

produce the exact proof of this, and also furnish the requisite

figures which I have found to be verified in hundreds of

preparations.

As regards the S-shaped germinal streak, which is well

known to all investigators of Aphides, the inferior curve of

this letter (which is turned to the left) represents the cephalic

hood^ the same structure, it may be said in passing, which

Huxley, entirely mistaking the relative position of the Aphis-

germ, characterized as the abdominal hood.” The upper curve

(turned to the right) represents the rudiment of the abdomen,

and the intermediate part contains the material for the forma-

tion of the head and thorax.

"iTe limhs take their origin from a special superficial layer,

the so-called limh-plate (“ Extremitatenplatte ”), as to the

origin of which Metschnikoff has not got at the truth eitlier

in Simulia and Coriaoa or in Aphis Roses (see his paper, loc.

cit. pp. 400, 427, and 448). For the Aphides I have

^ Zeitschr, f. wiss. Zool. Bel. xvi. (18G6), pp. 448 and 450,

57

of the Viviparous Aphides.

succeeded in proving that the so-called “ Extremitatenschicht ”

is a part of the blastoderm which at a very early period enters

into an intimate fusion with the true germinal streak. The

details of this are treated of in my memoir.

Besides the two vertex-plates (Huxley’s procephalic

lobes ”) which originate from the primitive lateral plates, we

have also to distinguish in Aphis-embryos a median plate^

which is produced from the ventral part of the cephalic hood.

I would name it the mandibular plate., as the two mandibles

are formed from it.

The first and second maxillce originate from that part of

the limb-plate which overlies the procephalic portion of the

germinal streak, or is amalgamated with the latter. The

arrangement of the three pairs of buccal organs is of such a

kind that a hexagon is formed, the outer angles of which are

constituted by the first maxillae.

Later on the rudiments of the mandibles and first maxilla?

are enclosed in the depth of the head, and from them originate

the retort- shaped bodies"''' (of Metschnikoff), which secrete

the chitinous stylets of the rostrum. In the mature embryo

we perceive two such bodies on each side, not one only, as

Metschnikoff’s figures show. By the demonstration that it

really is by the transformation of the mandibles and maxillce

that the retorU shaped bodies are produced, the parts of the

mouth of the Aphides are first brought into homology with the

corresponding organs in other insects.

According to Metschnikoff, as is well known, the

mandibles and first maxillae are completely retrograded, and

the ‘‘"bodies” secreting the piercing seta? originate quite

newly. This was a priori very improbable, and observation

gives quite another result. By carefully crushing half-

mature embryos the actual conditions may often be very

beautifully brought into view.

Witlaczil, who has treated of the anatomy of the Aphides

in much detail in a recent memoir gives a very full descrip-

tion of the structure of the retort-shaped bodies as studied by

him by means of sagittal and transverse sections through fully-

developed animals.

For orientation in many difficult points of the development

of Aphides, e. g. as to the question whether or not the Mal-

pighian vessels are present in these animals at any time, I

have turned to the allied group of the Coccidae, and not

without success. Thus I distinctly saw in Coccus hesperidum

that the brown masses of substance corresponding to the

secondary vitellus in Aphis Rosce arrange themselves very

* ‘ Zur Anatomie der Apliiden ’ (Wien, 1882).

58

Dr. W. Dybowski on the

early in the form of two long cords, which open close together

into the intestine in the region of the rectal section. Accord-

ing to this observation T do not hesitate to adopt Witlaczil’s

view, according to which the green cell-mass in the abdomen

of the viviparous Aphides (which is likewise arranged into

two cords) represents the Malpighian vessels of other insects.

In a very pale Aphis-embryo I was able clearly to detect,

besides the very distinctly marked dorsal vessel, the point of

convergence of the two cords, but without distinctly seeing the

point of discharge (as in Coccus hesperidum) .

The egg of the viviparous Aphides, for which I have here

and there, for convenience’ sake, employed the antiquated

term pseudo vum,” therefore presents exceedingly interesting

and very distinctly observable developmental processes, which

cannot be sufficiently studied. In recent times Dr. Arnold

Brass* * (of Leipzig) and Dr. Ludwig Willf (of Bostock)

have occupied themselves with the earliest stages of develop-

ment. The last-mentioned gentleman has also already an-

nounced the publication of a work upon the later stages of the

Aphis-embryo. In course of time many other workers will

certainly have to be registered for this highly interesting

subject.

VII. — Notes on the South-Russian Spongillidse.

By Dr. W. Dybowski |.

Dr. P. T. Stepanow, Professor of Zoology in the University

of Charkow, has had the kindness to send me some specimens

of the freshwater sponges for scientific investigation. The

sponges preserved in the Museum of the above University

and those collected by Prof. Stepanow himself are from the

following localities : —

1. From the river Udy (a right affluent of the Siewiernyj

Daniec), Gov. Charkow.

2. From Lake Lebiedin (Circle Lebiedin), Gov. Charkow.

3. From the river Kolomak (left affluent of the Worska, a

branch of the Dnieper), Gov. Poltawa.

* Das Ovarium und die ersten Eiitwicklimgsstadien des Eies der vivi-

paren Aphiden,” in Zeitschr. f. Naturwiss. Bd. iv. (1882).

+ Zur Bildung des Eies und des Blastoderms bei den viviparen

Aphiden.” in Arbeiten des Zool. Instit. zu Wiirzburg, 1883, Heft 3.

I Translated by W. S. Dallas, F.L.S., from the ‘ Sitzungsbericlite der

Naturforsclier-Gesellschaft bei der Universitat Dorpat,’ Bd. vi. pp. 507-

515 (1884).

59

South-Rusfiian Spongillidae.

4. From the river Siewiernyj Daniec, Gov. Charkow.

5. From Lake Wielikoje (Circle Lebiedin), Gov. Charkow.

Offering my best thanks to the sender of these sponges,

I now communicate the results of my investigations upon

them.

These results are as follows. The Spongillas of these five

localities represent three species, namely : —

A. Spongilla lacustris.

B. Meyenia {Ephydatid) fluviatilis»

C. Dosilia (?) Stepanowii^ n. sp.

The first two of these * I have only briefly described ; but

the third, as a form hitherto unknown to science, has received

as accurate and detailed a description as possible.

Spongilla lacustris^ Carter, occurs in very thin lamellse,

coating in spots or cones the leaves of Acorus Calamus and

Quercus sp. ; they are evidently quite young and undeveloped

specimens, in which I have consequently found no gemmules.

The skeleton-spicules are 0*224-0*130 millim. long, 0*010-

0*002 millim. thick t* The size of the parenchyma-spicules

varies between 0*050 and 0*060 millim. in length, and 0*002

and 0*004 millim. in thickness. These spicules perfectly

agree with those described by me {loc. cit.).

Among the spicules such deformities frequently occur as I

have already described and figured {1. c.) ; but among the

most peculiar are the clavate and pin-shaped spicules, the

heads of which are sometimes quite smooth and rounded, but

sometimes variously misshaped, or furnished with small

spines J.

Localities. Daniec, Kolomak.

The Meyenia fluviatilis^ auct., agrees perfectly with that

described by me (Z. c. p. 13). I have before me four frag-

ments of a large, fully developed sponge and a couple of

smaller ones. In all numerous gemmules are present. The

fragments belong to cushion-like sponges ; the smaller sponges,

on the contrary, are growing round a stem, 8 millim. thick, of

Arundo sp.

Localities. Lake Lebiedin, river Udy.

* See Dybowski, “ Stiidien iiber die Siisswasser-Scliwamine des Russi-

schen Reiches/’ in Mem. Acad. Sci. St. Petersb. ser. 7, tome xxx. no. 10,

p. 6, tab. i. figs. 4, 6, 7, and p. 13, tab. i. fig. 3, and tab. ii. fig. 9.

t In my memoii’ {1. c.) p. 10, column 1, line 4 of the measurements,

0-009 is erroneously printed instead of 0-002, and in column 3, line 6, 0-028

instead of 0-002.

X The figures necessary for the more ready intelligence of all the de-

scriptions here given, I have prepared with the aid of a Hartnack’s prism,

and will publish them when opportunity serves.

60

Dr. W. Dybowski on the

Dosilia (?) Stepanowiij n. sp.

This Spongilla, from Lake Wielikoje, is of very peculiar

interest. So far as I know, no form similar to our sponge

has hitherto been described among the European Spongillidae,

at least, I have been unable to find any notice of it in the

literature accessible to me ; on the other hand, among the

exotic (American and Asiatic) Spongillidae, 1 find analogous,

and, it seems to me, nearly allied forms.

The genus Dosilia^ Gray* * * §, which Carter f places in his

“ Meyenia^'' possesses two species — Dosilia plumosa (from

Bombay) and D. Baileyi (from New York). The latter

appears to me the form most nearly allied to our sponge J.

To justify and support my opinion I will here give as

accurate a description as possible of our sponge, and then

place side by side the most prominent characters of the two

(Kussian and American) sponges, so as to facilitate for other

authors the comparison of the two sponges with one another.

Description. — Of the sponge under consideration I have

before me six small spirit-specimens, all of which are defec-

tive and do not enable us to form any definite notion of their

form. They are chiefly shapeless masses, growing round

various foreign bodies (such as leaves and stalks of grasses,

fragments of bast, very thin twigs, and even quills of a small

wing-feather). The specimens in spirit are pale tawny, and

look not unlike soaked bread. The skeleton-spicules are long

and slender acerates with acute ends, that is to say, they have

the form of the spicules proper to the SpongillsB with smooth

spicules in general ; but they are somewhat smaller, their

length being 0*200-0*104, and their thickness 0’065-0*004

millim. The surface of the skeleton-spicules is, however,

not smooth, but furnished with very short, acute, and exceed-

ingly scattered spines §.

* J. E. Gray, “ Notes on the Arrangement of Sponges, with the De-

scription of some new Genera,” in Proc. Zool. Soc. Lond. 18G7, pp. 550-

553, pis. xxvii. & xxviii.

t Carter, History and Classification of the known Species of Syon-

gillai' in Ann. & Mag. Nat. Hist. ser. 5, vol. vii. (1881), pp. 78-107,

pis. V. & vi.

t Neither of the two species mentioned is known to me by autopsy. I

have drawn my conclusions as to the affinities of our sponge only from

the statements in the literature, and must therefore for the present abstain

from a certain and final decision.

§ The above-mentioned spines are so small and inconspicuous that

they may very easily be overlooked. They are most conveniently observed

with the light of an oil-lamp and with the objective no. 8. When they

have once been observed they may quite easily be recognized with objec-

tive no. 4. The spines appear most distinctly at the periphery of the

spicules.

61

South-Russian Spongillidge.

The skeleton-spicules of the sponge under notice constitute, ^

as it were, a transitional form between the smooth and the

spiny Spongillid spicules*. The parenchyma-spicules are

also remarkably peculiar and characteristic. They are small

acerates, measuring 0*040-0*050 millim. in length and 0*025-

0*010 millim. in thickness, and furnished with spines ; in form

they generally resemble those of Spongilla lacustris^ but the

form and arrangement of the spines are quite different from

those in the spicules of the latter Spongilla, Here the spines

occur in three different forms : the middle portion of the

spicule is beset with long, obtuse, vertically projecting spines,

but the two ends present small, pointed booklets, while towards

the middle of the spicule there exist pointed erect spines.

The long straight spines of the middle section are frequently

also covered with small acute spines ; the free end of the

spine is sometimes rounded off, sometimes truncated, and

sometimes furnished with a knob or a T-shaped rod. The

gem mules I have not been able to find, but I found in the

parenchyma of the sponges numerous amphidisci.

The spindle-shaped amphidisci are very long ; their dimen-

sions are as follows : —

millim.

Total length 0’040-0’028

Thickness of the shaft 0*002-0‘004

Diameter of the disk 0'012-0-010

The shaft is furnished on its surface with large perpen-

dicular spines.

The two terminal disks (disci) are furnished at the mar-

gins with deep notches ; the teeth thus formed have a per-

pendicular position.

The actual form of the disk, as well as the number of teeth

on each disk, I have been unable to ascertain.

If we summarize the most important characters of our

sponge and compare them with those of Dosilia Baileyi^ it

appears that the two sponges must belong to the same genus,

but are specifically different.

These characters are as follows : —

Dosilia Baileyi^ Carter (I, c, p. 95).

Coating, surface smooth. Structure friable, crumbling.

Skeleton-spicule curved, subfusiform, gradually sharp-pointed,

smooth. Flesh-spicule minute, curved, fusiform, gradually

sharp-pointed, covered with erect obtuse spines throughout,

extremely small towards the extremities, and extremely long

and perpendicular about the centre of the shaft. Statoblasts

* See Dybowski, 1. c. tab. i. figs. 3 & 5.

62

On the South-Russian Spongillidae.

globular ; aperture infundibular ; crust, which is thick and

composed of granular cell-substance, charged with birotulate

spicules consisting of a long, straight, sparsely spiniferous

shaft wdiose spines are large, irregular in length, conical and

perpendicular, terminated at each end by an umbonate disk of

equal size, deeply but regularly denticulated, w^hose processes

arc claw-like and turned inwards, arranged perpendicularly,

with one disk resting on the chitinous coat and the other form-

ing part of the surface of the statoblast.

“ Locality. New York. In a stream on the Canterbury

Road, West Point.”

Dosilia Stepanowii^ n. sp.

Surrounding ; surface smooth.

Skeleton-spicules long, pointed, and covered with small,

acute, but scantily distributed spines.

Parenchyma-spicules small, pointed, and covered with

spines. Spines in the middle section long, obtuse, and per-

pendicular ; at the two ends small, acute, hook-like ; towards

the middle small, acute, perpendicular. Amphidisci spindle-

shaped, their shaft long and furnished with a few large per-

pendicular spines.

The two terminal disks are toothed at the margins. The

teeth have a perpendicular position.

Locality. Lake Wielikoje.- •

In conclusion, it may be mentioned that I found in the

parenchyma of the Spongillidae just investigated by me some

siliceous corpuscles, which it seems to me may be small paren-

chyma, or coating spicules of still unknowm Spongillidae.

From this I conclude that other unknown Spongillida3 must

certainly occur in the waters of the Government of Charkow

and of South Russia generally.

Would it not be advisable on the part of the University of

Charkow to make a prize-problem of the investigation of the

South-Russian Spongillidse ? It would be a very grateful

theme for a pupil of that University, the solution of which

might advance the knowledge of a group of animals which is

still but little known not only in Russia but also in the rest of

Europe.

Further, I may call attention to the fact that very numerous

Diatoms and Algae are present in the parenchyma of the

Spongillid^, so that these plants, otherwise so difficult to

discover, are to be sought in the interior of these sponges.

Postscript. — The Spongilla from Lake Hertha (in the

island of Riigen), kindly communicated to me by Dr. Braun

of Dorpat, proves, from my investigation of it, to be a Spon-

Mr. E/. Eosenstock on Heterocerous Lepidoptera, 63

gilla lacustids, auct. It agrees perfeetly with the speeimens

of that species obtained from the Ludwinow estate (see

Siissw.-Schw. d. Euss. Eeiches, p. 6). This fact is of interest

as furnishing a small contribution to the zoogeography of the

Spongillse, especially as, so far as I know, no Spongillse were

previously known from that locality.

VIII. — On the Synonymy of some Heterocerous Lepidoptera,

By Eudolph’Eosenstock, B.A.

I INCIDENTALLY discovered and noted the following synonyms

while systematically studying the collection of Lepidoptera

in the British Museum. They are for the most part redescrip-

tions by the late Mr. Walker of speeies previously described

either by himself or other authors.

1. Noctuites.

Poaphila congesta, Walk. Vene- = Anthophila erecta, Walk. San

zuela. Domingo.

Remigia tiiangularis, Walk. N, = Toxocampa costimacula, Walk,

India. Sylhet.

2. Pyralites.

Hypena disclusalis, Walk, S. =

Africa.

Manmatha confisinalis, Walk, —

Loc. ?

Pyralis dispansalis, Walk. San =

Domingo.

Lepyrodes lepidalis, Walk. Cey-"'

Ion, N. India.

Stenia pipleisalis, Walk. Sierra t

Leone. r'""

Hymenia meridionalis, Walk. S.

India. J

Botys hortalis, Walk. Bogota, =

Santarem.

strictalis, Walk. N. Ame-^

rica.

oUiusalis, Walk. U. S. I _

America. | “

ofellusalis, Walk. Loc.

philealis, Walk. Venezuela. =

senippialis, Walk. Bogota. I _

codrusalis, Walk. Bogota. |

Hypena senialis, Gu4n. Central

Africa.

Anthophila semipurpurea, Walk,

Loc. ?

Carcha hersilialis, Walk. San

Domingo.

Samea {Gum.) sidealis. Walk.

Sierra Leone. (This is evi-

dently an Old-World species

of wide range.)

Botys marialis, Walk. San Do-

mingo.

flavidalis, Walk. N. Ame-

rica.

lycialis, Walk. San Do-

mingo.

dorisalis, Walk, Villa

Nova.

64 Mr. R. Rosenstock on Heterocerous Lepidoptera,

Botys semizebralis, Walk. S. In-

dia.

convectalis, Walk. S. In-

dia.

suspicalis, Walk. Ceylon.

memmialis, Walk. Loc.

ogmiusalis, Walk. San

Domingo.

cinctipedalis, Walk. Geor-

gia.

Ebulea heronalis, Walk. Hon-

duras.

Spilodes helvialis, Walk. U. S.

America.

Botys gnomalis, Walk. San

Domingo.

peleusalis, Walk. San Do-

mingo.

orontesalis, Walk, jpga,

Venezuela.

= Botys amyntusalis, Walk. Cey-

lon.

= neoclesalis, Walk. Cape.

= campalis, Walk. Jamaica,

San Domingo.

= gastralis, Guen. San Do-

mingo.

= oxydalis, Guen. U. S.

America.

= acastalis, Guen. Hondm'as.

= apertalis, Guen. N. Ame-

rica.

= Omiodes humeralis $ , Guen.

San Domingo.

= c?, Guen. San Do-

mingo.

= simialis, GuAn. Cayenne.

The following species placed by Walker under. pos-

sess the generic characters of Guenee’s genus Omiodes.^ which

appears to have a wide distribution : —

Botys ceresalis, Walk. San Do-

mingo.

jasonalis, Walk. San Do-

mingo.

helicitalis, Walk. San Do-

mingo.

pbiletalis, Walk. Santarem.

Botys orpbnealis, Walk. Lee.

bianoralis, Walk. Japan.

pharaxalis, Walk. More-

ton Bay, Australia.

3. Geometeites.

Tepbrina confiniaria, San = Psamatodes nicetaria, G'wew. San

Domingo. Domingo.

(Walker intimates the possible identity of these two species.

Cat. xxiii. n. 971.1

Sterrha participata, Walk. Na-

maqua Land.

Aspilates proxantharia, Walk.

S. Africa.

? biferaria, Walk. S.

Africa.

Mergana bilineata, Moore. Dar-

jiling.

= Sterrha plectaria, Guen. (Phal.

pi. viii. fig. 7). S. Africa.

= Aspilates occupata, Walk. S.

Africa.

= justaria, Walk. Namaqua

[ Land.

= Sarcinodes carnearia, Guen. In-

dia.

The genera Mergana and Auxima of Walker are synony-

jnous with Sarcinodesj Gudnde’s single genus of Asiatic OEno-

chromidae. Auxima and Sarcinodes are absolutely identical,

and Mergana differs according to Walker in having two

instead of four spurs to its hind tibiss. The number of spurs,

Miscellaneous.

65

liowcvcr, is probably variable even within the same species ;

nor can it be a sexual character, as out of two male speci-

mens of Menjana equilinearia in the collection, one has two,

the other four tibial spurs.

I submitted all the synonyms enumerated above to the

consideration of Mr. Butler, who kindly endorsed their

correctness.

MISCELLANEOUS.

The System of the Monactinellidoe. By Dr. B. von Lendenfeld.

The rich collections of Australian sponges in the museums at

Adelaide, Christchurch, and Dunedin, which were placed at my

disposal by Dr. Haacke, Dr. J. von Haast, and Prof. Parker, as well

as the material collected by myself among the Australian shore-

sponges, inelude about 500 species, of which I have only been able

to identify a few with forms already described. I have easily

recognized among my specimens a number of the species accurately

described by Selenka and Marshall, but have had little success in

the identification of the species from the Australian region described

by English and American authors.

As was very justly foreseen by 0. Schmidt, it is not practicable to

regard the system of the sponges established upon the Mediterranean

fauna, and enlarged through the Atlantic forms, as universally

applicable ; uniting intermediate forms make their appearance

where, from known facts, one would have suspected no relationship.

However, the new forms furnish further proofs of the correctness

of Zittel’s system, and I have taken this as the foundation of my

investigations.

The Calcispongiee are few and insignifieant. Hexactinellidae and,

singularly enough, Tetractinellidae also are almost entirely defi-

cient. Of the latter group I have obtained two specifically different

individuals. As Myxospongiae are also extremely rare (three

species), the whole mass of the Sponges is distributed in the two

groups of the Monactinellidac and Ceraospongise.

I have carefully examined the Monactinellidae especially, and will,

in what follows, bring together the most important systematic

results of this work.

Although I worked upon sponges at home for a long time under

F. E. Schulze’s guidance, and have also paid much attention to them

in Australia, the investigation of so great a number of forms as has

lately been at my disposal has compelled me to arrive at a clear

idea of what is to be understood as a species among sponges. In

the siliceous sponges it is hero, as elsewhere, merely the form of

the spicules, and never their arrangement, that behaves conserva-

Ann. (U Mag. N. Hist. Ser. o. Vol. xiv. 5

GG

Miscellaneous .

lively, and consequently can be applied to the establishment of the

higher systematic groups. O. Schmidt’s principle of division there-

fore applies also to the Australian sponge-fauna, divergent as it

may be.

(). Schmidt has so far accepted a polyphyletic pedigree for the

Chalinidfo as to derive the sponges with axial siliceous spicules in a

network of horny fibres in part from the horny sponges and in part

from the llenieridae. I am in a position to describe a continuous

scries of forms which lead from the Ceraospongiae to the true Renie-

ridae without any horny matter. There are all steps between the

skeleton of the Chalcaenidae, consisting of spicules arranged in

bundles and combined into thick cords, and the tissue of the typical

llcnieridae, with its loose triangular meshes. Thus I possess a

whole series of sponges the skeleton of which consists of parallel

cords, representing the main fibres of the horny sponge. These

cords consist of a dense mass of siliceous spicules, and are united

with each other by single spicules of the same form, representing

the uniting fibres. They are consequently separated from each

other b}' the length of a spicule. In some of these sponges the

main fibres are feeble and often bent, so that the looser Renierid

tissue is formed by the weakening first of the uniting fibres and

then of the main fibres of the Chalcaenidae. The Australian Renie-

ridae, however, show no relationship with the Myxospongiae, from

which they must have been derived, if not from the Chaliuidae.

Our sponges show that the non-ceratose Monactinellidae represent

the terminal members of a series which starts from the Cerao-

spongiae. The spicules of the Monactinellidae are either uni- or

biradiates. (In this the biacuates ac^ and ae ac, Vosmaer, are re-

garded as biradiates, and the obtuse-ended, tubercular, and pin-like

forms tr ac, tr° ac, tr ac sp, Yosmaer, as uniradiates.) In agree-

ment with r. E. Schulze, I regard the pluriradiate form as phylo-

genetically older.

I regard the spicules formed in the parenchyma of the sponge,

the flesh-spicules, as in every respect essentially different from those

which were deposited in the axes of the horny fibres, originally

as biradiates, in the ancestors of the Chalinidae. By reduction of

the number of rays the obtuse-ended spicules have thus been deve-

loped from the biradiates, and from these again the pin-like and

tubercular forms. In many cases a gradual disappearance of the

horny substance takes place pari passu with the transformation of

the spicules, while in other cases again the spongioline disappears

without the spicules changing their form.

If we now consider the Mouactinellidm from this point (for the

present without reference to the fiesh-spiculcs), we obtain the

following classification, in which they form as a whole an order

Monactinellidae ” in the class of Sponges : —

Misctlla neo ua .

f)7

SuherUidce.

Uniradiatos without perceptible horny substance.

Chlathridce.

Network of horny fibres with imbedded obtuse spicules.

Ecldspidee.

llenieridiB.

Biradiates without percep-

tible horny substance.

Network of horny fibres

with imbedded uni- or

biradiates and projecting

tubercular spicules.

Chalccenidce,

Network of horny fibres with dense masses of coarse

imbedded biradiates.

ChalarchidcB.

Network of horny fibres with scanty, axial, extremely

delicate biradiates.

Ceraospongiae.

The Chalarchidae and Chalcaenidae, at least the Australian forms,

are so different from each other, that for the sake of uniformity I

have preferred to divide the Chalinidae of authors into these two

main sections. 0. Schmidt’s numerous families of Chaliuidae would

in this scheme appear as subfamilies.

Among the Chalarchidae we find sponges resembling Euspongia,

Cacosponyia^ and Spongelia. Possibly therefore the pedigree of the

Chalarchidae may be polyphyletic, different members of the family

being descended from the above-mentioned horny sponges.

I will here mention a peculiar intermediate form which combines

characters of the Chalarchidae with those of Spongelia. The skele-

ton of this sponge consists of a ladder-like network of horny fibres.

The radial main fibres are entirely filled with sand, and bear no

spicules. The tangential uniting fibres, which are not much

weaker and do not anastomose with each other, are entirely filled with

axial, evidently self-formed biradiates, and bear no foreign bodies.

The Echispidae for the most part coincide with Gray’s Echino-

nemata.

C8

Miscellaneous.

I have found an interesting form which so far unites this family

with the ChlathridoD, that we find in this sponge projecting, obtuse,

perfectly smooth spicules in great quantity, but no tubercular

spicules.

Among the other three families which include 0. Schmidt’s

groups of the same names, we should then have the Desmacidonidae*

to distribute.

I have proposed the foregoing classification without reference to

the flosh-spiculcs, the multifarious anchors, hooks, &c. of the Des-

macidonidse, and without taking into account the siliceous stars in

the cortex of many Monactinellidae.

I regard it as preferable not to adduce the flesh -spicules in

the formation of the principal groups, but rather to employ them as

generic characters. Although I have long been inclined to this

view, I have hitherto refrained from bringing it forward, as autho-

rities like Yosmaer and 0. Schmidt lay great stress upon the sepa-

ration CvSpecially of the anchor- and hook-bearing sponges and their

collocation to form a whole. In the same way, however, that 0.

Schmidt united sponges with and without siliceous stars in the

family Gumminese, I think we may bring together Monactinellid

families with and without anchors &c., provided they agree in the

structure of the fibrous skeleton.

I believe that these flesh-spicules are of coenogenetic origin. 0.

Schmidt has demonstrated that in part they contain much more

organic material than those siliceous structures which occur in the

corals. Their coenogenetic character is especially proved by the

extraordinary multiplicity and variability of their forms. In sponges

which show themselves to be nearly allied by the character of their

fibrous skeleton we often find anchors in one and nothing siliceous

in the parenchyma of another ; while, on the other hand, hooks

occur in very different sponges. We usually meet with them in

sponges the fibres of which contain Monactinellid spicules. But I

possess a well-preserved spirit-specimen of a sponge from Port

Phillip, which is a true Hircinia, and the fibres of which contain no

siliceous spicules, while in its parenchyma, besides the filaments,

great quantities of S-shaped double hooks are to be found.

The same applies to siliceous stars. These may sometimes occur

and sometimes bo deficient in nearly allied forms, while, on the

other hand, they are found in very different sponges.

If we represent the system of the sponges in the form of a genea-

logical tree, taking the above conceptions as the foundation, wo

arrive at the following conclusions : —

1. From the Myxospongia3 originates a series of forms, the cen-

tral members of which resemble the Spongidse. From the sides of

this series branches are given off, at the extremities of which stand

the Aplysinae and Hirciniae. The Chalarchidae and Chalcicnida) are

placed in the upper part of the scries, from the end of which tlie

llenieridse, Suberitidae, and Echispidae radiate like an umbel. The

* 0 course I can here refer only to the most important groups.

Miscellaneous, 69

Guramincac have branched off between the Myxospongiac and the

Spongidac.

2. In all the forms of this series, from llalisarca to Suherites or

lieniera, wo meet with the tendeney to form liesh-spicules.

3. The flesh-spicnles are quite independent of the rest of the

skeleton, and occur in two types, Monactinellan (anchors &c.) and

Polyactinellan (stars &c.).

4. When another skeleton was already formed by production of

fibres, when the flesh-spicules originate they remain small and

unimportant, and in this case it is of no consequence whether the

fibrous skeleton consists of horny substance (Jlircinia), connective

cords (Gummineae), or siliceous cords (Desmacidonidae).

5. When there was no fibrous skeleton when the flesh-spicules

were formed they attained considerable dimensions, and on their

own part formed connected frameworks. Both the Monactinellan

and the Polyactinellan forms occur in these sponges. The anchor-

spicules of the Tetractinellidae perhaps belong in part to the former,

and the structures in Tetractinellidae and Hexactinellidae originating

by reduction of the many rays to 4 or 6 to the latter group. The

Plakinidae unite all these with Halisarca.

From the series of fibrous sponges which culminates in the non-

horny Monactinellidao branches are given off at many points in the

same direction, all parallel to that powerful but homologous branch

which contains the Hexactinellidae and Tetractinellidae. — Zoologischer

Anzeiyer, No. 164, April 7, 1884, vii. p. 201.

On Orbulina universa. By M. C. SenLUMBEEGEK.

Several naturalists have already paid attention to the genetic

relations which appear to exist between the OrhuUnce and the Globi-

gerince, which are so abundantly distributed in our seas. Pour-

tales^ was the first to indicate the presence of a Glohigerina in the

interior of OrhuUncG dredged in the Gulf-stream. Dr. A. Krohn t

made the same observation upon living Orhulince taken at Madeira.

These two observers J concluded that the Orbulina gives origin to a

Glohigerina, which, increasing in size, finally bursts the sphere

which encloses it and escapes to lead an independent existence.

Carpenter §, in his classical work on the Foraminifera, opposes this

opinion by a series of irrefutable arguments and retains the two

genera Orbulina and Globigerina founded by D’Orbigny.

Recent researches upon the embryogeny of the Foraminifera have

* Silliman’s Journal, July 1858; reprinted in this Journal, ser. 3, vol. ii.

p. 235.

t Referred to in a paper by Prof. Max Schultze, in the Arch. f. Naturg.

I860, p. 287 ; translated in this Journal, ser. 3, vol. vii. The point is

discussed at pp. 311-313.

t Krohn simply observed the fact and communicated it to Max

Schultze.

§ Introduction to the Study of Foraminifera, 1862.

70

Miscellaneous,

led me to interpret the facts observed in a manner quite different

from that which had been regarded as admissible. In examining

the sands from a deep dredging (4255 metres) executed by the

‘ Talisman ’ at the Canaries, I found a great number of Orhulince of

all sizes, and so clean that when soaked in chloroform and immersed

in Canada balsam they became perfectly transparent. It is then

observed that among the smallest of only 320 in diameter, and

those of medium size, some are empty, while others have their cavity

occupied entirely or in part by a succession of globular chambers

arranged in a trochiform spire, like those of certain Glohiyerince.

The large Orhulince^ attaining nearly I millim. in diameter, are

almost always empty.

These interior chambers are more easily distinguished after the

removal of a portion of the Orhulina^ or when entirely separated

from their envelope. We then find that their exceedingly delicate

plasmostracum is pierced by distant perforations ; the chambers of

the first two turns of the spire are smooth ; the following chambers

bear fine scattered spines, which, upon the last ones, are prolonged

to the inner wall of the Orhulina^ to which they attach themselves

These chambers communicate with each other and with the interior

of the OrhuUna by a small semilunar aperture, situated below and

opposite to fhe turn of the spire. In the largest Orhulince the spire

does not include at the outside more than sixteen chambers.

Now all Glohiyerince, as Carpenter points out, even when young,

have a comparatively thick plasmostracum, very closely placed per-

forations, one or several apertures widely invading the chambers,

and a rugose exterior, in consequence of the great number of spines

which cover its surface. Hence between the interior chambers of

the Orhulince and the Glohiyerince there is only a resemblance of

form.

On the other hand, we find many small Orhulince in which the

last or last two interior chambers form projections upon the sphere;

but then these protuberances are surrounded by a plasmostracum

as thick as the rest of the envelope. The interior chambers there-

fore do not quit the OrhuUna ; further, if they did so we ought

never to meet with large empty Orhulince.

From these facts we can draw only one logical conclusion, namely,

that we have before us a case of dimorphism analogous to those

which M. Munier-Chalmas and myself have already indicated in the

Nummulitest, the Miliolidae t, and many other genera of perforate

and imperforate Foraminifera§.

The single chamber of the OrhuUna is the homologue of the

initial chamber of the other Foraminifera ; when it remains empty

* This fact was observed by Pourtales.

t Bull. Soc. Gdol. Fr. ser. 3, viii. p. 300; ‘Annals,’ ser. 5, vol. xi.

p. 336.

X Oomptes Rendus, 1883, pp. 862 and 1508 ; ‘ Annals,’ ser. 5, vol. xii.

p. 67.

§ Feuilledes jeunesNaturalistes, 14*’ annee; Congresde Rouen, p. 520

(1883).

Miscellaneous.

71

it is of the form A ; with tho series of interior cli.ambers it is of the

form Ih But it is iiecess<ary to remark that, as among the OrhidincK

wo meet, on tho one hand, with large empty individuals, and, on tho

other, with small individuals, some empty, some with interior

chambers, wo cannot assume in this case, as has been indicated for

Miliolida), an absorption of the large embryonal chamber.

Tho case of the Orhidinfe is in favour of our first hypothesis, and

seems to demonstrate that the dimorphism of tho Foraminifera is an

initial character, tho result of two original forms. — Comptes EenduSy

April 21, 1884, p. 1002.

On the Ascidian Oenus Bhopalea. By M. L. Boule.

Philippi first described (Muller's Archiv, 1843), under the name

of lihopalea neapolitana, an Ascidian that he had collected in the

Bay of Naples, and of which he has given a short anatomical de-

scription, but sufficiently accurate as to most of the details of organi-

zation that he has noticed. In his memoir ho approximated this

new genus Rhopalea to the Clavellince. Since Philippi’s time, so

far as I know, no other naturalist has studied this curious form of

Ascidian ; Traustedt does not notice it in his work upon the simple

Ascidiaof the Bay of Naples (Mittheil. aus der zool. Stat, zu Neapel,

1883), and Herdman (Tunicata collected by the ‘ Challenger ’), re-

ferring to the description given by Philippi, places it alongside of

Ecteinascidia, in the family Clavelinidae.

The Rhopalece are very abundant on the shores of Marseilles on

the bottom surrounding the Zosterce, in the muddy sands collected

by the currents at depths of from 25 to 60 metres ; hence I have

been enabled to observe numerous individuals and to make a regular

investigation of them. The body, of an average length of 8 to 10

millim., of a nearly pure white colour, is divided into two parts

— one anterior, of triangular form, free, containing the branchia,

and bearing the two siphons (buccal siphon with eight or nine, and

cloacal siphon with six papillae) ; the other posterior, of irregular

form, adherent to the ground, and incrusted with debris of various

kinds, containing the mass of the viscera. These two parts are

joined together by a slender region of considerable length, through

which the rectum, filled with substances destined to be rejected,

may be distinguished ; the general aspect much resembles that of a

somewhat stout and very large Clavelina.

The thick tunic is of a soft consistency around the anterior part

of the body, firm and resistant around the posterior part. In this

latter region the fundamental substance of the tunic encloses nume-

rous vacuolar cells, while it contains none in the anterior region.

The dermis (mantle), which is thin, bears small muscular bundles,

most of which run in the direction of the length of the body ; around

the siphons there also exist a certain number of annular muscular

bundles. Philippi erroneously regarded as a peritoneum the portion

of the dermis which surrounds the posterior visceral mass. The

fundamental web of the branchia, the apertures of which are oval

72

Miscellaneous.

and regularly formed, is folded longitudinally ; the folds are very

small, invisible to the naked eye, and in this respect resemble those

of the branchia of the Fhallusice. The transverse branchial sinuses

are united to the longitudinal sinuses by an anastomotic branch,

which does not project into the interior of the branchial cavity

beyond the longitudinal sinuses, to produce papilhe like those pos-

sessed by most of the simple Ascidia. The dorsal raphe is consti-

tuted by a series of papillae ; it terminates in front, not far from the

vibratile organ, and in this region the pericoronal groove sends

towards it a small prolongation analogous to that which that groove

bears in the Cionm. The peribranchial cavity does not communi-

cate with the empty spaces left between the viscera in the posterior

region of the body, lacunae which may be regarded as forming by

their union a reduced general cavity; it is stopped immediately

behind the branchia by a peritoneal lamina, like that of Ciona.

The other organs are constituted nearly in the same way as in

the donee. The only important differences relate to the arrange-

ment of the sexual organs, which are collected into a single mass

surrounding the intestinal cavity, and to the greater length of the

oesophagus and rectum. With the exception of these not very

important distinctions, the digestive tube, the nervous system, the

hypoganglionic gland and its excretory duct, the heart, and the

principal sinuses &c. present the same fundamental structure as in

the donee.

The relations of the Rhopalece with the other forms of Ascidia are

multiple. By their general facies, it is true, they approach the

Clavelinidse ; but we cannot place them in that family, for they do

not reproduce by gemmation and possess a more complex organiza-

tion. We must class them among the Phallusiadae, and regard

them as establishing a close connexion between the simple and

aggregated Ascidia ; by their viscera situated behind the branchia

and a certain number of less important characters, they are more

nearly allied to the donee than to the true Fhalliisiee; but they

nevertheless approximate to the latter by the presence of longitu-

dinal folds in the branchial wall. The affinities of the lihopaleee are

therefore numerous, and they form as it were a bond of union

between several different groups ; in the general arrangement of

their organs they show a certain resemblance to the Clavelince.,

while at the same time they are simple Ascidians very nearly allied

to the donee and also ])resenting some relations to the Fhallusiee. —

Comptes Eenchis, May 19, 1884, p. 1294.

On the Frocess of Digestion in Salpa.

By Dr. Cn. 8. Dolley.

The author remarked that, preliminary to giving tlie full re-

sults of a somewhat extended study of the histology of iSa/pa, he

desired to make a few remarks in reference to certain statements

Miscellaneous,

73

recently made by Dr, A. KorotnofF of Moscow which he considered

erroneous in so far as they indicate the presence of a huge amoeboid

cell or plasmodiiim, in the oesophagus and stomach of ^aVpa, func-

tioning as a digestive organ. Dr. Korotueff describes this cell as

arising from the repeated division of a single cell which early in

the life-history of the animal is separated from the intestinal wall.

This giant cell or plasmodiiim, acting like a huge rhizopod, carries

on a form of parenchymatous digestion of the food taken by the

animal, passing the resulting chyle into the walls of the intestine by

means of its pseudopodia. Now by reference to an article by

Metschnikoff “ On Intracellular Digestion in Invertebrates ” (in the

‘ Quarterly Journal of Microscopical Science’ for January 1884), it

will be seen that such a form as Korotneff describes has never been

met with, and his description stands alone and anomalous, both as

regards the situation and size of the digestive plasmodium and as to

the method of its formation, for in all cases in which such struc-

tures have been found in Invertebrates they have always arisen by

the fusion of separate cells, not from the repeated division of one cell.

In a large number of series of sections made by the new “ribbon ”

method, the speaker was not only unable to find “ the lumen oblite-

rated ” by the peculiar structure of the wall of the intestine de-

scribed by Korotneff, but in a model of the visceral nucleus made

after Born’s “ Plattenmodellirmethode ” the lumen of the entire

intestinal canal is shown to be completely free throughout. He

did, however, get sections which gave pictures almost identical with

those portrayed by Korotneff, i. e. the lumen filled with what he

describes as a large nucleated granular cell, containing various food-

particles, and he could trace this so-called “ cell,” not only back

into “ the portion of the intestine lying next to the stomach,” but

through, the rectum into the cloacal chamber, and through the

oesophagus into the branchial sac. He accounts for it as follows -

The endostyle of Salpa has been very carefully studied by Hermann

Fol, who demonstrated, by means of carmine suspended in water,

that it threw out a constant stream of mucus when excited by the

presence of nutritive material in the same water, with a reflex

action, like a salivary gland. The mucus is, by an arrangement of

cilia, spread out like a curtain over the inner surface of the branchial

sac, when it acts as a means for catching the food-particles from the

ingurgitated water. By the action of ciliary bands bordering the

groove of the endostyle, the mucus is swept towards the oesophagus,

and as it approaches this, it is, by means of the stiff cilia on the

sides of the gill, twisted into a thread, and carried by a continua-

tion of the aforesaid bordering bands through the oesophagus into

the stomach. Now in studying a series of sections of a Salpa which

had had abundant food, we found as we approached the oesophagus a

mass of material answering to the description of Korotneff’s “ rhizo-

pod.” It takes staining readily and may be traced backward into

* Ueber die Knospung der AnchiniaP in Zeitschr. f. wiss. Zoologie,

Bd. 40, Hft. i. (1884). •

Ann, & Mag, N. Hist, Ser. 5. Vol, xiv. 6

74

Miscellaneous,

and through the oesophagus, stomach, and intestine. As the sections

api)roach the rectum, however, the mass gradually ceases to take

staining, and is much more distinctly marked out from the intes-

tinal wall, having had all the organic matter digested out, and

consisting only of the inorganic remains, which do not stain. The

alimentary matter of Salpce is composed of animal and vegetal

elements in nearly equal proportions, and the microscope reveals

the calcareous shells of Foraminifera, the beautifully sculptured

frustules of Diatomaceae, keen siliceous needles, and the sharp

armatures of minute Crustacea.

In the fore part of the intestinal canal, the food-mass, staining

almost as readily as the wall of the gut itself, seems to merge into

the ill-defined epithelium of the latter, and it is scarcely possible

to say where the food-bearing mucous thread eeases and the intes-

tinal epithelium begins, especially as this latter has a rugous ar-

rangement. That we have here to do with a form of digestion

entirely anomalous and unprecedented, he could not believe, and

begged leave to differ from Dr. Korotneflf on this point. Fol and

others have recognized the endostyle as a sort of salivary gland,

and have traced its food-laden mucous thread into the stomach of

the living animal, while the speaker had been able to trace the

same thing in well-preserved specimens. He had also several series

of sections from animals which must have been without food for

some time previous to death, in which the lumen of the intestine is

not only free of food, but of any obliterating mass of cells or plas-

modium. The only protoplasmic bodies not food are certain Grega-

rm«-like organisms adhering to the walls of various parts of the

intestine, and which he took to be parasites. These give on section

the appearance of the large “ scattered cells, entirely free from

their surroundings,” which Korotneflf figures and regards as ‘‘ analo-

gous to the great stomach-cell of Anchinia.'” The first opportunity

would be taken to examine these structures in living Salpce ; but

he was now forced to conclude that Dr. Korotneff has endowed the

food-bearing mucous thread with a power it does not possess, that

Salpa does not exhibit any unusual form of intracellular digestion,

and that there is no immediate cause on its account for questioning

the high genetic place occupied by the Tunicates. — Froc, Acad.

Nat. Sci. Philad., April 15, 1884, pp. 113-115.

On a Species of Tachina occurring on the Tracheal System of Carabus.

Dy M. K. Cholodkowsky.

In the summer of the year 1882, when I was examining

various species of the genus Carahus for purposes of comparative

anatomy, I found on the abdominal stigmata of some specimens of

Carahus cancellatus some peculiar small whitish bodies which pro-

jected freely into the body-cavity of the beetle. These bodies were

of an oval foi’m and about 1 millim. long. On closer examination,

after cutting out the stigma with a small piece of skin and with

the tracheal stem starting from the stigma, the following proved to

Miscellaneous,

75

be the case. The oval whitish body had one end turned towards a

thick tracheal stem close to the stigma, and tliis end of it was

pushed into a brownish chitinous cup wliich surrounded it, but the

narrow base of the cup was attached to the trachea. After removing

the whitish body from the cup it could be seen that at the bottom

of the latter there was a small aperture leading into the trachea.

From the margins of the cup extended irregular translucent chiti-

nous deposits which surrounded the whitish oval body. On micro-

scopic observation of the whitish oval body annulation was observable

upon it ; tubular organs (alimentary canals) shimmered through

from the interior ; at the end turned towards the body-cavity (of

the Carahus) sharp booklets were observed, and at the opposite

extremity two respiratory apertures. From these characters a

young Tachina-lsiTysi was easily recognized in these little bodies.

Soon after I obtained some specimens of Carahus cancellatus, each

of which was infested with several Tachina-lar\se already full-grown.

The larvjB were so large that they filled nearly the whole ventral

cavity of the beetle. The beetles infested by Tachince were dis-

tinguished by their sluggishness from those not so attacked, and

soon died in captivity. The chitinous cups which embraced the

hinder extremities of the large larvae were large and had an irre-

gular margin ; the translucent chitinous deposits which surrounded

the body of the larva were greatly developed and had the form of

irregular and in part confluent lobes. In short, chitinous patho-

logical structures surrounded the body of the larva just as inflamed

connective formations enclose foreign bodies which have got into the

body of a vertebrate animal. This fact is certainly in favour of the

conception of chitine as the physiological equivalent of the connec-

tive tissue in the bodies of insects. There is no doubt that the

deposition of chitine took place from the soft hypodermal layer of

the wall of the trachea. Fesides Carahus cancellatus I obtained a

specimen of Carahus glahratus^ which was also infested by numerous

larvae of Tacliina.

I did not succeed in rearing a single fly from any of these larvae,

partly because I had other purposes in view in the investigation of

the species of Carahus, but partly because the infested beetles did

not live long in confinement. I hoped to have obtained some in-,

fested Carahi in the summer of 1883, but none of the beetles of this

kind collected by me that summer contained Tac7im«-larvse. On

the other hand, I found a specimen of Harjpalus ruficovnis which

was literally stuffed with these larvae.

The occurrence of rac7im<?-larvae in the bodies of adult insects is

by no means a new fact. As long ago as the year 1828 Boheman

found the larvae of Uromyia curvicauda in Harpalus ruficornis and

H. aulicus *. Leon Dufour has described Hyalomyia dispar, which

is parasitic in Bracliyderes lusitanicus'\ ; and he also found the

larvae of Phasia in Pentatoma grisea and Cassida viridis and the

^ Stockholm Akademiens Handlingar, 1828, p. 164.

t Ann. Soc. Ent. Fr. 1852, p. 443.

X Ibid. 1848, p. 427.

76

Miscellaneous,

larva of Ocyptera hicolor in Fentatoma yrisea* * * §. As regards the

s})ecies of Carabus, Bo3^e, in the year 1838, reared Tachmce from

Carahas violaceas^ ccmcellatus, and clathratus f. The description of

this fly under the name of Tachina pacia is to be found in Zetter-

stedtj.

Since that time, so far as I know, no 7V(c7wna-larva3 have been

found in species of Carabus. The species Tachina pacta is very

little known and very doubtful. Our llussian dipterologist, J. A.

Portschinsky, to whom I am indebted for many references to lite-

rature, is of opinion that Tachina pacta is identical with Tachina

{Masicera) cinerea. Zctterstedt himself says of Tachina pacta that

it is “ valde similis et affinis Tachina cinereceF Schiner is of the

same opinion.

As regards the mode of penetration of the larva into the body of

the Carabus, we must, in all probability, consider that the fly de-

posits its egg in the stigma, and the larva, escaping from the egg,

bores through the wall of the trachea and gradually extends its body

into the bod}^-cavity of the beetle. During this time there are

formed around the larva, on the part of the hypodermal layer of the

trachea, chitinous deposits, which are strongest in the vicinity of

the abdominal wall at the hinder extremity of the larva, and here

form a brown cup, the margin of which, however, passes without

any sharp boundary into the translucent chitinous lobes which sur-

round the rest of the body of the larva.

A little while ago, Jules Kiinckel d’Herculais described a parasitic

fly {Gymnosoma rotundatum), the larva of which lives in the body

of Fentatoma §. In this case also the larva has its hinder extremity

turned towards the stigma, and this end is embraced by a chitinous

cup, called by Kiinckel “ le siphon.” Kiinckel, however, thinks that

“ le siphon ” is a secretion of the larva itself, and by no means a pro-

duct of the hypoderm of the infested insect. Kiinckel also describes

the mode of penetration into the body in a different fashion, namely,

that the fly sticks its eggs to the ventral segments of the Fentatoma,

and the escaping larva penetrates between the ventral segments

into the abdominal cavity and only by degrees becomes connected

with the stigma. However this may be with regard to the Fenta-

toma and Gymnosoma rotundatum, in Carabus the case is most

probably as I have suggested. The penetration of the larva through

the stigma is in this instance evidenced by the fact that even the

very smallest larva3 are attached to the stigma, and that they are

only met with on the stigma. — Zooloyischer Anzeiyer, no. 169, June 9,

1884, p. 316.

* Ann. Sci, Nat. tome x. p. 248.

t ‘ Kroyer’s Naturhistorisk Tidsskrift,’ 1838. See also Erichson’s

‘Bericht iiber die wiss. Leistungen im Gebiete der Entomologie im Jahre

1838,’ Berlin, 1840, p. 93.

t ‘ Diptera Scandinaviee,’ tome iii. pp. 1038-1039.

§ Ann, Soc. Ent. Fr. ser. 5, tome ix. (1879).

THE ANNALS

AND

MAGAZINE OF NATUEAL HISTOEY.

[FIFTH SERIES.]

No. 80. AUGUST 1884,

IX. — Notes on Species o/Ascodictyon and Rhopalonaria/roDi

the Wenloch Shales, By George Robert Vine.

In the ^ Annals and Magazine of Natural History ’ for June

1877, Prof. H. Alleyne Nicholson and Robert Etheridge, jun.,

published their joint paper on Ascodictyon^ a new Provi-

sional and Anomalous Genus of PalaBozoic Fossils.” In that

paper (pp. 463-468) the authors describe forms from the

Devonian (Middle, of Ontario) and Carboniferous strata (of

Scotland). In a paper read before the Geological Society and

ultimately published in their journal Notes on the Polyzoa

of Wenlock Shales,” Feb. 1882), I placed upon record

(p. 54) the name Ascodictyon filiforme^ Vine, as a provi-

sional one. Since the publication of that paper I have been

closely engaged on the study of Silurian and Carboniferous

Ascodictyaj and I find that the forms that I originally placed

under the above name may conveniently remain. The or-

ganism, however, is such a peculiar one, and my opportunities

of studying its varied aspects so singularly fortunate from the

possession of a large series of specimens, that I make no

apology for adding further details of the species to the brief

notice already referred to.

The genus Ascodictyon is Palgeozoic, though not peculiarly

Ann. & Mag. N. Hist. Ser. 5. Vol. xiv. 7

78

Mr. G. H. Vine on Species of

so. In examining a series of Cretaceous Polyzoa in the

possession of Miss E. C. Jelly, one specimen appeared to me

to ])clong to the genus.

In describing an American Silurian form — in some respects

similar to forms found in the Wenlock Shales — Mr. E. O.

Ulrich (Journ. of Cincinnati Soc. Nat. Hist. April 1879, pp. 18,

19) has established a genus under the name of lihopalonaria.

The species of the genus, B. venosa^ Ulr., the author places with

the Crisiidse, remarking, that the form has only been observed

incrusting Streptelasma corniculum. On account of the great

delicacy of the fossil, the fronds themselves are rarely found ;

but instead we find a series of impressions on the exterior

coat of the Streptelasma^ which very well represents the fronds

and cells of the same.” A specimen of the species described

by Mr. Ulrich is before me. It is from the same Cincinnati

rocks ; as I shall have to refer to the genus again, I have

thought that it might be more satisfactory to make reference to

an actual specimen than to the mere description of the same.

Ascodictyon, Nicholson & Etheridge, Jun.

Ascodictyon, Nich. & Etli. jun., Ann. & Mag. Nat. Hist,, June 1877,

pi. xix. ; “ Notes on Polyzoa of the Wenlock Shales,” Vine, Quart.

Journ. Geol. Soc. Feb. 1882, pp. 52, 5o.

1. Ascodictyon JiUf or me j Vine.

Ascodictyon filiforme. Vine, Notes on Polyzoa of the Wenlock Shales,”

op. cit. pp. 54, 55 (merely referred to in the above).

Organism filamentous, forming linear, contorted or clustered

threads, adherent to shells, stems of crinoids, fragments of

trilobites, but rarely to corals. Filamentous threads hollow,

but surrounded by delicate calcareous walls ; the hollows filled

with a dark brown granular mass. Lagena-\\kQ divergences

developed on the sides of the thread, sometimes as single

vesicles, otherwise as groups of vesicles. Peculiarly clustered

stellate fibres are also formed at unequal distances.

Locality, Buildwas beds, generally distributed throughout

the whole of the washings, but more abundant in nos. 36

and 38.

This curious organism begins its existence as a mere speck

upon stone or shale or stem, which forms the nucleus of a

colony. From this delicate filaments are developed (fig. I. 1 «,

o})^ sometimes in two or three, at other times in four different

directions ; these vary in thickness, but the average size, both

in breadth and depth, may be taken as measuring between

and 2^-0 inch. The threads are sometimes, but rarely,

white, more generally of a dark brown tint. Viewed as

79

Ascodictjon and Rhopalouaria.

opaque objects ou shell-fragments they appear like fine hairs

laid in lines across the surface, or contorted, crossing each

other at ditferent angles, or running in parallel lines ; but it

would be useless to direct attention to any s})ccial feature in

their inodes of growth, for they vary considerably ; but one

Fiir. L

1. Ascodicty 071 Jiliforme,Y me, Two filiform threads run-

ning almost parallel. The point of origination is a} ; the portion

marked a is nearly similar, but wanting the nucleus ; single

vesicle.

2. A. radiciforme, Vine, inch thick. One of the filiform contracted

threads, placed here for comparison,

3. Thread of A. filiforme, drawn from a transparent specimen, showing

dark brown pulp.

feature which appears to me to have something to do with the

development of the colony must not be lost sight of. Occa-

sionally some of the filaments bifurcate, and before bifurcation

takes place the organism contracts at intervals, and out of this

contracted portion a new thread originates. It must not be

supposed, however, that this simple explanation is, on the

whole, a plainly satisfactory one. The two different sorts of

threads, though apparently allied, give rise to two distinctly

separable colonial growths in the after stages of their exis-

tence. Yet in the earliest, or initial stages, it is not easy

to distinguish the difference between them. It will be better,

however, to keep the two forms distinct.

In a few cases I have been able to reduce the thickness of

the shell to which a typical A, filiforme is attached, and have

mounted the specimen in balsam on glass. It is then seen

that the filament is hollow, and the central parts filled with a

dark brown granular matter. This granular matter, which I

7*

80

Mj% G. K. Vine on Species of

sliall call tlic pul]), is sometimes eontinuous, at other times

sliglitly separated from the adjoining mass, when the grains

ap|)ear like a row of beads dotting the centre of the threads

(figs. I. & II. 3 to 5). Surrounding the pulp are delicate and

trans})arent walls, which are distinct and clearly defined. In a

few cases the filament bifurcates and the pulp in the undivided

portion se])arates at the node and passes into the divisions

thus formed. Ileal development takes place at irregular inter-

vals. On the sides of the thread there is a slight protuberance

of the wall, which increases in size until a roundish knob or a

lagena-like vesicle is formed. Into these vesicles, which may

be either single or in groups, the pulpy mass passes ; but to

wdiat extent the colony increases beyond the A.-siluriense

stage, I am unable to say ; although I believe we may safely

regard this species, at least, as the ultimate outcome of the

colonial development of A. jiliforme^ Vine.

4; 5. Ascodictyon jiliforme (transparent and semitransparent), showing

vesicles and pulp.

6. Ditto, showing clusters of vesicles, passing into

7. A. siluriense, Vine.

At this stage of inquiry it may be well to ask, whether the

name A. siluriense^ Vine (Wenlock Polyzoa, op. cit. p. 52),

should be suppressed. If the Silurian name is suppressed

the Devonian name [A. stellatum^ Nich. & Eth.) must be

suppressed also, unless the one name may be allowed to

embrace both the type and the varieties. In opposition to this

view, I think the wiser course would be to allow the forms to

retain their present names as given below, for the simple

reason that it would be perhaps impossible, or almost impos-

sible, to make another collection similar to my own unless the

Fig. II.

81

Ascodictyon and Rliopalonana.

slinles were searched with the same minute care that I have

bestowed upon those furnished to me by Mr. George Maw.

Then, again, M. siluriense and A. radiciforme were the firstfruits

of my labours; and A. jiliforme was tlie result of closer exa-

mination. As with me so with others, because the matured

forms will, I fancy so at least, be considered by the student of

micropalaBontology as by far the most important as initial

stages in an inquiry like the present one. Another eonside-

ration is with me of much greater weight than any previously

given. It may be, after all the care that I have exercised in

thus tracing the origin and development of a colonial growth,

that some few facts or fossil illustrations may have been over-

looked, and it may be necessary at some future time to limit

the type now characterized as A. jiliforme. If this should

happen, the suppressed names would have to be restored.

In his observations on A. stellatuin Prof, Nicholson remarks

{op. cit. p. 465), that in its youngest stage tlie organism pre-

sents itself simply in the form of scattered oviform or pyriform

calcareous vesicles attached to the exterior of foreign bodies.

When mature it consists of similar vesicles combined into

clusters.’’ I do not doubt the accuracy of Prof. Nicholson’s

observations, though I cannot, on the whole, endorse them from

my own labours : all the vesicles are united to the filiform

thread, though in a few isolated instances “ apparently ” they

are not so. I have examined a large series of A. filif or me for

the purpose of putting the observations of Prof. Nicholson to

the test, and I am consequently unable to confirm his views.

In his description of fig. 6, pi. xix. (pp. citi) , the author says,

“ Four young vesicles (?) of the same [A. stellatum) &c. the

vesicles are not foraminated, as in the other figures (2 to 5,

pi. xix.) , and in this special feature the Ontario vesicles are allied

to, though not identical with, those shown in fig. II. of the

present paper as gradational stages in the development of A .

jiliforme j Vine.

2. Ascodictyon stellatum^ Nidi. & Eth., jun.,

var. siluriense j Vine.

—Ascodictyon stellatum^ Vine, Quart. Journ. Geol, Soc. Nov. 1881,

P-

= Ascodictyon stellatum, var. siluriense, Vine, Q. J. G. Soc. Feb. 1882,

p. 52.

The details of this species ha.ve been given in the works

referred to above.

82

Mr. G, E. Vine on Hjpecies of

3. Ascodictyon radiciforme^ Vine.

=1 Ascodicfyon radians?, Vine, Quart. Journ. Geol. Soc, Nov. 1881,

p. GIO.

=■ Ascodictyon radiciforme, Vine, Q. J, Geol. Soc. Feb. 1882, p. 53.

When I wrote tlie details already given in the papers re-

ferred to above I did not possess the fine suite of specimens

which has enabled me to extend my remarks on the origin,

growth, and apparently final (?) development of this peculiar

group of organisms, found as yet only adherent to other

organisms in the Wenlock shales.

In speaking of A.filiformej I incidentally referred to a single

form apparently related to the species, but which I said it

would be best to keep separate. I have now to consider the

relationship of that form, but which, not being the earliest in

the development of the species, I will defer till later on.

In fig. III. 5 I have sketched the fragment of acrinoid stem,

magnified about 2 diameters, on which one of the finest of my

colonies of the earliest stages of this species {A. radiciforme)

is attached. I do not, however, found the whole of the evi-

dence which I am about to bring forward on a single specimen.

I have corrected both the positive and the negative evidence

by appeals to between sixty and seventy other specimens ; but

as this one affords me evidence of continuous growth, I have

built up my description mainly on it. And here I must be

allowed to say that, contrary to my general plan in drawing,

the whole of my sketches are drawn by the eye, and not by the

aid of the camera lucida ; but every figure is a faithful delinea-

tion of the original.

In fig. III. 1, 2, and 3 I have drawn a fibre which is conti-

nuous on the fragment of crinoid stem already referred to. At

2, a and a', we have two nuclei, either of which may be

referred to as the originating nucleus of the colony ; but it

will be best to speak of them separately. In fig. III. 2, «, a kind

of false stellate cluster is formed ; but this I regard as only an

offshoot ; the true nueleus is at A, and at this the central part

only. It appears to me that the central nucleus originates by

the combination of minute spores, which up to the present

have escaped my observation ; but immediately after the

combination, delicate prolongations of the central mass or

radii are sent off which do not in every case produce fibrous

threads. At a one of these minute combinations is formed,

but only one of the rays is apparently developmental *.

* It will be understood by the paljeontologisttliat in restricting- myself

thus I only take the evidence presented to me. I have not the least

doubt but that what I say of one might be said of all the rays ; but I

have only the one evidence to rely upon — the positive.

Ascodictyon and Rliopalonaria. 8H

This extends for a short distanee, wlien anotlier, or pseudo-

combination, takes place, the result of which is shown, so far

as I arn able to trace it ; but in that portion which recedes

towards a" the fibre is slightly contracted at certain distances.

Fig. IIT.

1 to G. Varied forms of Ascodictyon radiciforme, Vine (already

described in text).

In fig. III. 1 the fibre is white, undoubtedly hollow, with here

and there a broken surface, showing the dark brown pulp (or

matrix) below. In fig. III. 3, which is a continuation of fig. III.

2, at a" we find the continuation of the contracted fibre and

84

Mr. G. R. Vine on Species of

other features which it may be well for the reader to refer tOy

the stellate cluster being a continuation of the fibre at 3*. In

following the above remarks it will be easily understood that

1 attach very great importance to the method of formation and

the character of the tongue-like vesicles which form the radii

of this anomalous species, A. radiciforme. Sometimes after

single vesicles are formed, at other times after a combination

of vesicles^ the organic matter of the thread or fibre undergoes

other changes, to which reference has not been made. A group

of vesicles combined as in fig. III. 3^, some of the cells, and

occasionally the w^hole of them, will contract tov/ards the

centre or separate from the nucleus, forming club-like cells.

In nearly all the instances where I have seen this mode of con-

traction the cells have a very delicate covering of calcareous

matter, and are foraminated either in a single line along the

centre or indifferently over the whole surface ; the lattery

however, is a very rare occurrence. I have given figures of

the club-like cells (fig. III. 4 a, 4Z>), showing the direction and

positions of the foraminated surfaces ; but in fig. III. 4 I have

shown the basal attachment of one of the fibres, which shows

that the under surface was more densely foraminated than

the upper. The irregular fibre shown in fig. III. 6, though

not strictly speaking a portion of a stellate cluster, is likewise

foraminated,

In the ^ Journal of the Cincinnati Society of Natural

Eistory,’ 1879, vol. ii. pi. vii. figs. 24 and 24 a, Mr. E. O.

Ulrich describes and figures a very peculiar fossil adherent

to the exterior coat of Streptelasma corniculum. For the

species a new genus is founded- — Rhopalonariaj from rliopalon^

a club- — which Mr. Ulrich places in the family Crisiidie ; and

he says that the genus is related to Hippothoa.j but in the

form and arrangement of the cells they differ widely.” There

is not, however, any relationship to Hippothoa in the species

described ; but as the cells are somewhat club-shaped, it may

be well to accept the genus if more fully defined.

In the species, R. veiiosa^ Ulr., the cells are uniserial,

long, acutely elliptical, and joined together at their contracted

ends. . . . Cell-mouths not clearly determined, hut appear to

he situated near the middle of the celV j". In my own speci-

men of tliis species I cannot detect any cell-mouths, but the

other characters are well defined by the author.

In the Ludlow rocks and also in the Wenlock shales there

are several fossils that might have been conveniently placed

in this genus if my specimens did not throw some little light

t Italics mine.

85

Ascoclictyon and E-hopalonaria.

U])on tlicir origin and development. In fig. IV. 1 have given

sketches of four distinct types of very common forms found in

the shales. In fig. IV. 1 we have the rarest of these, showing

elongated cells very similar to some of the cells of Stomato-

'pora elongata^ Vine (fig. IV. 2), only tliat the cells are in an

opposite direction to what (apparently V) is the case in 8.

elongata. I cannot, however, detect any orifices in these cells

such as we have in S. elongata.

Another form, very common indeed, is shown in fig. IV. 3.

Only that the cells are not club-shaped, I suggested for this

type R. hotellus^ as descriptive of its peculiar sausage-like

character. But even of this type better evidence is afforded by

Fig. IV.

1, 3, 4, 5. Rhopalonaria, Ulricli : species described in text.

2. Stojnatopora elongata^ Vine.

fig. I V. 4, in which both the hotelloid and the rliopaloid charac-

ters are shown in one colony. Some of the cells in the last figure

have, one would suppose at first sight, cell-mouths. This is

not so ; those cells that are shown thus are a little more cal-

careous than the others ; the walls are broken at this part,

and the dark brown matter is shown below. I have some

few specimens of the last two types, and the whole of the

walls or outer covering is destroyed, and the hotella-Y[kQ

matrix is still adherent to the fragments to which the original

organism was fixed.. In fig. IV. 5 we have a fourth and

80 Mr. G. II. Vine on Siyecies of

last type, wlilcli is similar in some respects to U. venosa^

Ulrich

Jn addition to my own observations I have the evidence

of a most careful observer as to the existence of Rhopalonaria

in the Ludlow rocks. In 1881, J. D. Longe, Esq., of Chel-

tenham, sent me a series of sketches (fig. V.), with the

MhoimloJiaria, Ulricli, from the Ludlow rocks. Figures supplied by

J. 1). Longe, F.G.S., Cheltenham.

following remarks : — I also enclose a sketch of a very

abundant, encrusting, creeping ‘ stoloniferous ’ form, which I

have on shells {Sjoirifer) from some U])per Silurian bed . . . .

probably Ludlow.” These beautiful forms are different from

any known to me in the Wcnlock shales, and their publica-

* Not similar to his figures, hut similar to some cells seen on the spe-

cimen of Ulrich’s species in my own cabinet.

87

Ascoclictyon and Rliopalonaria.

tion with the above details may be the means of lielping others

in their researches among the micaceous and otlier shales of

the Ludlow series of rocks. My labours are entirely confined

to the Wenlock series and some only of the shales over the

Wenlock Limestone.

I do not think it necessary to speak more fully now of the

genera Ascodictyon and Ilhopalonaria. It is important, how-

ever, in vindication of my remarks, to make some reference to

the various opinions on the organisms which Prof. Nicholson

gives in the section of his paper entitled Systematic Position

and Affinities ” [I. c. p. 466). He says, in the first place, that

Dr. Strethill Wright (to whom Scotch specimens of A. radians^

Nich. & Eth., were submitted for examination) was unable

to throw any light upon their nature.” Prof. Huxley, to

whom the same specimens were submitted, after considerable

hesitation, suggested that they might be Protozoa. Mr.

H. B. Brady, after a protracted examination of both the

Scotch and the American forms, has arrived at the conclusion

that they cannot be referred to the Foraminifera. Some of

the American specimens [A. fusiforme^ N. & E., and

stellntum^ N. & E.) were kindly submitted by Mr. H. B.

Brady to the Kev. Thomas-Hincks, who suggested that they

were possibly allied to the recent Anguinarice. Neither Prof.

Nicholson nor Mr. Etheridge expresses any positive opinion

as to their systematic position or affinity. The difficulties

encountered by these various authorities when speaking of this

remarkable group are valuable so far in helping to establish

the uniqueness of the t}q)es submitted to them ; but none of

the suggestions help to throw light upon their nature and

affinity. Yet I have respected the whole of the remarks, and

have compared specimens of the fossil species with specimens

of every known living type suggested as probable” by these

authors, but without any definite results. Perhaps it would

be wise to pause here, for it is not for me to suggest possible

affinities when so many experts have failed. Yet 1 cannot

allow the paper to pass out of my hands without making a

suggestion, which may possibly share the same fate as the

others.

There are not, so far as I am aware, any Cyclostomatous

Polyzoa which may be considered as truly stoloniferous.

Some of the Hydrozoa are; but I know of none whose

stolons are adherent to stone or shell, such as are found in

these ancient rocks, neither am I aware that the stoloniferous

Ctenostomatous Polyzoa are adherent to stone and shell, like

Ascodictyon or Rliopalonaria. Yet it seems to me that we

have, in Ascodictyon fiUf or me at least, primitive representatives

88 On Species (^Ascodictjon awe? Ehopalonana.

of the stolonifcrous Vesiculariida3, such as Vesicularia and

Bowerhanhia^ or, possibly, some member of the more humble

race of the Entoprocta. Barrois has already, in his paper “ On

the Embryogeny of the Cyclostomatous Polyzoa ” (Ann. &

Mag. Nat. Hist. Nov. 1882, p. 402), spoken of a pro-Bryozoan

race, composed of “ free swimming organisms.” May Asco~

dictyon be the attached, or larval form, of some of the as yet

unknown pre-Upper- Silurian types of organic life, polyzoan

or otherwise ?

There is, however, another suggestion which may help to

throw some little light on the development of A. radiciforme^

Vine, though we cannot hope by the comparison to explain away

all the difficulties which surround the subject. I refer now

especially to some remarks contained in a paper by Mr. George

Busk, E.B.S., entitled “Notes on a peculiar Form of Poly-

zoa closely allied to Bugula^ —Kinetosldas^ Kor. & Danielsse i ”

(Quart. Journ. Microsc. Soc. vol. xxi. new ser.). After

speaking of the development of the various species of Kineto-

sldas^ Mr. Busk says, “ I have yet scarcely adverted to the

most remmrkable feature of Kinetoskias^ viz. the peduncle or

stem, which appears to exist in all species. . . . The mode of

formation of this part of the zoarium, which is undoubtedly

the homologue of the bundle of separate radical tubes so

commonly met with among the Polyzoa, is extremely curious

and interesting, and, at the same time, in some points as yet

more or. less obscure, as, in fact, may he said respecting the

mode of formation and development of the more ordinary form

of radical tubes *.

“ In the more common forms they are cylindrical, jointed,

chitinous tubes, with rather thick walls and with very scanty

contents, beyond a few granular particles and irregular

threads, representing, as it would seem, the remains of an

endosarc, with which, in order that their progressive increase

in length, and occasionally complicated branching &c., may

be effected, we must suppose the tube to be furnished. In

fact it is otherwise impossible, without assuming the presence

of a germinal material, to account for the fact that even after

the tubes have attained a considerable length the extremity,

or a considerable part of the tube, may undergo great changes

in form, as is seen in the production of hooks and other means

of ensuring adhesion to foreign bodies, changes showing a

most extraordinary adaptability to circumstances. Not the

least remarkable of these adaptations is the division of tlie

extremity of the tube into a multitude of very minute tubular

* Italics mine.

89

On two new Species Walckeiiacra, Blackw.

filaments, cacli of wliicli may be traced into independent con-

nexion with small foreign bodies.” Had Mr. Busk been

writing of Ascodictyon instead of species of Kinetoskias^ lie

could not have given more faithful descriptions of some of the

specimens found in the Palaeozoic rocks. The suggestive

inference to be drawn from these remarks, and others that

might have been given, is that the dark brown masses (pulp

&c.) in the various species of Ascodictyon are probably the

remains of endosarc in these once living filaments and semi-

tubular and bulbous tubes.

ddiere is just one other point in Mr. Busk’s paper to which

I will direct attention in conclusion, because it will help us

to understand and appreciate at its proper value Ascodictyon

and the abortive or ‘‘ blind cells ” of Rhopalonaria : — That

the radical and connecting tubes, like the avicularia and vibra-

cula, represent modified zooids, is, I believe, generally admit-

ted ; nor can it be denied in this case {Bugula &c.) that each

successive joint or internode is a distinct zooid.” And in a

note the author says, “ In Bicellaria and in Notamia it may

almost be said that the inhabited part of the zooecia is simply a

dilatation at one part of the internode of a radical tube, which

is continued to the ultimate extremity of the branch.”

X. — Descriptions of two new 0/ Walckenaera, Blackw.

By the Rev. O. P. Cambridge, M.A., C.M.Z.S., &c.

[Plate IV.]

Among a large number of Spiders sent to me for identification,

during the last three or four years, by Major-General A. W.

M. van Hasselt, from Holland, are two of the curious genus

Walckenaeraj Bl., which I believe to be undescribed. Their

discoverer having kindly permitted me to do so, I now sub-

join descriptions and figures of these novelties.

Family Theridiidae.

Genus Walckenaera, Bl.

Walckenaera Hasseltiij sp. n.

Adult male, length of inch, or f of a line.

Gephalothorax rich black-brown.

Legs yellow, tinged with orange-brown. Perhaps i.i some

examples they would be bright orange-yellow.

90

Ilcv. 0. P. Cambridge on tioo

Abdomen jet-black, thinly clothed with short fine yellowish

hairs.

The upper part of the caput is slightly raised above the

ordinary level, and presents a flattish summit, the middle of

which forms an oval distinctly marked by a surrounding

groove ; the hinder slope or occipital portion looked at in

profile is short and abrupt. The profile resembles in this

respect pretty nearly that of W. Beckii^ Canibr., and a not

very strong, small, narrow-oval, slightly curved indentation

or fovea runs backwards from just above each lateral pair of

eyes a little below and parallel with the upper margin of

the caput ; in the middle of the ocular area are a few short

-strong upturned hairs. The cephalothorax is short, almost

round, and the thoracic region is somewhat flat and its sur-

face rugose, while that of the caput is smooth and glossy.

Eyes small and very indistinct, placed on the fore part of

the caput, one pair (the hind centrals) on the anterior edge of

the raised portion in a transverse line about a diameter apart

from each other ; immediately beneath them is a slight trans-

verse indentation, and just below each extremity of it is a

lateral pair placed obliquely, and midway between the two

lateral pairs (and contiguous to each other) is the fore central

pair.

The height of the clypeus is about two thirds of that of the

facial space.

Legs slender, not very long nor very unequal in length,

1, 4, 2,»3, furnished with very short fine hairs only.

Palpi short, similar in colour to the legs, excepting the

radial and digital joints, which are dark yellow-brown. The

radial joint is similar in length to the cubital, but much

stronger ; it is of a rounded spreading form and has a short

bifid apophysis bent abruptly inwards at its fore extremity on

the outer side. The digital joint is rather large, oval. Palpal

organs highly developed, prominent, with several spines and

corneous processes ; a very long, slender, filiform spine issues

from the middle of their outer side and curves round beneath

with a long, free, sinuous, exceedingly slender, hair-like point,

and another short black spine is curved in a circular form at

their extremity.

Falces small, conical, directed strongly backwards, and of

a deep yellow-brown colour.

Maxillw and labium of the ordinary form common to the

genus, and similar in colour to the falces.

Sternum large, short, heart-shaped, convex, and of a glossy

deep black-brown colour.

An example of this distinct spider, which I have great

91

naio Bpecies of Walckcnacira, Blachw.

])lcasm-c in naming after its discoverer, was sent to me in

1880 by Major-General A. W. M. van Ilassclt from the

neighbourhood of the Hague, Holland. This species appears

to be nearly allied to W. sor didata ^ Thor. [W. atra^ 131.), a

spider I have never seen ; but the description of it is not

sufficiently close to the spider now described to justify the

conclusion that the two are identical. The profile also is

much like that of W. erytJiropus^ Westr. ; but the caput is

less elevated and the palpi totally unlike. It is also allied to,

but I thiidv quite distinct from, W. elegans^ Cambr., a Bava-

rian species (P. Z. S. 1872, p. 766, pi. Ixvi. fig. 23).

Walchenaera nemoralwides^ sp. n.

Adult male, length of an inch ; adult female yV of

inch.

In size and form, as well as in the coriaceous punctured upper

surface of the abdomen, this little spider closely resembles IF”.

7iemoraliSj Bl. ; but the colour of the cephalothorax and abdomen

is blacker and that of the legs is a clearer yellow than in that

species. The two may also be more readily distinguished by

the form of the radial joint of the palpi. The apophyses of

this joint are in a similar position, but the outer (tapering)

one is much larger, longer, and more prominent, being double

as long as the joint, slightly curved, projecting outwards at

right angles to it, and very slightly hooked at the point. The

other apophyses are very similar to those of W. nemoralis^

but proportionately larger and of a different form, that in front

being constricted near the middle.

In IF. nemoralis the outer apophysis is not only shorter and

less strong, but consists apparently of two parts, a basal por-

tion, prolonged and ending with a very fine, sharp, somewhat

thorn-like addition.

Examples of this spider have been sent to me at different

times during the last two years by Maj.-Gen. A. W. M. van

Hasselt from Holland. Mons. Simon, on examining one of

these which I forwarded to him, considered it to be only an

example of IF. nemoralis^ Bl. ; but the differences above

noted (as well as some other minor ones) are so constant that

I do not feel the smallest doubt of its being specifically

distinct. I have received also one example of the typical

IF. nemoralis^ BL, $ , from Holland ; but the species now

described has not yet been found in Great Britain.

The female resembles the male in colours, but the occiput

is simply a very little gibbous when seen in profile, and the

height of the clypeus is rather less than half that of the facial

space.

92

Prof. F. J. Bell on Pentastonuim polyzonura.

EXPL.VNATION OF PLATE IV.

Fi(j. 1. Walchenaera Tlasseltii. a, spider, magnided ; b, ditto, in profile,

without legs or palpi ; c, slightly perspective view of caput ; d,

caput, from in front, showing the position of the eyes ; e, left

palpus, from in front and rather inside j /, natural length of

spider.

Fi(/. 2. Walchenaera nemoralioides. a, spider, magnified, cf ; b, ditto,

in profile, with legs and palpi removed ; c, caput, from in front,

showing the eyes ; d, left palpus, from in front and rather on the

inner side ; e, radial joint of palpus \ ^ in profile, without

legs or palpi ; h, natural length of spider ; k, genital aperture, 2 ■

Fif/. 3. W alckena'era nemoralis, Bl. Part of palpus of S •

XL — A second Note on Peiitastomura polyzonum.

By F. Jeffrey Bell, M.A.

In the sixth volume of the current series of the ^ Annals ’

(pp. 173-176) I published a short note on the rediscovery of

t\\Q Pentastomum polyzonum of Harley, two female specimens

of which had been acquired by the British Museum in 1880.

Lately we have received other specimens which formed part of

the collection of the late Dr. Edwards Crisp, but are without

any indication of origin* and not in first-rate condition.

A short time since an interesting essay on the structure of

Pentastomum was published by Mr. W. E. Hoyle in the

^Transactions of the Eoyal Society of Edinburgh’ (vol. xxxii.

pp. 165-191), in which he describes a new species {P, pro-

tells) , and gives an account of its anatomy.

Mr. Hoyle was fortunate enough to have examples of both

sexes of the parasite, and he describes the male as being

13-17 millim. in length, and as having sixteen or seventeen

annuli. Of the two specimens which formed the basis of my

former note neither was male ; of the seven specimens now

received one is a male, and I have been able to observe that

it, while measuring 36 millim. in length, has only seventeen

rings, and that the most anterior of these are much less pro-

minent than they are in the female. In addition, therefore, to

the numerous points of similarity indicated by Mr. Hoyle, we

have another in the smaller number of annuli in the male

than in the female. Another point is to be observed in the

* Although a careful search has been made in Dr. Crisp’s collections,

there are no indications of the Pentastomum annulatum of Baird, which

did, I believe, on the dispersal of the Zoological Society’s museum collec-

tions, pass into the hands of Dr. Crisp. It is greatlyho be wished that

this type should be found.

/]//// K' jl'hifi Nat Htst S F). Vof H /V IV

op Cambridge del.

P Carter litli.

Few Species of WcJclceiicLera BL

Mintcm Pros amp

LIBRARY

OF THE

UWiVERSITV OF ILLINOIS

Mr. 11, llitclicock oti the Causes of Variation.

fact that of the six female specimens now before me two have

twenty, while the others have only nineteen annuli ; in other

words, the result to which I was led {tom. cit. p. 176), a good

deal to my surprise, as to the great value of the number of

rings in the body, is a little shaken, although it falls in rather

with one’s general experience as to the specific value of num-

bers such as these. It is to be noted, further, that the two

females with twenty annuli measured respectively 75 and

80 millim., or less than three with nineteen rings, whieh

measured 90, 95, and 105 millim. ; a specimen of 46 millim.

in length had nineteen rings.

The fact that the male has seventeen annuli, while that of

P. j^rotelis has sixteen or seventeen, and the discovery of the

fact that the female of P. polyzonum is not absolutely limited

to nineteen rings, diminishes the gap that separated the two

speeies, Mr. Hoyle being apparently inelined to give as mueh

importanee as I did to the seeming constancy of the number

of rings in the female.

While these eonsiderations, then, tend to the union of the

speeies P. protelis with P. polyzonum^ the faet that the two

animals, the small earnivore and the voracious snake, do

live in the same area gives a clenching force which, to my

mind, is almost irresistible.

XII. — The Causes of Variation. By KOMYN Hitchcock’^.

The reeent studies of Dr. W. B. Carpenter upon Orhitolites\

are of special interest, owing to the remarkable manner in

which the stages of variation and development have been

traeed. The monograph by Dr. Carpenter, published in the

Keports of the ^ Challenger ’ Expedition, was the subject

of some remarks reeently made by the writer before the

Biological Society of Washington, in which an effort was

made to explain how such a simple sarcode organism as the

animal Orhitolites has been led to produce a shell of complex

form. Dr. Carpenter regards it as the expression of a not

understood progressive tendency along a definite line towards

a higher speeialized type of strueture in the ealcareous fabric.”

This, however, is merely a statement of the facts observed,

and in no wise assists in their explanation. Elsewhere it

may be gathered from the author’s words that he regards the

* From the ‘American Journal of Science’ for Ji% 1884, pp. 49-52.

t Phil. Trans, part ii. (1883).

Ann. & Mag. N. Hist. Ser, 5. Vol. xiv.

8

Mr. Li. Hitchcock on the

\)i

complex shell as significant of a plan so definite and obvious

as to exclude the notion of ^ casuaV or ^ aimless ’ variation.”

The facts seem capable of a somewhat different interpreta-

tion, which seems more in accord with our present knowledge

of simple organisms, and quite sustaining the views of

Darwin that plan,” in the sense used by Dr. Carpenter,

should be superfluous. For if there be an inherent tendency

to variation among these organisms, as Dr. Carpenter seems

to believe, how do we explain the persistence of the original

Orbitoline type, 0. tenuissima^ Biologists seek to discover

the causes of variations which they observe ; but it seems not

less important that the persistence of types should also be

cx})lained. 0. tenuissima is a very ancient species, and

surely any inherent tendency to change would have mani-

fested itself during the long period of its existence, even under

unfavourable conditions.

Tlic observations I have to offer may be said to relate

entirely to change of environment ; but their tendency is to

demonstrate that the changes observed in the shells of this

family are not due to any inherent tendency resulting in a

definite plan, but that they are due to causes easily understood.

It is far from my intention to deny a definite plan of growth

to these organisms. But plan of growth does not imply that

there have been causes acting within the organism — special

tendencies of the protoplasm toward higher structure. It

seems to be such an assumption that has led Dr. Carj)enter to

speak of a ^Ciot understood” progressive tendency, &c. Jn

my opinion the causes of such progression as can be observed

are easily understood ; and the plan of growth becomes a

natural consequence of these causes, which are purely physio-

logical, and independent of any supposed tendency to varia-

tion. While Dr. Carpenter, on the one hand, seems to regard

variation as due to an inherent tendency of the protoplasmic

body, the writer, on the other hand, attributes it entirely to

the more or less favourable conditions of life of the different

species. Moreover, I am quite unable to understand how any

inherent tendency to variation impressed upon tlie sarcode

could fail to find expression in some differentiation of the

sarcode, which in the cases in question has not been ob-

served.

The same view seems to be held by 0. Schmidt, wlio, in

his ^ Grundziige einer S})ongien~Fauna des Atlantischen

Gebietes,’ alludes to Dr. Carpenter’s previous studies, and

compares the changes observed in the Sponges and Forami-

nifera. He says the changes in tlic latter are found in the

general habit of tlie form and the variable grouping of

Causes of Variation.

95

the chamber-systems, while among the Sponges the variation

is in tlie microscopic detail. One may speak of the micro-

scopic form of Foraniinifera, but not of microscopic elements.”

Tlie complexity of the shell is merely in the multiplicity of

chambers and the manner of their intercommunication. The

process of growth, even in the complex 0. complanata^ is in

all respects identical with that in other species, and in no

essential feature differs from that of Peneroplis. What Dr.

Carpenter designates as a higher specialized type of struc-

ture ” does not represent an advanced degree of specialization

in any part ; nor can we discover any advantage to the organ-

ism arising therefrom. It is true there is an advance in com-

plexity ; but unless accompanying this there is an evolution

in function, or unless it results from some effort of adaptation

which confers some benefit upon the organism, it seems not

proper to regard complexity of shell-structure as a proof of

biological advancement.

Seeking for an explanation of the cause of the increased

complexity of shell-structure, so beautifully illustrated in the

Milioline family, the writer was led to the conclusion that it is

entirely due to the favourable conditions of life and the abun-

dance of food available. It is true, as already said, this may

be regarded as a mere statement of the influence of environ-

ment causing variation ; but a careful consideration of the

subject will show that there is a broad distinction between

environment as a cause of variation and adaptation to environ-

ment j for in this case we are unable to perceive any benefit

to the organisms arising from their adaptation to changed

conditions.

If it be said we can seldom discover the benefits supposed

to be derived from adaptation, it may be answered that it is

usually possible to infer how the changes observed may prove

beneficial. In the case under consideration, however, an

examination of the changes that have taken place does not

indicate any possible benefit to the organism. The multipli-

cation of chamberlets necessitates very intimate intercommu-

nication for the transference of food and the continuation of

the processes of life. The organism is not thereby better

adapted to its surroundings, but is made more dependent

for its existence upon the continuance of the favourable

conditions under which it has developed. The advance in

complexity — the multiplication of chamberlets — would only

be possible under the most favourable conditions, for all the

nutriment received by the interior segments must be col-

lected by the sarcode at the margin of the shell, and the

necessary food could only be obtained where the supply was

8*

DO Mr. II. Hitchcock on the Causes of Variation.

abundant. It may be conceived tliat if 0. complanata were

placed in situations less favourable as regards food it would

die of starvation, owing to the quantity of inner sarcode

requiring nourishment, while 0. tenuissima needs only more

favourable conditions as regards food and, perhaps, tempera-

ture to become as highly complex in structure as the last-

mentioned species. As a further proof of the influence of

environment leading to changes which cannot be regarded as

speeial adaptations, in the usual meaning of the word, the

forms of 0. complanata found on Fiji reef are especially

characterized by thick plicated margins, as though growth .

proceeded with too great rapidity to produee symmetrical

disks, and these forms are associated with the largest repre-

sentatives of the species.

The distinction above referred to seems an important one,

which, if it has already been recognized, has not been promi-

nently brought forward in the writings with which I am

familiar. Before the Biological Society the subject was

briefly considered in the following words : —

“ Begarding the subject from this point of view, we are

led to examine more closely the relations between the spiral

and the eyelical methods of growth. Their intimate relation

is only noticeable when we observe how one has been derived

from the other. When the spiral growth of OrhicuUna pro-

duces a complete circular disk, further spiral growth becomes

impossible ; and if we concede that the extrusion of the

sarcode to form successive chamberlets is due to nutrition and

growth, the cyclical plan then becomes a necessity. In this

way it may be supposed cyclical growth originated, purely a

’ result of nutrition, not by adaptation to environment, but as

a result of it ; not because such growth is or ever was better

adapted to the conditions of life.

We find here a steady course of variation a result of

physiological processes, independent of those external causes

to which we are accustomed to attribute such changes. Tliese

variations, as successively produced, have been perpetuated

through inheritance, until the plan of growth has, in some

species, totally changed. Herein, therefore, we may find an

indication of how the plan of growth originated, and a sugges-

tion that the inscrutable laws which govern the progress of

evolution may each have beginnings equally simple, and not

beyond the range of human insight to discover. Evolution in

this case seems not to be a result of a definite plan of growth,

but the plan of growth is the result of physiological pro-

cesses. However great and important the influences of

environment and selection may have been in the production

1)7

On the Veriehrate Zoology of Persia.

of genera and speeies, perhaps tlie attraetiveness qf the idea

and the ease with whieh it enables us to dimly understand

many biogenetie problems permits us to lose sight of other

iiiHuenees more obseure, but of equal importanee in the history

of life.”

ddiis view of the subjeet seems to derive still further

support from the geographieal and bathymetrieal distribution

of the speeies. Without entering into a lengthy diseussion

of this part of the subjeet, it may be said that as a rule the

more complex species are found in the warmer waters under

conditions most favourable to the activity of nutritive pro-

cesses. As an example, the very large specimens of

O. complanata from Fiji reef may be taken. On the other

hand, the ancestral form 0. tenuissima still inhabits the colder

and deeper waters, retaining the simple characters of its earliest

known condition.

XIII. — Additions to the present Knowledge of the Vertebrate

Zoology of Persia. By James A. Murray.

Since the publication of Mr. Blanford’s valuable work on

the Zoology of Persia (1876), giving a complete list of the

animals inhabiting that country, nothing, I believe, has been

published as an additional contribution, except a single paper

in the Proc. Zool. Soc. for 1881, which added five species to

the already large list of reptiles ; these are Againa persica^

Scincus conirostidsj Hydrophis temporalis^ Catachlcena dia-

demaj and Hydrophis cyanocincta^ the first three being newly

described species.

The Kurraehee Museum, having now rather an extensive

collection of Mammals, Birds, and Beptiles from Eastern

Persia — very kindly made for the institution by Mr. W. D.

Gumming, of the Persian Telegraph, during the past three

years — and having also acquired a collection, comprising thirty-

six species of Beptiles and seven Mammals (also from Persia)

— made, it is said, by a member of some foreign exploring

commission in 1876-77 — I am enabled, after careful examina-

tion of these materials, to add a few more species to the

existing knowledge of the Vertebrate fauna of the country.

For the collection said to be made by a member of some

foreign commission, the institution is indebted to Mr. Poss-

man, also of the Persian Telegraph. Although this collection

dates as far back as 1876-77 the specimens are in an excel-

08

Mr. J. A. Murray on the

lent state of preservation ; eaeli specimen has been carefully

labelled, giving the date, year, and localities of capture, the

latter being chiefly Bushirc, Tanjistan, and Charbagh, near

Bushirc.

Among Mammals there is nothing new, but a few species,

of the occurrence of which Mr. Blanford seemed to doubt and

which arc comprised in the collection, are noted below.

1. Bhinolophus ferrum-equinumy Sclireb.

ltJmwh2)1ms ferriim-eqtiinmn, Sclireb., lUf. E. Pers, ii. p. 19.

IcT: Bushire, 26. 11. 83.

2. Ursus tliihetanuSy F. Cuv.

TJrms, sp. ?, Blf. E. Pers. p. 47.

Vrsus (jedrosianus, Blf. J. A. S. B. xlvi. pt. 2, p. 317 5 P. A. S. B. 1870,

p. 4.'

Mr. Blanford records this from Bcloochistan, on the assu-

rance of the natives of the country. Major Mockler very

kindly procured for me three skins with skulls of the animal

inhabiting the Bcloochistan hills. These with two other skins

from the Sind hills were those of Lrsus tliibetanus. Tliere

is now a live specimen in the Kurrachee Zoological Gardens,

from the Sind hills.

3. DelpMnus plumheusy Duss.

Not recorded in Zool. E. Pers. The Kurrachee Museum

has two skulls from Lingah.

4. Bipus Blanfordiy sp. nov.

Diqms macrotarsus ?, Wagner, Blf. E. Pers. p. 74.

1 J', 1 ? adult (pregnant), and two adolescent: Bushire,

July 1882.

Four juv. : Tangak, May 1877.

One adolescent, Tanjistan, June 1877.

These agree in every particular with Mr. Blanford’s de-

scription. The long black tuft of hair beneath the hind feet

is very characteristic, also the broad whitish band across the

upper part of the thigh and the rufescent fawn thigh-patch.

These dilferences being constant, the Persian form must be

considered distinct ; and Mr. Blanford having first character-

ized the species, I have much pleasure in associating his name

with it.

I must, however, add the following particulars to the de-

scription given by Mr. Blanford : — Mamma3 8— one pair

under the throat in front of the fore legs, one pair behind

Verlehvate Zoology of Persia.

tlic fore legs, and two inguinal pairs. In adults the taii is

unicolorous up to the peneil of hairs and of a pale isahelline

colour, while the black portion of the pencil in adolescents is

a dark brown, tinged slightly with rufescent ; the back is

mncli darker in colour in adults, owing to the bases of the

hairs, which are of a dark ash-colour, showing through ,* claws

horny. Molars of upper jaw all biplicate, inside and out : of

lower jaw, 1st biplicate on both sides, 2nd triplicate without

and biplicate within, 3rd higMcate outside and rounded within.

The following'are dimensions of an adult pregnant female

and an adolescent male: —

Adoles-

Adult 5, cent cJ.

in. in.

Length from tip of nose to root of tail 6*25 3*75

Ditto of tail 7*2 6*0

Ditto of pencil at end of tail 0*75 0*56

Total length 13*2 10*31

Height of ear from upper margin of outer conch 0*7 5 0*5

Ditto from base of skull 0*82

Breadth of ear laid flat 0*62 0*44

Length of tarsus, foot, and claws 2*62 2*12

Longest whisker (reaching to beyond the axil

of the thigh) 3*25 3*0

Skull, from upper edge oi foramen magnum to

end of nasal bones 1*25

Skull, from lower edge of ditto to front of upper

incisors 1*12

Breadth across hinder part of zygomatic arches 0*87

Ditto, between orbits 0*5

Length of lower jaw from condyle to inner

base of incisors 0*72

Teeth-line, upper jaw 0*18

Teeth-line, lower jaw 0*18

Space belM'^een inner edge of 1st molars 0*18

Across tympanic bones 0*03

5. Dipus Lqftusi.

Dipm Loftusi, Blf. East Pers. p. 75.

1 d' Bushire, 6. 12. 83.

1 $ juv. : Tanjistan, Nov. 1876.

1 ? and four foetal young : Nov. 1876.

The foetal young of this species have the whiskers fairly

well developed, and the tail is less than half the length of the

liead and body.

Length of largest specimen (spirit) : —

in.

Head and body. 5*37

Tail 6*0

Tarsus, foot, and claws 1*95

100

Mr. J. A. Murray on the

6. Lagoniys rufescens^ Gray.

Lagomys rufescens, Blf. East Pers. p. 83.

1 ^ : Bushire, 4. 1. 84.

This agrees well with the description of it by Mr. Bkn-

ford, except that the chin, throat, and underparts are a silky

yellowish white, as are also the fore and hind feet and the

soles of the feet. The longest whisker is white at its extre-

mity, and the lower series of 4-5 white througliout. Tiiis is

recorded from Afghanistan, Northern Persia, and Mesopotamia,

but not from S.E. Persia.

Length 7*5 inches.

Among Birds I have to add : —

1. Falco peregrinatorj Sund.

Falco peregrinator, Sund., Elf. East Pers. p. 103.

Four live birds were netted at Bushire and sent to me for

the Eev. Mr. Watson, who trained them for the quarry. All

belonged to the atriceps type.

2. Circus macruruSy Gmel.

2 J* : Bushire,

1 d : Eao, Shat-el-Arab.

3. Hyjwcolius ampelinuSy Bp.

1 , 1 $ : Bushire, 13. 9. 83.

According to Mr. Gumming this species passes throiigli

Bushire in November. Its range extends N.E. to Eao, on

the Shat-el-Arab, as far as at present known, soutliward to

Sind. It breeds in the country. Mr. Gumming and Mr.

Betts have taken the eggs at Eao. This is not recorded from

E. Persia by Mr. Blanford.

4. Ardeola leucoptera.

Several specimens from Bushire. Breeds in June and

July.

5. Sterna fuliginosa^ Gmel.

2 ? , Bushire, in breeding-plumage, received witli ten

eggs ; and 1 7 taken on the Astola Island, S. Persia.

6- Pelecanus onocrotalus and P. crispus.

Both common in the Persian Gulf. The first has been

found breeding at Eao, or rather 50 miles west, on a mud

island surrounded by a large marsli, wljciicc Mr, (himmiiig

obtained live eggs.

Vertebrate Zoolofjy of Persia.

101

7. Phamicopterus minor j GcofFr.

A male sent to me from Lingali by a Mr. Belelier, a pas-

senger to Bussorali, with a note to the eflfeet tliat it was shot

out of a floek of larger ones^ evidently P. antiquorum.

Among Reptiles the eolleetion eontains several speeies not

reeorded from Persia.

1. Stellio nuptuSj ys^x. fuscus.

StelHo nuptus, var. fuscus, Blf . East Pers. p. 320.

2^,1 ? : Bushire, July 1876.

Mr. Blanford reeords this from Jalk and Kalagan, in

Belooehistan, and not from Persia.

2. Centrotrachelus Asmussi^.

Centrotrachelus Asmussi, Straucli, Blf. East Pers. p. 337.

3 Bushire, July 1883.

Largest speeimen 23’5 inehes in length. I have had a live

speeimen sinee August 1882, still in exeellent eondition. The

animal is extremely lively during the hot hours of the day up

to 4 o’eloek ; after this hour it sleeps soundly, curling itself

in a corner of the box in which it is kept. Its means of

defence is the spiny tail it possesses, which it lashes like a

whip when disturbed. It is extremely fond of having cold

water thrown on its body ; it then appears much pleased,

standing high on its fore legs, with head erect, turning it

round, upwards, and looking with each eye, and extending

the loose skin of its body to double the usual size. Whether

the animal burrows for itself or occupies the burrows of field-

rats &c. is a question, as, although it has a foot of soft sand

in the box (without a bottom), on the bare ground, it has

never yet attempted to burrow.

3. Hemidaetylus Cocteaui^ D. et B.

Three : Charbar, Beloochistan, June 1880.

Two : Charbagh, near Bushire, August 1876.

One : Bushire, August 1883.

Four: Tanjistan, July 1877.

The non-entry of this species by Mr. Blanford is evidently

an omission.

^ ? a loricatus, Blf. ^ Eastern Persia/ ii. p. 340, described from the

neighbourhood of Bushire.

102

Mr. J. A. Murray on the

4. Gymnodactylus hrevi'pes.

Gymnodacfylus hrevipes, Blf. East Pcrs. p. 344.

Tlircc : Busliire, Oct. 1883.

One : Tanjistan, Aug. 1876.

T1 lis has been recorded from Aptar, near Bampur, in Be-

loocliistan. Mr. Cumming’s collection from Busliire contains

well-marked specimens of this species.

5. Gymnodactylm scaher^ Biipp.

Thirteen : Bushire, June, July, August, 1883.

Seven: Tanjistan, September 1876.

In form the counterpart of Gymnodactylus petrensisy

Murray (Vert. Zool. Sind, p. 362).

Kostral broader than high and cleft above. Upper

labials 10-12 ; lower labials 8-10. Pupil vertical ; first

labial and three small shields behind rather smaller than

those covering the muzzle ; interorbital space and occiput

with large conical tubercles interspersed, a few also on

the muzzle and a line of 3-4 in front of each eye. Two

pairs of chin- shields, the first largest and in contact. Back

covered with granular scales and sharply-keeled trihedral

tubercles, the latter as^ large as or slightly larger than

the vertical ear-opening, and arranged across the middle of

the trunk in 14 longitudinal rows ; between the hind limbs

the number is six. The tubereles on the sides of the body

are rather smaller and subcarinate. Scales across the middle

of the abdomen in 18-20 rows. Preanal pores 5-6. Outer

surface of limbs with large trihedral tubercles. A pair of

tubercles on each side of the sacral region. The fore limb

laid forward reaches the end of the snout ; laid back it reaches

the axil of the hind limb. The hind limb laid forward

extends beyond the axil of the fore limb. Tail verticillate,

with three rows of sharply- keeled trihedral tubercles on each

side to within an inch of the tip, beyond which it is covered

with irregularly- arranged imbricate scales. Subcaudals

distinct, single, about 44-54 ; a few of the anterior ones bifid.

Length 4‘5 to 5 inches, of which the tail is 2*5 to 2*75.

Colour greyish brown, with three longitudinal rows of

dusky subquadrate spots on the back ; in some specimens one

more row of rather indistinct spots on each side. Tail with

10-12 dark bands above.

Hah, Bushire and Tanjistan, in Persia; Fao, in Southern

Mesopotamia, at the head of the Persian Gulf, on the banks

of the Shat-cl-Arab, and Charbar, -in Beloochistan.

Vertebrate Zoology of Persia. 103

Collected by Mr. W. D. Cumming, to whom I am in-

debted for a large collection of reptiles, tish, &c. from Bushirc

and Fao, in Southern Mesopotamia.

The synoptical table below will show the differences between

this species and the other allied forms of Gymnodactylus.

6. Pristurus rupestris.

Pristurus rupestris, Blf, East Pers. p. 850.

Seventeen specimens from Bushire and Tanjistan, 1876-77.

7. Ceramodactylus Borice.

Ceramodactylus Borice^ Blf. East Pers. p. 353.

I have seven specimens of this lizard from Tanjistan, nearly

300 miles further north-east of Bunder Abbas, where Marquis

Doria’s single specimen was obtained.

8. Ceramodactylus affinis.

General form of Ceramodactylus Borice^ but of a more

robust habit. The nostril is placed rather behind the outer

hind angle of the rostral, instead of immediately above and

between the suture of the rostral and first labial. The three

shields behind the nostril are flat, and not distinctly swollen ^

as in C. Borice. The mental is rather of a different shape,

having slightly concave instead of straight sides, and a very

* In a continuous line on both tliiglis.

104

Mr. A. Murray on the

convex hind margin. Upper labials 11, with a number of

smaller ones behind, searcely larger than the granular scales

of the tympanic region. Tail not attenuate, as in G. Doriwj

but ends rather abruptly ] it is three fifths the length of the

head and body, while that of G. Dorice is nearly the length

of the head and body. The head is short and thick, and

presents a very different aspect from that of G. Dorice] its

height is about a fifth less than its greatest breadth, or a fifth

less than that of G. Dorice^ both in height and breadth. The

fore and hind limbs are very mueh stouter and shorter than

those of G. Dorice. The fore limb laid forward, the tips of

the fingers reaeh only to the end of the snout, while in G.

Dorice the tips of the fingers extend beyond the snout by the

length of the foot. The hind limb laid forward does not

extend as far as the axil of the fore limb, while that of G.

Dorice reaehes nearly to the ear-opening. Fingers and toes

mueh shorter and more robust.

Length, head and body 2*62 inches, tail 1*37.

Colour : instead of the spotted character of the markings of

G. Dorice,^ this species is banded, the bands being rather

curved_, with the concavity in front. There are four bands,

one on the oeciput, one behind the shoulder, another on mid-

trunk, and the fourth on the loins. The tail has 7-8 dark

bands. The ground-colour is apparently ochraceous. There

are 2-3 dark oblique streaks below the eye along the labials.

Sides of the body slightly darker than the back.

Hah. Tanjistan, in Persia. Two specimens among seven

of G. Dorice j collected in 1877.

9. Rhagerrhis productuSj Ptrs.

Head moderate, distinct from neck. Snout rather de-

pressed and produced beyond the mental. Tail short * anal

bifid. Eyes rather large; pupil circular; rostral produced,

obtuse in front, reverted on to the uj)per surface of the head,

105

Vertebrate Zoology of Persia.

and forming a triangular suture witli the prefrontals ; below,

the rostral is deeply eoneave. Nasals 2. Nostrils situated

in the anterior portion of the postnasal, whieh is reeeived in

the hind, nearly quadrangular, eavity of the prenasal. Loreal

1, square. Preoeular 1, reverted on to the upper surface of

the head, but not touching the vertical. Postoculars 2.

Upper labials 8 ; lower labials 11, 6 in increasing series from

the mental, and 4-5 other smaller ones about the size of the

adjoining scales. Sixth upper labial the largest ; the fifth

under the eye. Mental narrow, angular and pointed behind,

slightly convex in front. Mental groove distinct. Two

])airs of subequal chin-shields. Two pairs of frontals ; ver-

tical elongate, obtusely triangular in front and triangular

behind, where it is received in the triangular concavity of the

occipitals. Occipitals rounded behind, convex laterally, and

longer than their greatest width ; temporals variable, gene-

rally 2-3 alongside the occipitals, and in some split into small

plates. Scales in 17 rows, all, even the head-shields,

minutely punctulated with brown (only seen through a

Coddington lens). Ventrals slightly angulated.

Colour (spirit-specimens) ochraceous, with equally distri-

buted dusky spots in five rather oblique series. A small

dark brown spot under the eye ; another less distinct above

it (not present in all specimens), a third obliquely behind the

eye, followed by a large temporal spot, also oblique, and

extending beyond the last labial. Lower parts unspotted,

pale yellow.

Hah. Bushire, 1 , 2 ? ; Tanjistan, 3 (^ , 2 $ .

Length of largest specimen 46 inches, of which the tail is

7 inches ; length of smallest specimen 15 inches, of which the

tail is 2*5 inches.

10. Bufo viridiSj Laur.

The following is a description of specimens from Bushire : —

Crown of the head flat, smooth, and devoid of osseous

ridges ; orbitals elevated and covered with liorny-tipped

tubercles ; interorbital space equal to the width of the eyelid.

Hind crown, back, and the upper surface of fore and hind

limbs covered with rather close-set horny-tipped tubercles.

Parotoids oblong, flattened above about their middle, and

separated from each eye by a deep groove. Length of each

parotoid equal to the distance between its front edge and the

nostril. Tympanum distinct, about one half the size of the

eye. Under surface of the body smooth, without any trace

of tubercles. Fore feet with a large mesial palmar pad and a

smaller one at base of first finger. Laid side by side the

10()

Mr. J. A. Murray on the

first finger is very slightly longer than theseeond and as long

as the fourth ; third finger longest. Tarsus with a mesial

and lateral longitudinal row of distant tubereles, and a euta-

neous fold on the inner side extending to the inner metatarsal

tubercle. The tubercle on the outer edge scarcely so promi-

nent as the inner one, which is elongate. Soles of the hind

feet faintly tuberculate. Toes half-webbed ; the tip of the

first reaches the second joint of the second toe. Hind limb

long ; laid forward alongside the body the metatarsal tubercle

reaches the eye, and one half of the foot extends beyond the

snout.

Length 3 inches ; hind limb from anus to tip of second

finger 4*2 inches.

Colour yellowish, a dark spot on each eyelid; another

oblique one from the hind edge of the eye to the tympanum,

and a third very small one on each nostril ; fore and hind

legs with 2-4 transverse blotches on their upper surface.

Sides of the first and upper surface of first and second fingers

black. Under surface pale yellowish. Tips of toes slightly

swollen and of a brownish colour.

These specimens come near to Bufo oUvaceuSy Blanf. (East

Pers. p. 434, pi. xxviii. fig. 3), but differ from it by having the

dorsal surface closely set with horny-tipped tubercles, a flat

instead of a concave crown, by its under surface being smooth

and not tuberculate, and by its shorter hind limbs. From B,

vulgaris it is distinguished by its longer hind limbs, having

a cutaneous tarsal fold, a distinct tympanum, and no dark

band below the parotoid.

XIV. — Additions to the Reptiliayi Fauna of Sind.

By James A. Mureay.

Since the publication of my work on the ^ Vertebrate Zoology

of Sind,’ a collated descriptive account of all the species of

mammals, birds, and reptiles (including several new species)

known to inhabit the province, some little interest appears

to have been aroused in zoological inquiries, which has

resulted in the Kurrachee Museum acquiring several collec-

tions of reptiles from hitherto unknown localities in Upper

Sind.

Among these are four species from the barren sandy wastes

of the frontier districts, collected by my indefatigable corre-

107

Hep til Ian Fauna of Hind.

s])oiulcnt Mr. F. Glcaclow, of the Forest Department, three of

wliieh 1 believe are undescribed forms. These are : —

1. Melanochelys pictus.

Head two thirds as broad as long, its greatest length 3

inclics. It is covered with skin, divided into plates ; a long

central one above a single broad frontal ; a superciliary on

each side, and a small subtriangular plate behind in suture

with tlie sides of the central plate. Temples covered with

numerous irregular-shaped plates. Upper jaw with a small

festoon on each side, the groove in the middle of the jaw

rather deep. A plate in front of the eye in suture with the

sides of the frontal ,* another nearly as large under the orbit,

and a third about twice the size of the latter behind the eye.

Anterior half of neck covered with small subimbricate plates

in transverse series. Shell oblong-ovate, elevated, much

arched, nearly half as high as long, nodosely tricarinate, the

costal Carina being much nearer the vertebral carina than the

marginal plates. Length of shell over curves 14 inches;

breadth over vertebrals 11*75 inches. The sternum is bent

upwards from the suture of the pectorals with the postgulars ;

greatest length of sternum to point of furcate projection of

anal plates 12 inches. Anals deeply notched posteriorly, the

distance between the projecting ends being 1*5 inch and the

depth of notch 1*37 inch. Width of sternum at axillary

incision 6 inches, at inguinal 4*75. Gulars together broader

than long, their hind margin received 'into the subtriangular

concavity in front of the postgulars, which are as broad as

long. Pectorals very narrow, each 3x 1*5 inches, the suture

between them about equal in length to that of the postgulars

and slightly more than half of that of the abdominals. Abdo-

minals nearly rectangular, winged beyond the inguinal

incisions, and forming a suture on each side with the inguinal

plate and sixth marginal. Postabdominals longer than broad,

the length of their suture together slightly less than that of

the abdominals ; transverse sutures of postgulars with abdo-

minals and abdominals with postabdominals straight ,* suture

of postabdominals with anals concave; the suture together of

the anal plates is shorter than the suture of a single one with

the postabdominal. Nuchal plate oblong (0*75x0*5 inch).

Vertebrals hexagonal ; first somewhat bell-shaped, convex in

front, straight behind (except a concavity mesially to receive

an apophysis of the second vertebral), sinuately concave on

each side in its anterior half and convex lower down. Second

and third vertebrals hexagonal, as broad as long ; fourth

108

Mr. J. A. Murray on the

similar, but concave behind in suture with the convex front

of the fifth, which is about twice the size of the other verte-

brals, broader than long (2*37 x 3*37 inches), with the sides

sloping outward to the last marginal ; its suture with the

caudals is straight. Caudal notched, the suture of both plates

a little more than half the length of the fifth vertebral. Tail

short, 1*5 inch in length. Costals large, four in number,

nodosely carinate on their upper margin, but not extending,

or scarcely seen on the fifth vertebral. The first costal is

subtriangular and largest, convex in front, in suture with one

half of the first and the whole of the upper margin of the

second, third, and fourth marginals ; second costal in suture

with the fifth and sixth marginals, its greatest breadth about

two thirds its greatest length (4x 2*75 inches) ; third costal

in suture with the seventh, eighth, and anterior third of the

ninth ; it is nearly the size of the second in length and

breadth ; fourth costal smallest and forming sutures with the

posterior two thirds of the ninth marginal, entire tenth,

mesially the apex of the eleventh, and the sides of the fifth

vertebral. Marginals variable in size, not serrated poste-

riorly, but slightly dilated at the eighth, ninth, and tenth.

Feet anteriorly covered with imbricate scutes, posteriorly, or

higher up, subimbricate, the scutes much smaller. Toes

short, strong, and webbed to the claws ; they are covered with

annular scute-like plates. Claws strong and hooked. Sides

of the legs fringed with large scales. Colours : all the scales

on the tarsi and feet with a yellow spot. Head black, with

large spots, blotches, and streaks of yellow ; a patch on each

side of the snout, also on each side of the nostrils ; one under

each eye, another at the maxilla on the labial margin, and

two behind each eye. There are also some large blotches on

the tympanic and temporal regions, and three on each side of

the lower jaw. Shell olive or greenish brown, the marginals,

lower part of costals, and vertebrals with pale yellowish

blotches and streaks of irregular shape. Sternum pale

yellow, with linear transverse lines, very close together on the

abdominal plates, and forming a large patch. The gulars

and postgulars are not marked.

Mr. Gleadow, obtained this species in the Sind

in the Kushmore Talooka, Upper Sind.

It differs from all the described forms of Melanochelys^ first

by its greater size, next by the size and shape of the vertebrals

and costals, and lastly by the markings of the shell and the

spotted character of the head and feet.

Reptiliati Fauna of Sind,

109

2. llemidactylus Icushmorensis,

Head rather depressed. Rostral grooved above, slightly

wider than high. Upper labials 10 ; lower labials 8. d.V'o

pairs of chin-shields, the first only in contact. Muzzle covered

with granular scales. Nostrils between the rostral, first

labial and three small shields behind about equal in size to

tliose covering the muzzle. Grown of the head interspersed

with numerous rounded tubercles. Back with rounded

tubercles arranged in twenty-two longitudinal series across

the middle of the body; a few tubercles between the hind

limbs are subtrihedral. Tail verticillate, each verticil armed

laterally with three rows of rather elongate subtrihedral

tubercles, except on its posterior third, where they are re-

placed by imbricate scales. Fore and hind limbs on their

upper surface studded with round tubercles. Toes covered

with imbricate scales. Claw on thumb well developed.

Scales on the throat about one third the size of those on the

abdomen, across the middle of which they are arranged in

longitudinal series ; the anterior half irregularly and

minutely 1-3 crenulate, less conspicuous on the posterior

lialf.

Femoral pores 10-12 on each thigh. Under surface of

tarsi covered with large imbricate scales. Subcaudals single,

44-46. Middle toe with six pairs of plates and an odd one

at each end.

Colours neutral grey or brown, with three rows of squarish

dark blotches, forming either longitudinal or obliquely trans-

verse interrupted bands ; a few smaller spots on the sides. A

dark streak through the eye with a pale line above it. Scales

on the under surface of the body freckled with 1-3 dark

spots ; many, especially on forward part of body, without

them. Tail with 14-15 dark bands. Pupil vertical.

Hob. Upper Sind, Kushmore and Thool Talookas.

Two only of six specimens with unreproduced tails. Length

4 to 4*25 inches. Type from Bhaner, Upper Sind frontier.

Differs from all the other species of the genus in having a

greater number of dorsal tubercles, also femoral pores, except

//. GleoAowi^ and fewer abdominal scutes, except H, triedrusy

and in having rounded tubercles.

The following Table will sufficiently show the differences

between it and the other allied species of the genus : —

Ann, <b Mag, N, Hist. Ser. 5. Fol. xiv.

9

no

On the Iie2)tiUan Fauna of Sind.

3. Gymnodactylus scaherj Riipp.

Two specimens collected at Sukkur and one at Laki, be-

tween Sliikarpoor and Sukkur, by Mr. F. Gleadow (see my

preceding paper, “ On Additions to tlie Fauna of Persia,” for

description of this species).

It YQ^\iiQ,QF^Gymnodactylus 2)et't'Gnsis in Upper Sind.

4. A contiophis paradoxa.

AcontioFiis paradox a y Giiiitli. Proc. Zoob Soc. 1875, p. 232.

This snake was known from a single specimen only, the

loeality of which has only now been ascertained. I count

ventrais 180, and subcaudals 52. My specimens are greyish

brown ; a dark line from behind the eye to the nape, a

subovate dark patch on the occiput and a border along the

margins of the occipitals from the anterior half of the super-

ciliaries. A dorsal series of quadrangular dark spots with

white interspaces, nearly of the same width to within an inch

of the end of the tail, where they become smaller and more

faint, and gradually disappear.

Total length of larger of two specimens 14‘25 inches, of

which the tail is 2T0 inches.

* From Giiutlier and Tlieobald s works on Reptiles of Rr. Iiid.

t In a nearly continuous line.

Oil a new Species o/* Lycopoditcs, Ooldenherg. Ill

Hah. Upper Sind, Tliool Talooka, at Zimgipoor, frontier

districts.

Mr. F. Gleadow states tliat the two specimens of this snake

were dug up from depressions a quarter-mile apart in a con-

siderable area of blown sand forming hillocks 20 to 30 feet

high.

XV. — On a new Species of Lycopodites, Qoldenherg (L,

Stockii) the Galciferous Sandstone Series of Scotland.

By R. Kidston, F.G.S.

[Plate V.]

The genus Lycopodites., as originally employed by Brong-

niart* and most of the older writers, did not contain any plant

which was really entitled to the name, in so far as it was

used to infer their closer affinity to the recent Lycopodium

than that held by the genus Lepidodendron ; and Brongniart,

in his later writings, discarded his genus Lycopodites^ as sub-

sequent investigations had shown him that his original view

of the plants he included in it was founded on an erroneous

notion of their true nature f-

Hence, when Goldenberg resuscitated the genus Lycopo-

dites^ it was used by him in an entirely different sense from

that to which it had been applied by previous writers, and in

fact was a new genus though under an old name.

To enable us more clearly to appreciate the light in which

Goldenberg regarded his genus Lycopodites^ I quote his intro-

ductory remarks regarding it.

Lycopodites, Goldenberg. 1855 J.

Branches with leaves placed spirally or in verticils,

orangia placed in the axils of the leaves or forming ter-

minal cones.”

In the genus Lycopodites we place the true herbaceous

Lycopods of former ages, which agree in all essential points

so exactly with recent Lycopods that, at the most, they can

only be regarded as a subdivision of the genus Lycopodium.

The fossil plants included by Brongniart and others under

this name are probably only young twigs of Lepidodendron

* ‘ Prodrome,’ p. 83 (1828), and ^ Classification des vegetaux fossiles,’

p. 46 (1822).

t Tableau d. genres de Veget. foss. p. 40 (1849).

I ‘ Flora Sarajpontana Fossilis,’ Heft i. pp. 9, 10 (1855).

9^

L12 Mr. R. Kidston on a neiv Syecies of

or Conifeise. This view, which we here accept, has been

})ointed out by Brongniart in his last work (‘ Tableau des

genres de vegdteaux fossiles, considdres sous le point de vue

de leur classification botanique et de leur distribution geolo-

gique ’), and is thus stated by him : —

“ ‘ The plants really analogous to the recent Lycopods are

very few in number in the fossil s'tate.

I do not know even one which, by its dimensions and

the disposition of its leaves, may be compared with certainty

to the species of the genus Lycopodium properly so called ;

the greater part of the plants which I have designated or

which have been indicated as Lycopodites^ are probably either

the upper portions of young branches of Lepidodendron or the

brandies of Conifers.

“ ‘ Thus the greater part of the Lycopodites with dichotomous

branches from the Carboniferous formation appear to belong

to the first class ; those species with distichous pinnate

branches evidently belong to Conifers of the genus WalcMa.

The greater part of the species from more recent forma-

tions, as the Lias and Oolite, belong to this latter group ; such

are in particular Lycopodites Williamsonis and patens.

^ Among these there is, however, one specie'^, which has

all the characters of a Lycopod, or perhaps more the character

of the genus Beiaginella. This, the Lycopodites falcatusj L.

& II. has lately been rightly separated, and from its delicate

and dichotomous branches, apparently distichous leaves (but

which are probably opposite and unequal), has all the appear-

ance and essential characters of the numerous species of the

genus Beiaginella.

^ I know no species which resembles the true Lycopods, as

at present defined, nor the genus Tmesipterisd

We have therefore in Lycopodites the addition of a new

genus of fossil plants to the Carboniferous flora, which,

according to Brongniart, is at present only known by one species

from the Oolitic formation of Englandf. For the plant-remains

which hitherto have been described and figured from the

Carboniferous formation under the name of Lycopodites

Bronniij longifolius^ &c., belong, as we have seen, to quite

other genera of plants, as they do not exhibit any of those

points which form the principal characteristics of club-mosses.

“ Many years ago 1 found plant-impressions in the Carbo-

niferous rocks of this neighbourhood (Saarbriick) which, in

the character of their growth, showed a great similarity to our

herbaceous Lycopods. At the Meeting of the Natur-

* ‘ Fossil Flora/ pi. Ixi.

t Lycopodites falcatus, L. & H.

Ljcopodltcs, Goldenherg (L. Stockil). 113

historisclien Vercins clcr prcussisclicn Illicinlande at Kreuz-

iiacli I exhibited a pretty eomplete example, and also later,

in the Transactions of this Society, made some preliminary

remarks on the occurrence of such plants in the Carboniferous

formation. 1 succeeded later in discovering several other

species, of which some even bore distinctly their organs of

fructification. It then appeared that, as regards the position

and form of the fruit, these fossil remains agree completely in

all essential points with our living Lycopods.

The Lycopodites which we are about to describe may, like

our recent club-mosses, be placed in two subdivisions, accord-

ing to whether the sporangia are seated in the axils of the

leaves or form terminal cones.”

The six species described by Goldenberg are classed under

these two heads : —

A. Sporangia placed in the leaf-axils.

1. Lycopodites denticulatuSj Goldenberg.

2. elongatuSy Gold.

B. Sporangia forming terminal cones.

3. Lycopodites primeevusy Gold.

4. leptostachyusy Gold.

5. macrophylluSy Gold.

6f taxinusy L. & H., sp.

In regard to his last-mentioned species, which he identifies

as Knorria taxinoy L. & H., its claim to belong to the genus

Lycopodites rests on very slender grounds.

His figure only shows a small portion of a stem IJ inch

long and about i inch wide, with spirally arranged leaf-

scars, in general form very like those of L. Stochii (PI. V.

fig. 1) ; but this single character, as shown in Goldenberg’s

figure, appears of too little importance to be of generic value.

But apart from the question as to the systematic position

of Goldenberg’s L, taxinuSy it is clearly not the same fossil

as that named Knorria taxina by Bindley and Hutton.

The specimen, from which the last-mentioned author’s

plate is taken, is preserved in the “ Hutton collection,” New-

castle-on-Tyne. Tliis I have compared with their figure,

which, I am sorry to say, is not a very correct representation

of the fossil. I believe Bindley and Hutton’s plant is merely

a small stem of Gordaites, certainly a quite different plant

from Goldenberg’s I^ycopodites taxinus.

In the same year (1855) in which Goldenberg described

114

Mr. E. Kidstoii on a new Sjjecies of

his specimens, Geinitz figured and described another species,

Lycopodites G uthieri * * * § .

All these plants Scliimper has placed in the recent genus

Lycopodiumf Both Goldenberg and Geinitz considered them,

if not identical with Lycopodium^ extremely closely related to

that genus.

Schimper’s opinion of the systematic position of these plants

has been adopted by Kenault|, who has described a small

Lycopodiaceous stem under the name of Lycopodium puncta-

tum^ ; another, in the same communication, is described by

Ad. Brongniart as Lycopodium Renaultii.

Both these small stems, though perhaps closely related to

Lycopodium^ possess some structural differences which we are

inclined to regard as of sufficient importance to exclude them

from the recent genus in which they are placed.

Benault, in his very interesting paper, has pointed out the

presence of areolated vessels in his fossils, which he admits

do not occur in recent Lycopodium. It is quite possible that

the specimens described by Benault and Brongniart are stems

of plants similar to Goldenberg’s Lycopodites ; but as they are

not identical in structure with any known Lycopodium.^ I

think it better to place them in Lycopodites than in Lyco-

train of reasoning inclines me to reject the genus

Lycopodium for Goldenberg^s species, notwithstanding their

undoubted close relationship to it. If we place them in

Lycopodium^ we commit ourselves to the opinion that the

fossils are identical in all essential structural peculiarities with

their recent representatives — an opinion not at present satis-

factorily substantiated by proof. I therefore propose to rein-

state the genus Lycopodites.^ Goldenberg, for the following

species : —

A. Sporangia placed in the axils of the leaves.

1. Lycopodites denticulatusj Gold.

2. elongatus^ Gold.

B. Sporangia forming terminal cones.

3. Lycopodites primcevus^ Gold.

4. leptostachyuSj Gold.

* ‘ Die Versteinerungen der Steinliolilenformation in Sachsen,’ p 32

pi. i. fig. 1 (1855). ^

t Schimper, Traite d. paleont. veg4t. vol. ii. p. 8, pi. Ivii. (1870).

i Cours d. botan. foss. p. 74, pi. xii. figs. 9, 10 (1882).

§ Ann. des Sciences Naturelles, 6® ser. Bot., vol. xii. pp. 178-182

pis. xii.-xiv. (1869).

podium.

The £

115

Lycopoclltes, Goldcnherg (L. Stocldi).

5. Jjycojyodites macropliyllus^ Gold.

G. Gutbierij Gdpp. (=L. stachyyynandroldes^

Gutbier).

7. StocJcUy Kidston, ii. s.

C. Sporangia unknown.

8. Lycojwdites punctum^ Renault, sp.

9. Renaultii^ Ad. Brong. sp.

10. (?) taxinus^ Gold, (not L. & H.).

The above list contains all the plants, as far as I am aware,

which are entitled to be placed in Lycopodites^ Goldenberg ;

but since Brongniart and Goldenberg wrote on this subject,

notwithstanding the warnings given by these two authors,

many Lepidodendroid twigs have been hgured and described

as Lycopodites. Most of these belong, I believe, to Lepido-

dendron Sternhergii and Lep. rimosum^ and probably to other

species of the same genus.

I have seen specimens of these two Lepidodendra with small

delicate branches springing from stems of considerable size,

which could be specifically identified with described species of

Lycopodites. These small lateral branchlets are merely the

result of very unequal dichotomy*.

Whatever view may be taken of these Lepidodendroid

twigs, or so-called Lycopodites^ it will, 1 think, be admitted

on all hands that they do not find a suitable place in Golden-

berg’s genus Lycopodites.

Lycopodites Stochiij Kidston, n. s.

Description. Cone terminal, composed of a number of oval

sporangia ; leaves arranged in whorls, dimorphic (?), the

larger leaves ovate-cordate, acuminate with a strong central

midrib, the (?)smaller leaves transversely oval.

Remarhs. The specimen from which the above description

is taken, is about 4 inches long; of this, the cone, which is

imperfect at its apex, occupies 1| inch. The leaves are mostly

displaced, but the form of many of them is well shown.

Unfortunately the state of preservation of the fossil is not

all that could be desired ; but it is sufficiently distinct to

enable one to give a description by which it can easily be

recognized.

In PI. V. fig. 1 is given a careful sketch of the plant.

The cone consists of a number of oval bracts : some of

them appear reniform in shape, but the fossil is so much

compressed that the individual contour of the sporangia

As an example, I give on PI. Y. fig. 5, some small branchlets oiLepid.

rimostim from the Upper Coal-measures, Timsbury, Somerset.

116 On a new Species c^ Lycopodltes, Goldenherg.

cannot be well made out. The leaves shown on the sides of

the stem are ovate acuminate, with a very distinct middle

nerve (fig. 2).

An interesting point in the fossil is the occurrence of a ver-

ticalrow of curiously formed leaves (?), entirely different from

those just described. One of these (a, fig. 1) is shown en-

larged at fig. 3. This curious structure has very much the

appearance of a sporangium ; but the occurrence of a terminal

cone and of sporangia situated in the axils of the leaves of the

same species, is altogether unknown in any Ly copod, either

fossil or recent.

The most perfect of these curious structures (whether leaves

or sporangia) appears to have three inflations (fig. 3) ; but

I am rather inclined to think that this appearance has been

caused by its being pressed against the stem, and that we have

under consideration a leaf and not a sporangium. In this case

we have merely a dimorphic condition of leaves, such as occurs

in those fossils already described by Goldenberg and Geinitz,

and is common in the recent genus Selaginella. In this

example, in no case are they exhibited so clearly that one can

positively affirm they are leaves ; but I believe this to be

their true nature notwithstanding their sporangium-like form.

In the enlarged sketch (fig. 3) the dark dentate margin

has no connexion with the supposed leaf, bat only a small

broken piece of carbonaceous matter, which probably represents

the cortex.

The leaves appear to have been arranged in whorls of 6 or

8, as shown by the cicatrices on the enlarged portion of the

stem (fig. 4).

My thanks are due to Mr. T. Stock, Edinburgh, who has

submitted this fossil to me for examination and description,

and after whom I have pleasure in naming it.

I believe the opinion generally current regards the genus

Lepidodendron as the ancestor of our herbaceous Lycopods ;

but I am rather inclined to believe that the genus Lepido-

dendron has entirely disappeared, and that our recent Lyco-

pods are the descendants of Goldenberg’s Lycopodites.

Horizon. Calciferous Sandstone series (Culm of Stur).

All the species of Lycopodites^ Goldenberg, previously de-

scribed have been derived from the Coal-measures; hence

the discovery of the genus so far down in the Carboniferous

formation is of considerable interest.

Locality. Glencartholm, Eskdale, Dumfries.

^y^nri. Ma^ Nah.Hi^b. iS.5, Vol 14, FI. V.

» M®Fa,rlane &. Ers"kiTie. LifcliT* EHinf

1 “ 4' . LyCOPODITES StOCKII. K-id,sboTh. n.s.

5. LePIDODENDRON RIMOSUPT. jS beT^nle.T>^.

LIBRARY

OF THE

UJilVERSiTY OF ILLINOIS

On the Families of existing Lacertilia.

117

EXPLANATION OF PLATE V.

Fi(j. 1. Lycopodites Stockii, Kidstoii, n. sp., nat. size', a, h, c, d,f. Sporan-

gium-like leaves (?) ; e. Keniform sporangia of terminal cone.

Fiy. 2. Leaf, enlarged, seen on fig. 1, y.

Fiy. 3. Sporangium-like leaf (P), enlarged, seen on fig. 1, a.

Fkj. 4. Small portion of stem, enlarged, showing leaf-cicatrices.

Fiy. 5. Fpidodendron rimusurn, Sternberg.

XVI. — Synopsis of the Families of existing Lacertilia,

By G. A. Boulenger.

Whilst engaged in a revision of the Lizard-collection in the

British Museum, I have felt the necessity of a thorough syste-

matic rearrangement of the order Lacertilia. The classifica-

tions proposed by Dumeril and Bibron and Gray, and now-

still generally in use, with slight modifications, are, on the

whole, as unnatural as can be, and founded to a great extent

on characters of pholidosis and physiognomy. Physiognomy

is worth nothing as a guide in the formation of higher groups ;

as to the characters afforded by the scales I have convinced

myself that they are very deceptive, and ought to be taken

into consideration in the definition of families only when

accompanied by other characters. Like Cope, whose lizard-

families* I regard as the most natural hitherto proposed, I

shall Jay greater stress on osteological characters and on the

structure of the tongue. Special importance must also be

attached to the presence or absence, and the structure, of

dermal ossifications on the head and body, and these will be

found to correspond with many other characters. Bocourtf;

to whom is due the merit of having pointed out their syste-

matic importance, did not realize the very great progress

made by means of that character, the modifications of which

he so ably illustrated, for he still maintains the artificial

group Scincoi’diens, in spite of the objections of Cope, whose

views are evidently confirmed by the researches of the French

herpetologist.

The order Lacertilia, as restricted by Gunther J, may be

divided into two primary groups only, the Cham^eleons on

the one hand, and all the other Lizards on the other. The

Amphisbsenians, which by nearly all recent authors are sepa-

* Proc. Acad. Pbilad. 1864, p. 224, and Proc. Am. Assoc. Adv. Sc.

xix. 1871, p. 236.

t Mission Scient. Mexique, Kept. p. 476 (188]).

f Phil. Trans. Roy. Soc. clvii. 1867, p. 625.

118

Mr. G. A. Boulcnger on the

rated as a suborder, or even as an order, I include among the

true lizards, and regard them as a degraded type of the

Teiidm, with which they are to some extent connected by the

Chalcides and their allies. The principal characters which

have been put forward in favour of their separation are : —

(1) absence of interorbital septum ; (2) absence of columella

cranii ; (3) very short mandible, causing the quadratum to be

nearly horizontal ; (4) division of the occipital condyle ; (5) ab-

sence of postorbital and fronto-squamosal arches; (6) absence

of scales. These characters, which are mostly negative, are

not all constant throughout the group, and many will be found,

to a greater or less degree, to be characteristic of all strongly

degraded, burrowing forms, such as Amelia near the

Anguidse, Andytrops {Typhline) and Dibamus near the

Skinks, &c. The importance of these characters justifies our

placing the Amphisbgenas in a separate family ; but, in my

opinion, not in a higher group, for the following reasons : —

1. The absence of interorbital septum also occurs in

Ophiognomon among the Teiidse, and there is every grada-

tion between the skull of that genus and that of higher

members of the same family : besides Amelia and Dibamus ^

which belong to totally different families, also possess the

same negative character.

2. The columella disappears gradually with the interorbital

septum ; it is hardly distinguishable in Ophiognomon and

totally absent in Aniella and Dibamus.

3. The aberrant lower jaw, not in itself a very important

character, is not even constant, the genus Blanus differing in

that respect as much from the typical Amphisbmna as from

a typical Lizard.

4. The division of the occipital condyle, also a character

the importance of which ought not to be exaggerated, is not

even constant, the Acrodont Amphisbgenians forming ex-

ceptions.

5. The absence of postorbital and fronto-squamosal arches,

which occurs in the most diverse groups of Lizards, cannot be

regarded as more than a family character.

6. The naked integuments (if we may apply this term to

the skin of the Amphisbsenians with its soft scales) are not

special to the group, but occur also in Geckos ; and they arc

so closely approached by those of some Cercosaurine and

Chalcidine Teiidas as to render any sharp distinction im-

possible.

On the other hand, characters such as are afforded by the

tongue, which in all Amphisbaenians is in every respect

similar to that of the Cercosaurine and Chalcidine Tciidiv,

119

Families of existing Lacertilia.

flic preaiial pores of most Ampliisbseriians, and the anterior

limbs of Chirotes are indieative of affinity to the Teiida?.

llespeeting the latter, it may be remarked that in the other

Lacertilia which dis])ense with the limbs, the fore pair dis-

ajipear before the hind pair, and this holds true for the Ojdii-

dians, the less modified type still showing rudiments of pelvis,

whilst not one preserves any thing of the pectoral arch. A

reverse process obtains in the Teiidge and Ampliisbaenida3.

I have already put forward my objections to recognizing

the suborder Nyctisaura*.

Having separated the Chama3leons, we are in presence of

the large suborder of true Lizards. This I have divided into

twenty families, which I regard as perfectly natural groups.

But there is great difficulty in arranging these families in a

line. Two characters seem to demand special attention — those

of the lingual papilla3 and the clavicle, as, excepting the

Geckos and Eublepharidge, they exactly correspond, i. e. the

forms with smooth or villose tongue have a slender, non dilated

clavicle, whereas those with scaly tongue have the clavicle

strongly dilated proximally and generally enclosing a fora-

men.

Order LACERTILIA.

Suborder I. LACERTILIA VERA.

A. Tongue smooth, or with villose papillce ; clavicle dilated, loop-shaped

proximally ; 7io postorhital or fronio-squamosal arches.

Earn. 1. Geckonid^. Vertebrae auipliicoelian ; parietal bones distinct.

Earn. 2. Eublephaeid^. Vertebrae procoelian ; parietal single.

B. Tongue smooth or with villose papillce ; clavicle not dilated

proximally.

Earn. 3. UEOPLAXiDiE. Vertebrae amphicoelian ; interclavicle minute;

no postorbital or postfronto-squamosal arches.

Earn. 4. Pygopodid^. No postorbital or postfronto-squamosal arches ;

pre- and postfrontal bones in contact, separating the frontal from

the orbit.

Earn. 5. Agamid^. Postorbital and postfronto-squamosal arches pre-

sent ; supratemporal fossa not roofed over by bone ; tongue thick ;

acrodont.

Earn. 6. Iguanid^. Postorbital and postfronto-squamosal arches pre-

sent; supratemporal fossa not roofed over by bone; tongue thick;

pleurodont.

Earn. 7. Xenosaueid^. Postorbital and postfronto-squamosal arches

present ; supratemporal fossa not roofed over ; anterior portion of

tongue retractile.

Earn. 8. Zonueid^. Postorbital and postfronto-squamosal arches com-

plete ; supratemporal fossa roofed over ; tongue simple.

Earn. 9. Anguib^. Postorbital and postfronto-squamosal arches present ;

supratemporal fossa roofed over ; body with osteodermal plates

Ann. & Mag. Nat. Hist. (5) xii. 1883, p. 308.

120

Mr. G. A. Boulenger on the

witli irregular, arborescent, or radiating channels ; anterior portion

of tongue retractile.

Fara. 10, Anikllid^. No interorbital septum, no columella cranii, no

arches.

Fam. 11. IIklodehmatid^. Postorbital arch present, postfronto-squa-

mosal arch absent j pre- and postfrontals in contact, separating

the frontal from the orbit.

Fam. 12. VAiiANiDiE. Postorbital arch incomplete ; postfronto-squa-

mosal arch present j supratemporal fossa not roofed over j nasal

bone single ; tongue deeply bifid, sheathed posteriorly.

C. Tongue covered ivith imbricate scale-like papillce or with oblique plicce ;

clavicle dilated proximally, frequently loop-shaped.

Fam. 13. Xantusiid^. Parietals distinct ; postorbital and postfronto-

squarnosal arches present ; supratemporal fossa roofed over.

Fam. 14. Teiid^. Postorbital and postfronto-squamosal arches present ;

supratemporal fossa not roofed over ; no osteodermal plates.

Fam. 15. Amphisb^nid^. No interorbital septum; no columella

cranii ; no arches ; premaxillary single.

Fam. 16. Lacertid^. Arches present ; supratemporal fossa roofed over ;

premaxillary single ; no osteodermal plates on the body.

Fam. 17. GEERnosAURm^. Arches present ; supratemporal fossa roofed

over ; premaxillary single ; body with osteodermal plates with

regular channels (a transverse one anastomosing with perpendicular

ones).

Fam. 18. SciNCiDiE. Arches present ; premaxillary double ; body with

osteodermal plates as in the preceding.

Fam. 19. Anelytropid^. Premaxillary single; no arches; no osteo-

dermal plates.

Fam, 20. Pibamid^. Premaxillary double ; no interorbital septum ; no

columella cranii ; no arches ; no osteodermal plates.

Suborder II. KIIIPTOGLOSSA.

Fam. 21. Cham^leontid^.

The Geckonidce and EuhlepharidcBj which differ from all

other families in combining a dilated clavicle with a simply

papillose tongue, are well distinguished from each other by

the vertebrae, which are amphicoelous in the former and pro-

ccelous in the latter. As characters of minor importance may

be mentioned the coossification of the parietal bones in the

Eublepharidae, while they remain distinct in the Geckonidae,

which are also distinguished, constantly I believe, by having

one bone less in the mandible, the supra-angulare having

coalesced with the angulare.

Next come the Uroplatidce^ which are now for the first time

separated from the Geckos. Although agreeing in most

respects with the latter, their sternal apparatus differentiates

them widely ; the clavicle is slender, not at all dilated, and

the interclavicle is reduced to a minute bone. Except the

chamaoleons, all other lizards in which the })ectoral arch is not

121

Families of existing Lacertilia.

rudimentary have a large interclavicle. To this very im-

portant character is added another ; the nasals are united into

a single bone, a peculiarity which is found elsewhere only in

the Varanidte among recent lizards. A single genus, Uro-

gilateSj from Madagascar, is known.

After the Uroplatidje I have placed the Pygopodidee

( == Pygopidge + Aprasiadm + Lialisidm of Gray), which family

is now based on new characters. They were formerly arranged

with or near the Scincoids,” a view which cannot be main-

tained, since that group was an assemblage of forms having

totally ditferent affinities, and Scincoids ” will now be found

scattered through the following families : — Anguidse [Anguisj

Dijdoglossus, &c.), Aniellidse, Teiidm {GymnopfithalmuSj &c.),

Scincidae, Anelytropidge, and Dibamidge. The skull of the

Pygopodidge in its simplicity of structure approaches that of

the Geckos, and the parietal bones remain distinct in all the

genera except Lialis ; the bones of the lower jaw are still

more reduced in number, the angular, supra-angular, and arti-

cular having coalesced, a character by which they approach

the snakes. The affinities of this little group are very obscure,

and a complete investigation of their anatomy is highly

desirable.

The two closely allied families Againidce and Iguanidoe

remain as before.

The Xenosauridee must be regarded as intermediate between

the Igiianidge, with which Peters was inclined to associate

them, and the Anguidge, near which they are placed by Cope.

The Zonuridoe correspond only in name with the Zonu-

ridge of Gray and most other authors. The members of

Gray’s Zonuridge will be found in the following families : —

Anguidge [Gerydionotus^ FseudopuSj &c.), Lacertidge [Tacky-

dromus)^ Gerrhosauridge. They have, like the Anguidge, a

villose tongue, though not retractile at the end, a slender

clavicle, and in some the body even presents bony plates,

which are destitute of symmetrical canals. As here under-

stood, the Zonuridge comprise the genera Zonurus^ Flaty-

saurusj and Ghamcesaura.

The Anguidee correspond to Cope’s Anguidge and Gerrho-

notidge, the differential characters of which latter group seem

to me insufficient for family separation. As Cope has shown,

this group is perfectly natural, though containing “ Chalci-

doid” and “ Scincoid ” forms, and an excellent illustration

of how misleading it is to trust only to external characters.

The “ Scincoid ” forms correspond to Bocourt’s Diplo-

glossidge.

The family Aniellidce was also established by Cope. I

would regard it as a degraded form of the Anguidge.

122 On the Families of existing Lacertilia.

The llelodermatidoBy as already shown by Cope, liave the

greatest affinity to the Anguidge, from which they are, how-

ever, well distinguished by the structure of the skull. The

grooved teeth might be given provisionally as another family

character. It would be highly important to have some infor-

mation on the osteological characters of Steindachner’s Lan-

thanotidge, as there is reason to suspect they will enter this

family.

The Varanidce^ which come last in the series of alepidote-

tongued lizards, remain characterized as before, and form a

perfectly isolated group.

We have next a series of families characterized by the

peculiar scale-like lingual papillae and the proximally dilated

clavicle.

The Xantusiid(B are closely allied to the Teiidse, but

distinguished by the different skull and scarcely incised tongue.

The Teiidce form a very natural group, comprising the

Cercosaurid^e, Chalcidida3, Chirocolidge, Anadiadae, and part

of the Gymnophthalmidae of Gray and the Tretioscincidae of

Bocourt. It thus contains Lacertoid,” Chalcidoid,” and

Scincoid forms of the Dumerilian system, all passing into

one another by insensible gradations and all agreeing in the

structure of the skull, tongue, and pectoral arch. All are

confined to the New World, whereas the analogous family

Lacertidee is restricted to the Old World. As mentioned

above, I regard the Amphisbasnidae as strongly degraded

forms of the Teiid^.

I establish a family Gerrhosauridm for Gerrliosaurus^

which was formerly associated with the Zonuridae, but which

agrees closely with the Scincidae, from which it is to be dis-

tinguished by the coalesced premaxillaries. Although the

arrangement of the scales of the body is different from what

we see in the latter family, the underlying dermal bony plates

are precisely similar in their symmetrical canals.

The Scincidee correspond to Cope’s Scincidm^ Sepidae, and

Acontiidae, and to Bocourt’s group Aspidoscinciens, less the

Biploglossidae.

The Ajielytropidce^ a small family so named by Cope and

synonymous with the Typhlinidae of other authors, are a

degraded type of the Scincidae, having completely lost the

cranial arches — which, in some forms of the latter group,

show a tendency to disappear — and also the osteodermal plates.

The Dihamidce^ characterized for the first time, and com-

prising only the genus Dibamus^ go still further in the direc-

tion of degradation, and are exactly analogous in this series

to the Aniellidge in the other series.

Mr. A. G. Butler on a new Species of Pscudacra3a. 123

XVII. — Description of a new Species of Pseudacraaa from

Natal. By ARTHUR G. Butler, F.L.S., F.Z.S., &c.

The genus Pseudacroia is one of the most interesting groups

of butterflies, the species of which mimic the various forms of

Acrcea^ Planema^ &c.

The present species is in the collection of Mr. Walter de

Ilothscliild, and was captured in Natal by Mr. Peter Colville,

after whom I have much pleasure in naming it.

The nearest allies of P. Goloillei are P. Trimenii and P.

BoisdiLvalii^ between which it is in some respects interme-

diate ; it appears to me to resemble Acrcea horta rather than

the groups copied by its two allies.

Pseudacrcea ColviUei^ sp. n.

(S . Al0e anticae area interno-hasali riifa, nigro maculata ; area api-

cal! cinerea, subhyalina, venis strigisque internervularibus nigris ;

limbo externo nigrescente ; alee posticae rufae, area basali nigro

maculata ; limbo externo nigro, rufo maculato ; corpus nigrum,

fulvo alboque macnlatum.

Primaries with the basi-internal half reddish fulvous (pro-

bably carmine-red when fresh), with black markings, exactly

as in P. Trimenii ; apical half smoky semitransparent grey,

with black veins and internervular streaks nearly as in P.

Boisduvaliij but without the transverse <-shaped markings

near the base of the median branches : secondaries nearly as

in P. Trimenii^ but with large oval red spots on the black

border, as in the female of P. Boisduvalii ; form and expanse

of wings corresponding with those of the latter species.

Natal (P. Colville). Coll. W. de Pothschild.

Although P. Trimenii is subject to slight variation in

colouring, the secondaries sometimes exhibiting a snow-white

patch from the three round black spots crossing the median

vein, as in Acrcea acara^ there can be little doubt that the

present species is far too distinct from it for a mere sport ; its

resemblance to an entirely different form of Acrcea^ its diffe-

rent outline, and other characters show it to be clearly a

separate species. Neither is it any more remarkable that two

Pseudacrceas of the same group should occur in Natal than

that the corresponding forms of Acrcea should independently

exist there ; the only strange thing is, in spite of the rarity of

the species in this genus, that the present sjDecies has not

already been described.

The allied P. Boisduvalii inhabits Western and South-

western Africa.

124

M. de Quatrefages on

X VIII. — Moas and Moa-hunters. By A. de Quatrefages

When I published, in the ^Journal des Savants,’ a first

article upon New Zealand and its inhabitantsf we had re-

ceived in Europe only the first three volumes of the ^ Trans-

actions ’ in which are brought together the works of the

learned societies of New Zealand. At tliat time I had to

express my regret that this collection contained only a single

very short note relating to the large sliort-winged birds desig-

nated by the common name of Moas This deficiency has

since been filled up. The succeeding volumes have brought

us numerous memoirs, in which the various questions raised

by the history of these birds are treated of. My present

object is to give a general summary of these researches, which

are very interesting in many respects, avoiding too technical

details, for which I can only refer the reader to the writings of

MM. Haast, Hochstetter, &c., and especially to those of Sir

Bichard Owen, which have been classical since their appear-

ance.

I.

We must first of all refer to the most striking feature of the

New-Zealand fauna.

The travellers who first landed upon this distant country §

were surprised at finding there of Mammalia only a domestic

dog and a rat, which the natives hunted as game. Since then

two bats of different genera have been discovered ||. The

researches of geologists have extended to palaiontological times

the results furnished by the study of the living animals, and

have even rendered them still more strongly marked. No

fossil mammal has yet been discovered through the whole

* Translated by W. S. Dallas, F.L.S., from the ‘ Annales des Sciences

N aturelles/ ser. 6, tome xvi.

t In January 1873, The present memoir has also appeared in the

same journal (in the numbers for June and July 1883).

} “ Address on the Moa,” by the lion. W. B. Mantell (Trans, and

Proc. of the New Zealand Institute, vol. i. p. 18). Mr. Mantell alone has

occupied himself with the general history of the Moas, But it is only

just to add that in the same volume Dr. Ilaast gave a memoir, entirely

technical in its nature, in which he published the results of measurements

made upon very numerous bones {ihid. p. 80).

§ New Zealand was discovered by Tasman on the 13th December, 1642.

It was forgotten and in a manner lost for more than a centijry, and was

rediscovered by Cook on the 6th October, 1769.

II Scot()2')hilus iuherculatus (Gray), identical with an Australian species,

and MysUwina tuherciilata, which has hitherto been found only in New

Zealand. (Note communicated by M. Alphonse Edwards.)

Moas and Moa-hunters.

125

extent of the lands composing New Zealand. This rendered

the exceptions which 1 have just indicated even still more

sii iking. How are we to interpret the existence of these four

isolated species, each representing one of the subtypes of

the class, and not preceded by any other belonging to the

fundamental group ? We have here a strange fact with no

analogy elsewhere. Nowhere else do we see a whole class of

animals entirely wanting in the fossil faunas and only repre-

sented in the existing fauna by an insignificant number of

species belonging to distinct orders. On the contrary, there

always exist more or less close affinities between the past and

the present in the animal creation. We know that these

relations are even every day invoked as so many arguments

in favour of the transformist doctrines.

The New-Zealand fauna therefore presents a unique excep-

tion to one of the most general facts hitherto ascertained.

Now it is very difficult to admit the existence of exceptions of

this kind. One is therefore naturally led to inquire whether

some accidental phenomenon has not intervened here to mask

the natural facts — wliether this dog, this rat, and these bats

really belong to the New-Zealand fauna, whether they are

not simply colonists introduced, it matters not how, into a

country to which they were originally strangers.

The presence of the Chiroptera might easily be ascribed to

a fact of accidental dissemination resulting from a few blasts

of wind, as has been demonstrated at the present day in the

same regions But that of the two terrestrial mammals has

long remained unexplained.

To solve this curious problem of zoological geography Sir

George Grey has had to discover, translate, and publish the

historical songs which have furnished equally precise and

curious information upon the first origin of the Maoris.

Through him we have learned that on quitting Hawaiki for

the new country discovered by Ngahue the emigrant chiefs

brought with them the plants and animals of which experi-

ence had taught them the utility. The dog and the rat figure

in the list of these treasures of the colonist tj and still attest

^ Zosterops lateralis (Latham), a bird originally inhabiting Australia,

has been carried in this way to New Zealand and into Campbell Island.

It did not exist in the Chatham Islands until 1861. At this time it

appeared suddenly after a storm Rapport sur Fexposition faite au Mu-

seum des objets d’histoire naturelle recueillis par MM. de L’Isle et Filhol,”

par A. de Quatrefages, ‘ Archives des missions scientifiques et litteraires,’

tome V. p. 24).

t ‘ Polynesian Mythology,’ 1855 ; ‘ The Emigration of Turi,’ pp. 212,

214 ; ‘ The Emigration of Manaia,’ p. 228. I have analyzed these docu-

ments and all those relating to the same set of ideas in a work entitled

Ann, & Mag, N, Hist, Ser. 5. Vol, xiv. 10

126

M. de Quatrefages on

the exactitude of the traditions for the knowledge of which we

are indebted to the late Governor of New Zealand. They

did not originate in those islands, but they were imported

there

The Mammalia which did not occur in the natural fauna of

this archipelago were to a certain extent replaced by birds

belonging to a peculiar type, represented elsewhere by a very

small number of species, but which here acquired an abso-

lutely exceptional development. 1 refer to birds with rudi-

mentary wings and with loose-barbed feathers, incapable

‘ Les Potyn^siens et leurs migrations/ accompanied by four maps. I only

remind tbe reader that Hawatki here mentioned is one of the Manaia

Islands, and probably Armstrong Island or Bouroiiti of our atlases.

* New Zealand now possesses mammals which the Europeans have

introduced there, and the acclimatization of which has not been without

its inconveniences. Our common rat has almost entirely destroyed the

rat imported by the colonists from Hawaiki, the Kiore of the Maoris.

As a matter of course the mouse has accompanied it. Our cat has re-

turned to the wild state in this island, and it is probably one of these

animals that has been taken for an indigenous species of otter supposed

to have been once seen. Our rabbit has multiplied there, as in Australia,

to such an extent as to become a plague in the cultivated lands ; so much

so indeed that a few years ago the Acclimatization Society of Paris

received an application for a certain number of weasels, for which 100

francs a pair Avere offered. They were to have been set free, in the hope

that they would multiply and Avage war against the rabbits. But it is

easy to understand that the remedy might have been worse than the

disease. As to the pigs, introduced by Cook in 1769, they are now so

numerous and commit such ravages that hunters are expressly engaged

to destroy them. Hochstetter tells us that in tAventy months three men,

hunting over an extent of 250,000 acres, killed no fewer tlian 25,000 wild

pigs, and undertook to kill 16,000 more upon the same ground (‘NeAv

Zealand/ p. 162). These wild pigs will speedily bring about the complete

extinction of the last wingless birds of the country {Apteryx)^ of Avhich

they destroy the nests.

In other respects the acclimatization of foreign animals has proceeded

in New Zealand with astonishing rapidity. Fourteen species of birds,

coming from Europe, Asia, and America, have made themselves a new

home in this maritime country. The colonists have transported there

not only the sparrows and larks, but also the pheasant and the Califor-

nian quail. All these new comers have driven before them the indige-

nous species, the representatives of which are becoming scarcer and scarcer,

while several of them appear to be menaced Avith a speedy extinction.

We may notice in passing that the invasion of NeAV Zealand by foreign

plants has been no less general and no less fatal to the indigenous vege-

tation. Our cereals and our vegetables everyAA^here replace the batatas

and cause the eradication of the ferns on the roots of which the Maoris

fed. Flven our weeds, transported here involuntarily, have multiplied to

such an extent as to extinguish those of the country. “ In the plain of

Christchurch,” writes M. Eilhol, ‘‘ it is in vain to seek — we can no longer

find — a Polynesian plant ; one might fancy oneself in Beauce ” Eapport

sur I’exposition,” &c., loc. cit.).

Moas and Moa-hunters,

127

of fliglit and more or less analogous to the ostrieh or the

cassowary*. Four or five species of this group still exist

in New Zealand. They are known to the natives by the

common name of Kiwi^ and have been placed together by

naturalists in the genus Apteryx f. They vary in size from

tliat of a fowl to that of a turkey. But the number of extinct

species is far more considerable, and among them are some of

tnily gigantic proportions. It is these vanished species that

are called by the common name of Moa^ borrowed from the

language of the Maoris |.

The first investigations upon this curious chapter of orni-

thology date from 1830. The illustrious English anatomist,

Richard Owen, had received from a Mr. Rule the middle

portion of a femur, and from the examination of this single

imperfect specimen he drew conclusions which every thing

has tended to confirm §. More abundant and more complete

* Beyond New Zealand the ornithological type in question is only

represented by four species, each having a very different area of habitat

and isolated from the others by vast spaces. These are the ostrich

(Struthio camelus, Linne), which inhabits almost the whole of Africa

and the warm parts of Asia on this side of the Ganges •, the nandou, or

American ostrich (Rhea mnericana, Lath.), which inhabits South Ame-

rice from Brazil to Patagonia ; the emeu, or helmeted cassowary (Casu-

arius emeu, Lath., Struthio casuarius, Linne), found only in the Indian

Archipelago and principally in the forests of Ceram ; and lastly the emou,

or helmetless cassowary {Casuarius novcB-holkindice, Lath.), which appears

to have been spread over the whole of Australia, but which the European

colonists drive back more and more and will not fail to destroy. [It will

be remarked here that M. de Quatrefages refers only to four species instead

of four genera. The number of species is considerably greater, there being

three known species of Rhea, two species of Dromceus, and nine of Casu-

arius. The last-named genus has a much wider range than is indicated^

above, extending from the Indian Archipelago to the island of New

Britain and to Northern Australia. — Tn.]

t These species are Apteryx australis, A. Mantelli, A. Oioenii, and A.

Haastii. A fifth species of large size perhaps exists in the desert regions

of the Middle Island. It has even been described by Verreau, a French

natural-history traveller. But he had never seen more than a single skin

covered with feathers, of which a Maori chief had made himself a mantle,

(Note communicated by M. Alphonse Edwards.)

X The Kiwis were in existence at the same time as certain species of

Moas. Their bones have been found mixed together in caves and also

among the kitchen refuse, of which I shall speak hereafter. The Moas,

moreover, are not the only birds the species of which have disappeared

from New Zealand. Owen has shown that this is also the case with two

Rallidse, of which he makes the genus Aptornis. Dr. Haast has described

the remains of a large bird of prey, which he has named Harpagornis

Moorei', it is perhaps the Weka spoken of in certain Maori traditions (see

“ Notes on Harpagornis Moorei,^'’ by J . Haast, in Trans. &c., vol. iv.

p. 192, pis. X. and xi.).

§ Sir Richard Owen made his first communication on this subject to

the Zoological Society of London on the 13th November, 1839. He fol-

10*

128

M. de Quatrefages on

materials soon enabled him to recognize five distinct species,

which he united into the genus Dinornis,

Subsequently this number gradually increased to thirteen,

and there have been found in these representatives of an

extinct fauna more and more marked differential characters.

This is so much the case that Dr. Julius Haast, the eminent

New-Zealand geologist, has thought it right to form of them

four genera, themselves divided into two groups or families

It is easy to see that these palseontological discoveries con-

firm the observations which I have just been making and

bring New Zealand under the general rule. This austral

country has never produced Mammalia. To make up for

this the type of the short- winged birds has been developed

there with an abundance and a variety of secondary types

such as we meet with nowhere else. There is complete

accordance between its fossil and its recent faunas, and these

faunas, precisely by the exceptional character which is

common to them, furnish another proof of the universality of

the laws which everywhere bind together the past and the

present of the animal world f-

lowed out his researches chiefly by the aid of the materials sent by Mr.

W. Mantell. The results have appeared in the ‘ Transactions of the

Zoological Society ’ for 1844 and following years.

* The following is Dr. Haast’s classification, which, however, only

includes eleven species : — I. Family Dinornithid^ : genus Dinornis, in-

cluding D. maximus, D. rohustus, D. inyens, D. struthioides, D. yracilis ;

genus Meionornis, including M. casuarinus, M. didiformis. II. Family

Palapterygid^ : genus Falaiiteryx, including P. elephantopus, P. cras-

sies; genus Dury apteryx, including E, gravis, E. rheides (Proc. Philos,

Inst, of Canterbury, March 1874 ; Address by J. Haast, president j Trans-

actions, &c., vol. vi. p. 426). Dr. Haast, from considerations drawn

especially from size, seems disposed to think that he has himself united

under the name of Meionornis casuarinus two species which will have to

be distinguished hereafter. He makes analogous remarks with regard to

Ealapteryx elephantopus (p. 429). Prof. Hutton, Director of the Museum

at Otago, has criticized Haast’s classification, and denied some of the

facts relied upon by his confrere. With Owen, he thinks that the Moas

form only a single natural family, that of the Dinornithidae (‘ Transactions

&c., vol. ix. p. 363). Owen and M. A. Edwards only admit the two

genera Dinornis and Ealapteryx, the former tridactyle, the latter Iiaving

a fourth digit, which is short and directed backward.

t The preceding observations apply not only to the history of the New-

Zealand fauna, but affect the history of man himself. By themselves

they suffice to refute a theory recently put forward by M. P. A. Lesson

in a book in other respects filled with important facts and documents, of

which three volumes out of four have appeared, namely ‘ Les Polynesiens,

leur origine, leurs migrations et leur langage ’ (Paris, 1882). The author

assumes that the whole of Polynesia, Tahiti, the Sandwich Islands, the

Samoa and Tonga Islands. &c. has been peopled by means of migrations j

Moas and Moa-hunters.

129

The number of Moa-bones gathered by the scientific men

or by simple amateurs inhabiting New Zealand is very con-

siderable, and it is only just to acknowledge the generosity

with which these scientific treasures have been communi-

cated to those whom they might interest, and even divided

among the naturalists of the whole world. All the great

museums of Europe and America now possess more or less

complete specimens of these strange birds. Mr. Mantell,

who was one of the first to interest himself in the question,

has sent to Sir Eichard Owen more than 1000 specimens

When the learned geologist of the ‘ Novara,’ M. Hoch-

stetter, jvished to dig for himself in the marshes and bone-

caves, he found everywhere the most earnest cooperation. It

has been the same with our countryman, M. Filholt* *

It is to the kindness and liberality of our New-Zealand

confreres j and especially of MM. Haast and Hutton, that we

but instead of accepting the Malay Archipelago as the starting-point of

the race, he makes Polynesian man originate in New Zealand. He thus

reverts to the old doctrine of autochthonism, of which the magnificent

work of Mr. Hale had already demonstrated the untrustworthiness, and

at the same time he places the cradle of the Polynesian islanders upon

the land which afibrds least support to any hypothesis of this kind. I

have already briefly examined M. Lesson’s theory, and indicated how,

independently of the data furnished by the study of the faunas, the his-

torical documents which we owe in part to that author himself, but

principally to Sir George Grey, Thomson, Shortland, &c., do not allow

of our accepting it Hommes fossiles et hommes sauvages,’ p. 483). I

shall return to this subject when M. Lesson’s work is completed.

* Hochstetter, loc. cit. p. 182.

t MM. Filhol and de L’Isle were attached as naturalists in 1874 to the

expeditions sent out to observe the transit of Venus at St. Paul and

Campbell Islands, under the command of Admiral Mouchez and M. Bou-

quet de Lagrye. Both of them brought back important collections. But

M. de I’Isle, being prevented by illness, could not realize all that was

promised by his known zeal. More fortunate, M. Filhol fulfilled his

mission in a remarkable manner. After having thoroughly explored

Campbell Island, he went twice to New Zealand, of which he traversed

the principal provinces, He afterwards visited the Fiji Islands, New

Caledonia, and the Sandwich Islands, and returned to France by San

Francisco. From wherever he went he brought back remarkable collec-

tions and most interesting observations. Prof. Hutton, Director of the

Museum of Otago, gave M. Filhol for our museum numerous bones of

Moas and two complete skeletons, one of Palapteryx elephantopus, the

other of P. crassus (“ Rapport,” &c., loc. cit).

On his part Dr. Haast has sent us, besides a great number of separate

bones, four nearly complete skeletons, which could be mounted, namely

of Pinornis crassus, giganteus, elephantopus, and didiformis.

The museum also possesses a model in plaster of the magnificent

Pinornis ingens, collected and reconstructed byHochstetter and figured in

his book, pp. 187 and 188.

130

M. de Quatrefages on

are indebted for the magnificent specimens which now figure

in the museum. I shall not be reproached for having dwelt

upon these facts, and here publicly thanking men who so

worthily understand and practise scientific confraternity.

II.

This abundance of materials has enabled us to form a

pretty complete idea of what the Moas were. We have been

able to reconstruct entire skeletons of several species, and thus

to judge of their size and their proportions. On the whole,

and notwithstanding the secondary differences which distin-

guish them, all these birds, as I have already said, resemble

the ostrich or the cassowary. The head is small, and nothing

in it indicates the existence of a solid crest analogous to that

which distinguishes the emeu SJJasuariu^^ and has obtained it

the name of the helmeted cassowary. The very long neck, at

first slender, gradually thickens as it approaches the trunk, as

in the cassowary. The skeleton of the body is robust. The

sternum alone is comparatively very small and flat. The

reduction of this bone, so highly developed in flying birds, is

explained here by the smallness of the wings, which are truly

rudimentary. On the other hand, all that portion of the

skeleton connected with the hinder limbs has acquired excep-

tional dimensions. The pelvis is massive ,* the bones of the

thigh, the leg, and the metatarsus have enormous heads, and

the body of the bone itself is comparatively much thicker

than in the living representatives of the type. These cha-

racters are particularly marked in Palapteryx ele^pliantopus.

This bird was a little smaller than our ostriches, and never-

theless in it the metatarsus presents a circumference nearly

double that of the same bone in the ostrich and the Casso-

wary *.

The size varied in a very noticeable manner in the different

species of Moas. The smallest [Meionornis didiformis) was

only 3 or 4 feet (0'97-l*30 m.) in height f. These were

therefore very inferior to the ostrich, the size of which varies

from 6 to 7 feet (l*95-2*27 m.). But Paiwpteryoc ingens

was precisely of this same size ; Dinornis rohustus was 8 to 9

* Hochstetter, loc. eit. p. 138,

t I borrow all these numbers from Hoclistetter’s table of measure-

ments {loc. cit. p. 198). That learned traveller seems to have judged of

the size not by measuring the distance from the beak to the extremity of

the feet, but by supposing the bird in repose in its position of equili-

brium, the neck inclined forward and presenting a double curvature, just

as he has represented Falapteryx myens, the entire skeleton of whicli is at

Vienna Qoc. cit. p. 188).

Moas and Moa-huntors.

131

feet (2*60-2*92 m.) high, and Dinornis maximm raised

its head to 9 or 10 feet (2*92-3*25 m.) from the surface

of the ground. Thus it exceeded by nearly a metre our

largest ostriches. According to Thomson, cited by M. Al-

phonse Edwards in an unpublished work which he has been

kind enough to communicate to me, there even existed indi-

viduals 13 to 14 English feet (4*0-4*25 m.) in height.

By comparing a great number of bones of adult individuals

of the same species. Dr. Haast has ascertained that tliey always

formed two series of slightly different size. He has attributed

this inequality to sex, and, guided by what oecurs in the

Apteryx^ he regards the larger bones as having belonged to

females

Besides the osseous remains of Moas, there have been dis-

covered fragments of eggs and even some entire eggs, most

of which, unfortunately, have been broken. But it has been

possible to restore a considerable number t* These eggs, of

a pale yellow colour^:, are dotted over with hollow points and

little grooves §. Their volume was considerably greater than

that of the eggs of the ostrich, but without equalling in this

respect those of ^pyornis 1|. In one of them the bones of

* Address Transactions ’ &c. vol. vi. p. 4il8).

t Mr. Mantell alone has reconstructed a dozen of these eggs, which he

has for the most part divided between the British Museum and the

Museum of the Royal College of Surgeons. Among these specimens,

which testify so strongly to the address and patience of the author, there

are some which contain no less than 200 or 300 pieces brought together

(“ On Moa-heds,” ^Transactions ’ &c. vol. v. p. 94).

X [The fragments of egg-shell which accompanied the York specimen

were of a dark green colour ; the pale yellow specimens must have been

bleached. — Tn.]

§ On the Microscopical Structure of the Egg-shell of the Moa,” by

Capt. F. W. Hutton Transactions ’ &c. vol. iv. p. 166, pi. ix. figs. 1-5).

The shell of the egg, about 1*75 millim. (0'07 inch) in thickness, consists

of two layers. The outer one is formed by lamellae parallel to the sur-

face, the inner one by a kind of prisms perpendicular to the former.

Other observers speak of these eggs as being perfectly smooth. It may be,

perhaps, that the little grooves in question here are due to the action of

grains of sand driven by the wind. We know, in fact, that this action is

exerted even upon rocks much more resistant than egg-shells, and this

fact has been demonstrated precisely in New Zealand.

\\ JEpyornis maximus inhabited Madagascar. It was destroyed by the

hand of man, but we do not know at what period. The eggs and some

bones were described for the first time by Isidore Geoffroy Saiut-Hilaire

(‘ Oomptes Rendus,’ 1851, tome xxxii. p. 101, and Ann. des Sci. Nat.

ser. 3, tome xiv. pp. 206 and 213). M. Alphonse Edwards, having

received fresh materials, has produced a very complete memoir upon this

species (‘ Recherches sur la faune ornithologique des lies Mascareignes

et de Madagascar,’ p. 85, 1873). From the investigations of this natu-

ralist it appears that ALiiyornis approached the Moas, although presenting

characters proper to make it the type of a family probably including three

132

M. de Quatrefages on

a young foetus were found, and Dr. Hector has been able to

compare them with those of an embryo of the emou \I)rom(2u^

of the same age It is interesting to find that, even at this

period of life, the principal differential characters are distinctly

marked, and that the pelvis, the bones of the leg, &c. are

much more voluminous in the Moa than in its near relative

from New Holland.

Lastly, isolated feathers of M oas belonging to various parts

of the body have been collected from time to time at different

places, as also even portions of the skeleton, to which muscles,

tendons, shreds of skin, and feathers still adhered in a remark-

able state of preservationf. I shall revert hereafter to the

consequences to be drawn from these facts. I only speak of

them here to complete the description of these birds.

Prof. Hutton has investigated the feathers found in two

localities in the midst of bones of Moas. These feathers

belonged to the same species. They were as fresh and their

colours were as bright as if they had just been pulled out.

But all were broken except a single one, of which he gives a

figure J . The total length is 16 centim. (6 inches) . The tube

is only 5 or 6 millim. (| inch) long, and bears two very slender

species. In particular the bones of the metatarsus were still stouter and

more massive than even in P. elejyhantopus. Its height must have been

about 2 metres. Its eggs, several of which are in existence, have a capa-

city of more than 8 litres, and represent in volume six ostrich eggs and

148 hen’s eggs.

* On recent Moa-remains in New Zealand,’’ by James Hector (‘ Trans-

actions’ &c. vol. iv. pi. vi. figs. 3, 4). The same plate gives drawings of

the eggs of the moa and of the emou {Drommis), reduced to one third

the natural size (figs. 1, 2). — Letter from Mr, T. M. Cockburn Hood to

Dr. Hector Transactions’ &c. vol. vi. p. 387).

t Address on the Moa,” Extracts, by the Hon. W. B. Mantell (Trans-

actions &c. vol. i. p. 19) ; ‘‘ On some Moa-feathers,” by Capt. F. W.

Hutton {ibid, vol, iv. p. 172) ; On recent Moa-remains in New Zealand,”

by James Hector, M.D., F.R.S. (ibid. vol. iv. p. 110). Analogous facts

are often referred to in other memoirs, and I shall have to return to them.

[It is curious that throughout the articles published in New Zealand on

the Moa, there is scarcely any mention of the finest and most interesting

Moa-skeleton in existence, namely that of Dinornis robnstus in the museum

at York, and M. de Quatrefages also says nothing about it. And yet

this specimen not only is a nearly perfect skeleton of an individual bird, but

its bones were in part united by ligaments and covered with skin, whicli

bore the bases of feathers : parts of the skin of the foot were preserved,

and much dried muscle was attached to some of the bones. With the

skeleton were some bones of young chicks and a few fragments of gi-eon

eggs. — See Thomas Allis, in Journ. Linn. Soc. vol. viii. pp. bO & 140 ;

R. Owen, On Dinornis, nos. ix. & xi.,” Trans. Zool. Soc. vols. v. & vi. ;

and W. S. Dallas, Proc. Zool. Soc. 1865. — Te.]

X Loc. cit. pi. ix.

Moas and Moa-hunters.

183

stems, the barbs of which, alttiougli fiirnislicd with barbules,

remain separate from each other. These barbs, which arc at

first very short, attain a length of about 2^ centim. (1 inch),

and the feather terminates with a rounded margin. The

first two thirds from the base are of a more or less reddish

brown, which gradually passes into black, while the rounded

extremity is of the purest white. Prof. Hutton points out

that these characters approximate the Moas to the A.merican

and Australian brevipennate birds rather than to the African

ostrich * * * §.

As a matter of course, however, all the Moas had not the

same plumage. The discoveries of Mr. Taylor White in this

respect furnish a confirmation of what was easy to foresee.

In the cave of Mount Nicholas he found feathers of a pale

yellowish-brown colour, darker at the margins. Some were

of a blackish brown. The feathers from another cave, near

Queenstown, were of a reddish brown and marked with a dark

brown streak towards the extremity of the stem t- Thus we

know, at least partially, the plumage of three species of MoasJ.

The feathers which I have just described came no doubt

from the middle or posterior region of the body. The

precious specimen described and figured by Dr. Hector shows

the modifications presented in this respect by the anterior

dorsal region and the neck §. This includes seven vertebras,

the first dorsal, and the six lowest cervicals, united by their

ligaments, and retaining upon one side their muscles and

integuments. The author believes that the neck of this Moa

was 18 English inches in circumference at its base.

Upon the portion of this specimen corresponding to the

first dorsal vertebra, the skin is seen to be covered with large

conical papillse which nearly touch each other, and give the

whole the appearance of a rasp. A certain number of these

papillas bear double-stemmed feathers of a reddish-chestnut

colour, furnished with barbs like those of the preceding-

feathers, but the longest of which are at the utmost 2 inches

long. The papillse diminish in size and the feathers in length

on arriving at the level of the cervical vertebrae. Soon the

feathers appear to be reduced to mere hairs, and they disappear

entirely upon about half the specimen. There the papillae

are much smaller and are distinctly separated from eacii

other.

* Loc. cit. p. 173.

t Loc. cit. p. 114, pi. V., with five figures.

X ‘‘ Notes on Moa-caves in the Wakatipu District,” by Taylor White

(‘ Transactions ’ &c. vol. viii. p. 97).

§ Note added to the preceding by F. W. Hutton (ibid. p. 101).

134

M. de Quatrefagea on

Taking into account these various data and the characters

distinguisliing the Brevipennes inhabiting the other regions of

the globe, we may form a very precise idea of what these

large species of Moas were like. They presented the general

form of the emeu [ = Cassowary], but upon a mueh larger

scale. Like this, they had the greater part of the neck naked ;

but they were destitute of the characteristic crest, and in this

respect resembled the emou \promo2us~\. Very probably the

legs were naked and the body was covered with silky plumes,

in which darker or lighter and more or less reddish tints of

brown predominated, variegated with black and white, at

least in some species.

Documents, to which I shall have to return further on,

enable us to complete this picture, and make known to us the

mode of life of these strange birds^. The Moas were slug-

gish and stupid animals, as is shown by a proverb which is

repeated at the present dayf. They were essentially seden-

tary and went about in pairs accompanied by their young.

No doubt they sometimes disputed the field on which they

were seeking the same food, for the Maoris still, in speaking

of a struggle between two pairs of combatants, say — Tw^o

against two, like the Moas.” Their nests were formed of

various dried grasses and fragments of ferns simply brought

together into a heap. They ate various species of plants

growing upon the borders of the woods and marshes, the

young shoots of certain shrubs, &c. ; but their principal food

appears to have been the root of a species of fern which they

dug up either with the beak or with the feet. To assist in

the trituration of these articles of food, the Moas, like many

other birds, swallowed small pebbles, which, when rounded

and polished by friction in the stomach, acquire a peculiar

aspect and are still called Moa- stones by the natives, who

know them well |. But this very polish rendered them unfit

for the service which the bird expected from them, and then

he disgorged them just as do the ostrich and the emou§

Letter from Mr. John White to Mr. Travers Transactions ’ &c.

vol. viii. p. 81). Mr. Travers informs us that his correspondent has

occupied himself for more than thirty-five years in collecting all possible

information upon the past of the Maoris, that he has been initiated by

their priests into all the mysteries of indigenous knowledge, so that he

knows the history of their race better than the natives themselves.

t Extracts from a letter from F. L. Maning, Esq., relative to the extinc-

tion of the Moas (‘Transactions,’ &c. vol. viii. p. 102). The author

translates the Maori proverb by the words, “as inert (ngoikae) as a

Moa.”

J ‘ liochstetter,’ p. 180.

§ “ Note on Discovery of Moas and Moa-hunters’ Remains at Patana

River, near Wangarey,”' by J. Thorn, jun. (‘Transactions,’ &c. vol. viii.

'Moas and Moa-huntera.

135

[Dromaius]. These stones were not always of the same nature,

and varied with the localities* * * * §.

III.

The details that I have just given assume not only that

man and the Moas were contemporaneous, but also that the

disappearance of the latter is of recent date. Such, in fact, is

the conclusion to which we are led by the results of a regular

inquiry pursued in New Zealand for nearly forty years by a

great number of investigators and distinguished naturalists.

Nevertlieless, until the last few years it was quite permissible

to harbour doubts on the subject. One of the most autho-

ritative of New-Zealand geologists. Dr. Julius Haast, has

pronounced most absolutely in a very different sense. While

accepting as demonstrated the coexistence of man and the

Moas at a very distant epoch answering to our prehistoric

times, he denies that the Maoris themselves ever knew these

great birds f.

On the other hand, Mr. W. Mantell, to whom his nume-

rous researches justly give an authority upon this point, has

distinctly and repeatedly expressed the opposite opinion, and

believes that these large Brevipennes were hunted and exter-

minated at a comparatively recent epoch by these very

Maoris j;.

Lastly, Mr. Stack, who is accepted by his confreres as a

very competent judge, has adopted an intermediate opinion.

He regards the belief in the recent destruction of the Moas as

inadmissible, but does not wish to throw it back into a very

distant past§.

p. 85). A certain number of these Moa-stones have been collected and

appear in the museum at Auckland, and no doubt in other collections in

New Zealand.

* Haast, loc. eit. p. 73.

t “ Moas and Moa-hunters : Address to the Philosophical Institute of

Canterbury, 1871,’’' by Julius Haast Transactions,’ &c. vol. viii. p. 66,

1872). Dr. Haast has maintained his original opinion in other memoirs^

and in the work that he has published under the title of ^ Geology of

the Provinces of Canterbury and Westland, New Zealand,’ 1879.

J “ On the fossil Remains of Birds collected in various Parts of New

Zealand by Mr. Walter Mantell of Wellington,” by Gideon Algernon

Mantell, LL.D., F.R.S. (Quart. Journ. Geol. Soc. vol. iv. p. 225, 1848);

“Address on the Moas:” Extracts by W. B. Mantell Transactions ’ &c!

vol. i. p. 18, 1869). However, in this latter paper Mr. Mantell seems

disposed to throw further back the epoch of the destruction of the Moas

in consequence of the obscurities in the traditions which he has been

able to collect upon the subject. Mr. White’s letter already cited, and

to which I shall revert, fully answers this objection.

§ “ Some Observations on the Annual Address of the President ” bv

the Rev. J. W. Stack Transactions ’ &c. vol. iv. p. 107). ’

136

M. de Quatrefages on

To show how the question has become elucidated, and

justify the point of view which I have adopted, it is neces-

sary to enter into some details.

The bones of Moas have been met with under the most

different conditions of deposition. Sometimes they simply

rest upon the suface of the ground, or are scarcely covered

by a few centimetres of sand*. But usually they are found

buried at various depths in the sands of the seashore, in the

alluvia of the rivers, in the marshes, and also in caves. The

quantity of these remains accumulated in restricted spaces is

sometimes very remarkable.

In digging canals for the drainage of a marsh at Glenmark

there were obtained the remains of 144 adult and 27 young

birds t. I might cite many other examples, but I confine

myself to a summary of the details given by Mr. Booth of

the discovery made by him at Hamilton, in a small half-dried

lagoon f .

Having been informed of the discovery of a few bones, he

opened a first pit of 4 feet square, and obtained from it

fifty-six femora with a proportionate quantity of other bones.

Regular diggings were then organized. It was ascertained

that the deposit to be worked formed a sort of irregular

crescent, measuring 40 feet from one point to the other, and

18 feet across at the middle, with a depth of from 2 to 4 feet. In

this restricted space were collected about 3-J tons (more than

8500 kilogrammes) of bones, and those who took part in the

work estimated the number of Moas accumulated in this

estuary at more than 400.

These bones were very unequally perserved. A great

number fell into paste at the least contact. Hence they had

not all been deposited at the same period. But in consequence

of the conditions of interment the Hamilton swamp could not

furnish certain data as to the relative age of the deposits. It is

otherwise with the caves, which were scientifically excavated

* Dr. Hector, loc. cit. p. 115,* Dr. Haast, he. cit. p. 103 ; Kev. J. W.

Stack, loc. cit. p. 109; Rev. R. Taylor (‘ Transactions ’ &c. vol. v. p. 97).

Those bones which were seen in great numbers scattered over the ground

have rapidly disappeared. Mr. Stack endeavours to explain their per-

sistence for centuries by saying that the Maoris carefully preserved the

jungles, which, on the contrary, the European colonists have caused to

disappear. The latter, by destroying this shelter, have facilitated the action

of atmospheric agents and have thus brought about the disappearance of

these bones, which had hitherto remained intact. 1 think it useless to

point out what, in this interpretation of the facts, has but little founda-

tion and is opposed to daily experience.

t Haast, loc. cit. p. 89.

[ “Description of the Moa-swamp at Hamilton,” by D. IS. Booth

Transactions ’ &c. vol. vii. p. 123, pi. v.).

Moas and Moa-hunters.

137

by Iloclistetter. Here very distinct layers, sej)arated by a bed

of stalagmite, contained different species. At the top was

Meionorms didiformis^ at the bottom Palapteryx elephantopus.

The bones of the former seemed to be still fresh, those of the

latter were semi-fossilized. This diversity of aspect corre-

sponded with differences of chemical composition, itself con-

nected with more or less advanced alteration. The quantity

of organic matter found in the bones of Moas which have

been analyzed has proved very variable. It is sometimes

only 10 per eent. ; but sometimes, also, it rises to 30 per cent.

— a proportion almost exactly the same as is met with in

fresh bones of the ostrich*.

Hochstetter, arguing from his own personal observations

and from some previously known facts, approximated to the

opinions of Dr. and Mr. W. B. Mantell. He thought that

the extinction of the Moas could not be thrown back

several thousand years f. He regarded their existence as

alone eapable of explaining the development which the popu-

lation of New Zealand had attained:];, and attributed the

origin of anthropophagism to the deficiency of animal food

resulting from their extermination §. He consequently iden-

tified the existing race of Maoris with the hunters of the

Moas.

To sustain a very different doctrine Dr. Haast especially

appeals to geology. The bones of Moas, he says, occur

principally in the beds which were formed during the glacial

period or immediately after it|l. Having himself collected a

certain number of these bones situ^ it seems to him to be

demonstrated that these large birds represented, in New Zea-

land, the gigantie quadrupeds which inhabited the northern

hemisphere during the post-Pliocene period. Hence he does

not hesitate to refer the existence of the Moas to an epoeh as

far from the present time as that of the mammoth, the rhino-

ceros, cave-lion, and cave-bear, the remains of which are found

in European Quaternary deposits ; and he asserts that if the

Moas survived these times, geologically so different from ours,

they were nevertheless speedily annihilatedl[.

It will be seen that Dr. Haast seems to assume not only

the analogy of the glaeial phenomena which took place in

New Zealand and in Europe, but also their contemporaneity.

We have to do here with geology proper, and questions of

this kind are out of my province. Nevertheless, even accept-

ing these two propositions as true, and reasoning by analogy,

* Hochstetter, loc. cit. p. 190. f Ihid,

I ■(: Zoc. cit. p. 194. § Loc. cit. p. 196.

j| Loc. cit. p. 68. ^ Loc. cit. p. 7o.

138

M. de QuatrefagGS on

we might at once raise some grave objections to the conse-

quences which Dr. Haast draws from them with regard to the

antiquity of the extinction of the Moas.

It is very true that the great Mammalia mentioned by Dr.

Haast no longer exist and are known to us only by their

remains. But with them lived other species which survived

them, and are even still living. The monks of St. Gall still

ate the urus in the fifteenth century ; the reindeer, in Ballasts

time, descended in winter to the shores of the Caspian Sea ;

the aurochs and the elk still inhabit Poland ; the chamois,

the ibex, and the marmot are close to us. Why should all

these species of Moa have been condemned to perish with the

geological period that witnessed their appearance?

Dr. Haast no doubt would object to me that the European

Mammalia of which I have mentioned the names, and others

which it is useless to enumerate, have generally migrated

either in longitude or in altitude. But, even without bringing

tlie action of man into play, this change of habitat was im-

posed upon them by the transformation of the nature of the

climate. This had become continental^ instead of insular as

it was in glacial times. In New Zealand this was not the

case. Whatever may have been the movements of elevation

or depression of its land*, it remained isolated in the middle

of the sea, and its climate cannot have varied, at least in the

lower regions, except within very narrow limits. Dr. Haast

himself, although starting from other data than those indi-

cated by me, insists upon considerations of the same kind,

and shows very well that, in this great island, the extension

of the glaciers by no means involves the existence of a climate

much more rigorous than that of the present day f. The

general conditions of existence remaining the same, what

reason can the New-Zealand paleontologist give for regarding

the extinction of all the Moas as necessary ?

In all his writings, published to the present day, which

have come to my knowledge. Dr. Haast maintains the general

opinions indicated abovej. It would seem that they have

* Tlie ^ Transactions of the New Zealand Institute ’ contain several

memoirs explanatory of the glacial phenomena of which New Zealand

was the theatre. I need not dwell upon these, and I shall only call

attention to those of MM. Travers and Dobson, who, in expounding their

own views, have summed up those of their confreres (see “ Notes on Dr.

Haast’s supposed Pleistocene Glaciation of New Zealand,” by W. F. L.

Travers, vol. vii. p. 409, and On the Date of the Glacial Period,” by

A. Dudley Dobson, ibid. p. 440). But, upon this question, Dr. Haast’s

work upon the geology of the Provinces of Canterbury and Westland

should especially be consulted.

t Loc. cit. p. 72.

% Besides the ‘‘ Address,” above cited, Dr. Haast has published, in the

Moas and Moa-hunters.

139

for him the value of axioms capable of serving- as a criterion —

so much so that ])ositive or negative facts have no value in

his eyes, or rather cannot really have taken place, except

they agree with his tlicory. If we speak to him of more or

less complete skeletons found upon the ground side by side with

a little heap of Moa-stones, which would seem ito indicate

that the bird died upon the spot and has never been buried,

he declares that he cannot believe that these bones could have

resisted the influence of atmospheric agents for hundreds if

not thousands of years * *. If we speak to him of the recol-

lections preserved by the natives with regard to the existence

of the Moas, their external characters, their mode of life, and

the means employed in killing them, he replies, that the most

civilized Europeans have no traditions relating to the mam-

moth and the rhinoceros, and that an inferior race which has

attained only to a condition corresponding to that of our

neolithic populations cannot have preserved any relating

to an epoch separated from them by an immense number of

years f. He adds that distinguished men have vainly

inquired into the traditions in question J. He refers particu-

larly, like Hr. Colenso, to the fables which, in New Zealand

as everywhere else, have become mixed up with the recol-

lection of actual facts in the memory of peoples §. He con-

nects what is said of the Moas with vague reminiscences of

Cassowaries brought by the Maoris from their original

country ||, or with information furnished by occasional emi-

grantslj. The examination of the ovens^ exactly like those of

the present islanders, and of the remains of repasts contain-

ing bones of Moas, furnishes him with a demonstration of the

contemporaneity of certain men with those birds ** ,* but the

former, in his eyes, were an absolutely savage population,

knowing only how to chip and not to polish stone. If some

‘ Transactions of tke New Zealand Institute/ the following memoirs

upon the same subject: — in vol. iv. 1872, Additional Notes,” p. 90;

Third Paper on Moas and Moa-hurters,” p. 94, pi. vii. ; in vol. vii.

1875, “ Eesearches and Excavations canned on in and near the Moa-bone

Point Cave, Sumner Road, in the Year 1872,” p. 64 ; Notes on an

Ancient Native Burial-place near the Moa-bone Point Cave, Sumner,”

p. 54, pis. iii. & iv. ; “ Notes on the Moa-hunter Encampment at Shag-

Point, Otago,” p. 91 ; Results of Excavations and Researches in and

near the Moa-bone Point Cave, Sumner Road (Postscript),” p. 528.

Dr. Haast has also maintained his theory and the consequences which

he derives from it in his book entitled, ‘ Geology of the Provinces of

Canterbury and Westland, New Zealand,’ 1879.

* Address,” p. 71. t Ihid. p. 75.

t Ibid. p. 76 6^ spqq. § Ibid. p. 76.

11 Ibid. p. 77. 51 Ibid. p. 106.

** Ibid. p. 82.

140 M. de Quatrefages on Moas and Moa-Jmnters.

polished hacJies have been found mingled with the ancient

kitchen-middens, this, he declares, is because they were lost

or intentionally hidden in modern times, long after the hunters

of the Moas had disappeared* * * §. The latter, he says repeat-

edly, never had any thing in common with the Maoris who

occupied New Zealand at the time of the arrival of the Euro-

peans.

I think I have sufficiently indicated the mode of reasoning

and the nature of the arguments employed by Dr. Haast. I

shall not follow him here into the discussion of a number of

subjects upon which he touches, but which are only indirectly

connected with the principal question. However, I think I

ought to quote literally the conclusions with which he termi-

nates his third memoir f : —

1. The different species of the Binornis or Moas began

to appear and flourish in the post-Pliocene period of ISew

Zealand.

2. They have been extinct for such a long time that no

reliable tradition as to their existence has been handed down

to us.

^^3. A race of Autochthones ^ probably of Polynesian origin J,

was contemporary with the Moa, by whom the large wingless

birds were hunted and exterminated.

“4. A species of wild dog was contemporaneous with them,

which was killed and eaten by the Moa-hunters.

‘‘ 5. They did not possess a domesticated dog.

^^6. This branch of the Polynesian race possessed a very

low standard of civilization, using only rudely chipped stone

implements, whilst the Maoris, their direct descendants§, had,

when the first Europeans arrived in New Zealand, already a

high state of civilization in manufacturing fine polished stone

and weapons.

Moa-hunters, who cooked their food in the same

manner as the Maoris of the present day do, were not can-

nibals.

“8. The Moa-hunters had means to reach the Northern

Island, whence they procured obsidian 1|.

* Ibid. pp. 85, 104.

t Third paper, ‘ Transactions ’ &c. yoI. iv. p. 106.

X It is difficult to understand the association of ideas which Dr. Haast

here wishes to express.

§ Here, again, Dr. Haast’s idea is not easy to understand. Throughout

he carefully distinguishes the existing Maoris from the Moa-hunters.

Here he seems to regard the former as being the grandsons of the latter.

II Dr. Ilaast’s investigations were made principally in the province of

Canterbury, which is in the South Island.

implements

‘‘7. The

On the Presence of Eyes in the Shells of the Cliitonidgc. 141

9. They also travelled far into tlie interior of tliis island

to obtain flint for the raanufaeture of their primitive stone

implements.

“10. They did not possess implements of nephrite (green-

stone) *.

“ 11. The polishing proeess of stone implements is of eon-

siderable age in New Zealand, as more finished tools have

been found in sueh positions that their great antiquity eannot

be doubted, and whieh is an additional proof of the long ex-

tinetion of the Moas.”

Thus Dr. Haast here appears to be absolute in every thing,

and it is with an appearanee of absolute eertainty that he asserts

or denies faets. But we shall see that he has himself been

obliged to go baek over some of these propositions and to

reeognize that some of them are not well founded. Never-

theless the general convictions of the learned geologist have

not been shaken on this account, and we shall have to inquire

whether this persistence is justified,

[To be continued.]

XIX. — On the Presence of Eyes and other Sense-Organs in the

Shells of the Chitonidse. By H. N. Moseley, M.iV.,

F.B.S., Linacre Professor of Human and Comparative

Anatomy in the University of Oxford.

On examining a specimen of Schizochiton incisus, preserved

in spirit amongst a number of other animals dredged by

Captain W. Chimmo, R.N., in the Sulu Sea, in H.M.S.

^Nassau’ in 1871, and by him presented to the Anatomical

Department of the Oxford University Museum, I was asto-

nished to remark on the shells certain minute, highly refracting,

rounded bodies arranged in rows symmetrically ; they struck

me at once as resembling eyes, and further examination proved

that such is really their nature. On searching for eyes on the

shells of other Chitonidga I found them present in the majority

of the genera, differing, however, in each genus more or less

in structure and arrangement.

The eyes in the Chitonidas are entirely restricted to the

outer surface of the shells on their exposed areas (tegmentum),

not extending at all on to the lamina3 of insertion (articula-

* It is with this stone, often called jade, that the Maoris fabricated

their stone clubs, haches, and various ornaments. It was of great value

in their eyes, and often plays a part in their legends. Upon this point I

have given some details, borrowed from Sir George Grey, in a book en-

titled ‘ Les Polyn^siens et leurs migrations.’

Ann. & Mag. N. Hist. Ser. 5. Vol. xiv.

11

142

Prof. II. N. Moseley on the

mcntum), and never being present on the girdle or zona,

wliicli is occupied, as is well known, by various calcareous

structures, some of wliich have been carefully investigated by

Reincke *.

In the case of all the intermediate shells the eyes are con-

fined to the arose laterales, or to the line of demarcation

between the arose laterales and the area ventralis, which latter

is usually entirely devoid of them.

The eyes, which are mostly circular in outline as seen on

the shell-surfaces, measure about 5 of an inch in diameter

in BcMzocliiton incisus^ of i^oh in Acanthopleura spini-

gera^ and in CorepMum aculeatum (in which they are oval in

outline) of an inch by about -4^. In Enoplochiton they

are smaller still and only with difficulty seen at all.

The eyes appear, when viewed by reflected light with a

low power of the microscope, as highly refracting, convex,

circular spots, looking as if made of glass or crystal ; they

are surrounded and set off by a narrow zone of dark pigment,

which is the margin of the choroid seen through the superficial

shell-substance. In the centre of each convex spot is a

smaller circular area, somewhat darker, caused by the outline

of the iris, but showing a brilliant speck of totally reflected

light, due to the lens.

The entire substance of the tegmentum in the Chitonidas

is traversed by a series of branching canals, which are occu-

pied in the living condition of the animal by corresponding

ramifications of soft tissues, accompanied by abundance of

nerves. The nerves and strands of other soft tissue enter the

substance of the tegmentum along the line of junction of its

margin with the upper surface of the articulamentum. A

narrow area, perforated all over by pores, so as to have a

sieve-like appearance, here intervenes between the two com-

ponents of the shells, and in some shells the actual margin of

the tegmentum itself is perforated. In the case of the inter-

mediate shells, in most genera tli^ere are a pair of slits (incisura3

laterales) , one on either side, in the lateral lamina of insertion ;

these slits lead to two narrow tracts in the deeper substance

of the shell, which follow the line of separation between the

area centralis and the arese laterales of the tegmentum. These

narrow tracts are permeated by numerous longitudinal canals

which lodge each a specially large stem of soft tissue and

nerves, which ramifies in the substance of the tegmentum.

Corresponding with this tract on the under surface of the shell

are a series of minute openings leading into it, through which

further strands of soft tissue, possibly mostly nervous, pass

* “ Beitrage zur Bildimgsgescliiclito dcr Staclieln &c. im M antclraiido

der Cbitoneii,” Zeitsclir. fiir wiss. Zonl. r>d. xvii. S. 805.

Presence of Eyes in the Shells of the (Jliituiiidic. 143

from the surface of the shell-bed into the shell, to give the

general network of soft tissue. In tlic anterior and posterior

shells there arc usually a considerable number of such marginal

slits, each with a corresponding tubular tract and ramifying

strands of soft tissues.

The network of soft tissues contained in the canals within

the tegmentum ramifies towards the shell-surface and termi-

nates there either in eyes or in peculiar elongate bodies, which,

ap})arently, are organs of touch. These latter are long, some-

what sausage-shaped bodies, which terminate at their free

extremity in dicebox-shaped plugs of transparent tissue, which

show a somewhat complicated structure.

The tegmenta of the shells of most Chitonidas are perfo-

rated at the surface by circular apertures or pores of two sizes,

arranged in more or less definite patterns with regard to one

another and sometimes with regard to the eyes also.

The end plugs of the sense-organs above described lie in

these larger pores. From the sides of the sausage-shaped

sense-organs are given off more or less numerous fine strings

of soft tissue, which, diverging, pass to the smaller pores

above described and there terminate in very small plugs, just

like those of the larger similar organs, but less complex in

structure.

The eyes are evidently to be regarded as having arisen as

modifications of some of the organs of touch above described.

They are ‘connected with the same network as terminal

organs of its ramifications in the same manner, and have

points of resemblance to them which are convincing as to the

liomogeny of the two. The soft struetures of eaeh eye lie in a

more or less pear-shaped chamber excavated in the substance

of the tegmentum. The stalk of the pear, which forms the

canal for the passage of the optic nerve, is directed always

towards the free margin of the tegmentum, whence the nerve

reaches it. One side of the bulb of the pear is closely applied

to the outer surface of the tegmentum, and here its wall is

pierced by a circular aperture, which is covered by the cornea.

The cornea is calcareous; it resists the action of strong

boiling caustic alkalies, but collapses at once when treated

with acid. Probably some soft tissue is present in its sub-

stance, but I have been unable as yet to find it.

The cornea in sections shows itself to be formed of a series

of concentric lamellee ; its substance is continuous with the

general calcareous substance of the tegmentum at its margins.

The pear-shaped eavity of the eye formed by the shell-

substance is lined by a dark brown pigmented choroid mem-

brane of a stiff and apparently somewhat chitinous texture.

This membrane exactly follows the shape of the cavity, but

144

Prof. H. N. Moseley on the

by projeeting beyond the margin of the cornea all round forms

an iris of less diameter than the latter.

A perfectly transparent, hyaline, strongly biconvex lens is

fitted in behind the iris- aperture. The lens is composed of

soft tissue, and dissolves in strong aeetic acid, gradually but

completely, showing a fibrous structure in the process.

The optic nerve at some distance from the eye is a com-

pact strand ; but within the very long tube continuous with the

choroid — the narrow part of the pear — its numerous fine fibres

are much separated from one another and loose. The retina

is formed on the type of that of Helix^ and not, as might

have been expected, that of the dorsal eyes of Oncidium. It

is not perforated by the optic nerve, but is composed of a

single layer of very short but extremely distinct and well-

defined rods, with their extremities directed towards the light.

Beneath them is a layer or several layers of nuclei amongst

the ultimate ramifications of the nerve.

Not all the fibres of the nerve entering the eye- cavity

proceed to the retina. A large number of the peripherally-

placed fibres pass outside the retina all round, and, perforating

the choroid at its outer margin, end at the surface of the shell,

all round the area occupied by the cornea. They terminate in

small ])lugs of tissue, corresponding to those minor organs

of touch universally distributed over the shell in the smaller

pores already described — being, in fact, exactly similar and

identical structures with these. They apparently form a sen-

sitive zone round each eye, and they arise from the optic nerve

just as do the other minor sense-organs from the nerves of

the larger organs of touch. The choroid sacs of the eye

show a curious open fold or gutter leading from the bulb

superficially along the stalk of the pear, recalling curiously

the choroid fissure.

In some genera of the Chitonidae eyes are entirely absent.

This is the case with the genus Chiton, The shell in Chiton

is perforated ; the usual small and large pores and the small

and large touch-organs are present, but I have as yet found

no trace of eyes. I have examined especially (7. magnificus

and C. marmoratus. In Molpalia^ Maugina^ Lorica^ and

Ischnoehiton there appear to be also no eyes so far as a cur-

sory examination has yielded evidence to me.

The arrangement and forms of the eyes vary much in diffe-

rent genera, and will probably prove of great value in classi-

fication, which has hitherto proved so difficult a problem.

The genus Schizochiton is distinguished by having the

mantle deeply notched posteriorly in correspondence with

a deep median notch in the posterior shell. In Schizochiton

incisus the eyes are restricted to single rows traversing the

Presence of Eijes in the Shells of the Chitonkhv?. 145

lines which in the intermediate shells separate theeentral from

the lateral areas, and which correspond in position with the

marginal slits and the courses of the principal nerves. There

arc six rows of eyes on the anterior shell, two on each of the

intermediate shells, and six on the posterior shell — twenty-

four rows altogether, with an average of about fifteen eyes in

each, or in all 360 eyes. In the specimen examined all the

rows except one have the eyes arranged in a single straight row

at regular intervals, but at the base of one row there are, as

an exception, two eyes side by side. There are also a very

few irregularly scattered eyes on the lateral area, showing that

the condition here existing is probably derived from one in

which the eyes were more ancestrally diffused.

In Acanthopleura spimger the eyes are irregularly scat-

tered around the bases of the tubercles with which the surface

of the tegmentum is covered, and are confined, in the specimens

I have examined, to the region of the margins of the shell

adjoining the mantle. The eyes in this species seem to be

very liable to be broken or to flake off, in consequence of the

decay of the surface-laminae of the tegmentum. Hence those

remaining on old specimens are those probably most recently

formed by the mantle at the margin of the tegmentum. In

decalcified tegmenta of some species I have seen eyes thus

apparently in process of formation and not yet completed.

In some specimens to be referred apparently to this species I

have been unable to find any eyes at all. It will be necessary

to examine a series of specimens of various ages to discover

whether the eyes are originally more widely extended over

the shell-surface or always marginal only in this species.

In a large Corephium aculeatum^ the exposed shells of

which w^ere densely covered by a green alga, immense numbers

of eyes were found when the alga was scrubbed off, and at the

newest margin of the shell not yet encroached upon by the plant.

The eyes are very small and their corneas are oval in outline,

the long axis of the oval being directed vertically parallel

with the height of the shell. The two kinds of pores are

arranged in vertical parallel lines with great regularity, the

large pores occurring at intervals in the lines of smaller pores.

The eyes are never placed on the tubercles, wdth rows of which

the shell is covered, and which are possibly contrivances for

protecting the eyes from being rubbed and destroyed.

The eyes are present in enormous numbers. I estimate the

numbers present on the anterior shell alone at 3000, counting

only the younger ones, which are in good condition, near the

free margin of the tegmentum, and not the older eyes, more

or less destroyed by the boring of the shell by algae and

animals on the rest of the area.. On the remaining shells, at

146 On the Presence of Eyes in the Shells of the Ohitoiiida3.

a moderate estimate, reckoning, as before, oidj the eyes in

tolerable condition, there must be at least 8500 eyes.

In Tonicia mannorata the eyes have the peculiarity of being

sunk in little pit-like depressions of the shell-surface. This,

no doubt, is a contrivance for preventing them from being

worn off, and the result is that they arc all retained complete

in large old specimens. They are arranged in single, straight,

radiating rows on the anterior and posterior shell, disposed

with considerable symmetry. There are thirty-four such

lines on the anterior shell in one specimen, containing about

eighteen eyes each. On eacli lateral area of the intermediate

sliells there are from two to four similar rows of eyes, with a

few eyes grouped irregularly also. In some forms placed in

the British-Museurn collection as species of Tonicia^ there are

no eyes present ; these possibly will be found to require to

be placed in a separate genus.

In Ornithochiton the eyes are not sunk so deeply in pits,

but are disposed somewhat as in Tonicia^ though the rows

are not so regular. In Chitonellus there are no eyes and

but a scanty supply of organs of touch.

I have been unable to trace the nerves supplying the shells

and eyes directly to their source, although I have no doubt

that they proceed from the parietal (branchial) nerve, from

whieh I have traced numerous offsets proceeding in the

required direction.

I have searched in vain for any similar eyes in the shells

of Patella and allied genera. The tegmentary part of the

shell of the ChitonidaB appears to be something sui generis^

entirely unrepresented in other Mollusca. Its principal func-

tion seems to be to act as a secure protection to a most exten-

sive and complicated sensory apparatus, which in the Chitonidae

takes the place of the ordinary organs of vision and touch

present in other Odontophora, and fully accounts physiologi-

cally for the absence of these in the group. In some respects

the arrangement of the hard and soft parts curiously resembles

that existing in the Brachiopoda.

It is most remarkable that these eyes should have been

missed hitherto by all writers on the shells of the Chitonida3.

The fact is due, no doubt, to their minuteness and to the fact

that they are not very easily seen with a powerful lens in the

dried condition of the shell in most instances. In order that

they may be made most conspicuous the dried shell should

be wetted with spirit, and a lens as powerful as Hartnack’s

no. 4 objective be used.

Dr. W. B. Carpenter* observed the perforate structure of

the tegmentum in Chiton^ but did not apparently investigate

* ' Oyclopaedia of Anatomy and Physiolog-y/ artiolo Slu'll/’ p. /5G5.

Miscella)ieous.

U7

the contained soft structures, lie writes, “In the exter-

nal layer, which scerns to be of a delicate fibrous texture, but

which is of extreme density, is perforated by large canals,

which pass down obliquely into its substance, without pcnc'

trating, however, as far as the middle layer.”

My father-in-law. Dr. Gwyn Jeffreys, has pointed out to

me that Costa* figures what are evidently the eyes on one of

the intermediate shells of a very small species of Chiton

[Tonicia] ruhicundus. They are figured as mere black dots

and referred to as fine punctuations, but their arrangement is

correctly shown.

The late Dr. Grayf, in his well-known paper on the struc-

ture of the Chitons j wrote: — “ The greater number of species

have a part of the valve which is not covered by the mantle,

but exposed. This exposed part consists of a perfectly dis-

tinct external coat, 'peculiar^ I helieve^ to the shells of this

family. The outer coat of these valves is separated from the

lower or normal portion by a small space, filled by a cellular

calcareous deposit, which is easily seen in a section of the

valves.”

I have prepared drawings illustrating the arrangement and

structure of the eyes and other sense-organs in the shell in

various genera of Chitonidge, and hope to publish them with a

more complete account of my results in the coming winter.

I beg to express my best thanks to Dr. Gunther for giving

me every facility in making use of the British-Museum col-

lection. Dr. Woodward kindly went over the fossil Chitons

in the Pala3ontological Department with me, but we could

detect no traces of eyes in any of them. This is remarkable,

since the ancient forms of the group appear to be allied to

Schizochiton.

MISCELLANEOUS.

On the Suhmaccillary in Masticating Insects.

By M. J. Chatix.

The maxilla in masticating insects is supported by a basal piece

the functional importance of which cannot be disputed, but which

possesses a still greater interest from the point of view of the mor-

phology of the parts of the mouth and even of the appendicular

organs considered generally. Nevertheless it has hardly been even

mentioned by a few writers, among whom we must cite Kirby and

Spence, who gave it the name of the cardo (hinge), a term happily

enough representing its mode of articulation ; BruUe gave it the

name of suhmaxillary, which I here retain, so as not to introduce

any neologism into an exposition already full of details.

* ‘ Fauna di Napoli : Auimali molli, Chitone,’ taf. iii. fig. l,e.

t J. E. Gray, On the Structure of the Chitons," Phil. Trans. 1848.

148

Miscellaneous.

In order to acquire a sufficiently exact knowledge of the funda-

mental characters of the submaxillary and of the variations which

it may present, it is indispensable to multiply the objeets of inves-

tigation and to select them with care, not limiting observations to

a few common species which have been almost exelusively studied.

Oligotoma Saundersii may be taken as a starting-point for this

series of analytieal and comparative investigations. Its submaxillary

in fact is very simple ; it has the appearanee of a small piece trans-

versely developed and rising slightly on its internal surface, where

a prominence, which will soon become more strongly marked in

other types, is sketched out.

In ^dipocla cinerascens the form is already eonsiderably modified,

chiefly as regards the configuration of its lower surface. This is not

only destined to limit the submaxillary towards its base, but it has

also to provide for the articulation of the maxilla considered as a

whole ; the ginglymus, scarcely represented in Oligotoma by slight

sinuosities, here gives rise to the formation of deep cavities which

impress a peculiar physiognomy upon this region of the submaxillary.

Entomologists have long since indicated the genus (Edipoda as one

of those in which the maxilla is most firmly articulated with the

head. It will be seen that this remark fully agrees with the results

of anatomical analysis.

In Decticus verrucivorus the general aspect undergoes further

changes, the origin of which must be sought in the inner and outer

surfaces, but no longer on the basal surface. Each of the lateral

surfaces commences with an inferior tuberosity; then comes an

excavated middle part, surmounted by an upper portion, which

is very prominent, especially at the outer surface. From this

results a most singular form, which can only be correctly interpreted

when we examine the submaxillary isolated and freed from the

surrounding parts.

This dissection, always delicate, is particularly difficult in Gryllus

domesticus, the submaxillary of which x>resents an appearance which,

more than in the preceding types, justifies the name selected by

Kirby and Spence ; the depressions and articular facets of the infe-

rior and superior surfaces, the orientation of the piece and its rela-

tions, all concur here to form a regular hinge.

On the other hand, the articulation of the maxilla is very feebly

constructed in Phasma japetus, in which several of the characters

proper to Gryllus domesticus are effaeed. This tendency is still more

strongly marked in Mantis rdigiosa\ the submaxillary, chiefly

developed vertically, becomes in that species almost abnormal, and

in its general conformation greatly resembles some maxillaries.

In the great green grasshopper (Locusta viridissima) it bettor

displays the double part assigned to it, of securing the articulation

of the maxilla and forming for it a sufficiently solid base to support

the whole organ, and thus to seeond or even replace the maxillary.

Thus the inferior surface is deeply excavated, while the transverse

dimensions become more appreciable.

The relative proportions of the different parts of the submaxillary

Miscellaneous,

14y

are so profoundly modified in HydropMlus picem that wo have some

difficulty in recognizing them, especially in a rapid examination.

The inferior surface is undulated and the outer surface rather short ;

the inner surface presents a marked obliquity and bears a tuberosity

which claims our more particular attention, because this arrange-

ment, indicated in Oligotoma Saundersii &c., tends to become

general in many other masticating insects.

The mandibles, as is well known, play the most active part in

the division and mastication of food ; but the maxillae also assist in

the operation to a variable extent according to the species, and the

inferior projection of the inner surface from this point of view

acquires particular importance. It did not escape Latreille, who

sometimes mentions it under the name of molar. It is pretty con-

stantly met with, but it presents frequent modifications. I confine

myself to indicating the following : —

In Carahus auratus this prominence occupies an intermediate

position between the lower and the inner surface ; in Forficula

auricularia it becomes conical and represents a lacerating rather

than a grinding tooth ; in Blaps producta it seems to be Wanting,

but its absence is compensated by a peculiar arrangement : the

submaxillary considerably exceeding the maxillary, especially

within, the inner surface of the submaxillary comes to project at

the base of the maxillary, and may thus in its entirety fulfil the

function generally reserved for the “ molar” above indicated.

Although reduced to their essential points, the preceding descrip-

tions suffice to show on the one hand all the interest that attaches

to the morphological study of the submaxillary, and on the other

the variations presented by this piece, which is too often overlooked,

but the correct interpretation of which is indispensable in the com-

parative investigation of the appendicular organs in the Arthropoda.

— Comptes Bendus, July 7, 1884, p. 51.

On a new Type of the Class Hirudineae.

By MM. PoiKiEK and A. T. de Eochbrtjne.

As the crocodile lives in the water, says Herodotus, the interior

of his mouth is covered with Bdellas (Lib. II. Chap. Ixviii. p. 94,

ed. Muller). The translators of the Greek historian, down to

Scaliger, understood the word fideWewv to refer to leeches ; since

then several have asserted that these animals were Diptera of the

genus Culex. The scientific researches of one of us during a pretty

long sojourn in Senegambia enable us definitely to settle a still

controverted question, and to prove that the BdeUas of Herodotus

must be referred to the class Hirudineae.

The remarkable type under consideration lives attached not only

to the buccal mucous membrane of Crocodilus vulgaris, cataphrac-

tus, and leptorhynehus, but also to the lingual papillae of Gymno-

plax cegyptiacus and to the interior of the pouch of Pelecanus crispus

and onocrotalus.

In its general form and the presence of branchial tufts on each

Ann. (fc Mag. N. Hist. Ser. 5. Vol. xiv. 12

150

Miscellaneous.

Bide of the body it approaches, at the first glance, the genus Bran-

chellion ; hut by the peculiarities of its organization it dilfers from

all known forms. We shall refer particularly to the following: —

Bifjesiive apparatus. — The first part of the digestive tube presents

the characters of that of the leeches with a proboscis — an exscrtile

proboscis, followed by an oesophagus with very thick muscular

walls, of which the lumen of the canal shows a transverse lozenge-

shaped section. The diameter of this organ goes on regularly in-

creasing as far as the level of the first segment provided with

branchiae. At this point it opens into a ver} wide intestine with

thin walls, presenting seven pairs of lobes, which ramify in the

digitate branchial tufts borne by the segments of this region. The

intestine is then continued into two long caeca, extending to the

hinder part of the body of the animal. Eetween these caeca passes

the very slender rectum, which bears laterally four pairs of very

sinuous tubes, placed between the dorsal wall and the caeca.

As appendages of this digestive tube, which is so remarkable for

its prolongations into the branchiae, we must mention some large

unicellular glands with finely granular contents placed on each side

of the oesophagus, the very long excretory ducts of which penetrate

into the walls of that organ, in which they ascend to a greater or

less distance, and finally open into the internal cavity. These are

the salivary glands.

Numerous glandular cells, probably hepatic, cover the walls of

the lobate intestine.

Generative organs. — The male genital apparatus consists of four

pairs of ovoid testes, situated in the last four segments with branchiae.

The epididymes, placed in the second branchiferous segment, form

two cellular masses, in the interior of which the deferent ducts make

numerous circumvolutions. These ducts, on issuing from the epi-

didymes, unite in the median line to form a short unpaired spermatic

duct, which penetrates into a large muscular sac, into which the

very large penis can enter. The male aperture is situated in the

eighth segment, or that which precedes the branchiferous segments.

The female apparatus is formed by two very long pyriform

ovaries and two slender oviducts opening into a very small matrix ;

the female aperture is situated in the ninth segment.

Circulatory apparatus. — The circulatory, like the digestive appa-

ratus, presents some remarkable peculiarities. The dorsal vessel

furnished with sacs of the proboscis-bearing leeches does not exist.

There are instead two pairs of lateral vessels, superposed, which

send forth ramifications into the branchial tufts. In the digitations

of these branchiae these ramifications ai e placed in communication

with each other by numerous transverse circular canals.

The superior lateral vessels, which we may regard as arterial,

communicate with each other in each segment by an annular vessel

which sends forth fine ramifications to Ihe surface of the skin.

Anteriorly these two vessels unite a little above the eyes, and emit, in

front and into the thickness of the tissues, branches which unite with

others, emitted by an anterior ring proceeding from the ventral vessel.

At the posterior part of the body of the animal these two lateral

MisceUaneous,

151

canals bifurcate and unite with each other by the branches tlius

formed ; at this point these vessels emit rmmorous branches, which

ramify upon the inferior surface of the disk and flow into a double

circular vessel which runs along the margin of this disk.

llesides these lateral vessels, the circulatory apparatus includes a

median ventral vessel enveloping the nervous s}stem. At the ante-

rior part this vessel gives origin to a ring, the ramifications of which

unite with those proceeding from the two superior lateral vessels ;

at the hinder part this ventral vessel passes above the canals which

unite the lateral vessels, and gives origin to numerous ramifications

which open into tho circular vessels of the margin of the disk.

Nervous system. — The nervous system, which is very like that of

Cle]ysine., besides the cerebrum and the posterior mass, consists of

eighteen ganglia, each formed of two pairs of large lateral vesicles,

and two rather smaller ventral vesicles placed one behind the other.

Each ganglion emits on each side a single nerve, which shows itself

further on.

The eyes, two in number, are very large, of an orange colour and

cup-like shape.

The integuments, especially in the anterior part, are very rich in

large glandular cells with granular contents.

The very peculiar arrangement of the circulatory and digestive

apparatus, as we have just described them, appear to us to combine

a set of characters sufiicient to authorize the formation not only of

a genus but also of a family. This family, the position of which

seems to be indicated in the neighbourhood of the Ehynchobdellidae,

we shall designate by the name of Lophobdellidse, derived from the

word Lophohdella (from Xd^os, a tuft, and /3deXXa, a leech), which

we propose as the name of the genus. The species from Sene-

gambia and the African rivers may bo inscribed under the name of

Lophohdella Quatrefagesi. — Coinptes Rendus, June 30, 1884, p. 1597.

On a new Type of Elastic Tissue observed in the Larva of Eristalis.

By M. H. Yiallanes.

There are few naturalists who have not had occasion to observe

the singular movements of the respiratory tube which terminates

the body of the larvse of Eristalis. This tube, which is composed,

like a telescope, of cylinders fitted one into the other, can, at the

j)leasure of the animal, be greatly shortened or lengthened to seek

the air at the surface of the water. The elongation of the respi-

ratory tube is effected by means of the contractions of the body,

which drive the cavitary liquid into it. Its shortening is produced

by special muscles and by elastic bands lodged in its interior.

It is to the structure of these latter parts, which, at least so far

as I know, have not been investigated, that I wish now to call

attention. Each of these elastic bands is a single cell, but con-

structed in such a way as to perform the part of a thread of india-

rubber. One of these elements, examined in a half-retracted state

and in the blood of the animal, presents the following characters : —

its cell-body is fusiform ; one of its extremities is attached to the

152

Misaellaneous.

neighbouring integuments, the other drawn out into a long pro-

eess, which is likewise attached to the inner surface of the respi-

ratory tube. The cell and its prolongation are lined with a thick

but very elastic membrane. In the centre of the cell-body we

observe a very large spherical nucleus ; this is surrounded by an

abundant protoplasm which fills the whole cell as well as its process.

It must be noted that around the nucleus the protoplasm is opaque

and strongly granular, while elsewhere it is transparent.

In the interior of the element that we have just described there

is developed a long elastic fibre, exactly similar in physical pro-

perties to the elastic fibres which are observed in the cervical ligament

of a mammal for example. It appears, in fact, under the aspect of

a perfectly cylindrical refractive thread, rectilinear when stretched,

curled and coiled up when left to itself ; further it may be remarked

that it is unalterable by acetic acid and by potash.

In the cell that we have described above the elastic fibre is

coiled upon itself a great number of times around the nucleus,

in the granular part of the protoplasm, and extended in a straight

line into the prolongation of the cell, at the extremity of which it

terminates. The elastic fibre is attached by one of its ends to the

terminal extremity of the prolongation ; by the other it amalga-

mates with and attaches itself to the protoplasm of the cell by means

of a sort of branched enlargement.

When traction is applied to the prolongation of the cell the latter

stretches out entirely, and at the same time the coiled portion of the

fibre is unrolled ; if it be left to itself it shortens, at the same time

that the fibre coils up again in the cell-body.

The facts just described seem to me to be interesting upon various

accounts. In the first place they prove once more to what degree

of complexity a simple cell may attain ; in the second, they seem to

me to throw a new light upon the morphology of the elastic tissue,

since they show us that in this tissue the active part, the elastic

fibre, may be developed either in the intercellular substance (Verte-

brata) or in the protoplasm of the cells themselves, as I have just

described in Erisialis.

I may remark that striated muscular tissue presents analogous

variations, since we see its active parts, the fibrillse, sometimes be-

longing really to the protoplasm of distinct cells (striped muscular

fibres of the heart), sometimes developed at the expense of the

fundamental undivided substance which separates the musculo-

genous cells (alar muscles of insects).

Thus it would seem that one and the same tendency presides

over the advance of the elastic tissue and that of the muscular

tissue, since in both cases, in proportion as the advance is

produced, we see the mechanically acting parts (elastic fibres,

striped fibrils) quitting the protoplasm of the cells to M'hich they

belonged originally, to be developed in the intercellular substance

and thus become the undivided property of neighbouring cellular

elements. — Comptes Bendus, June 23, 1884, p. 1552.

THE ANNALS

AND

MAGAZINE OF NATURAL HISTORY.

[FIFTH SERIES.]

No. 81. SEPTEMBER 1884.

XX. — On some Peculiarities in the Geographical Distribution

and in the Habits of certain Mammals inhabiting Continental

and Oceanic Islands'^, By G. E. DOBSON, M.A., F.R.S.

The geographical distribution of Mammals inhabiting con-

tinental and oceanic islands has been lately so ably treated

of by Mr. Wallace, in his work ^ Island Life/ that I do

not purpose entering upon the subject from a general point

of view, but will limit my remarks to some peculiarities

of distribution which have attracted my attention while

engaged in the special study of certain Mammalian orders : I

refer particularly to the Chiroptera and Insectivora.

It is an interesting fact, not hitherto noticed, that many of

the most characteristic species of the Chiropterous fauna of

Australia have their nearest allies, not in the Oriental, but in

the Ethiopian Region, thus contrasting remarkably with the

avifauna. The peculiar genus Ghalinolobus is represented

only in Africa south of the equator and in Australia, a single

species extending into New Zealand. Again, the species of

the subgenus MormopteruSj which belongs to a genus {Nycti-

nomus) of world- wide distribution, are limited to the same

* Read before tbe Biological Section of the British Association for the

Advancement of Science at the Montreal Meeting, August 29, 1884.

Ann. (k Mag. N. Hist. Ser. 5. Vol. xiv. 13

154 Mr. G. E. Dobson 07i certain Mammals

zoological regions, being fonnd only in Africa south of the

equator, Madagascar, the Mascarene Islands, Australia, and

Norfolk Island. The presence of a species of this genus in

Norfolk Island and its absence from New Zealand is very

remarkable, for, as I pointed out for the first time about ten

years ago, one of the two known New-Zealand bats, namely

Chalinolohus tuber culatus^ is also common in Australia.

The species of the extraordinarily specialized genus Mega^

derma have their head quarters in the Oriental and Ethiopian

regions ; yet the largest species, not only of the genus, but

also of all known insectivorous bats, namely M. gigas^ lately

described by the writer from Central Queensland, has its

nearest ally, not in any of the Oriental species, but in M. cor

from Eastern Africa. Another very remarkable leaf-nosed

bat, the type of my genus Tricenops^ found in Madagascar,

Eastern Africa, and Persia, but unknown in the well- searched

Oriental region, has its nearest and only ally in Rhino-

nycteris aurantia of Australia, the type of another very

peculiar genus. Finally, Australia agrees much more closely

with Madagascar and the Mascarene Islands than with the

Oriental region in the species of the large genus Pteropus^

for, while species of the section of which Pt. vulgaris of Mada-

gascar is characteristic are well represented in the former

regions, they are absent from the latter. Furthermore it is

noticeable that, while 80 per cent, of the species of the genus

inhabit the Australian region and Madagascar with its

islands, a single species only has found its way to the great

continent of Hindustan and to Ceylon.

How can we account for this resemblance of the Australian

and Ethiopian regions in certain very peculiar species of bats

while their birds differ so conspicuously ?

In the first place, to account for the presence of closely

related species in both continents, it is necessary to postulate

the existence of some land connexion, not necessarily con-

tinuous, between them ; but that such connexion was not by

way of India appears evident from the absence of such species

in that country or in the islands connecting it and the Malay

peninsula with Australia.

W e are therefore obliged to suppose that at a comparatively

recent period a chain of islands connected these continents,

the islands being sufficiently far apart to prevent the entrance

of terrestrial mammals, yet near enough to permit of the

occasional passage of some of the flying species ; still it must

be remembered that, in estimating such distances, the narrow

strait between the Comoro Islands and the coast of Africa,

about 180 miles wide, has sufficed to limit the western distri-

inhahiting Continental and Oceanic Islands. 155

bution of tlie flying-foxes, for not a single speeies of Pteropm

is known from Africa, though they abound in Madagascar

and the Comoro Islands. On tlie other hand, the insectivorous

bats, with much greater powers of flight, are very similar in

Madagascar and Africa.

But it may be urged that such propinquity of islands to

one another and to tliese continents would also permit inter-

change of the avifaunas.

To this the following reply may be made : — That the

existence of a complete chain of islands separated by suffi-

ciently narrow straits may have existed for a short period

only, the completeness of the chain being, perhaps, dependent

on some volcanic group, which may have disappeared as sud-

denly as it came into existence. Under such circumstances

bats would be much more likely to establish themselves

successfully in the new continental lands open to their migra-

tions for the following reasons : — (1) that the food of both the

frugivorous and insectivorous species is of a more general

character than that of birds, few of the species of which are

so omnivorous, within these limits, as the bats ; (2) that the

nocturnal habits of the bats would enable them to escape ob-

servation from enemies always sure to recognize the presence

of solitary individuals.

It may now be urged that if we acknowledge the effect of

such circumstances in favouring the distribution of bats, we

ought then to expect to find more bats than birds in all

oceanic islands. Such an objection may be easily disposed

of when it is remembered that volant insects are very scarce

in all oceanic islands, whereas they are abundant in all con-

tinents, and, furthermore, that a straggling bird on arrival at

an oceanic island would encounter far fewer enemies than it

would meet in a continent, and, owing to its power of seeking

its food on foot as well as on wing, would also be much more

likely to survive than the thoroughly aerial bat.

It is, I believe, to a great extent, on this very principle, that

the Chiropterous fauna of New Zealand is so limited ; that,

as yet, two species of bats only, represented apparently by

few individuals, are known from these islands, while in the

British Isles, wffiich about equal them in extent, there are

eight times the number of species, and, probably, a far greater

proportion of individuals. The striking paucity of winged

insects which, in other countries of corresponding climate,

form wholly the food of the bats, has evidently, in a great

measure, not only caused this remarkable difference, but, as I

pointed out some years ago for the first time, has led to a

change in the structure of one of the two species comparable

13*

156 Mr. G. E. Dobson on certain Mammals

to that of no other species of bat. This species, Mystacina

tuberculata^ has the claws of the pollex and toes remarkably

elongated, very acutely pointed, and provided at the base of

each with a small talon projecting from its concave surface

near the base ; the wings are peculiarly folded so as to occupy

the least possible space, and they and tlie interfemoral mem-

brane are preserved from injury by being encased, when so

folded, in a specially thickened part of the wing and inter-

femoral membrane, analogous to the thickened part of the

anterior wings in Hemiptera and to the elytra of the Coleo-

ptera ; furthermore, the plantar surface of the foot, including

the toes, is covered with very soft and very lax integument

deeply wrinkled, and each toe is marked by a central longi-

tudinal groove, with short grooves at right angles to it, as in

the species of the Gecko genus Hemidactylus. All these

peculiarities of structure must accompany some corresponding

peculiarities in the habits of this species. There can be little

doubt that the denticles at the bases of the claws of the

thumbs and toes give additional grasping-power to these

organs, and this, taken into consideration with the peculiar

manner in which the wings and interfemoral membrane are