aig yey, ae ayes tate aoe Bah satel ‘ owe bier Shee ae py, ps rit) s ii ce Ey ay Wives ? Fe ea PS ve UBe Mar See : | AM ri; ‘ as; f, ‘4 i iid airy Ay, AQT , ay i Pay Prue bX ; Men a Ta ie prot war ine 52 a THE ANNALS AND MAGAZINE OF NATURAL HISTORY. INCLUDING ZOOLOGY, BOTANY, ann GEOLOGY. (BEING A CONTINUATION OF TILE ‘ANNALS’ COMBINED WITIE LOUDON AND CHARLESWORTII'S ‘ MAGAZINE OP NATURAL HISTORY.’) CONDUCTED BY ALBERT C. L. G. GUNTHER, M.A., M.D., Ph.D., F.B.S., WILLIAM CARRUTHERS, F.R.S., F.L.S., F.G.S., AND WILLIAM FRANCIS, Ph.D., F.L.S. a » x0 LONDON: PRINTED AND PUBLISHED BY TAYLOR AND FRANCIS. SOLD BY SIMPKIN, MARSHALL, HAMILTON, KENT, AND CO., LD. ; WHITTAKER AND CO.: BAILLIERE, PARIS: MACLACHLAN AND STEWART, EDINBURGH: HODGES, FIGGIS, AND CcO., DUBLIN: AND ASHER, BERLIN, 1893. “Omnes res create sunt divine sapientix et potentia testes, divitiz felicitatis humane :—ex harum usu bonitas Creatoris; ex pulchritudine sapientia Domini ; ex ceconomid in conservatione, proportione, renoyatione, potentia majestatis elucet. Earum itaque indagatio ab hominibus sibi relictis semper sxstimata ; & veré eruditis et sapientibus semper exculta; malé doctis et barbaris semper inimica fuit.”—Linnavs. “Quel que soit le principe de la vie animale, il ne faut qu’ouvrir les yeux pour voir qu’elle est le chef-d’ceuvre de la Toute-puissance, et le but auquel se rappor- tent toutes ses opérations.’—Bruckner, Théorie du Systéme Animal, Leyden, 1767. wee a FM a fees. e ee whe sylvan powers Obey our summons; from their 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, AU, all to us unlock their secret stores And pay their cheerful tribute. J. Taytor, Norwich, 1818. y ’ CONTENTS OF VOL. XI. [SIXTH SERIES.] NUMBER LXI. Page I. On some Points in the Morphology of the Arachnida (s. s.), with Notes on the Classification of the Group. By R. I. Pocock, of the British Museum (Natural History). (Plates I. & IL) ........ 1 IL. Descriptions of Thirteen new Species of Terrestrial Mollusea from South Africa. By James Cosmo Metvitt, M.A., F.LS., and Joun Henry Ponsonsy, F.Z.S. (Plate IIL)............+- 19 Ill. Additional Notes on the Origin of the Trachere from Seti- parous Glands. By Henry M. Bern ARD, M.A. Cantab. (from the Huxley Research Laboratory) ..........2-.0--eceeoeececneres 24 IV. On the Terminal Organ of the Pedipalp of Galeodes and the Discovery of a Homologous Organ on the Pedipalp of Phrynus. By Henry M. Bernarp, “MLA. Cantab. (from the Huxley Research Re TER NRE Nere set ota ale Gad ence enna wee ape OP eee Fe oe aes 28 V. On the Parley olOey: of = River-Lamprey. By Px. Ows- NE ea ee on sa eel aie nin epee cg oo oe aig Os 30 VI. Notes on Apteryx Haastii. Bythe Hon. WALTER Roruscritp 483 VII. Some Observations on the Mouth-orzans of Diptera. By ere OINV ATER HOUBEY 2°50 os ijeyidiv as dale» caine mawinlalaw o's ss 45 VIII. Description of a new Baboon from East Africa. By Oxp- MMMREDMEUOMEAS 1m Ceira tiie; S's 3]s Pate esr otis om supe me nibinge Sela vie se Weale eee 2s 46 IX. On a new Cephalolophus from Mount Kilima-njaro. By SreerrmBmE SE TIOWUNG: £45455 5/55 0s aie winieleiwe « Sek ia So t's wipes 44 © 48 X. On the Mexican Representative of Sciurus Aberti. By OLp- HURTS UT OWU SIS aD ie neh clare o's 5 \acaye eye sian n! oie shecal uaa; o so > Mo oe a 49 XI. Note on the History of the so-called Family Techonide. By ArtHuR Denpy, D.Sc., F.L.S., Melbourne University .......... 50 XII. Description of a new Species of the Cicadan Genus Pecilo- wearer, Seay Wi oy, LIDAR AE. ta Ss ae harah wecclen wiv ans » aed’ 52 XIII. Notes on two Genera of Coreid@ found in Madagascar. By OE ea aig ese oe ip a ee ar a ca anh aa 53 XIV. On the Phy malay | of the Rattle of Crotalus durissus. By A. £. Froxristow, M. yh cD REL cee eer 54 New Books:—Catalogue of Eastern and Australian Lepidoptera Heterocera in the Collection of the Oxford U niversity Museum. —Part I. Sphinges and Bombyces. By Col. C. Swrnnog, F.L.S. &c.—Fossil Plants as Tests of Climate. By A. C. WATE Ds WP coe ic bales noc bate SOR ite taboo 58—G0 _ iv CONTENTS. Page Comparative Researches upon the Organization of the Brain in the principal Groups of Arthropods, by M. H. Viallanes ; On the Circulation of the Blood in young Spiders, by M. Marcel Causard ; A Contribution to the Knowledge of the Anatomical Structure of the Sexual Organs in the Galeodide, by A. Birula, of the Zootomical Institute of the University of St. Petersburg ; On Two Species of Myzostoma parasitic upon Antedon phalan- gium, Miiller, by M. Henri Prouho..........0+- +200 eee 62—70 NUMBER LXI. XV. Natural History Notes from H.M. Indian Marine Survey Steamer ‘ Investigator, Commander C. F. Oldham, R.N., com- manding.—Series II., No. 7. An Account of the Collection of Deep- sea Asteroidea. By A. Aicock, M.B., Surgeon-Captain I.MLS., late Naturalist to the Survey. (Plates IV.—VI.) ........0.0ee0eesees 73 XVI. Report upon the Myriopoda of the ‘ Challenger’ Expedition, with Remarks upon the Fauna of Bermuda. By R. I. Pocock, of the British Museum (Natural History). (Plate IX.) ............ 121 XVII. The Influence of Light on the Coloration of Crustaceans. By A=. MALARD 0. uci cc cee eee nes vaames nes o¥ coe shimeipe ee 142 XVIII. Observations upon Ameaba, with especial reference to the existence of an apparent Micro-nucleus in that Organism. (Prelimi- nary Communication.) By Joun E. 8. Moors, A.R.C.S, (from the Huxley Research Laboratory, R. Coll. Sei. Lond.), (Plate XIi.).. 149 XIX. On the probable Sensory Nature of the “ Appendix ” of the Antennz of Coleopterous Larve. By Caries J, GAnan, M.A., of the British Museum (Natural History) ...........+eseseecees 154 XX. Classification of the Pelecypoda: Fischer’s Families re- arranged in accordance with Pelseneer’s Scheme. By B. B. Woop- WARD, F.G.S., F.R.M.S., of the British Museum (Natural History). 156 XXI. Notes on Apteryx Haasti. By H.O. Forsrs .......... 159 XXII. Natural History Notes from H.M. Indian Marine Survey Steamer ‘Investigator,’ Commander R. F. Hoskyn, R.N., com- manding.—Series II., No. 1. On the Results of Deep-sea Dredging during the Season 1890-91. By J. Woop-Mason, Superintendent of the Indian Museum, and Professor of Comparative Anatomy in the Medical College of Bengal, and A. AxLcocx, M.B., Surgeon I.M.S., Surgeon-Naturalist to the Survey. (Plates X.& XL) .... 161 XXII. Aglia tau, a connecting-link between the Ceratocampide and Saturnide, and the Type of a new Subfamily, Agline. By ALPHEUS S. PACKARD .......... gs aaa as e's b ob iie'e 1 Stee 172 XXIV. Contributions towards a General History of the Marine Polyzoa, 1880-91—Appendix. By the Rev. Tuomas Huncks, S00 a od?
(Plate XR fios: 47) °. oe a sad d's abs eae 377
LXII. On some allied Pentatomide, with Synonymical Notes. By
re ER iS are aia Fe igre eo AO Ie Se ai NE Die bac ere aaa 389
LXIII. On a new Beetle from Japan (Omaliidz). By G. Lewis,
2 Pa Aine SS eee eee er Ars perenne tre rece 394
LXIV. Notes on the Genus Acronycta of Authors and its Position
in the Classification of Heterocerous Lepidoptera. By A. G. Bur_er,
a Ee eae nit alae nis sins ae S #561 males were OOO
LXV. Notes on some Mexican Oryzomys. By OLpFIELD THomas. 402
~_ LXVI. Description of a new Species of Perognathus from Colorado.
PePOUMERTE DCE ROMAN 1 lias eos pee Sadie peceweene dees 405
LXVII. Description of a new Species of Tree Trap-door Spider
from Trinidad. By R. I. Pocock. (Plate XIX. figs. 1-3.) ...... 407
New Book :—The Faunaof British India, including Ceylon and Burma.
a mete, VOL L. By GF, ELAMPESON | os 0c 5 eee ary db ante ee 410
New Observations on the Affinities of the different Groups of Gastro-
pods (Expeditions of the Yacht ‘Hirondelle’), by M. E.-L.
Bouvier; On the Branchial Sense-Organs of the Patellide, by
Dr. J. Thiele, of Dresden ; On Cirripedes and other Crustaceans
commensal with Mediterranean Turtles, by MM. E. Chevreux
ND AO AMBER Gc 2,2 o5:ccciu.pig wa loatinns Hoviing wean nen KA ee 411—414
NUMBER LXVI.
LXVIII. On new Species of Histerid@, and Notes on others. By
bree bbs). PE late Ake AC. se ries Sets orodn we eine ee oe ae 417
LXIX. On new and little-known Tessaratomine of the Order
Bennie SUN. Ls PISPANT 220 see nes ce ss +s + v'sleSe ens 450
LXX. The Range of Placostylus: a Study in Ancient Geography.
eg ere PCE ean rate. a rhein os we'd «(a 2,0 shee apa 435
LXXI. Note on Mesoplodon bidens. By the Hon. Watrer
ERO HE SE LEO a Bee athe i ip ig G 2a, d's! di 0% os 0.0: xh 0.9.0, mialao pheael olan 439
LXXII. A Contribution to the Knowledge of the Genealogy and
Classification of the Crustacea. By Prof. Kari. Gropnen, of Vienna. 440
Vill CONTENTS.
Page
LXXIII. Report upon the Stomatopod Crustaceans obtained by
P. W. Basset-Smith, Hsq., Surgeon R.N., during the Cruise, in the
Australian and China Seas, of H.M.S. ‘ Penguin,’ Commander W. U.
Moore. By R. I. Pocock, of the British (Nat. Hist.) Museum.
(Ente ek kee Ne alate ee etna cke nett Canine) a cktis once sore eee 473
New Book :—Eléments de Paléontologie. Par Féi1x Bernarp &e. 479
Proceedings of the Geological Society ......0...-eseeececenneces 481
Notes on Cheropsis liberiensis (Morton), by Henry C. Chapman,
IVES ie 4 icitscs as wis..o 8 an oe etesee eas Aero oe. eke eee ee 481
LG ES ahr re PPE ene Mir sats etn om ae err mn NRE Ay OG Tae On & 484
PLATES IN VOL. XI.
B)
Prate I. | Morphology of the Arachnida.
aT.
Ill. New South-African Mollusea.
IV.
v, New Deep-sea Asteroidea.
VI.
ola New Scottish Entomostraca.
IX. Myriopoda of the ‘ Challenger’ Expedition
Se Psathyrocaris fragilis.
AA
XII. Structure of Amceba.
XIII
sl } Limnocnida tanganjice.
XIV.
XV. Mesoplodon bidens.
XVI. New Exotic Iulde.
XVII. Skull of Pisodus Oweni.
XVIII. New Land-Shells from the Philippine Islands.
XIX. New Trap-door Spider.—New Longicorn Coleoptera.
XX. New Species of Histeridee.—Stomatopod Crustaceans from
the Australian and China Seas.
THE ANNALS
AND
MAGAZINE OF NATURAL HISTORY.
{SIXTH SERIES.]
a aeaadedesscadeers per litora spargite muscum,
Naiades, et circiim vitreos considite fontes:
Pollice virgineo teneros hic carpite flores:
Floribus et pictum, dive, replete canistrum.
At vos, o Nymphe Craterides, ite sub undas ;
Ite, recurvato variata corallia trunco
Vellite muscosis e rupibus, et mihi conchas
Ferte, Dez pelagi, et pingui conchylia succo,”’
N. Parthenii Giannettasii Ec), 1,
No. 61. JANUARY 1893.
I.—On some Points in the Morphology of the Arachnida
(s. s.), with Notes on the Classification of the Group. By
R. I. Pocock, of the British Museum (Natural History).
[Plates I. & II.]
SINCE it is generally admitted that the ancestor or ancestors
of the Arthropoda must be sought for in animals resembling
the Annelidan worms in the complete segmentation of the
body, it seems clear that a species in which the metamerism
is highly developed is, ceteris paribus, more primitive than an
allied form in which it is obscurely manifested. The two
common decapod crustaceans, the crab and the lobster, furnish
a good example of the truth of this maxim, the latter animal
with its gangliated nerve-chord, its long, segmented, and limb-
bearing abdomen, being unquestionably more nearly related to
the primitive form or ancestor of the Decapoda than the crab.
Consequently in tracing the phylogeny of this group of crus-
taceans we should conclude that the Brachyura are the descen-
dauts of the Macrura, and that as such they should occupy a
higher branch of a genealogical tree.
The truth of this, however, is so very obvious that the only
excuse to be offered for its restatement is the circumstance
that not all authors have borne it in mind in dealing with the
Ann. & Mag. N. Hist. Ser. 6. Vol. xi. 1
Vili CONTENTS.
Page
LXXILL. Report upon the Stomatopod Crustaceans obtained by
P. W. Basset-Smith, Esq., Surgeon R.N., during the Cruise, in the
Australian and China Seas, of H.M.S. ‘ Penguin,” Commander W. U.
Moore. By R. I. Pocock, of the British (Nat. Hist.) Museum,
PE GTS a eka sip ese ss aah eee Rees Moke hos: xe 475
New Book :—E£léments de Paléontologie. Par Férrx Bernarp &e, 479
Proceedings of the Geological Society ......6...ceseeecsseeenees 481
Notes on Cheropsis liberiensis (Morton), by Henry C. Chapman,
WE racks 3:5 8k von oe Sm ae eele lee kia ene eh ieee 481
(Dit, eaten a ereeaws Bc ee ee RS en Ra it SoS 484
PLATES IN VOL. XI.
)
PLATE a Morphology of the Arachnida.
a,
Ill. New South-African Mollusca.
IV.
v, New Deep-sea Asteroidea.
VI.
il New Scottish Entomostraca.
IX. Myriopoda of the ‘ Challenger’ Expedition
aoe Psathyrocaris fragilis.
XI. i
XI. Structure of Amceba.
XII.
E Limnocnida tanganjice.
XIV.
XV. Mesoplodon bidens.
XVI. New Exotic Iulide,
XVII. Skull of Pisodus Oweni.
XVIII. New Land-Shells from the Philippine Islands,
XIX. New Trap-door Spider.—New Longicorn Coleoptera.
XX. New Species of Histeridee.—Stomatopod Crustaceans from
the Australian and China Seas,
THE ANN-ALS
AND
MAGAZINE OF NATURAL HISTORY,
(SIXTH SERIES. ]
eesanevansceaysance per litora spargite muscum,
Naiades, et circiim vitreos considite fontes:
Pollice virgineo teneros hic carpite flores:
Floribus et pictum, dive, replete canistrum.
At vos, o Nymphe Craterides, ite sub undas ;
Ite, recurvato variata corallia trunco
Vellite muscosis e rupibus, et mihi conchas
Ferte, Dez pelagi, et pingui conchylia succo.”’
N. Parthenii Giannettasti Ecl, 1,
No. 61. JANUARY 1893.
I.—On some Points in the Morphology of the Arachnida
(s. s.), with Notes on the Classification of the Group. By
R. I. Pocock, of the British Museum (Natural History).
[Plates I. & IL]
SINCE it is generally admitted that the ancestor or ancestors
of the Arthropoda must be sought for in animals resembling
the Annelidan worms in the complete segmentation of the
body, it seems clear that a species in which the metamerism
is highly developed is, ceteris paribus, more primitive than an
allied form in which it is obscurely manifested. The two
common decapod crustaceans, the crab and the lobster, furnish
a good example of the truth of this maxim, the latter animal
with its gangliated nerve-chord, its long, segmented, and limb-
bearing abdomen, being unquestionably more nearly related to
the primitive form or ancestor of the Decapoda than the crab.
Consequently in tracing the phylogeny of this group of crus-
taceans we should conclude that the Brachyura are the descen-
dauts of the Macrura, and that as such they should occupy a
higher branch of a genealogical tree.
The truth of this, however, is so very obvious that the only
excuse to be offered for its restatement is the circumstance
that not all authors have borne it in mind in dealing with the
Ann. & Mag. N. Hist. Ser. 6. Vol. xi. i
2 Mr. R. J. Pocock on the
phylogeny of the Arachnida. This at least seems to be borne
out by the phylogenetic tree of this group that was published
by Dr. Thorell in 1877*. From this tree it appears that the
Scorpions branch off from the Pedipalpi, the latter from the
Aranee, the Aranee from the Opiliones, &c., almost the
lowest branch of the Arachnid trunk being assigned to the
Acari.
Perhaps, so far as complexity of structure is concerned, a
Scorpion stands higher than a sheep-tick, so, no doubt, does
an active free-swimming lobster rank above a sluggish Maioid
crab. Nevertheless no one would probably on these grounds
place the Maiide in a phylogenetic tree at a lower level than
the Astacide. But the reasons that lead us to consider the
Macrura nearer than the Brachyura to the ancestor of the
Decapoda point also to the conclusion that of all Arachnids
the Scorpions most resemble the primitive type; for in these
animals the metamerism of the body is more fully expressed
than in any other order of Arachnida, all the twelve somites
of the abdomen being well developed with dorsal and ventral
representatives, the anterior six of them bearing permanent
or transitory appendages; furthermore, the nerve-chord is
furnished with a series of ganglia and the heart is divided
into a greater number of chambers.
Considering, then, on these grounds the primitiveness of
the Scorpion’s structure, we may imagine that the immediate
ancestor of all the Arachnida was constructed somewhat as
follows t :—The body was composed of eighteen somites, the
anterior six of which were provided with large appendages
set apart for locomotion and the prehension and mastication
of food; the terga of this cephalothoracic region were fused
to form a single shield or carapace, supporting a submedian
and a cluster of lateral eyes on each side, and the ventral
surface of the carapace, at least in its posterior half, was
protected by a sternal plate. Hach of the succeeding six
somites bore a pair of small ventral appendages, and the
generative aperture opened upon the sternal area of the first
of these somites. The posterior six somites had lost their
appendages, were probably narrower than the rest, and con-
stituted a limbless caudal termination to the body, the last of
them being furnished with a single plate, articulated above
the anal aperture.
The Scorpions have departed from this hypothetical type
in the following particulars :—The otherwise useless posterior
* Actes Soe. Ital. Sci. Nat. xix. p. 86.
+ Cf. in this connexion Prof. Lankester’s detinition of the class Arach-
nida in Quart. Journ. Mier. Sci. xxi. p. 647 (1881).
Morphology of the Arachnida. 3
five somites are converted into the so-called tail; the poste-
rior four pairs of appendages of the abdominal region have
disappeared in connexion with the development of the lung-
books, the second pair become the tactile sexual organs or
pectines, and the first in all probability constitute the genital
operculum *. Moreover the generative aperture has moved
forwards between the cox of the last pair of cephalothoracic
limbs, and the enlargement and ingrowth of the coxe of this
region have more or less obliterated the sternum.
_ The Arachnida which structurally come nearest the
Scorpions are the Pedipalpi. There are three existing very
distinct types of this order, Thelyphonus, Schizonotus }, and
Phrynus ; the first-named being the most Scorpion-like of
the three may advantageously be considered first. Great,
however, as is the superficial likeness between this genus and a
Scorpion, the differences are in reality very considerable. In
the first place the whole abdomen isimmensely reduced in length
by the shortening of the somites along the longitudinal axis,
the three posterior alone being abruptly narrowed to consti-
tute a small tail-like support for the filiform multiarticulated
telson ; in the second place a deep constriction separates the
cephalothorax from the abdomen. But more important than
all this is the disappearance of the two posterior lung-sacs and
the obliteration of the sternite and appendages of the second
abdominal somite by the enlargement and backward extension
of the sternite of the first, behind which the generative organ
opens. Moreover this first sternite, in addition to obliterating
the second, encroaches largely upon the third and fourth,
reducing them to narrow chitinous bands, the result being
that the pulmonary sacs that are situated in the third and
fourth somites open in front of their sterna, or, as it is usually
expressed, behind the first and second sterna f.
* I am not aware that the evidence of the appendicular nature of the
genital operculum is absolutely conclusive.
+ Aname proposed by Thorell to replace Nyctalops of Cambridge,
which was preoccupied.
t This at least seems to me to be the probable nature and extent of
the changes that have affected this region and these parts of the body.
I do not see otherwise how to account for the anomalous position of the
aperture of the pulmonary sacs behind the first and second sternites, when
these sacs belong to the third and fourth somites.
If this view and the one expressed below as to the derivation of the
Arane from the Pedipalpi is correct, it seems that the two abdominal
sternites of the spider Liphistius and the opercula of the lung-sacs of the
Mygalomorphe are the homologues of the first and second sterna of the
Pedipalpi. In this case the anterior lung-sacs belong to the somite that is
represented by the second sternite in Liphistius and by the aoe pair of
1
4 Mr. R. I. Pocock on the
Schizonotus presents a strong general likeness to Thely-
phonus in the form of the body, structure of limbs, &e, The
telson, however, consists of a single short segment, and the
carapace being rather shorter, has left a larger space between
its hinder border and the first abdominal tergite; but this
area, otherwise unprotected, is covered by asingle or a paired
sclerite, which may be either a special development or one of
the original cephalothoracic tergites. In any case the result
of this segmentation of the carapace is the power to flex the
two posterior somites in a vertical plane, the joint being
situated between the fourth and fifth appendages. There
appears usually to be a single pair of respiratory stigmata
situated behind the first sternite, as in Thelyphonus. ‘The
posterior pair that are developed in Thelyphonus appear to be
functionless, but upon the third, fourth, and fifth sterna
(morphologically the fourth, fifth, and sixth) close to the
posterior margin and behind the muscular impressions a pair
of dusky patches are visible. ‘These appear to be some
internal organs seen through the semitransparent cuticle, and
I believe they are the homologues of the three posterior pair
of lung-sacs of the Scorpion *. In one species described by
Dr. Thorell—S. Cambridgii—two eyes are present; but
most interesting of all is the disappearance of the respiratory
stigmata and the fusion t of the first two sterna of the abdo-
men, which leaves only ten for this region. ‘This species is
of interest in connexion with the possible derivation of the
Solifuge from this group.
The third group of Pedipalpi—the Phrynide—are of
interest inasmuch as in all the characters that they depart
trom the Thelyphonide they approach the true Spiders or
Aranee. The whole body is very much shorter than in the
opercula in the Mygalomorphe, while the posterior lung-sacs belong to a
somite which has no free sternite. This view is entirely opposed to
Macleod’s hypothesis respecting the derivation of the lung-books of the
spiders.
- These structures haye not, I believe, been previously noticed, but
they are certainly visible in both the specimens of. this genus that I have
seen. For affording me an opportunity to examine these examples I am
indebted to the kindness of the Rey. O. P. Cambridge, who with great
liberality sent to me the types of the two species he has described. That
named tenwcaudata is referable to the so-called genus Tripeltis of
Thorell, having the posterior cephalothoracic tergite paired.
+ Dr. Thorell’s words describing this feature are “. . . nec limitem inter
scuta ventralia lm et 2m, nec spiracula certo discernere potui” (Ann.
Mus. Genoy. xxvii. p. 560, 1889). This author ascribes only eleven terga
to the abdomen. I cannot but think, however, that he overlooked aa
first small tergite, which is present in the two specimens seen by me.
Morphology of the Arachnida. 5
Thelyphonide and wider. The telson has disappeared.
The shortness of the abdomen is brought about by the reduc-
tion in size of the three posterior somites, the terga and sterna
being represented by very short, closely approximated, trans-
verse sclerites. Moreover the anterior two tergites are also
much smaller than in Thelyphonus. The increase in the
width of the cephalothorax is accompanied by a recession of
the coxe, especially of the fourth and fifth appendages, from
the middle line, leaving a membranous space between the
anterior and posterior sternal pieces; this membrane is
chitinized from distinct centres. Furthermore, the coxe of
the second pair of appendages are freely movable, and not
fused, as in Thelyphonus.
The Aranez or Spiders agree with the Phrynide in having
a distinct constriction between the cephalothorax and abdomen,
in having the coxe arranged radially round the sternum, those
of the second pair being freely movable as maxille ; in having
two-jointed mandibles and normally eight (median and lateral)
eyes. Moreover in lower forms there are two pairs of lung-
sacs, asin Phrynus. Again, although a marked difference
between the two groups is the absence of segmentation in the
adult Spiders, yet it is interesting to note that during their
embryonic condition the ventral surface of the abdomen seems
at one time or another to be divisible into eleven sternal areas,
as in the adult Phrynus. The first of these is apodous and
represents probably, I think, the genital sternite; the second,
third, fourth, and fifth bear a pair of appendages each, while
the last six are without appendages, as in the Scorpion. The
anterior two pairs of appendages disappear in connexion with
the formation of the breathing-organs; the fate of the poste-
rior two, however, is quite unlike anything met with else-
where in the Arachnida, for they take on the form of dwarfed
limbs and constitute the spinning-mammille, which, with
their appropriate glands, are the most characteristic features
of this group. ‘Their presence constitutes the greatest struc-
tural break between the Spiders and Pedipalpi. In the
higher forms of Spiders the posterior lung-sacs are replaced
by tracheal tubes. In the lowest, i. e. Liphistius, the upper
surface of the abdomen is furnished with a series of nine
tergites, the posterior of which are very much reduced in size,
as in Phrynus, and the spinning-appendages retain their
primitive position close behind the lung-sacs.
Another order of the Arachnida, namely the Pseudo-
scorpiones, also seems to me to be tolerably nearly related to
the Pedipalpi. It is not unusual to associate this group with
the Scorpions, as Mr, Cambridge has done in his article in
6 Mr. R. I. Pocock on the
the ‘Encyclopedia Britannica ;’ but it is difficult to find
grounds to justify this classification.
In the form of the cephalothoracic appendages, especially
of the second pair, there is certainly a close similarity between
the Scorpions and the Pseudoscorpiones; but in the structure
of the abdomen the difference between the two groups is
very great. But the same cannot be said of the abdomen of
the Pedipalpi and the animals now under discussion ; for the
two groups resemble each other in the absence of the pectines
and in the presence of only two pairs of respiratory stigmata.
Moreover, in such a form as Garypus litoralis the same
number of somites can be made out in the abdomen as are
seen in this region in the Pedipalpi, namely twelve. Further-
more, there is the same inequality in the number between the
tergites and sternites, the former being one in excess of the
latter *. From the posterior somite forwards the dorsal and
ventral sclerites correspond plate for plate until the third
tergite and second sternite are reached. Here the corre-
spondence ceases, there being but a single genital sternite for
the first and second tergites, exactly as in the Pedipalpi.
The stigmata are situated in the third and fourth abdominal
somites, but they have taken up a more lateral position than
in the Pedipalpi.
In some other forms of this group one of the tergites has
disappeared, so that the abdomen is described by systematists
as being furnished with only ten of these plates. Moreover
the sterna similarly may be reduced to nine.
So far as the embryological history of these animals is
concerned, it is especially interesting to note the presence of
four pairs of provisional appendages attached to four of the
anterior segments of the abdomen. In the absence of evidence
to the contrary it seems justifiable to conclude that these
appendages are the exact homologues of the four pairs seen in
the Aranex, a group which we have seen to be apparently
nearly related to the Pedipalpi.
The next group to be considered is the Opiliones. The
animals of this order agree with the Pseudoscorpiones in the
tracheal nature of their respiratory organs and in the absence
of a “waist” between the cephalothorax and abdomen. This
last feature has led to very remarkable results in the forward
migration of the generative aperture between the coxe of the
posterior limbs, in some forms even to a position just behind
* The last somite has not, so far as [am aware, been previously recog-
nized as such, It is, except in distended specimens, almost entirely con-
cealed inside what is apparently the last, namely the eleventh, but which
is in reality the last but one.
Morphology of the Arachnida. 7
the mouth. This procession of the abdominal sternites is
accompanied by the obliteration of the anterior pair of tracheal
stigmata with the sternite that supported them. ‘The remain-
ing stigmata are situated on the first free sternite, and their
presence fixes this plate as belonging morphologically to the
fourth somite *. Since, however, there is an immense range
in structure within the Opiliones, especially touching the
number and distinctness of the segments of the abdomen, it
is not always easy to compare the somites with those of the
Pedipalpi or Chelonethi.
In Stylocellus, however, a genus in which the segmentation
of the abdomen is well developed, there are nine distinct
tergites and eight sternites, and since the presence of the
stigmata on the first sternite points to it morphologically as
the fourth, we have almost the full complement of somites
from the fourth backwards. Moreover, between the last
tergite and sternite there is a single anal sclerite which may
represent the twelfth tergite. ‘The first free tergite has no
free sternite to correspond to it; it appears to belong morpho-
logically to the third somite, since it immediately precedes
the tergite that covers the sternite upon which the tracheal
organs open. In this case the dorsal elements of the first
two somites have disappeared, and on the ventral side the
first three sternites appear to be represented in a general way
by that part of the intercoxal area of the cephalothorax
which lies behind the generative aperture.
In most forms of Opiliones, however, the segmentation of
the body is less well expressed than in Stylocellus. This
genus appears in this respect, as well as in the small number
of supernumerary maxillary sclerites, to be one of the most
primitive of the order. In the higher forms the abdomen is
much more reduced in length, and even when distinct tergites
and sternites are strongly developed, as in e. g. the Laniatores,
their number may fall as low as five; while in others, such as
Phalangium, the segmentation is represented merely by folds
of the integument, the tergal and sternal plates being un-
developed. ‘This reduction in the number of the dorsal and
ventral plates is brought about by their fusion with each
other or with the cephalothorax.
That the Acari are nearly related to the Opiliones seems
evident from a comparison between the two orders. For, as
Dr. Thorell says, “ from the Acari the Opiliones are scarcely
in all cases distinguishable by any other external character
than the structure of the abdomen, which, in the Opiliones, is
* If the third, it is not easy to explain the disappearance of the second
pair of stigmata.
8 Mr. R. I. Pocock on the
evidently segmented at least at its posterior end” *, Some
of the free-living Acari, e. g. the Trombidiide and Holothy-
ride, with their pediform, claw-tipped palpi and chelate
mandibles, bear so strong aresemblance to Stylocellus, that with
an example of this genus in hand it is impossible not to
think that the Acari are descended from the Opiliones f.
As for the Solifuge, their position is not so clear. They
seem to show affinities with the Pedipalpi through the
Tartarides (Schizonotidee) and also with the Pseudoscorpiones,
as Simon long ago suggested f.
In the first place the carapace, instead of consisting of a
single shield, is usually described as being divided into three
distinct sclerites, the posterior two of which are regarded as
the tergites of the posterior two somites of the cephalothorax.
The rest of the body, the abdomen, consists of only ten
somites, the tergites and sternites apparently corresponding
exactly in the first nine, while the last is a iia plate
perforated mesially by a vertical slit—the anus. The genera-
tive aperture is, as usual, situated upon the ventral surface
of the first, while the second and third bear the apertures
of the tracheal breathing-organs.
It is thus clear that these Arachnida fall into the same
section as the Pedipalpi, Aranez, Pseudoscorpiones, Opiliones,
and Acari, inasmuch as they are all devoid of the pectines
which are so characteristic of the Scorpions. So, too, does it
seem likely that the first sternite corresponds to that of the Pedi-
palpi, Aranez, &c., in which case it will represent, according
to our theory, the first enlarged abdominal sternite of the
Scorpion. Then the two following sternites bearing the
apertures of the breathing-organs will belong to the third and
fourth somites respectively, and the breathing-organs will
correspond in number and position with those of the Pedi-
palpi and Pseudoscorpiones. In this case one of the somites
has disappeared behind the fourth. In all probability the
missing one is the twelfth, and we can imagine that it has
vanished from view inside the eleventh, almost exactly as in the
Pseudoscorpiones. ‘Then the eleventh somite will resemble
that of the last-named group in consisting of a single plate,
* Bih, Sy. Vet.-Akad. Handl. xvii. no. 9, p. 8 (1892).
+ In an interesting paper, recently published in vol. xxiy. of the
Journ. Linn. Soc., Zoology, my friend Mr. H. M. Bernard has advanced
reasons for showing that the Acari are derived from the Spiders. Without
now venturing to discuss in detail the views put forward in this paper,
I will merely say that in my opinion the conclusion arrived at would have
been nearer the truth if the word “ Opilionid ”’ had been substituted all
through for “ Araneid.”
t ‘Les Arachnides de France,’ vii. pp. 9, 10 (1879).
Morphology of the Arachnida. 9
which results from the fusion of the tergite and sternite.
The other missing tergite is, I suspect, the first.
Concerning the Palpigradi, an order established by Thorell
for the genus Kenenia of Grassi, I can say very little, never
having had the good fortune to examine a specimen of this
group. ‘This curious form seems to lie somewhere between the
Pedipalpi, especially the Schizonotide, and the Solifuge. As
in the last the abdomen is said to consist of ten segments, but
the last three are narrowed to support an antenniform telson
as in Thelyphonus. Moreover, the cephalothorax appears
to be segmented very much as in the Solifuge and Schizo-
notide. Unfortunately nothing is known of the breathing-
organs of Kwnenia, except their reputed absence ; so it is
impossible to speculate further as to the true affinities of this
genus.
Respecting Gibbocellum, the systematic position of which
is a matter of debate, I can suggest nothing new. It is no
doubt referable either to the Opiliones or Pseudoscorpiones,
and probably to the former.
So far the structure of the abdomen has alone been con-
sidered. The cephalothorax it is not now my intention to
touch upon, and the homologies of the segments of the
appendages have lately been fully discussed by Gaubert.
Without either accepting or rejecting the opinion of this
author on the subject of the homology of the so-called patella
of the limbs, it may be interesting to state that the segmenta-
tion of the second pair or palpi is not to my mind satisfactorily
explained on the hypothesis that this segment results from
the subdivision of the ¢ébva. For throughout the class of
Arachnida—setting aside some aberrant groups—the palpi
curiously enough, considering the general plasticity of the
limbs, present the same number of segments—namely six.
This similarity in the number would lead one to think & priort
that the separate segments are numerically homologous each
to each throughout the class. So far as the Scorpiones,
Pseudoscorpiones, Solifugez, and Pedipalpi are concerned,
there can be very little doubt upon this point. But accord-
ing to Gaubert this is not the case with the Aranee and
Opiliones ; for according to this author the fourth segment
or tibia of the palp of e. g. Phrynus is homologous to the
fourth and fifth—the patella and tibia—of the palp of a Spider
or an Opilio. But if this is so, the last two segments of the
palp of Phrynus are represented by a single segment in the
two other groups. That such a double dissimilarity has
arisen, I am not at present prepared to believe.
These considerations, coupled with the great resemblance
10 Mr. R. I. Pocock on the
between the fourth segments of the limbs in Thelyphonus and
Scorpio—segments which, according to Mons. Gaubert, are
not homologous, that of Thelyphonus being a patella while
that of Scorpio is a tibia—seem to me to be serious obstacles
to an otherwise plausible theory.
So far as the mandible is concerned, there is not much that
need be repeated here. With the possible exception of some
Acari, the greatest number of segments for these appendages
is three. ‘Three are found in the Scorpiones, most Opiliones,
and the Palpigradi; the basal one, however, has disappeared
in the Pedipalpi, Aranex, Pseudoscorpiones, and some Opi-
liones. The Solifuge, too, are usually said to resemble the
Aranez in having two-segmented jaws; but in reality they
appear to have the primitive number, namely three; the
basal one, however, has hitherto never been recognized, on
account of its fusion with the cephalic shield. It is, never-
theless, a more or less distinctly defined sclerite to which the
rest of the jaw is articulated.
A further peculiarity in the mandibles of the Solifuge is
the inferior position of the terminal segment or fang with
respect to the apophysis of the penultimate segment, to which
it is opposable. Also in the male there is a peculiar process
on the penultimate segment. Peculiar processes are also
found on the same segment of the mandible in both sexes of
the Pseudoscorpiones ; the movable dactylus, too, is almost
inferior. Mainly on these two points of similarity, Mons.
Simon has expressed the belief that the affinities of the
Solifugee are with the Pseudoscorpiones—an hypothesis which
is further supported by the absence of a cephalothoracic
sternum and a close similarity in the number of abdominal
segments.
Another system of organs, which no doubt, if more were
known of their structure, would throw light upon the affinities
of some of the orders of Arachnida, is the eyes.
The ancestral form we believe to have been furnished with
two sets of visual organs, which differed in structure and
mode of development. These median and lateral eyes are
well shown in the Scorpiones, most of the Pedipalpi, and
most Aranee. But there is a marked tendency in many of
the ‘ higher’ forms to the disappearance of some or all of the
eyes. In the higher Opiliones the median only are retained.
This also appears to be the case in the Solituge. In the
Pseudoscorpiones, on the contrary, the median seem to have
disappeared and one or more of the lateral often retained. But
until the histology and embryology of these organs has been
worked out in this group, in the Acari and in Stylocellus,
Morphology of the Arachnida. 11
their true systematic importance cannot be understood. For
it is most interesting to note that Stylocellus, which is a true
Opilio, possesses two eyes on each side of the carapace, one
raised upon a tubercle, the other at the base of the tubercle.
This last eye has disappeared in the closely allied genus
Pettalus, which thus resembles the higher Opiliones in being
furnished with only two eyes, but whether or not they are
homologous to the two eyes of a Phalangium, I am not able
to say. If they are not, the fact will constitute a radical
difference between the Sironide (Siro, Pettalus, Stylocellus),
constituting Thorell’s suborder Anepignathi, and his Lania-
tores and Palpatores. Certainly the two eyes of Pettalus from
their position strongly call to mind those of the Pseudo-
scorpiones and of the unplaced genus Gzbbocellum of Stecker.
But until the exact nature of all these eyes has been deter-
mined by a study of their development and minute structure,
" very great taxonomic value can safely be placed upon
them.
Having thus passed in review the most important external
organs of the Arachnida, it seems to me that the best characters
for the classification of the class are to be found in the
- abdomen.
The following grouping of the orders of Arachnida will
serve to show briefly my present views as to their affinities :—
A. The embryo provided with six pairs of abdominal
appendages, the second of which persists in the
adult as the pectines. The adult with four pairs
of abdominal breathing-organs in the form of
lamellar tracheze ; the abdomen very long, the
posterior five segments compressed to form a
flexible tail; the post-anal sclerite furnished Subclass
with two poison-glands ; viviparous.,........ Ctenophora.
Order 1. ScoRPIONES,
B. The embryo not provided with more than four
pairs of abdominal appendages, the second of
which are never retained as external organs in
the adult. Not more than two pairs of abdo-
minal breathing-organs. Post-anal_sclerite
usually absent and never provided with poison-
glands; abdomen much shorter, with at most
the three posterior segments narrowed to form Subclass
Simi msnally OVIPATOUB. ./... 5.0. ec 2 cee ewes Lipoctena.
A. Cephalothorax and abdomen separated by a
deep constriction; the first abdominal stomite
or its remnants covering the apertures of the
generative organs and of the first pair of re-
spiratory stigmata. The breathing-organs,
except in some of the Arachnomorphous spi-
ders, in the form of lamellar trachez ; with
12 Mr. R. I. Pocock on the
rare exceptions, there are eight eyes arranged
in median and lateral groups ........++000 Caulogastra.
Order 2, PEDIPALPI.
5. ARANER,
B. Cephalothorax and abdomen not separated by a
deep constriction ; the first abdominal sternite
not acting as an operculum to the anterior
abdominal respiratory stigmata. Breathing-
organs in the form of tubular tracheze. Usually
a single pair of eyes, rarely two pairs.
a, A pair of ny pate stigmata between the
fourth and fifth cephalothoracic appendages.
The posterior two cephalothoracic somites
not covered by the cephalic shield. The
posterior legs with a series of racquet-shaped
tactile organs on the two basal segments ;
the trochanter of the two posterior pairs
of appendages bisegmented, &c. ...... .. Mycetophora.
Order 4, SoLiruGc»,
b. No stigmata between the coxee of the fourth
and fifth appendages. The cephalothorax
covered by a continuous shield. The pos-
terior legs without racquet-shaped tactile
organs, and the trochauters of all the legs
undivided G6. ids detededidvascoceves Holosomata.
Order 5, PSEUDOSCORPIONES,
6, OPILIONES.
7, ACARI,
This classification differs considerably from that proposed by
Prof. Lankester in his article “Zimulus an Arachnid” already
mentioned. This author united the Scorpiones, Pedipalpi,
and Aranee in a group termed Aerobranchia, while the Soli-
fugee, Pseudoscorpiones, Opiliones, and Acari constituted a
corresponding group, Lipobranchia. This terminology, how-
ever, does not allow for the fact that nearly all the higher
Aranex are partially or wholly Lipobranchiate.
Moreover, although the Opiliones, Pseudoscorpiones, and
Acari were placed in the same section of the Class, it appears
from the genealogical tree published in the paper referred to
above that these three orders have had an independent origin,
the Acari being an offshoot from the stem of the Aranee, the
Pseudoscorpiones from that of the Scorpiones, and the Opi-
liones from the Solifuge. If this be so, it is not easy to see
upon what grounds the Lipobranchia can be considered a
natural group.
As for Dr. Thorell’s genealogical tree, enough has already
been said to show how materially this author’s opinions differ
irom my own. It may be added, however, that the Solifuge
are removed from the Arachnid phylum and attached to that
of the Hexapoda and Myriopoda, a view which to me is quite
unintelligible.
®
% *& * %
Morphology of the Arachnida. 13
It now remains to be seen if any beneficial results can be
ascribed to the structural modifications that we have endea-
voured to trace right through the Arachnida, starting with
the Scorpiones and ending with the Acari.
At the outset of this article it was concluded upon mor-
phological and embryological grounds that the ancestor of
the Arachnida was an animal composed of 18 distinct somites,
which were divisible into three categories, composed of 6
somites each. The appendages of the anterior six were of
large size and were set apart as organs for locomotion and
for prehension and mastication of prey; those of the succeed-
ing six were of small size, the posterior four of them disap-
pearing in connexion with the development of four pairs of
abdominal breathing-organs. The six somites of the last
category were without appendages, and constituted what may
be termed a caudal termination to the body. Now it seems
perfectly clear that an Arthropod of this description would be
most unfitted for terrestrial life on account of the clumsiness
of its build. In fact, being forced to drag along a long,
trailing, heavy, legless abdomen it could not be otherwise
than sluggish, and would consequently find no little difficulty
in gaining a livelihood by the capture of other terrestrial
Arthropods, which, if not otherwise protected, are usually
characterized by extreme activity. It would consequently be
an undoubted benefit to our hypothetical ancestor if the
caudal termination to its body could be either dispensed with
or turned to some account.
The latter end could be without difficulty attained by the
lateral compression of the segments, which would confer con-
siderable flexibility upon them. Moreover, since, as we have
seen, the last segment was furnished with a post-anal, pro-
bably pointed, sclerite, it seems clear that a formidable
weapon of attack and defence might be thus constituted. But
its greatest use would probably be to put a speedy end to the
struggles of prey that had been seized by either one or the
other of the prehensorial limbs. In this capacity its efficiency
would be greatly increased by the development of a poison-
gland in the telson. A concomitant advantage in the deve-
lopment of this “tail”? would be a loss in the weight to be
dragged by the limbs owing to the decrease in the size of the
segments,
In some such manner as this we may imagine that the
group of Scorpiones has been evolved. But it does not seem
probable that any other group of Arachnida has been derived
from them. For it appears hardly likely that any variations
tending to the obliteration of so useful an organ as the “ tail”
14 Mr. R. I. Pocock on the
would be preserved. But we can without much difficulty
imagine that the rest of the Arachnida are the descendants of
our hypothetical ancestral form ; but the line of their evolu-
tion is quite different from that taken by the Scorpions.
The disadvantage that this form would be under from the
great and useless development of its posterior abdominal
segments has already been pointed out, and we have seen
that the difficulty has been overcome in the case of the
Scorpions by the conversion of these segments into a light
and easily carried, flexible, destructive tail. But clearly
another method of dispensing with this cumbrous caudal
prolongation would be its suppression by the shortening of
the whole abdomen along its longitudinal axis. At first it
would still retain its full complement of segments, namely
twelve, and the last of them would be furnished with the
telson. But owing to the loss of flexibility in the abdomen,
this telson would be of no use as an organ of offence or de-
fence. Where it is retained, as in Thelyphonus, Schizonotus,
and Kenenia, it functions merely as an organ of touch,
being studded with tactile hairs, and to add to its efficacy in
this respect the posterior segments of the abdomen are nar-
rowed to form amovable supporting stalk forit. In Schizo-
notus it retains its original form as a single sclerite either
cylindrical or cordate in shape, while in Thelyphonus and
Kenenia it is a long multiarticulated flagellum. In these
three cases greater range of movement is conferred upon this
instrument, and its utility is thereby increased, by the in-
crease in the flexibility of the abdomen brought about by
the constriction between it and the cephalothorax. Further-
more in Schizonotus, in which the organ is very short, we
find still greater flexibility results from the secondary con-
striction which marks off the posterior portion of the cephalo-
thorax from the anterior. The result of this double constric-
tion is that the abdomen can be flexed right over the cephalo-
thorax.
In the Pseudoscorpiones and Opiliones the telson has
entirely disappeared. There is consequently no great need
for mobility in the abdomen, and no constriction appears be-
tween it and the cephalothorax. These two regions of the
body are thus perfectly continuous throughout their width,
and there is nothing to prevent the two regions from fusing.
In the Opiliones, as we have seen, this takes place to a very
great extent, the result being a decrease in the length of the
body, which no doubt can thereby be carried with considerably
less effort by the legs. In this connexion it is interesting to
note that when the body is relatively large in this group, as in
Morphology of the Arachnida. 15
e. g. Trogulus and Stylocellus, the legs are short, robust, and
presumably stronger; but when the body is smaller, owing to
a decrease in its length, as in Phalangium, the legs are ex-
tremely long and thin; so that I think there can be very
little doubt that in this last-named form and its allies the
body is shortened and lightened that it may be raised more
easily upon the long stilt-like limbs.
The structure of the Opiliones and Acari is most favourable
for shortening by the concrescence of the abdomen with the
cephalothorax, for not only is there no constriction between
these two regions, but the wide space that separates the cox
of the posterior walking-legs allows of the forward migration
of the anterior abdominal sternites beneath the lower part of
the cephalothorax. In the Pseudoscorpiones and Solifugz,
however, the concrescence cannot take place to the same ex-
tent, owing to the union in the middle line of the cox, this
union forming an impassable barrier to the forward move-
ment of the genital aperture.
In the Pedipalpi, as we have seen, as also in the Aranez,
no fusion between the cephalothorax and abdomen is possible,
owing to the constriction between them. So that in Phrynus,
where the body is much shorter than in Thelyphonus, we find
that the shortening of the abdomen is brought about by the
reduction in length of the anterior and posterior somites.
In connexion with the cephalothoracic limbs there is much
of interest to note.
In the Scorpions, in which the body is very heavy, and in
which the large prehensorial chele have to be carried aloft to
act as clumsy organs of touch, it is not surprising that four
pairs of limbs are required for purposes of locomotion. The
Pseudoscorpiones also have heavy prehensile and tactile chele ;
so, too, in this group there are four pairs of walking-legs.
But in the Pedipalpi, in which the body is lighter than in the
Scorpiones, it seems that three pairs of legs are sufficient for
locomotion, for the third pair of appendages fulfil the much
needed function of antenne. So, too, do many Spiders use
this same pair of legs as feelers and move with comparative
freedom on three pairs. The same is true of the Solifuge.
In both these groups, moreover, there are no heavy palpi to
be carried, and it must be a distinct advantage, so far as
agility is concerned, for the mandibles to be adapted for
seizing and killing prey ; for these appendages, although en-
larged for the purpose, must be very much lighter and more
easily carried than the unwieldy prehensorial palpi of the
Pedipalpi, Scorpiones, and Pseudoscorpiones.
16 Mr. R. I. Pocock on the
So far as internal organization is concerned, it is interest-
ing to note that the great development of the abdomen in the
Scorpions is correlated with a serially gangliated nervous
chord, a many-chambered heart, and four pairs of breathing-
organs. Inallthe other Arachnida in which the abdomen is
shortened, and its muscularity diminished, the nervous chord
is simplified by the disappearance of the ganglia, the cham-
bers of the heart are reduced in number, and the two posterior
breathing-organs atrophy.
It is evident that in the Scorpions the posterior region of
the abdomen is the seat of great muscular activity. To repair
the muscular tissue and to absorb its waste products a rich
supply of blood is required, and the oxygenation of this blood
will be more efficiently performed by four pairs of lung-saes
than by fewer. But in all other Arachnida the abdomen is
little more than a vehicle for carrying generative and alimen-
tary glands, so that, seeing that in the Scorpions four pairs
of lungs are sufficient, it is not surprising that the rest of the
Arachnida have been able to dispense with two pairs.
Again, with regard to the position of the apertures of the
breathing-organs, it has already been pointed out that in
Thelyphonus and Phrynus, which retain the so-called lung-
books, these apertures lie in front of sterna of the somites con-
taining the lungs. A beneficial result of this arrangement is
that the sterna in front of the apertures form movable oper-
cula to them, so that they can be opened and closed at will.
In the Aranez, where the terga and sterna mostly atrophy,
we find the remnants of these sterna retained as the opercula.
In the higher Araneze (Dipneumones) the posterior pair of
pulmonary sacs are replaced by tracheal tubes, the apertures
of which in most forms take advantage of the continuity of
the integument of the lower surface of the abdomen to migrate
to its hinder end in the wake of the spinning-appendages.
In the Solifuge the apertures of the abdominal trachez are
very small. In most forms, e. g. Solpuga, they are visible
on the surface of the sterna, but in Galeodes they have
moved posteriorly until they lie behind the sterna and are
thus capable of being closed. ‘his, as well as the small size
of the stigmata, must be an advantage to an animal living in
desert countries, where sand would be liable to block the
breathing-tubes.
Another interesting point in connexion with these animals
is that a second pair of breathing-organs occurs on the ventral
surface of the cephalothorax behind the coxe of the fourth
appendages, as if to compensate for the small size of the
abdominal stigmata. ‘The apertures of these organs must
Morphology of the Arachnida, 17
be less likely to get blocked by sand than those of the
abdomen, owing to their being removed away from the sur-
face of the ground, and only very much exposed when the
cox between which they lie are separated by the movement
in the vertical plane of the posterior half of the cephalo-
thorax.
In some Pseudoscorpiones, too, the stigmata have also
changed their position, but instead of moving backwards as
in the Solifuge they have migrated externally from the
sternal plates to the lateral membrane of the somites. In this
situation they are much more freely exposed to the air and
much less likely to be blocked by sana or earth or other
foreign bodies.
In the Opiliones the stigmata of the remaining pair of
breathing-organs retain their position in the middle of the
sterna of the somite to which they belong.
From what has been already said concerning the affinities
of the orders of Arachnida, it will be seen that the replace-
ment of pulmonary sacs by trachez has taken place indepen-
dently at least twice—once in the Dipneumonous Spiders,
and once in e. g. the Pseudoscorpiones. This fact goes along
way towards weakening the evidence of affinity between the
Opiliones and Pseudoscorpiones, not to mention the Solifuge,
on the grounds of similarity in their breathing-organs. For
the replacement may have been independently brought about
in the three cases. But however this may be with regard
to the three orders just mentioned, the fact that these tubes
have been developed twice in the same group bears very
strong evidence as to their efficacy as breathing-organs.
They must in fact be better adapted for their purpose than
the lung-book trachew. Perhaps the following considerations
may throw some light upon the matter. It seems that an
Arachnid furnished with tracheal tubes, such as Galeodes,
-must be considerably lighter and consequently more agile
than one, like a Scorpion, which possesses pulmonary sacs.
The loss of weight will be due to two causes : firstly, to the
fact that there will be a much greater supply of air inside the
body, owing to the ramifications of the air-tubes through the
tissues ; and, secondly, to the fact that the blood will be re-
duced in quantity, for there will no longer be need for a
rich supply of it, since the oxygen will be carried directly
to all parts of the body by the branches of the tubular
trachez.
Ann. & Mag. N. Hist. Ser. 6. Vol. xi. 2
18 On the Morphology of the Arachnida.
EXPLANATION OF THE PLATES.
Pruate I.
Fig.1. Diagram to show fundamental plan of Arachnid structure.
I.-VI., cephalothoracic oe c, carapace; m, median,
and /, lateral eyes ; 1-12, abdominal somites ; ¢, telson ; 2’, anus ;
a-¢, abdominal appendages ; g, generative aperture.
Fig. 2. Semi-diagrammatic figure of a Scorpion. Lettering as in fig. 1.
a, genital operculum; 8, pectines; 3'-6', sterna bearing the
four pulmonary sacs which correspond to y-¢ of fig. 1. (N.B.
The coxe of the posterior two cephalothoracic appendages have
been omitted so as to show the genital operculum and the
vectines.
Fig. 3. Fig. of a Thelyphonus. Lettering as above. w, waist or con-
striction between cephalothorax and abdomen; 1-4, anterior
four tergites of abdomen, represented on the ventral side by
three sternites, 1'-2', 3’, 4’; 10-12, posterior three abdominal
somites forming the movable stalk for the antenniform telson.
Fig. 4. Schizonotus. Lettering as in fig. 3. c', supernumerary cephalo-
thoracic tergite.
Fig. 5. Diagram of Phrynus to show the absence of the telson and the
reduction in size of the posterior abdominal tergites and
sternites.
Fig. 6. Diagram of the anterior five abdominal somites of Thelyphonus or
Phrynus, to illustrate the hypothesis that explains why the
pulmonary sacs of the 3rd and 4th somites open behind the Ist
and 2nd sterna. 1-5, abdominal somites; op, sternum of the
lst (? 2nd) forming the operculum of the anterior pulmonary
sacs and of the genital orifice; p', anterior, p?, posterior pul-
monary sacs.
Puate II,
Fig. 7. Liphistius desultor, to show the persistence of most of the abdo-
minal tergites and of the two ancerior sternites. The Ist ter-
gite forms the dorsal plate of the abdominal pedicle, and the
12th the dorsal plate of the anus. Similarly the 12th sternite
persists as the ventral plate of the anus. The appendages marked
e, (in fig. 1 persist as the spinning-mammillz, mm’, m’.
Fig. 8. Diagram of one of the Mygalomorphous Aranez to show the dis-
appearance of the external segmentation, the persistence of parts
of the sterna as respiratory opercula (s’, s*), and the migration of
the mammille to the posterior end of the abdomen.
Fig. 9. Diagram of a Dipneumonous Spider (Dysdera), to show the
replacement of the posterior pulmonary sacs by a tracheal
tube (s*).
Fig. 10. Diagram of another Dipneumonous Spider (petra), to show the
migration of the tracheal tubes to the hinder end of the
abdomen.
Fig. 11. Diagram of one of the Pseudoscorpiones (Garypus), to show the
terga and sterna of the abdomen, also the presence of the stig-
mata in the 3rd and 4th somites and the single generative ster-
nite (gy) corresponding to the 1st and 2nd tergites.
Fig, 12. Stylocellus javanus, one of the Opiliones, showing the disappear-
ance of the anterior abdominal somites (after Thorell ; slightly
modified). 3-12, the tergites ; 4-11, sternites; s, stigma on the
4th sternite.
On new Mollusca from South Africa. 19
Fig. 13. The same from below (also slightly modified from a figure given
by Thorell).
Fig. 14. Gonyleptes (an Opilio) to show the fusion of the anterior 4 (? 5)
abdominal tergites with the carapace and the 4 (1-4) free ter-
gites and 5 (1'-5') free sternites.
Fig. 15. The same from below, for comparison with fig. 14.
Fig. 16. Holothyrus (one of the Acari), for comparison with fig. 13, to
show the jointed, chelate mandible, and pediform palpi I., I.
(after Thorell). a, anus; g, position of genital orifice.
Fig. 17. Diagram of one of the Solifuga, to show the segmentation of the
carapace, the basal segment of the mandible (b) fused to the
cephalic shield, the position of the stigmata on the 2nd and 3rd
(morphologically, I think, the 3rd and 4th) segments of the
abdomen, and (s) the position of the cephalothoracic stigmata.
N.B.—Where I have taken the liberty of utilizing figures published
by Dr. Thorell, I wish it to be understood that I have tested their accu-
racy by an examination of actual specimens.
II.—Descriptions of Thirteen new Species of Terrestrial Mol-
lusca from South Africa. By JAMES Cosmo MELVILL,
M.A., F.L.S., and Joon Henry Ponsonsy, F.Z.S.
[Plate III.]
THIS, our sixth contribution on the subject, we present in
fulfilment of our promise (Ann. & Mag. Nat. Hist. 1892,
vol. x. p. 237) to deal with new forms of Hnnea and Pupa.
bie now describe four of the former genus and seven of the
atter.
1. Helix (Pella) strobilodes, sp. n. (PI. III. fig. 1.)
H. testa obtecte umbilicata, conica, tenui, sordide alba, epidermide
sericco-cornea contecta ; anfractibus sex, supra angustatis, infra
ad basin rapide accrescentibus, paullum ventricosis; apertura
lunari; peristomate tenui, simplici, apud umbilicum triangulatim
reflexo.
Long. 7°75, lat. 5°50 mill,
Hab. 'Tharfield.
A somewhat conical, close-whorled shell, with simple mouth
and narrow umbilicus, allied to H. Loveni, Krauss.
Two specimens.
2. Helix (Patula) somersetensis, sp. n. (Pl. III. fig. 2.)
H, testa aperte umbilicata, subconico-depressa, levi, albo-cinerea ;
¥%
20 Messrs. J. C. Melvill and J. H. Ponsonby on
anfractibus quatuor (ultimo rapide accrescente), ad suturas im-
pressis, gradatulis; apertura ovali; peristomate tenui, simplici.
Long. 2, lat. 1°50 mill.
Hab. Somerset East (Miss M. L. Bowker).
A shell of simple character, with a superficial resemblance
to H. (Vallonia) pulchella, Mill. In the collection of
E. L. Layard, Esq.
Two specimens.
3. Pupa tabularis, sp.n. (PI. ITI. fig. 3.)
P. testa minuta, subdiaphana, cylindrica, brunneo-succinea, levi ;
anfractibus 5-6, ventrosulis, ad suturas compressis; apertura
rotunda; peristomate planato, minime reflexo, albescente, dente
parietali conspicuo, intrante.
Long. 3, lat. 1°50 mill.
Hab. Cape Town (R. I. Lightfoot, Esq.).
Recalls to some extent the British and European species
P. muscorum, Linn. The whorls are somewhat ventricose,
shining brown, and smooth; the peristome flattened, parietal
tooth large and conspicuous.
Several specimens.
4. Pupa dysorata, sp. n. (PI. II. fig. 4.)
P. testa minutissima, oblonga, levi, apice obtuso ; anfractibus 7-8,
ventricosis, ad suturas compressis ; apertura ovali; peristomate
albido, paullum reflexo, ad marginem columellarem incrassatum
unidentato.
Long. 2, lat. 55 mill.
Hab, Griqualand East (Z. R. Sykes, Esq.).
A microscopic, oblong, narrowed species of dull plain
substance; whorls ventricose ; peristome ovate, with one tooth
on the somewhat broadened columellar margin.
5. Pupa quantula, sp.n. (PI. III. fig. 5.)
P.testa minutissima, rimata, diaphana, brunnea, cylindriformi, apice
obtuso ; anfractibus sex, microscopice tenuiliratis, valde ventri-
cosis, preesertim tribus mediis; apertura ovato-rotunda ; peri-
stomate simplici, reflexo.
Long. 2, lat. *85, mill.
Hab. Port Elizabeth (J. Crawford).
Exceedingly minute, with simple mouth and remarkably
swollen ventricose whorls, especially the third, fourth, and
new Mollusca from South Africa. 21
fifth, and cylindriform in shape ; it is longitudinally striated,
but so minutely as to be almost invisible even with a fairly
strong lens,
Three specimens.
6. Pupa Sykesti, sp.n. (Pl. III. fig. 6.)
P. testa minuta, rimata, cylindrica, diaphana, apice obtuso ; anfrac-
tibus quinque, tumescentibus, ventricosis, undique confertim
exquisite longitudinaliter tenuicostatis; apertura ovata; peri-
stomate paullum reflexo, colorato, duobus dentibus munito, hoc
parietal, illo basali, oppositis, inconspicuis.
Long. -90, lat. -75 mill.
Hab. Griqualand Kast (HZ. R. Sykes, Esq.).
Another very small but exquisite little species. It is obtuse,
cylindrical, with tumid whorls; the teeth of the peristome
are with difficulty distinguished with a lens; they are oppo-
site to one another, one parietal, the other basal. The shell
is umbilicated, with the outer columellar margin of the lip
reflected to some extent over the umbilicus, and the tumid
whorls are everywhere ornamented with fine oblique longi-
tudinal strizx.
7. Pupa haploa, sp.n. (PI. III. fig. 7.)
P. testa oblongo-pyramidata, tenuissima, apice obtusato ; anfractibus
septem, ventricosis, undique longitudinaliter tenuistriatis, apud
suturas impressis ; apertura ovali; peristomate incrassatulo, sim-
plici.
Long. 1°70, lat. °75 mill.
Hab. Pretoria.
A very small oblong species, with simple and very slightly
thickened peristome.
8. Pupa pretoriensis, sp.n. (PI. LI. fig. 8.)
‘P. testa minutissima, tenui, cinereo-brunnea, apice applanato, con-
tabulato ; anfractibus quinque, valde ventricosis, undique longi-
tudinaliter tenuistriatis, tribus ultimis latitudine squalibus;
apertura rotunda; peristomate incrassatulo, simplici, apud mar-
ginem columellarem reflexo.
Long. 1, lat. ‘55 mill.
Hab. Pretoria.
Exceedingly small, with simple mouth, and peristome
slightly incrassate. ‘The whorls are five in number, very
ventricose, all longitudinally finely striate; apex extremely
flattened and broad.
22 Messrs. J. C. Melvill and J. H. Ponsonby on
9. Pupa griqualandica, sp. n. (PI. IIT. fig. 9.)
P. testa minutissima, rimata, tenui, rotundo-cylindrica, brunnea ;
anfractibus quinque, ventrosulis, apud suturas compressis, undique
confertim longitudinaliter tenuistriatis, ultimo anfractu in medio
dorsualiter juxta labium exterius depresso-sulcato; apertura ovato-
rotunda; peristomate incrassato, ad marginem labialem unisinuato,
plicato, plicis parietalibus duabus, valde prominentibus, recurvis,
tertia, marginem apud columellarem, dentiformi.
Long. 1, lat. *75 mill.
Hab. Griqualand East (2. R. Sykes, Esq.).
Extremely small, but with a wonderfully complicate mouth,
being furnished with three plaits, one of them being pro-
minent, straight, and parietal, the second also parietal, central,
somewhat recurved and very pronounced and distinct, ex-
tending nearly two thirds of the diameter of the orifice, in a
slightly oblique direction; the third plait is tooth-shaped and
towards the columellar margin. The shell is thin, i
longitudinally striate, with five whorls, which are tumid,
and compressed at the sutures.
Three or four specimens.
10. Ennea distincta, sp. n. (Pl. III. fig. 10.)
E. testa subrimata, cylindriformi, obtusa, pallide cornea ; anfracti-
bus octo, quorum tribus ad apicem brevibus, quatuor latitudine
fere uniformibus, ultimo ad basin paullum producto, undique
oblique regulariter densistriatis ; apertura ovato-rotunda; peri-
stomate nitente albido, reflexo, dentibus quatuor instructo, tribus
parvis labialibus, uno parvulo ad basin, plica columellari interna,
planata, subtus marginem intrante.
Long. 12, lat. 4:50 mill.
Hab. Middleburg, Transvaal (J. Crawford, Esq.).
A conspicuous and proportionately large species of Ennea,
of a delicate straw-brown colour, shiny white peristome, with
prominent parietal plait, and furnished with four small teeth—
one being close to the aforesaid plait, two at the labial inner
margin, one at the basal,—the columellar plait being deep-
seated and below the outer peristome; the shell is barrel-
shaped, very obtuse at the apex, and cylindrical, eight-
whorled, and beautifully obliquely finely striated.
Five specimens. Allied to Z. Albersi.
11. Ennea regularis, sp.n. (Pl. II. fig. 11.)
F. testa rimata, tenera, obtusa, subhyalina, ochreo-nivea, cylin-
new Mollusca from South Africa. 23
drica ; anfractibus sex, undique longitudinaliter oblique tenui-
striatis, ad suturas compressis; apertura ovata; peristomate
reflexo, plicis duabus deatibusque quatuor instructo, plica
parietali magna, valide intrante, columellari interna, mamme-
formi, dentibus duobus ad marginem labialem, tertio ad basin,
quarto parvulo columellari.
Long. 7, lat. 3°50 mill.
Hab. Cope’s Folly, Maritzburg (H. Burnup, Esq.).
A. neat, delicate, and chaste species, albeit of moderate
dimensions. It is white, with an ochraceous tinge, cylin-
drical, obtuse at the apex, six-whorled, the whorls being very
compressed at the sutures, so much so as almost to appear
channelled; the aperture is ovate; peristome shining white
and reflexed, furnished with a parietal plait of conspicuous
size and projection, also a columellar plait, internal and
nipple-shaped. Besides this there are four teeth, two labial,
the lower being the larger, one at the base, and one colu-
mellar.
One specimen.
12. Ennea perspicua, 8p.n. (PI. III. fig. 12.)
E. testa subrimata, pellucida, vitrea, tenui, cylindrica, apice obtuso ;
anfractibus septem, levibus, compressulis ; apertura ovata; peri-
stomate reflexo, biplicato et bidentato, plica parietali magna pro-
minente, columellari interna valde intrante, dente labiali con-
spicuo, basali parvo.
Long. 4, lat. 2°25 mill.
Hab. Transvaal.
Very small for the genus, but of beautifully shining and
transparent texture, with the whorls quite smooth and the
mouth with a complicated arrangement of teeth and plaits,
two of each—the parietal or sutural plait being thin, sharp,
and prominent, the labial tooth conspicuous and large, the
basal smaller ; the columellar plait very distinct and wholly
" internal.
Two specimens.
13. Ennea Colliert, sp.n. (PI, III. fig. 13.)
E, testa cylindriformi, pellucida, tenui; anfractibus septem, ventri-
cosis, ad suturas compressis, undique longitudinaliter suboblique
costulatis; apertura rotundo-ovata; peristomate albo, nitido,
duabus plicis et dente uno instructo, plica parietali multum
prominente, acuta, plica interna subtus marginem columellarem
lata, dente uno labiali.
Long. 7, lat. 4 mill,
24 Mr. H. M. Bernard on the Origin of the
Hab. Pretoria. In coll. Edward Collier, Esq.
A transparent shining little species, with regularly costu-
late whorls, ventricose, seven in number ; the shell is cylin-
driform in shape; peristome shining white, somewhat trans-
lucent, as is the rest of the shell, furnished with a promi-
nent acute parietal fold and an internal broad plica behind the
columellar margin of the peristome. There is also one simple
labial tooth.
Two specimens, one of which is slightly more elongate
than the other.
EXPLANATION OF PLATE ITI.
Fig. 1. Helix strobilodes. Fig. 8, Pupa pretoriensis.
Fig. 2. somersetensis, Fig. 9. griqualandica,
Fig. 3. Pupa tabularis. Fig. 10, Ennea distincta.
tg. 4. dysorata. Fig. 11. regularis.
Fig. 5. quantula. Fig. 12. —— perspicua.
tg. 6. —— Sykesit. Fig. 13. —— Collieré.
ig. 7. —— haploa.
IlJ.—Additional Notes on the Origin of the Trachee from
Setiparous Glands. By Henry M. Bernarp, M.A.
Cantab. (from the Huxley Research Laboratory).
In the last issue of the ‘ Zoologische Jahrbiicher’ (Bd. v.) I
brought forward some evidence in favour of the derivation of
the Arthropodan trachez from setiparous glands, and pointed
out how this derivation might be made to explain many of
the difficulties which at present beset this subject. I propose
here to bring forward a few more arguments tending in the
same direction.
(1) 1 endeavoured to show that in the Myriapoda, while we
could deduce the tracheee from the notopodial acicular glands,
the stink-glands which occur dorsally to the stigmata might
be the glands of the original parapodial sete. Mr. Pocock,
of the British Museum, kindly informs me that in the Chor-
deumide, which appear in many respects to be intermediate
forms between the lulide and the Polydesmide, the foramina
repugnatoria of these latter are not to be found, but sete
develop apparently exactly on the spot where these glands
ought to occur, and, further, that in the Polyxenide a great
tuft of seta grows in the same spot, apparently in place of
the stink-glands which are wanting, and, again, a still more
important fact, that in Scolopendrella, in which animal the
Trachee from Setiparous Glands. 25
stigmata are confined io the head, large sete spring from the
sides of the segments in the place where stigmata occur in all
the Chilopoda and Hexapoda. This curious retention of so
many primitive Annelidan characteristics on the part of the
Myriapods is after all what one might expect from animals so
clearly Annelidan in their rich segmentation.
(2) The arrangement of the stigmata, hairs, and pigment
in the pupa of the vapourer moth (Orgyta antiqua) is very
suggestive. The deep black pigment of the skin is relieved
at the sides of the abdominal segments by clear white or
whitish patches, running dorso-ventrally and nearly meeting
in the ventral middle line. The arrangement is shown in the
woodcut. On each side of the stigma is a very definite, more
Lateral view of an abdominal segment of a pupa of a female
Vapourer Moth, showing the “ parapodial scars.”
or less circular field covered with long sete. It is true that
the greater part of the dorsal surface is covered with long
sete, but no one can examine these setigerous areas under
the microscope without being convinced that they are distinct
regions. If this arrangement is compared with an Annelidan
parapodium, the similarity in the position and arrangement of
the sete is very striking. The stigma corresponds to the
opening of the acicular gland and the setigerous areas to the
setigerous regions of the parapodium on each side of the
aciculum. It was especially the aciculum of the dorsal para-
odium to which I referred the stigmata of the Hexapoda.
We have here, then, traces of a complete dorsal parapodium,
as shown in the figure, while beneath it is seen the dorsal
part of the ventral parapodium with a similar setigerous area
well marked off. More ventrally (not shown in the figure)
the white patch is continued, but without sete, almost to the
median line. It is the ventral parapodium which I have
suggested formed the leg of the Hexapoda.
Why these extraordinary “ parapodial scars,” as I propose
to call them, should suddenly appear on the pupa is a problem
26 Mr. H. M. Bernard on the Origin of the
I cannot pretend to solve. It may perhaps be referred to the
force of heredity coming into action as soon as the animal is
more or less withdrawn from the struggle for existence, as a
pupa may almost be said to be when encased in a cocoon.
(3) In order to get more light on the homologies of the
ventral row of structures in the Arachnids which I propose
also to refer to a row of setiparous glands, I have naturally
turned my attention to some of the less known orders. A
small Chernetid, apparently an Odbisium, has yielded me
unexpected results.
The stigmata of the tubular trachez on the second and
third abdominal segments are followed by a complete row of
segmental apertures running along each side to the end of the
abdomen. Their position corresponds exactly with those of
the stigmata, and I think it is impossible to doubt that they
are homologous with these latter. In this interesting Arachnid,
then, there are nine pairs of apertures on the nine posterior
abdominal segments. The two anterior pairs are stigmata.
The function of the other seven, for want of suflicient
material, I have not yet made out. It is well known that the
Chernetide spin webs, and there seems to be no very clear
idea where the glands are situated. Croneberg’s claim * that
the spinning-gland opens on the mandibles is probably correct.
I find a very distinct aperture on a small prominence behind
the point of the movable piece of the mandibles. In that
case these “stigmata” may be purely rudimentary and
functionless. If, on the other hand, these seven pairs of
apertures following on, and evidently homologous with,
stigmata prove to be the openings of spinning-glands (a point
I hope soon to investigate), we should have a remarkable con-
firmation of my suggestion that the lung-books or trachez
and the spinning-glands of the Araneids are homologous
structures as common derivatives from setiparous glands,
We learn also from these nine pairs of abdominal apertures in
Obisium that the limitation of the number of stigmata in
Scorpio is not original, ¢. e. inherited from a Limulus
ancestor, but is due to a secondary reduction of what were
originally segmental structures along the whole abdomen.
(4) The coxal gland of Galeodes opens, as Sturany +
suspected, between the third and fourth appendages, @. e. on
the coxa of the first leg. Dufour, who mistook it for a sali-
* “ Beitrag zur Kenntniss des Baues der Pseudoscorpione,” Bull. Soe,
Imp. Nat. de Moscou, t. ii. (1888).
+ “ Die Coxaldrusen der Arachnoiden,” Arb, Zool. Inst. Wien, t. ix,
(1891).
Trachee from Setiparous Glands. 27
vary gland, nevertheless correctly described its different
appearance in different Galeodide. In some it is a simple
long tube commencing in the region of the second leg,
running backward to form a tangled coil between the nerve-
cords in the region of the third and fourth legs. It then
bends forward again to open in the above-mentioned spot.
In others the proximal end of the gland divides up into a
sponge-like mass of anastomosing tubules, which, uniting with
those of the gland on the opposite side, form an amorphous
mass arching over the mid-gut. The course of the duct and
the position of the aperture agree almost exactly with that
described by Sturany for the coxal gland of Atypus.
The points which especially interest us here, however, are
the following. ‘The aperture of the gland on the coxa of the
first leg agrees almost exactly in position with the aperture of
the trachez on the coxa of the second leg, which suggests
their being homologous structures. The duct shows the
characteristic striped appearance, as if its wall were pitted by
countless pores. It stains very badly, and although it will
not stand boiling in caustic potash, resists the action of cold
caustic potash. ‘There seems to me to be little doubt that
chitin is present in its walls, although it nowhere forms a
definite lining to the tube. The retention of the gland in
Galeodes as an important functional excretory organ, while it
has degenerated in the Araneidx, although once well developed
in these latter, suggests a correlation between these coxal
glands and the spinning-glands. The development of the
spinning-glands to utilize and carry off excretory products
renders the coxal glands unnecessary. ‘This correlation is
the more probable if both glands can be referred back to
setiparous glands. The same may be said of the mandibular
poison-glands, which are well developed in the Araneids, but
absent in Galeodes.
Besides the fact mentioned in my first contribution on this
subject, that no difference can be discovered between the
trachez of the thorax and of the abdomen, it also remains to
be stated (what indeed has been known since 1848 *) that
the trachee of the two regions communicate freely with one
another. ‘The assumption that the abdominal trachee are
developed from lung-books while the thoracic trachee are
independent developmentst from ectodermal invaginations, and
therefore not in any way homologous with the former, seems
* Kittary, “ Anatomische Untersuchungen der gemeinen Galeodes und
der furchtlosen Solpuga,” Soc. Imp. Moscou, vol. xxi.
= t Korschelt and Heider, ‘ Vergleichende Entwickelungsgeschichte,’
p-_638.
28 Mr. H. M. Bernard on the
to me to ignore the fact that both structures arise in similar
positions, viz. at the bases of the limbs. The thoracic trachex
of Galeodes open posteriorly on the coxa of the second pair
of legs. In addition, then, to the extreme improbability of
the same structure—trachee—having had two independent
origins in the same animal, we have the further improba-
bility that the openings of the assumed independently deve-
loped thoracic trachex should bear apparently the same rela-
tion to the thoracic limbs as the lungs do to the embryonic
abdominal limbs.
These arguments, I think, lend considerable support to my
attempt to deduce trachez from setiparous glands. ‘The first
two points seem to show that the lateral row of stigmata,
spinning-glands, &c. found in the Myriapoda and Hexapoda
have actually been deduced from the acicular glands of dorsal
parapodia, the ventral parapodia forming the legs. Such a
confirmation of this part of my original suggestion leads us
almost naturally to conclude that the ventral row of trachee,
spinning-glands, &c. in the Arachnida have been developed
from the setiparous glands of the ventral parapodia. If so,
the legs of the Arachnids have been most probably developed
from the dorsal parapodia, while the ventral parapodia have
disappeared in the coxal joints, their setiparous glands, how-
ever, persisting as trachee &c. This origin of the limb in
the Arachnids is exactly what I have elsewhere endeavoured
to show must have been the origin of the legs of the Crus-
tacea. This would account for the great similarity between
the legs of Limulus and those of the Arachnids, and also for
their common possession of coxal glands. It would also
account for the traces of Crustacean characters found by
Jaworowski* in the developing limbs in the embryo of
Trochosa singoriensis t.
1V.—On the Terminal Organ of the Pedipalp of Galeodes
and the Discovery of a Homologous Organ on the Pedipalp
of Phrynus. By Henry M. Brernarp, M.A. Cantab.
(from the Huxley Research Laboratory).
THE remarkable protrusible organ at the tip of the pedipalp
of Galeodes has, since Dufour’s discovery, received but little
* “Ueber die Extremitaten, deren Driisen, und Kopfsegmentirung bei
Trochosa singoriensis,” Zool, Anz., May 1892.
+ Lregret to have omitted to mention in my former paper what had
been a very valuable work of reference to me, viz. Palmén’s ‘Zur Mor-
phologie des Tracheensystems,’ Leipzig, 1877, ‘
Terminal Organ of the Pedipalp of Galeodes &e. 29
attention. Dufour * himself assumed that it was a sucker-
like seizing-organ, and this view has again been expressed
quite recently f.
A close examination of it shows almost conclusively that
Koch’s ¢{ suggestion that it is a sensory organ, probably
olfactory, is correct.
We may at the outset remark, first, that such a minute and
delicate seizing-organ in an animal armed like (aleodes
would be ridiculously out of proportion, and, secondly, that a
highly specialized sensory organ might be expected in such
swift runners who hunt their prey.
The organ itself is essentially an invagination of the tip of
the pedipalp, forming a conical pit, the thin chitinous walls
of which are continued into the metatarsus as a tendon to
which a powerful muscle is attached. The external aperture
of the pit is a transverse slit, which closes like a pair of lips.
The upper lip, ¢. e. that on the dorsal or outer side of the
limb, is stiff and solid, and is opened by means of a long
tendon connecting it with a muscle far down in the meta-
tarsus. The opening of this lip like a lid leads apparently to
the slight protrusion of the organ, 7. e. by drawing it up with
it. I could find no other mechanism to explain it.
Within the pit one wall only, 7. e. the dorsal, is thickly
covered with fine sensory hairs, so regularly arranged that
the chitinous membrane from which they arise appears like a
fine network. The hairs are in evident connexion with a
deep sensory epithelium immediately under the chitinous
membrane. This epithelium runs a long way down the
(hollow ?) tendon. ‘The ventral wall of the pit is an exceed-
ingly fine chitinous membrane, against which, when the
aperture is closed, the tips of the hairs apparently rest.
If this is not enough to show that this organ is not a seizing
but a sensory organ, the discovery of a very similar organ in
the pedipalps of Phrynus on the inner surface of the most
distal claw effectually settles the question. In the adult
ail a sensory area runs longitudinally from the joint
along about half the length of the claw; but I was unable to
discover any chitinous invagination. In a young specimen,
however, which could be clarified, an invagination of this
same area was very clear. It occurred on the dorsal side of
* Dufour, “ Anatomie, histologie, et physiologie naturelle des Galéodes,”
Mémoires présentées 4 l'Institut de France, vol. xvii.
+ Bertkau, “ Ueber Sinnesorgane in den Tastern und in den ersten
Beinpaar der Solpugiden,” Zool. Anz., Jan. 1892.
¢ Koch, ‘‘Systematische Uebersicht iiber die Familien der Galeoden,”
Archiv fiir Naturgesch., 1842.
30 M. Ph. Owsjannikow on the
the area, and was so arranged that the sensory surface seemed
to dip away sideways under the cuticle of the claw, and could
then be followed under the microscope through the base of the
claw into the next joint. The lower ventral edge of the
exposed part of the sensory organ was protected by hairs
regularly arranged so as to slant over the delicate surface,
while dorsally, where the sensory surface disappears under
the cuticle, the slit-like opening is guarded by a regular row
of fine hyaline hairs, which rise from the sensory surface and
curve over the outer cuticle. Apparently similar hyaline hairs
can be focussed down in the lower parts of the invagination,
On both old and young curious hairs with heads like narrow
dentate leaves occur at each end of the sensory area*.
The finer histological details of this organ in Phrynus can
only be made out in young specimens when the chitin is not
too thick for sectioning. I reserve further details of the organ
in Galeodes for a comprehensive work which I am now pre-
paring on this animal.
The presence of this presumably olfactory organ in such
different Arachnidan types as Galeodes and Phrynus is of no
small interest. The clarified pedipalp of Scorpio showed no
trace of such olfactory organs. ‘The same must be said of a
Chernetid, apparently an Obistum, which had been boiled
in caustic potash. The pedipalp of a Telyphonus was
also searched in vain by clarifying rough sections. I
unfortunately had no young specimens of this latter at my
disposal. It seems to me not improbable that some traces of
such an organ might be found in very young animals con-
sidering the apparent aflinity which exists between Phrynus
and Telyphonus.
Whether the peculiar sexual organs at the end of the pedi-
palp of the Araneids had any original connexion with such a
sensory organ is a point well worth investigating.
V.—On the Embryology of the River-Lamprey.
By Pu. OwsJANNIKOW Tf.
MOovERN methods, including both serial sections and also the
new staining reagents, now enable us to prosecute a much
* As an index of the accuracy of Koch’s figures, the position of this
organ is marked by a group of hairs in the drawing of Phrynus ceylonicus,
plate 776 in vol. x. ‘ Uebersicht der Arachniden-Systems,’
+ Translated from the ‘Mélanges Biologiques tirés du Bulletin de
l’'Académie Impériale des Sciences de St.-Pétershourg, t. xiii. livr. i.
St. Petersburg, 1891: pp. 55-67.
Embryology of the River-Lamprey. 31
more precise investigation in the realm of embryology than
was possible in former years. Thus it may be readily under-
itood that, thanks to these new scientific processes, many
deficiencies which existed also in our knowledge of the
developmental phenomena in the Lamprey are supplied by
numerous fresh investigations. The new facts bring with
them new questions, which have a special interest for compa-
rative embryology. As the publication in their entirety of
my investigations upon the development of the Lamprey is
not immediately possible, since the exceedingly abundant
material from all stages of development must be arranged in
proper order, and since also the figures require much time, I
have determined to give a short provisional communication,
which is at the same time intended to complete the investi-
gations which I made upon this subject nearly twenty years
ago.
aa the ova of lamprey larve, which are from 70 to 90
millim. in length, the germinal vesicle is in many cases already
excentric in position. At this period of development, when
the contents of the ovum are in a semifluid state, the germinal
vesicle may alter its position. I have had occasion to observe
in a preparation three young ova in which the germinal
vesicle lay between the yolk and the egg-membrane.
The Graafian follicles are clothed with endothelial cells.
At the region where the pointed angles of the cells are in
contact with one another, which is usually at the upper pole
of the ovum, dark spots are to be seen after treatment with
nitrate of silver ; these may be regarded as entrances into the
lymph cavities.
The blood-vessels enter the Graafian follicle at its pointed
end, at which spot the active pole of the ovum also lies within
the follicular membrane. It is frequently to be observed that
the tip of the ovum does not quite touch that of the follicular
membrane, while everywhere else the latter is in close con-
tact with the ovum.
In all ova of lampreys which were ready to spawn when
they were removed from the ovaries I have observed the
germinal vesicle with the greatest distinctness in sections.
The germinal spot, however, was not present. On the other
hand, in very many ova which were squeezed out of the
female for the purpose of artificial fertilization the vesicle
could not be detected. In very few cases did it lie near the
active pole of the ovum as a small flattened disk of the vitel-
line membrane. It is hardly open to doubt that in the cases
in which the ovular nucleus is not visible its contents have
mingled with the elements of the yolk, for all the ova were
32 M. Ph. Owsjannikow on the
capable of being fertilized and in all the protoplasmic protu-
berance was noticeable directly after the act of fertilization.
In order that the several portions of the embryo shall be
more readily distinguishable from one another, it is necessary
for the preparations to be thoroughly stained. The prevalent
idea that the egg-membrane is impervious to stains is erro-
neous. I found that several staining-fluids penetrate the
ovum and the embryo. ‘The ova took the best stain, no
matter whether at first treated with Flemming’s fluid, alcohol,
or osmic acid, by being immersed for from twelve to four-
and-twenty hours in a strong saturated solution of hama-
toxylin. A longer immersion causes the yolk-granules to
become coal-black.
I possess a large series of sections in which there may be
seen all those processes which set in immediately after fertili-
zation, and were observed and described in living ova by
A. Miiller, Kupffer, Benecke, Calberla, and myself.
By treatment with osmiec acid the contents of the ovum
are instantaneously hardened, so that in sections we are
enabled to get a good view of the protuberance which arises
at fertilization as well as the protoplasm, which usually
appears on the withdrawal of the yolk from the egg-
membrane.
Moreover, in the serial sections we may study the most
varied forms of mitoses, which appear at the division of the
nucleus before the formation of the new yolk-segments.
The time occupied by the various metamorphoses in the
ovum depends upon the temperature at which the fertilized
ova are kept. This fact explains why the divisions of the
yolk which I previously described in the ova of the lamprey
proceeded more rapidly than they were found to do by Cal-
berla. Kupffer states that at Kénigsberg, when the tempe-
rature of the air stood at from 8° to 10° C, (46°-4 to 50° F.),
the larve were hatched on the sixteenth or seventeenth day,
and at Naples at the end of the eighth day. My larvee
hatched out on the ninth or tenth day at an atmospheric
temperature of about 16° R. (68° F.).
The first furrow is a longitudinal one, which consequently
has a meridional direction and divides the yolk into two
perfectly equal portions.
The furrow begins from the nucleus, which for a long time
lies almost entirely superficially in the neighbourhood of the
active pole.
The first segmentation proceeds exceedingly slowly. We
observe long fibres radiating from the nucleus, which are
Embryology of the River-Lamprey. 33
especially distinct and lie closer together at the spot at which
the furrow has begun to form.
Before the complete separation of the two segmentation
spheres several nuclei appear, and between every two nuclei
a spindle becomes visible. It is no doubt partly owing to
such a complete separation of the yolk-segments that in the
last days of development we find within one egg-membrane
two embryos entirely separate from one another, as I fre-
quently had occasion to observe.
The second furrow is likewise longitudinal, and it is only
the third that is the first equatorial one.
After fifteen hours the ovum is represented by a vesicle the
upper wall of which consists of a series of smaller cells, the
lower of much larger ones. Baer’s cavity has attained con-
siderable dimensions.
The ova of twenty-eight hours likewise exhibit at the roof
of Baer’s cavity a single layer of cells.
At the lower pole, however, there lies a group of larger
cells. ‘The transition between the smaller cells of the upper
pole and the larger cells of the lower is a gradual one.
In ova of fifty-five hours Baer’s cavity has reached a still
larger size.
‘lhe smallest cells lie at the upper pole and already have an
epithelial character about them. In many sections the ovum
has the appearance of a ring, which at one spot, namely at
the upper pole, consists of one layer, while it is bi- or tri-
laminate at the other. Sometimes there lies at the lower-pole
a small group of segmentation-spheres. Even at this period
a Rusconi’s pit is visible upon the under surface of the ovum.
The cells of the outer layer gradually pass into the floor of
the pit. The shape of this pit leads us to suppose that it
has arisen by the invagination of the outer layer, as is actually
the case. ‘This structure is nevertheless a transitory one; at
any rate its appearance very soon changes. The largest cells
_ are found in the pit of Rusconi and at its margin.
When eighty hours have elapsed the outer layer has under-
gone still further modification; the cells have become still
more cylindrical and have almost entirely consumed the
yolk-granules. The size of the segmentation-cavity has in
the meantime greatly diminished. ‘The whole of the lower
half of the ovum consists of small round cells, which, how-
ever, are separated by a slight space from the still smaller and
much more closely packed cells of the epiblast. The anus of
Rusconi can at this time be seen with great distinctness at
the lower pole, though it has undergone a change in form and
has become narrower and longer. ‘The ovum itself has also
Ann. & Mag. N. Hist. Ser. 6. Vol. xi. 3
34 M. Ph. Owsjannikow on the
grown longer and has become pointed at oneend. This latter
appearance is due to the fact that during this period the
multiplication of cells proceeds with especial activity at the
upper lip of the anus of Rusconi at the point where the upper
layer passes into the lower. _
Since during this period—it may be some hours earlier,
though usually later—many highly interesting processes take
place in the ovum, we will now consider these more closely.
TuE FORMATION OF THE ALIMENTARY CANAL.
In earlier stages the anus of Rusconi took the shape of a
broad pit which had arisen by invagination of the epiblast.
I am speaking of an embryo which is at least some four-and-
twenty hours younger than that represented in Scott’s
fig. 10, a. At that period we actually havea gastrula before
us. Ina few hours, however, the number of the yolk-spheres
lying at the bottom of the ovum has considerably increased.
The diminution in the size of Baer’s cavity during this time
is not to be ascribed to the fact that the cells are thrust into
it by the formation of the enteric cleft, but to the increase in
the number of these cells.
The formation of the enteric cavity proceeds by a splitting
off of the vitelline elements from the undifferentiated cells,
which become the hypoblast (“von dem Driisenkeim ”),
starting from Rusconi’s pit, precisely as this process has often
been observed and described in the frog, axolotl, and sterlet.
In the animals just mentioned this process is easier to observe,
because the cells which adjoin the cleft contain pigment-
granules during their division and separation from the remainder
of the yolk. In some cases a streak of pigment precedes the
cleft. Since the ova of Petromyzon are entirely unpig-
mented, observation becomes somewhat more difficult. In
spite of this we can convince ourselves by the examination of
hundreds of preparations that the lower layer, the endoderm,
arises by the separation of a series of cells from the yolk. It
is further to be remarked that before the formation of the
enteric cleft the sections already show a special grouping of
the cells which are subsequently utilized as the elements of
the lower layer. During this period and also somewhat later
two layers, the ectoderm and endoderm, are present, which are
separate from one another and continuous only at the point of
flexure. The cells of the ectoderm are smaller than those of
the endoderm, as is indeed perfectly natural, since the deve-
panes of the former commenced earlier than that of the
tter.
Embryology of the River-Lamprey. 35
After this the chief centre of the developmental processes is
situated for a time in the hinder portion of the embryo. In
successful longitudinal sections, namely such as pass through
the middle of the ovum from front to rear, so that they bisect
the upper and lower lips of the anus of Rusconi, it can be
seen that the embryo already extends over half of the ovum.
It consists, as already stated, of ectoderm and endoderm, and
each of these layers exhibits only a single series of cells
throughout its entire extent. It is true that sections are to
be found in which the ectoderm has more than one series of
cells, but this is owing to the fact that the sections were taken
obliquely and that the knife, working more on the surface of
the ovum, has carried a larger strip of this with it. I have
found preparations in which the enteric cleft communi-
cated with the segmentation-cavity. The cells which were
situated at the periphery of the cavity, immediately beneath
the ectoderm, were utilized for the formation of the lower
layer.
Ova 118 HOURS AFTER FERTILIZATION.
Ova belonging to this period exhibit but little difference
from those of the preceding one. ‘They have become some-
what longer; the two lips, the upper and the lower, and the
plug which lies between them are still more distinct, and the
embryo itself has still further increased in length. In many
longitudinal sections each of the two layers continues to exhibit
a single layer of cells. On the other hand, many transverse
sections, if they have passed through the posterior end of the
ovum, exhibit a multiplication of the ectoderm cells beneath
the dorsal furrow. ‘The spinal cord is in process of forma-
tion: as Calberla has already shown, this structure is at first
solid, and it is not until later that the cells separate from one
another and give rise to the spinal canal.
Ova 126 HOURS AFTER FERTILIZATION.
This is one of the most interesting periods. The embryo
has become considerably longer. In very many transverse
sections of the ovum the embryo is cut through in two places,
at the anterior and posterior end. At the latter we observe
the solid spinal cord, beneath which is the notochord already
completely developed, and then the epithelium of the enteric
canal, which forms its inner wall. On both sides of the
notochord lie in part still solid rudiments of the protover-
tebre, Wolffian ducts, and lateral plates; the latter pass into
e
36 M. Ph. Owsjannikow on the
a single layer. A section taken through the anterior portion
of the embryo presents another picture. At this point the
central nervous system is thicker: a central canal has already
developed at some distance from the brain, the anterior end of
which is also still solid. At this region the notochord is not
present, though the cells of the endoblast immediately adjoin
the central nervous system. At some distance from the ante-
rior end of the embryo the enteric cavity is of a very con-
siderable size. It is surrounded above by a double layer of
endoblast cells, which are somewhat thrust downwards in the
middle by the spinal cord. On the other hand, on both sides
of this depression, to the right and left of it, the endoblast
forms an upward expansion of the cavity. If we now
examine the sections in order from behind forwards, we
observe two folds of the endoblast approaching nearer and
nearer to the middle line. In this manner two cavities
appear, the uppermost of which is the smaller. Finally the
folds completely meet, forming a ring round the smaller
cavity, which at last entirely disappears.
In the remainder of the body the formation of the notochord
appears to take place by a simple constricting off of the
endoblast cells lying beneath the spinal cord, as has already
been described and figured by Calberla. On both sides of
the notochord lies a double row of cells, which soon passes
into a single one. ‘These cells have separated from the
endoblast. ‘There is no space between the two rows; but in
spite of this we already recognize the elements which will go
to form protovertebre, Wolffian ducts, and lateral plates.
Moreover in many sections we find the protovertebre already
almost completely developed: the formation of a cavity in
them takes place later on. The epiblast, too, during this
pot is still exclusively unilaminate exeept in the median
ine, where the spinal cord has already developed. In the
sections it is still sometimes entirely separate from the ovum.
_ The development of all primitive organs is further advanced
in the posterior than in the anterior portion of the embryo.
The formation of the cavities and canals is effected either by
the separation of the cells from one another or, as is especially
the case in the region of the undifferentiated cells which
become the hypoblast, by the consumption of the yolk-lamelle,
so that in the place of the yolk-cells only empty envelopes
remain.
Development proceeds from the median line towards the
sides. After the formation of the spinal cord and the noto-
chord a cavity arises, first in the protovertebre, which separate
from the rest of the cells of the mesoblast, then in the Wolffian
Embryology of the River-Lamprey. 37
ducts, and finally in the lateral plates. It is asserted by
A. Shipley (fig. 11) that the Wolffian ducts or the segmental
organ develop before the protovertebra. I have seen no
evidence of this in my preparations. ‘The figure in question
is altogether too diagrammatic.
The protovertebral segments are not cubical, but are drawn
out into a point at the lower inner angle towards the notochord.
ON THE ORIGIN OF THE SENSE-ORGANS.
I have not observed the formation of the auditory apparatus
in embryos less than 160 hours old. There appears at the
side of the brain a pit-shaped invagination of the ectoderm
which is equally distinct in transverse as well as in longitu-
dinal sections. The cells which are situated in the centre of
the pit are somewhat longer than the remainder. After some
time the pit closes and becomes a capsule, the future labyrinth
of the auditory apparatus. At this period all cells, both of
the skin as well as of the nervous system, are still pretty
richly packed with yolk-lamelle. The intermediate space
between the auditory capsule and the brain is very incon-
siderable, and subsequently almost completely disappears.
The formation of the eye takes place during the two
hundredth to the two hundred and fortieth hour after fertiliza-
tion. At the side of the upper surface of the fore brain, almost
immediately behind the olfactory pit, we find a sac-shaped
expansion of the cerebral wall. It extends backwards and
downwards, and can be better seen in longitudinal sections,
though in order to obtain a complete representation of it both
classes of sections must be examined. At first the cavity of
the saccule is exceedingly narrow, but it subsequently becomes
somewhat wider, especially at its outer end.
The earliest rudiment of the paired eyes in the lamprey
presents more similarity to the rudiment of the third eye in
many animals than to that of the eyes of the Vertebrata in
general. For in this case we find a relatively long tubular
stalk, the outer and somewhat expanded end of which is only
utilized for the formation of the retina. The outer wall of
the eye-stalk grows thicker and becomes indented like a pit,
just as the retina is generally developed. The epithelium of
the skin, which lies opposite the rudiment of the eye, takes no
part in its formation. During this time no trace of the lens
can be discovered.
The formation of the nasal pit proceeds by a pit-shaped
invagination of the outer layer, as has already been frequently
described.
38 M. Ph. Owsjannikow on the
The whole of the earliest sense- organs appear to have arisen
in the same manner throughout the entire Vertebrate phylum.
THe HEART.
I have observed the earliest rudiments of the heart, or,
rather, of the venous vessel in embryos of the age of 133
hours. The longitudinal section taken from above down-
wards exhibits an arrangement somewhat similar to that
figured by A. Goette in his well-known work on the develop-
ment of the fire-bellied toad (fig. 37); for we observe that
the enteric cavity has attained its greatest dimensions in the
region of the head, which is in process of formation. Poste-
riorly towards the dorsal surface it becomes narrower and
forms a very slight indentation in the middle of the undiffe-
rentiated hypoblast cells, and then proceeds as a sac-shaped
depression in the yolk in the direction of the ventral surface.
The walls both of the enteric cavity and of the venous sinus
are clothed with cells of the endoblast, which as yet are far
from having assumed an epithelial character. They are large
and full of yolk-granules. At this period we have only the
cavity of the vessel before us, which subsequently becomes
constricted off from the intestine. The appearance of the
rudiment of the venous system alters very Nittle during the
next forty or fifty hours.
In embryos of 180 hours the body has attained a consider-
able length. In longitudinal sections we observe the
branchial cavity in the form of a long canal which already
possesses gill-slits. Below the branchial cavity there appears,
as in the previous stage, a depression passing off in a lateral
direction from the intestine. It has in the meantime become
somewhat longer and its lowerend less regular. As a matter
of fact it is continued in the shape of a cleft, which can be
traced a very long way backwards, while rifts may be
observed proceeding from it in all directions, which finally
lose themselves between the yolk-spheres.
It is evident from what is here seen that long before the
development of the heart we have a system of canals which
are filled with a fluid—the lymph. In all the cavities men-
tioned, in the cavity of the intestine, the branchial chamber,
&c. we find circular rings, a kind of membranes, which are
probably remnants of yolk-spheres which have undergone
dissolution. Isolated channels of this kind are to be met with
in which some few yolk-granules are still present, while others
are quite full of them. The granules are much smaller than
the neighbouring cells or yolk-spheres of the undifferentiated
Embryology of the River-Lamprey. 39
hypoblast. We may consider that the yolk-spheres during
the period which is now being described are on the average
from nine to ten times larger than the granules alluded to:
the size of many of them, however, is greater than this.
Further and more important changes, which already stand
in direct connexion with the formation of the heart, are to be
noted in embryos of from 200 to 220 hours, when they have
attained a length of from 2 to 3 millim. Beneath the
branchial cavity an expanded oval region is formed, just as
if the embryo was somewhat swollen at this point. The
longitudinal sections show that a wide cleft has here appeared
in the lateral plates on the right and left: the outer layer
applies itself to the skin, the inner one to the intestine. We
then have on both sides of the intestine two oval vesicles, the
greatest diameter of which lies in the vertical direction. The
formations just described vividly remind us of the pleural sacs
of the higher Vertebrates before the appearance of the lungs.
In certain sections they appear quite empty ; in others, taken
higher up, we find the rudiments of the segmental organs,
that is of the primitive kidneys.
In order to obtain a distinct representation of the develop-
ment of the heart, we must have recourse to transverse
sections. If we examine one of these from the cardiac region,
we observe round the cesophagus a wide free space, which has
been formed by the divergence of the two lateral plates: this
is the body-cavity. We employ this term in order to apply a
general idea to this cavity. ‘The intestinal fibrillar plate
(* Darmfaserplatte ”’), or splanchnic mesoblast, can be traced -
particularly well in stained sections. We observe a fold of it
on the right, as also on the left side, which passes round the
segmental organs and the upper half of the esophagus. When
the two layers meet beneath the closed cesophagus they assume
a downward direction, and finally pass into the somatic meso-
blast, which attaches itself to the ectoderm. At the spot at
which the right and left fibrillar plates have approached one
another—namely, where the two portions lying between the
splanchnic and somatic mesoblast are opposite to one another
—there is at first a narrow, and subsequently a much broader
cleft, which extends from above downwards. ‘This inter-
mediate space is the cardiac cavity. ‘The above-mentioned
portions of the intestinal fibrillar plates had fused together at
their upper and lower margins and become a closed tube,
which now projects freely into the body-cavity and is attached
to the intestine by a short band. Before the fusion of the
median portion of the intestinal fibrillar plates to form the
cardiac walls, there could already be observed on their inner
40 M. Ph. Owsjannikow on the
margin a special layer of flat cells. This internal layer of
cells separates from the external one ; and in the transverse
sections we have before us two tubes lying one within the
other. The internal one becomes the endothelial coating of
the heart—that is, the endocardium. The second, somewhat
stouter tube becomes the muscular tissue of the heart, which
on the outside is likewise clothed with endothelial cells. The
heart in Petromyzon arises from the folds of the intestinal
fibrillar plates, which alone furnish the material for all its
constituent parts. I would especially point out that the
elements of the intestine take no share whatever in the forma-
tion of the heart. A communication is established between
the cardiac cavity and those lymph-chambers which we
designated above as the commencement of the heart, and
which have transformed themselves into the veins. At this
period the veins have no walls of their own, and are in com-
munication with many canals and lacune.
Though the literature of the subject will not be fully con-
sidered until my detailed and illustrated paper is printed,
I cannot here refrain from mentioning that A. Shipley’s obser-
vations with reference to the heart also are highly valuable.
His figures (figs. 24 & 26) are faithful representations, albeit
somewhat diagrammatic. At the time of the earliest formation
of the heart the endothelium on the intestinal fibrillar plate
cannot be seen with the distinctness with which it is shown
in the figure.
Since the formation of the heart is one of the most difficult
processes which embryology has to study, I would like to add
a few more words to the results above described. According
to an observation by P. Meyer, as communicated in his article
* Ueber die Entwickelung des Herzens und der grossen Ge-
fiissstiimme bei den Selachiern,” the endocardium arises in a
manner different from that which we have explained. This
author states that when the intestine closes the two lateral
veins apply themselves together beneath it, and unite into one
vessel, to provide the heart with a single unpaired tube—an
“endothelial saccule.” The figures (Tab. 12. fig. 2, and
others) which P. Meyer has given us entirely support his
view ; and in the embryology of the lamprey also we actually
find a series of preparations which appear to confirm it. For
it is an exceedingly striking fact that the endothelial saccule
of the heart often hes at a great distance from the myocardium,
as if it has arisen independently and has been in no way con-
nected with the lateral plates. Moreover, we find the cardiac
cavity bounded by the lateral plates, without its being possible
to recognize distinctly the endothelium on their inner surface.
Embryology of the River-Lamprey. 4t
Then, too, preparations occur in which the lateral plates have
not yet completely surrounded the lower surface of the
intestine; their two layers lie close together and more
towards the periphery, and yet in the middle of the body-
cavity we find an endothelial pellicle belonging to the future
endocardium. In spite of all these appearances I consider
that I am bound to adhere to the opinion that the endocardium
has separated off from the lateral plates on the inner surface
precisely in the same manner as on the outer. I find myself
compelled to do this by the arrangements presented by quite
young stages of the embryos, measuring about 2°5 millim. in
length. ‘The endocardium is here visible in all sections taken
in the cardiac region, but always in the closest connexion with
the myocardium. Both membranes appear thick, and are
richly provided with yolk-granules.
Before closing this division of my paper, I must not pass
over in silence the statements of A. Goette, since they are at
variance with my observations. According to this author, the
endocardium is formed from endoderm. Since it is difficult to
suppose that so capable an investigator as Goette should have
been mistaken, the material investigated must have been of
such a kind as to admit of what was seen being interpreted in
a different fashion. Goette’s view might be supported by the
circumstance that in the neighbourhood of the rudiment of the
heart the cesophagus is rather strongly compressed at the sides,
and almost touches the ectoderm with its under surface. When
the formation of the heart has taken place, the lumen of the
cesophagus has diminished by one half. Isolated cells are
then not infrequently to be found in the upper angle formed
by the contact of the intestinal fibrillar plates. A constriction
of the cesophagus, as has sometimes been described, or a
splitting-off of a series of cells from its under surface for the
benefit of the endocardium, I have never seen, and I therefore
adhere to the assumption which I have stated above.
THE ENDOBLAST.
The branchial cavity arises by the separation of the cells of
the yolk at the anterior end of the embryo, in the manner which
we have had occasion to observe in the formation of the intestine.
The great thick head, consisting of a compact mass of cells, com-
mences to recede somewhat from the yolk, and becomes smaller
and narrower. Between it and the yolk the anterior portion
of the body of the embryo becomes continually more distinct
and longer. The internal changes proceed hand in hand with
the external ones, The cavity which has been formed, which
was at first more spacious at the anterior end, gradually also
42 On the Embryology of the River-Lamprey.
extends backwards in similar dimensions. The entirely un-
differentiated cells surrounding the cavity assume an epithelial
character. In the canal which becomes the branchial cavity
lateral evaginations appear, which extend from in front back-
wards. ‘The evaginations become so pronounced that the
endoderm cells come into contact with the epithelium of the
integument, which is finally absorbed. Opposite the evagina-
tion a slight depression is sometimes noticeable. At certain
spots it seemed to me that the epithelium of the future slit has
thrust itself so far outwards that the cells of the epidermis here
appear thinner. ‘The groove which arises on the under surface
ot the branchial cavity, in the neighbourhood of the first to the
fifth slit, and which subsequently becomes for the most part
closed off—the thyroid gland—has been described very fre-
quently, and I have nothing material to add, The main
features of the formation of the mouth, the union of the ecto-
dermal pouch with the endoderm of the branchial cavity, are
likewise well understood. If we examine a few longitudinal
sections, which were made parallel to the ventral surface of
the embryo, pretty much such as those which are shown by
A. Dohrn in his plate iv. (Bd. vii.), we are struck by the
similarity between the gill-slits and the two outer angles of
the oral pouch. It seems as though the latter are only the
end of the series of those slits. ‘The similarity appears not
only in the position, but also in the form.
The liver is formed by evaginations of the intestinal wall.
We observe that this is in connexion with canals, which soon
become converted into tubes. At the spot at which these pro-
cesses are taking place the intestine is less abundantly sur-
rounded by yolk-cells than in its posterior division. At all
events the yolk-cells take no part in the formation of the
liver. In spite of Kupffer’s statement that a neurenteric
canal is not formed in the Lampreys, I must most empha-
tically insist upon its occurrence in these animals, though it
is true that at present not a single satisfactory drawing of it
exists.
If we follow the spinal cord in longitudinal sections, we see
with the greatest distinctness how the end of it bends round
the notochord, and then runs for a space beneath it, to end as
a somewhat thinner cord in the upper wall of the end-gut
above the anal opening. My preparatious were stained with
hematoxylin, and the cells of the spinal cord (as also the
neurenteric canal) appeared very dark, in consequence of
which they stood out from the rest. ‘lhe cells were, more-
over, much larger than others lying in the neighbcurhood,
such as, for instance, those of the epithelium. ‘The shape of
The Hon. W. Rothschild on Apteryx Haastii. 43
the canal in question is somewhat obscured, owing to the fact
that a large venous vessel lies near it.
THe BRAIN AND THE GANGLIA.
In a provisional communication, which is not accompanied
by figures, very little can be said as to these organs. Some
days before the embryos are hatched the brain already
possesses several divisions. During the formation of the
optic vesicles the anterior wall of the fore brain is represented
by a very thin lamella. The hemispheres and the olfactory
lobes are developed afterwards. ‘The hypophysis originates
from the ectoblast, while in almost all other animals it is
formed from the cells of the endoblast.
As to the epiphysis, I have made certain statements in the
paper which has recently appeared on the third eye of the
Lampreys. All ganglia (of the head, as also of the trunk) are
produced frorn the cells of the ectoderm. ‘The cerebral ganglia
appear to arise at a very early stage. ‘They subsequently
separate from the epidermis, and appear to be related to the
mesoblast. In embryos of from 2 to 3 millim. in length,
however, we still find them in connexion with the integument.
The spinal ganglia arise later on, and not above the spinal
cord, but to the side of it. I have observed epithelial cells
extending inwards from the skin like a plug between the
protovertebre. At a subsequent period they lay near the
spinal cord, already separated from the skin, and forming
a roundish independent group. From the spinal cord there
proceeded a short fascicle of nerve-fibres—that is, a root—in
the direction of the ganglion-cells. A junction between them
was not as yet established.
In concluding this short communication I cannot refrain
from mentioning that the description and figures by A. Dohrn
of the cerebral ganglia (as, for instance, those of the tri-
geminus, ophthalmicus, facialis, vagus, and other nerves) are
in the highest degree true to nature. I possess a large series
of preparations, which show precisely the same picture as his
figs. 1, 2, & 4 of pl. 10, and especially fig. 6 of pl. 11
(Bd. viii. Heft 2).
VI.—WNotes on Apteryx Haastii.
By WALTER RorHscHILp.
Mr. Forpes has expressed it as his opinion that Apterya
Haast? is a natural hybrid between A.australis and A. Owenii.
I have paid much attention to this question as well as to the
study of this genus generally, having had during the last four
Ad The Hon. W. Rothschild on Apteryx Haastii.
years over seventy living Kiwis of four species, and having
examined some one bandied and sixty preserved specimens,
of which over one hundred belong to my collection (in spirit,
skins, and skeletons), belonging to five or, possibly, six
species. More especially of A. Haastit I have at present :-—
One adult male, alive.
One young male, alive.
Two adult males and four adult females, in skins.
Two young males, in skins,
Two chicks, in skins.
One adult female, stuffed.
Two adult males and two adult females, in spirits.
One chick, in spirits.
One adult male and one adult female, in skeletons.
One egg.
After an examination of these materials I am unable to
agree with Mr. Forbes’s views. A consideration of the
geographical distribution of the various species of Kiwis must
lead to a conclusion opposed to them.
Apteryx aoe Oeste igieeis
Haastit.
t . . South Island.
— australis.
—— maximus . . . .. Stewart’s Island.
—. Owenit.
As A. Haastii occurs only in the North Island, where
A. australis and A. Owen are not found, it is evident that
it cannot be a hybrid between them. However, I must men-
tion that on looking over my skins of A. Owend I find that
seven specimens from the west coast of the South Island are
larger and darker than those from the east coast, and there-
fore that it is quite possible that this larger race of A. Owenti
has been mistaken for the true A. Haastci. ‘This latter is
called “Fioa-Roa”’ by the natives, and not Kiwi.
Apteryx Haastii is the largest species of the genus, a
female in my possession being considerably larger than the
largest known female of A. maximus. It is further distin-
guished by its enormous beak, which, though not so long as
that of A. maximus, is very much stouter even than that of
the recently discovered extinct species.
As far as I can see at present another distinctive character
will be found in the cecum, which seems to differ not only in
the two sexes, but also in the various species of the genus.
Mr. C. O. Waterhouse on the Mouth-organs of Diptera. 45
VII.—Some Observations on the Mouth-organs of Diptera.
By Crarues O, WATERHOUSE.
Some two years ago I had occasion to study the mouth-parts
of Diptera, and among other books consulted Prof. Lowne’s
work on the Blow-fly. In his first edition of this book,
Prof. Lowne put forward the theory that the proboscis of the
Blow-fly is chiefly composed of the maxille, and not of the
labium as usually supposed, and in his new edition he keeps
to this view. Prof. Lowne has arrived at this conclusion from
studying the development of the parts in the embryo &c.
Prof. Lowne says: “ So far as I know there is no dipterous
or hemipterous insect in which there are any traces of
mandibles, and the parts so named are always a part of the
maxilla, and articulate with the palpigerous scale” (New
Edition, p. 151). The species in which I found the parts of
the mouth most perfectly developed is Pangonia longirostris,
and with this insect in view it is impossible for me to doubt
that the larger pair of lancets are really the mandibles, and
that in spite of the care and skill which Prof. Lowne brings to
bear on his investigations, he has nevertheless fallen into
someerror. Prof. Lowne says: “ the only reason for regarding
the terminal portion of the proboscis as a modified labium is
its position, and this is no evidence from a morphological
point of view” (p. 129). This is true, but, so far as | am
concerned, it is certainly not position on/y that influences me
in my determination. With Prof. Lowne and others I consider
the slender pair of lancets in Pangonia to be the maxille ;
this is clear enough from the presence of the maxillary palpi.
The larger pair I consider to be the mandibles, not on/y on
account of their position (although this is just what it ought
to be), but because they appear to have a distinct origin from
the more slender pair, and the structure of the base with its
produced basal angles is only a slight modification of the form
constantly met with at the base of the mandible. I would
especially compare it with the mandible of the Australian
Neuropterous insect Bittacus.
It is somewhat difficult to understand exactly what Prof.
Lowne does consider these larger lancets. He speaks of them
as “parts of the maxilla” without saying what part. If
they are parts of the maxille, I presume they are either the
lacinie or galez.
This, however, does not seem to be Prof. Lowne’s view, for
in speaking of Pulex he says, the pseudolabium is “formed b
the united gale of the maxilla” (p. 152) ; and if the “ pseudo-
46 Mr. O. Thomas on a new
labium ” of the flea represents the galew, then the pseudo-
labium of the Diptera is formed in the same way, and yet we
have still four Jancets in many Diptera to account for!
If, however, these lancets are the mandibles and maxille,
then the sheath of the proboscis would be the labium ; and,
apart from its position, it is frequently divided into three
key very suggestive of the mentum, ligula, and para-
osse.
‘i I hope Prof. Lowne will some day give us a clearer idea of
what he considers these “ parts of the maxille” to be.
When speaking of the mouth-parts of the flea, Prof. Lowne
incidentally mentions that the antenna in that insect is behind
the eye, which, he says, ‘is a clear indication that the simple
eye in the Fleas is not homologous with the great compound
eyes of insects, which are never in front of the antenne ”
(p. 152). When I read this sentence I could not help think-
ing of Prof. Lowne’s statement above quoted (p. 129) that
position is no evidence; and when one sees how completely
the relative position of the eye, ocellus, and antenna change
in such insects as Zryzxalis and Fulgora for instance, I
scarcely think it a convincing argument to say that the eye
of the flea cannot represent the compound eye of other insects,
simply because it is in front of the antenna. A trifle more
and the antenna of Fulgora would be behind the eye.
VIII.— Description of a new Baboon from East Africa.
By OLDFIELD THOMAS.
AMONG some Mammals obtained by Mr. F. J. Jackson in
Kast Africa is a fine male Baboon evidently allied to the
Abyssinian Papio* thoth, Ogilb. (with which I presume
P. degquera, Pucheran, is synonymous), but so different from
it in the character and colouring of its fur that I think it ought
to be separated from it at least as a subspecies, for which I
propose the term
* T am entirely unable to follow those naturalists who, in deliberate
defiance of the laws of priority, use Cynocephalus instead of Papio for the
Baboons. Even on the inadmissible but oft-quoted score of convenience,
it is surely a greater nuisance and source of confusion that some natu-
ralists or curators of large museums (¢. gy. the Leyden) should use Papio
and others Cynocephalus, than that those who are (and know they are)
wrong in using the latter should give it up once for all, and learn the
name which has an unquestionable claim to adoption.
Baboon from East Africa. 47
Papio thoth ibeanus, subsp. n.
Fur remarkably coarse and shaggy all over the body, and
even longer than in P. t. typicus. Colour a coarse mixture
of blackish and dull fawny white, without any of the brighter
yellow always present in the typical form, at least on the head
and limbs. Hairs on the crown of the head broadly annulated
with black, this part being nearly uniform yellow in the true
thoth. Colour of outer side of limbs quite like back, the
hairs slaty grey basally, ringed terminally with black and
dull fawny white. Chin and throat well-haired, whitish, at
first, anteriorly, unannulated, but gradually proceeding back-
wards blackish rings make their appearance, and on the chest
all the hairs are broadly annulated with black and white and
on the belly with black and dull fawn. Inner sides of fore
limbs like chest, but inner sides of hind ones a clearer and
less annulated fawn-colour. Upper surfaces of both hands
and feet like the outer surfaces of arms and legs, not getting
darker terminally.
Skull_—Owing to the fact that the typical and only avail-
able adult male skull of P. thoth is obviously deteriorated by
captivity, it would be useless to enter into a detailed com-
parison between the two skulls; but, comparing the Hast-
African skull with a fine Angolan skull of what I take to be
P. anubis, it is characterized by its much less salient facial
ridges, shorter and broader palate, markedly smaller and
narrower foramen magnum, and higher and more vertically
directed mandibular coronoid processes.
Dimensions of the type (an adult male skin) :—
Head and body (c.) 850 millim.; tail (c.) 610; hind
foot 182.
Skull: basal length 151; greatest length, exclusive of
occipital crest, 195 ; greatest breadth 114; gnathion to lower
edge of orbit 111; nasals, length mesially 64, laterally 68,
greatest breadth 15; height of orbit 25:5; breadth across
orbits outside at fronto-malar sutures 84; nasion to occiput,
exclusive of crest, 99; palate, length 97, breadth between
outer sides of ™-2 56°5, between inner sides of ™? 33; com-
bined lengths of upper true molars 37.
Hab. Lamu, East Africa.
Of other species of the genus besides P. thoth none of the
present group are known from East Africa at all, P. porcarius
being South African and P. anubis and P. sphina being
West African.
48 Mr. O. Thomas on a new Cephalolophus.
IX.—On a new Cephalolophus from Mount Kilima-njaro.
By OLpFrieLp THOMAS.
Mr. F. J. Jackson has presented to the British Museum the
head of a Cephalolophus obtained by him some years ago on
or near Mount Kilima-njaro. This head is clearly that of
the animal doubtfully determined as C. nigrifrons* in
Mr. F. W. True’s excellent account of Dr. Abbott’s Kilima-
njaro collection, and is also, I believe, specifically identical
with a skin obtained many years ago by Sir John Kirk at
Malindi, and, owing to its having lost the fur of its face,
erroneously referred by me to C. natalensis, Sm., in my recent
monograph of the genus f.
At Mr. Jackson’s suggestion I propose to name the species
in honour of Sir Robert Harvey, who was the first of their
party to shoot a specimen of this interesting species; and
taking first as my type the Kilima-njaro head only, it may be
briefly diagnosed as follows :—
Cephalolophus Harveyt, sp. n.
Colour of face and nape as in C. nigrifrons, t. e. ground-
colour chestnut, with a black blaze extending from the nasals
to the occiput, expanding on the forehead ; of cheeks, sides of
neck, and throat pale bay, and of chin white, as in C. nata-
lensis. Horns of male conical, very thick at the base, their
greatest basal diameter going barely two and a half times in
their length.
Dimensions of head :—Muzzle to eye 95 millim. ; to occiput
188 ; ear above crown 70; horns, length 79, greatest basal
circumference 77.. The dimensions given by Mr. True may
also be referred to. :
Hab. Mount Kilima-njaro.
But if, as I see no reason to doubt, the Malindi skin and
skull are also referable to this species, the following points of
difference from C. nigrifrons may also be noted :—
General colour much paler, almost exactly as in C. nata-
lensis ; under surface decidedly paler than upper; feet slen-
derer and more delicate throughout; hoofs not elongated t
* Proc. U. S. Nat. Mus. xv. p. 476 (1892).
+ P. Z. 8. 1892, p, 419. Owing to this specimen proving not to be
C. natalensis, it will be necessary in the account (J. ¢.) of that species to
delete all references to the male, including description of male horns and
measurements of skull, these being all based on this Malindi specimen.
}{ It is probable that the true C. nigrifrons is an inhabitant of swamps
and marshes, as its hoofs are so very markedly elongated as compared to
those of its congeners, The measurement above given is between 30 and
35 millim. in the type specimen.
On the Mexican Representative of Sciurus Aberti. 49
(greatest length of ridge of hoofs above, both fore and hind,
about 20 millim.).
Skull, as compared with an equally fully adult male skull of
C. nigrifrons, that of the type, rougher and more swollen in
the frontal region ; muzzle shorter and more conical, broader
across the posterior ends of the nasals ; parietal region longer ;
orbit smaller and more exactly circular; median notch of
palate but little surpassing the lateral ones ; bulle decidedly
smaller. As to skull-dimensions, those given in my mono-
graph as those of “C. natalensis (g)” (really this Malindi
skull of C. Harvey’) may be compared with those of the
C. nigrifrons on the page following.
This new species may be inserted in the synopsis of species
in the above-quoted monograph (p. 415) as follows :—
a‘. Darker markings shining black or
brown.
a’. Back uniform rufous, no dorsal
stripe.
a®. Ground-colour dark chestnut
above and below; hoofs elon- ,
Sa, AED oo aes oe 5. C. nigrifrons, Gr.
6°. Ground-colour pale bay, lighter
below ; chin white ; hoofs not
Cn 2 ee nee ans 5A. C. Harveyt, Thos.
6°. Back with a black dorsal stripe, &c.
X.—On the Mexican Representative of Sciurus Aberti.
By OLpFieLpD THOMAS.
In 1882 * two squirrels sent from Durango, Central Mexico,
by Mr. A. Forrer were referred by me to Sciurus Abertt,
Woodh., a species previously only known from Colorado,
Arizona, and New Mexico, and which I presumed to extend
southwards down the high lands of the Sierra Madre as far as
the point at which Mr. Forrer found it. Some doubt has
been recently thrown on this determination, and I have there-
fore now made a renewed examination of Mr. Forrer’s beauti-
ful skins. This examination leads me to the conclusion that,
although the identity of the Durango animals with S. Aberti
should still be maintained so far as regards the species, using
this word in the larger sense to which English zoologists are
accustomed, yet there is sufficient difference in their detailed
coloration to necessitate their being considered as belonging
* P.Z. S. 1882, p. 372.
Ann. & Mag. N. Hist. Ser. 6. Vol. xi. 4
BO Mr. A. Dendy on the History of the
toa distinct subspecies. That modern American zoologists
would probably separate this local form as a distinct species
I am quite aware ; but such a procedure would, as in other
cases, have the great disadvantage of suppressing all signs of
the evident relationship existing between the two forms,
without gaining any greater local exactness than may be
obtained by the free use of trinomial nomenclature.
Sciurus Aberti durangi, subsp. n.
Size and general characters of S. A. typicus*. Sides of
nose from muzzle to eye rufous, as opposed to the “ grey
cheeks” of typicus; a ring round eye paler rufous. ars
entirely without tufts (type killed in middle of October) ; no
chestnut or rufous spots round or behind their bases. Hairs
of chest and belly slaty grey basally, as compared to ‘ pure
white” in typicus. Whole under surface of tail inwards of
the submarginal black band coarsely grizzled grey, each hair
being broadly banded with black and white; in typicus the
under surface of the tail is “ wholly white.” Median part of
dorsal surface of metatarsus dark-coloured, proximally like
the lower leg, terminally with a rufous tinge; inner side of
metatarsus and tops of toes white. Descriptions of typzeus
simply say “ feet white.”
Hab. Ciudad, Durango, Central Mexico; alt. 8100 feet,
on the Sierra Madre.
Type B. M. 82. 3. 20. 16. Female, killed Oct. 14, 1881 ;
a second specimen killed Aug. 7. * Coll. A. Forrer.
XI.—Note on the History of the so-called Family Teichonide.
By Artuur Denpy, D.Sc., F.L.8., Melbourne University.
In reply to my note on “‘ The Discovery of the True Nature
of the so-called Family Teichonide,”’ published in the
‘ Zoologischer Anzeiger’ (no. 895), Dr. R. von Lendenfeld, in
no. 402 of the same journal, endeavours to prove that he
* Tt appears to me equally simple and exact, while far more euphonious
to use the word typicus for the typical subspecies, instead of doubling the
specific name, as is commonly done. The original author’s name should
of course be appended after typicus. The ugly sound of a double specific
name is always bad enough ; but if that method is adhered to, we may in
some cases have to employ three repetitions of the same word, e. g. Lutra
lutra lutra, Linn., instead of (if like generic and specific are admitted)
Lutra lutra typica, 1, or, most euphonious of all, j pee vulgaris typica
Erxl. This possibility of a treble repetition may well make us hesitate
before insisting on the same word being used for both species and typical
subspecies
so-called Family Teichonide. 51
arrived at his results quite independently of my work, and
refuses to recognize that [ have any claim to priority in the
matter.
He bases his defence upon a preliminary communication
entitled “ Das System der Kalkschwimme,” which he informs
us that he read before the Vienna Academy on the 8th of
January 1891, and which he accuses me of wilfully ignoring,
although, as a matter of fact, I have never seen the paper in
question up to the present moment.
My principal work on the “ Teichonide” was published in
January 1891, though written in Australia many months
previously ; hence, of course, it is possible that Dr. von Len-
denfeld may not have seen it before he wrote his preliminary
account, though that can be no excuse for not referring to it
in his complete work (‘Die Spongien der Adria’), which is
not even dated till April 1891, and not published till the
following December.
It is evident from Dr. von Lendenfeld’s writings that he
has never personally studied the group of sponges whose true
nature he claims to have independently arrived at, and hence
to an outsider it is difficult to see what could have induced
him so suddenly to abandon the family “ Teichonide ” and
adopt my views as to the relationships of the sponges com-
prised therein.
1 would therefore like to point out, for the benefit of those
who might otherwise be misled by Dr. von Lendenfeld’s
statements :—(1) That on November 14th, 1889, I communi-
cated a paper on “ The Pseudogastrula-stage in the Develop-
ment of Calcareous Sponges” to the Royal Society of
Victoria ; (2) that in this paper I gave a preliminary account
of the results of my researches on the anatomy of ‘ Teicho-
nella labyrinthica” and “ T. prolifera,’ stating distinctly
that “ 7. labyrinthica”’ is a true Syconand “ T. prolifera” a
typical Leucon ; and (3) that I sent a copy of this paper to
Dr. von Lendenfeld.
In short, my main conclusions as to the structure and
relationships of the “ Teichonide” (with the exception of
“Kilhardia Schulzei,”’ concerning which we both owe our
anatomical information to Poléjaeff) were published and
received in Kurope many months before the date on which
Dr. von Lendenfeld says he read the preliminary paper on
which he bases his remarkable claim to independent
discovery.
As I have pointed out previously, Mr. Carter himself was
the first to show that “ Zeichonella labyrinthica” is a true
Sycon ; but this statement of his had passed quite unnoticed,
52 Mr. W. L. Distant on a new Species of Poecilopsaltria.
and no correct account of the anatomy of the sponge in ques-
tion had appeared until the publication of my work. ;
I am aware that in his “ apn of the Australian
Sponges ” (Proc. Linn. Soc. N.S. W. vol. ix. p. 1140) Dr. von
Lendenfeld accepted the “ family” with some hesitation. He
says, “I have not seen any representatives of this family
myself, but am of opinion that they might perhaps be con-
sidered as colonies of Leucones or Sycones, as Marshall
asserted before Poléjaef’s essay was published.’ He also
says of “ Teichonella labyrinthica” (loc. cit. p. 1142), “The
anatomy of this sponge is totally unknown, s9 that its name
and position here are only preliminary.” Hownear Marshall’s
opinion came to the truth of the matter spongologists will be
able to judge for themselves. Since the publication of the
above remarks Dr. von Lendenfeld has repeatedly maintained
the “family Teichonide,” and continued to do so until the
very year in which my preliminary results were published.
In other words, from 1885 till 1890 Dr. von Lendenteld
stuck to the family; then, when the outcome of my
researches on 7’. labyrinthica” and “T. prolifera” had been
published and sent to him, he suddenly dropped the “ family ”
and adopted my views without giving any reasons for so
doing. Surely it is taxing our credulity too much to ask us
to believe that he arrived at his results independently !
Melbourne,
November 8, 1892.
XII.—Description of a new Species of the Cicadan Genus
Peecilopsaltria. By W. L. Distant.
THE very beautiful species here described was obtained by
Herrn P. Frey at Nossi-Bé, an island near the north-west
coast of Madagascar. Dr. Karl Brancsik, who has already
commenced to publish the results of this expedition (‘ Jahres-
hefte des naturw. Vereines des Trencsiner Comitates,’ 1890-
91, xii. u. xiv. Jahrg.), has submitted a specimen to me for
identification, which proves to be a perfectly new species.
Pecilopsaltria Brancsiki, sp. n.
¢+- Head and thorax above ochraceous, the lateral and
osterior margins of the pronotum greenish ochraceous.
lead with a cruciform spot at centre of front and a spot at
each bagal angle of same, a broken transverse fascia between
Mr. W. L. Distant on two Genera of Coreide. 53
the eyes, and a spot near anterior angles of vertex black.
Pronotum with two central lineate black spots, a similarly
coloured semicircular spot at centre of basal margin, and the
incisures blackish. Mesonotum with two tooth-like spots at
centre of anterior margin, followed by a smaller spot on each
side, a slender broken lateral fascia, and a spot near each
anterior angle of the cruciform elevation black. Abdomen
above castaneous, shaded with black, and greyishly pilose.
Abdomen beneath and legs ochraceous; a broad subapical
annulation to anterior femora, the anterior and intermediate
tarsi, and the bases and apices of the tibiae black.
Tegmina pale greenish opaque for more than half their
length, remaining area obscure hyaline where the venation
is fuscous ; the transverse veins at the bases of the second,
third, fifth, seventh, and eighth apical areas are narrowly infus-
cated; two small fuscous spots in upper ulnar area, some
narrow and obscure discal fuscous markings and a series of
small fuscous spots near the apices of the longitudinal veins
to the apical areas. Wings pale greenish, with the venation
and almost the basal half blackish.
The width of the head, including the outer margins of the
eyes, is a little more than that of the base of the mesonotum ;
the lateral angles of the pronotum are broadly and angularly
produced.
Long. ? 41 millim.; exp. tegm. 115 millim.
Hab. Nossi-Bé Island.
XI1I.—WNotes on Two Genera of Coreide found in Madagascar.
By W. L. Distant.
In the last published part of the ‘ Revue d’Entomologie,’
tome xi. p. 264, Dr. Bergroth has continued his criticisms on
the writings of his colleagues who study the order Rhynchota.
Much of this criticism is valuable, some unfortunately is mis-
leading ; for excessive criticism is as open to error as is too
prolific description. Amongst other proposed corrections is
the statement that my genus Parabrachytes ‘est identique au
genre Odontorrhopala*, Stal.” I should be very glad to
adopt this opinion of Dr. Bergroth but for the following
distinctive characters, which he strangely appears to have quite
overlooked or forgotten when he wrote: “ Les seules diffé-
rences sont offertes par les antennes .... et par le rostre.”
* Dr. Bergroth has altered the spelling of this generic name; as pro-
posed by Stal it was “Odontorhopala.”
54 Dr. A. E. Feoktistow on the Physiology of the
Odontorhopala.
Pronotum with the lateral angles
produced in foliaceous lobes.
Abdomen strongly and abruptly
dilated, especially at the fourth
segment in both sexes.
‘irst joint of the antenne longer
than the second.
Parabrachytes.
Pronotum with the lateral angles
moderately prominent and angu-
larly Shieh Sy
Abdomen moderately and regu-
larly dilated in both sexes.
First joint of the antenne shorter
than or subequal in length to the
second,
Other points of difference, but of a minor nature, are to be
found in the structure of the incrassated apices to the second
joint of the antenne.
I am now able to add the description of what appears to be
a second species to the genus Odontorhopala.
Odontorhopala Bergrothi, sp. n.
@. Allied to O. callosa, Stal, but differing from the
description of that species by the following characters :—
The clavus is not black, but concolorous with the corium ;
the second joint of the antenne is not black, but is ochra-
ceous, with the apical incrassated portion only black, and the
sixth as well as the fifth segment of the abdomen is outwardly
toothed.
Other characters not mentioned by Stal are as follows: —The
pronotum, clavus, and corium are not only coarsely punctate,
many of the punctures being black, but the lateral expanded
angles of the pronotum are ornamented with small black
tubercles on their upper surface which are continued in a more
obsolete manner across the disk. ‘The spinous tubercles to
the femora and first joint of the antenne are pale greyish in
hue.
Long. 20 millim.; max. abd. exp. 10 millim. ;
ang]. 10 millim.
Hab, Madagascar, Fianarantsoa.
exp. pronot.
X1IV.—On the Physiology of the Rattle of Crotalus durissus.
By A. E. Froxtistow, M.D.*
In June 1887 I received ten living specimens of Crotalus
durissus, which I have since been able to observe closely,
Owing to want of a sufficient quantity of suitable food (the
* Translated from the ‘Mélanges Biologiques tirés du Bulletin de
l’'Académie Impériale des Sciences de St. Pétersbourg,’ t. xiii. livraison 1,
pp. 1-4: St. Pétersbourg, 1891.
Rattle of Crotalus durissus. 55
animals refused to eat anything but very young rabbits),
I lost five of them in the course of the first six months. The
remainder are in good condition, and now (August 1888)
devour birds in addition to rabbits. They live in a large
den, provided with a cement floor and a spacious water-tank,
and permanently warmed by hot water, which renders it
possible to maintain the temperature of the air in the interior
at 77° to 81°°5 F. (20° to 22° R.). The snakes are provided
with living food in sufficient abundance, and are equally
lively in winter and summer.
This opportunity has enabled me to make accurate obser-
vations upon the growth, shedding, and renewal of the rattle.
Up to the present nothing is known with certainty as to the
physiology of this apparatus. Throughout America, for
example, the inhabitants estimate the age of a Rattlesnake
according to the number of rings in the rattle, and they
believe that a new ring is added annually (Brehm), Others,
again, consider that a new ring is formed at each ecdysis, and
soon. Brehm * points out that though it is true that R-ttle-
snakes which have been kept in captivity and observcd for
several years consecutively have been found to increase in
size, an increase in the number of the joints of their rattles
has not been perceived; on the contrary, these remained
unchanged for years. It is likewise not definitely stated by
any author how long a rattle may become. I am in the
happy position of having been able to make certain obser-
vations upon healthy specimens with good appetites, which
decide these questions.
Five of my snakes shed their long rattles independently at
different times, and I was now able especially to observe how
rapidly these redevelop. I would first, however, remark that
it is perfectly natural for the rattle of the Rattlesnake to fall
off periodically or at irregular intervals. The organ in
question consists of dead horny tissue, which is developed
into the well-known hollow “cones”: while these cones
partly enclose one another, to form the rattle, they are yet
only somewhat loosely connected together. Now it may be
readily understood that such a chain of cones, when it attains
a certain length, is greatly exposed to mechanical shocks, and
consequently may easily break off. Without any injury to
the snake itself, this chain may be also cut off or torn off by
force. This is indeed the simple reason why the rattle never
becomes particularly long, and rattles with from fifteen to
* Brehm, ‘Thierleben,’ Bd. vii., “ Kriechthiere,’ 1883, p. 491,
56 Dr. A. E. Feoktistow on the Physiology of the
eighteen joints are rare *. As a rule, the rattle only lasts
long enough to become eight- or nine-jointed.
When a rattle has fallen off, its last-formed joint (con-
sequently the first, counting from the base) at least is always
left behind on the tail; for it is firmly united with the latter,
until it is replaced by a new one forming underneath it.
This last-formed joint of the rattle has not the dusky-brown
colour of the older joints. It is rather of a horn-like trans-
parency, and only of a faintly yellowish tint. Showing most
distinctly through the thin walls of this joint of the rattle, we
see the horn-producing cone at the end of the tail as a white,
flattened, irregularly conical mass, with a deeply indented
tip.
As I have already stated, I was able to follow the repro-
duction of the rattles in the case of five of the Rattlesnakes,
which had shed these organs. So long as their rattles
remained short, the snakes were naturally also unable to
make a noise. But the joints were gradually replaced, and
in such a way that in all cases in the course of from three to
four months two new ones were already present, in addition
to the joint already mentioned, which was left behind, and
which was now terminal in position. Three-jointed rattles
like these produced a fairly loud sound. In the course of a
year the rattles developed into chains with from five to six
joints, and were then capable of producing the usual very
intense rattling sound. The reproduction of the rattles had
nothing in common with the ecdyses fT. It is well known
that the epidermis is shed without the rattle, separating close
to the margin of the latter, and, indeed, in such a way that
the end of the tail in the cast skin is represented by an
aperture with finely notched edges corresponding to the rows
of scales.
It consequently follows from my observations that a joint
of the rattle can be produced in the course of every two to
three months f, and I do not understand why other observers
have not noticed the growth of the rattle in captive Rattle-
* Rattles of forty-two joints, as figured by Seba, surely exist only in
the imagination !
+ Schlegel (‘ Essai sur la physionomie des serpens,’ t. ii. 1837, p. 557)
thought that a new joint of the rattle is formed at each ecdysis. This
ste has recently been expressed by Garman also. According to my
observations, however, the ecdyses succeed one another much more
rapidly (an ecdysis every six weeks) than the formation of the joints of
the rattle.
t This assertion of course implies the provision of artificial warmth
during autumn, winter,and spring. In the natural state the growth of
tne rattle evidently proceeds much more slowly.
Rattle of Crotalus durissus. 57
snakes. Probably the snakes were kept under conditions
unfavourable for their welfare, whereby the vital processes
were checked. Perhaps, too, the observations were not con-
ducted with sufficient care.
As regards the actual rattling, I can in general confirm the
words of Geyer: if the Rattlesnake is crawling slowly along,
it draws its rattle after it quite quietly along the ground; if,
however, it is endeavouring to escape, it raises the rattle at an
angle of about 60°, and at the same time rattles continuously.
When the snake is in pursuit of its prey, nothing is usually
to be heard of it. ‘This is not always the case, how-
ever ; on the contrary, I often saw Rattlesnakes pursuing the
rabbits which served them for food and making a lond
rattling, whereat, however, the latter showed no fear at all.
When irritated the Rattlesnake assumes the threatening
attitude which is admirably represented in Brehm’s ‘ Kriech-
thiere’ (p. 492), and is at the same time capable of rattling
literally for hours and continuously. The noise which it
thus produces can be with difficulty compared with any other ;
in any case it only very distantly resembles the “ chirping of
a grasshopper,” as is asserted by Brehm. A large powerful
Rattlesnake makes so loud a noise with its rattle that it is
impossible to undersiand the words of a person speaking in a
loud tone at a distance of three paces when the snake’s cage
happens to be between the individuals conversing. ‘The
powerless specimens which we usually see in zoological
gardens give no idea of this. If the vibrating rattle dips
into water, a peculiar sound arises which completely resembles
the hissing of red-hot iron when it is plunged into water.
Beneath the water the rattling is almost noiseless. If a
Rattlesnake is held with one hand behind the head, while the
other hand grasps the end of the tail just behind the rattle, it
becomes impossible for it to make a noise.
I have attempted to determine the number of vibrations
which the rattle makes per minute. A large Rattlesnake was
grasped by the neck, while an assistant thrust a needle
through the middle joint of its seven-jointed rattle in such a
way that it pierced the rattle in its greatest diameter—conse-
quently from above downwards, if we imagine the snake
lying quiet with its tail outstretched upon the ground. Now,
since the rattle (in the position in which we have supposed
the snake to be) is, in making a noise, moved trom left to
right and back again, the needle was able to trace curves of
vibration upon paper blackened with soot. As a registering
apparatus 1 used Dudgeon’s polygraph, with a strip of black-
ened paper which was made to slide rapidly forward by means
58 Bibliographical Notices.
of the clockwork. The tail of the snake was to a certain
extent fixed by my holding the snake with my hand in the
region in front of the anus. After much trouble I succeeded
in bringing the needle in a suitable manner into contact with
the strip of paper, and in obtaining curves of vibration, from
which the number of the vibrations per minute (the rapidity of
the progression of the strip of paper being known) could be
calculated with a fair degree of accuracy. In this manner it
was found that the movements of the rattle are composed of
great vibrations of the entire tail itself and of smaller vibra-
tions of the actual rattle, in such a way that the tail makes
seventy-five and the rattle, on the other hand, one hundred
and ten vibrations per minute. These are approximate
average numbers, since I was able to obtain only taulty
curves, because the rattle does not perform its vibrations
precisely in one plane. Movements kept up for hours with
rapidity like this are absolutely amazing. When observed
with the naked eye, only a blurred image is seen of the rattle
moving at this rate *.
BIBLIOGRAPHICAL NOTICES.
Catalogue of Eustern and Australian Lepidoptera Heterocera in the
Collection of the Ouford University Museum.—Part I. Sphinges
and Bombyces. By Col. C. Swrnuog, F.L.S. &c. Oxford: Claren-
don Press, 1892.
For the production of this volume and the arrangement of the moths
in the Oxford Museum so that they have become available for useful
study we are indebted to the entomological zeal of Col. Swinhoe,
his many years of study of the moths of the Indian region having
rendered him well fitted for the work he has volunteered to do and
so ably begun, a fact evidently appreciated by the University of
Oxford, as testified by the honorary degree they have just conferred
on him.
The volume includes all the Sphinges and Bombyces from the
Oriental and Australian regions in the collection of the late W. W.
Saunders, which was acquired by Prof. Westwood for the Oxford
* The following authors have written upon the structure of the
rattle :—Lacépéde, ‘Histoire des Serpens,’ vol. ii. pp. 390-420, pl. xvii.,
1789; Leuckart, ‘ Anatom.-physiologische Uebersicht des Thierreiches,’
1855; Czermak, ‘“ Ueber den schallerzeugenden Apparat von Crotalus,”
Zeitschr. f. wiss. Zoologie, Bd. viii. pp. 294 et seqq., 1857 ; Wymann,
“The Mode of Formation of the Rattle of the Rattlesnake,” Proceedings
of the Boston Society of Natural History, vol. viii. p. 121, 1861-62 ;
Garman, “The Rattle of the Rattlesnake,” Bulletin of the Museum of
Comparative Zoology at Harvard College, vol. xiii. No. 10, 1888,
Bibliographical Notices. 59
Museum about the year 1877; and it is a matter for the greatest
regret that the species from America and Africa, which would not
have swelled the volume to any great extent, could not have been
included at the same time, so as to make it complete in itself.
The collection was originally arranged and described, whilst still
in Mr. Saunders’s possession, by the late Francis Walker, his descrip-
tions being published in the British Museum Catalogue which he
was then engaged on; and here, as in all his other entomological
work, he fully exhibited his well-known propensities towards
describing the same species over and over again and placing them
in the most incongruous families and genera, so as to render his
description entirely unintelligible without inspection of his types.
In this state of chaos and almost entirely neglected by entomo-
logists the collection has been left till Col. Swinhoe took it in hand
little more than a year ago; and it has been his task to bring it to
London bit by bit and compare it with Walker’s types in the British
Museum and with the types of Jater authors, so that the correct
synonymy of the species might be arrived at; this has been most
carefully done, and little is left to be desired in this direction.
The not less arduous task of reducing the species to their proper
families and genera has also been ably carried out, though there
are some species still out of place; thus Balatea belongs to the
Zygenide, not the Aigeriide, and Bonia probably to the Tinege-
riidxe, whilst the affinities of the Epicopeinz are with the Uraniide,
not the Chalcosiinze, and Duga belongs to the Geometridae, not the
Lithosiide. With regard to the genera, as much has been done as
is well possible taking into consideration the state of utter con-
fusion in which the genera of tropical Heterocera are at present
involved, and many years must elapse before a comparative analysis
of the whole can reduce them to order. The drawing of the neura-
tion of new genera will be found useful, though that of one,
Chalioides, is not very correct, and in another, Platyzyg ygena, the
internal veins of the fore wing have been omitted ; whilst the eight
coloured plates will enable many of Walker’s species to be identified.
For the species the system has been adopted of describing the races
from each different district or island as distinct, as indeed is usual,
though we believe that in the near future this system will be
entirely abandoned and all these slight races be reduced to the rank
of subspecies ; and not till this is done will it be possible to see at a
glance the value of the names in any faunistic list or to deal with
the vast numbers of insects in such a manner as to give a compre-
hensive and intelligible classification of the whole of them; then
perhaps the 2,000,000 species, at which the number of existing
insects has been estimated by several of the late presidents of the
Entomological Society, may be reduced to some 400,000, or about
double those at present known, which seems the only chance for
systematists of avoiding a general migration to the lunatic asylums
of the country ! G. F. Hampson.
60 Bibliographical Notices.
Fossil Plants as Tests of Climate. By A. C. Sewarp, M.A.,, F.G.S.,
&e. Svo. 151 pages. C.J. Clay & Sons: London.
Tuts is the ‘Sedgwick Prize-essay” for the year 1892, and its
subject has been well chosen (1) as being worthy of, and indeed
requiring an earnest and comprehensive study of both recent and
fossil botany by the essayist, and (2) as a subject, a broad view of
which, based on trustworthy details, is a desideratum with geolo-
gists.
~" As is due to observers in the past, though results of value were
scanty even in the early part of the present century, careful refer-
ences to early works, and critical notes on the facts and opinions
therein stated, are given in the “ historical sketch ” (pages 1-32),
which, like the Essay itself, is based on Mr, Lester F. Ward’s
‘Sketch of Paleobotany,’ in the ‘ Fifth Annual Report of the U.S.
Geol. Survey’ for 1883-84, pages 363-453 (1885), and extended to
the present time.
A general knowledge of the present distribution of plant-families,
and of their relation to climatal conditions, must precede the con-
sideration of the coexistence of the ancient and extinct floras with
the successive geographical conditions of bygone lands and their
changing climates; Chapters II. and III., therefore, comprise per-
tinent remarks by trustworthy authors treating of the distribution
of plants, with reference to geography and topography, height of
lanl above the sea-level, the nature of the soil, and, lastly, low tem-
peratures, as in Greenland, Grinnell-land, Alaska, and Russian
Lapland.
In Chapters LY. and V. the influence of external conditions upon
the macroscopic and microscopic structures of plants is considered,
on the basis of published reports with respect to the habit and size
of plants in relation to climate; the form, position, and structure of
leaves ; their minute structure and its relation to external condi-
tions ; water-plants ; acclimatization and naturalization ; and minute
anatomy of fossil plants; also the annual rings in recent and fossil
lants.
a" By far the most interesting lessons in questions of ancient
climates have been taught by fossil plants in the high northern
latitudes of the Arctic regions,” hence Chapter VI. comprises the
notices and remarks made by various observers on (1) the Devonian
and Carboniferous plant-remains of Bear Island, Spitzbergen, and
Melville Island. (2) Upper Carboniferous of Siberia, Novaya
Zembla, and Spitzbergen. (3) Jurassic of Spitzbergen and Melville
Island (?). (4) Cretaceous of Spitzbergen and Greenland. (5) Ter-
tiary of Spitzbergen, Iceland, Greenland, Grinnell-land, Bathurst
Island, Banks’ Land, Prince-Patrick Island, and Mackenzie River.
(6) Quaternary of Spitzbergen.
So far as the fossil plants of the Arctic regions have been studied
©, Heer considered that they give no evidence of any difference in
the temperature of the Polar Regions from that of Central Europe ;
Bibliographical Notices. 61
and it may be taken conclusively that no direct evidence is afforded
in favour of the existence of climatal zones in Carboniferous and
Jurassic times.
The Tertiary Arctic plants were regarded as Miocene by Heer ;
but J. 8S. Gardner has adduced reasons to prove that they show a
passage from the Cretaceous to the Miocene, and that therefore
they may claim to be Eocene.
The difficulty of regarding the plants of the old Coal-measures as
definite indicators of climate is carefully dwelt upon. ‘heir several
structures and relative size cannot be taken without great caution
as indications of a tropical climate, nor of the presence of an excess
of carbonic acid in the air; nor indeed would a hot climate suit
the peat-like decomposition of marsh-plants. Certainly the coal-
plants had a very wide geographical distribution, fayoured by (1) the
absence of the highest plant-forms, and which would have been
strong opponents in the struggle for existence ; (2) their propaga-
tion by spores, being thereby widely disseminated. Nevertheless,
there are important differences in the plant-remains of distinct coal-
fields. Thus the Carboniferous flora of North America contains a
large number of species not represented in that of Europe.
In Chapter VII. the climate and vegetation of the Carboniferous
Period are reviewed according to evidences derived by geologists
from its plant-remains found in different parts of the world. Those
of the European and North-American Coal-measures are abundantly
noticed in the foregoing chapters, as they take their places under
the different structural or other relationships; and now (pp. 102,
&e.) that the probable existence of different climatic zones or
botanical provinces in the Coal-period has to be discussed, the Car-
boniferous (or Permo-Carboniferous) strata and plants of India,
South-Africa, and Australia are taken in order. 1. The Lower
Gondwana System in India, including the Panchet, Damuda, and
the Talchir series. 2. The Karoo formation in South Africa, or
rather the Ecca Beds lying just below it. 3. The Wianamutta,
Hawkesbury, Newcastle, and Muree beds of New-South-Wales.
4. Queensland. 5, Tasmania. These several regions and their
formations are tabulated (pp. 122, 123) after Feistmantel and
Waagen, to show more particularly the horizons of 1. Glossopteris,
2. Glacial phenomena, aud 3. Lepidodendron. The essayist is as
cautious in giving an opinion in this case as in all the other points
of discussion, but seems to accept the opinion expressed by Neu-
mayr, Blanford, and Feistmantel that the Glossopteris flora came in
with the cold climate indicated by the glacial conglomerates, and,
replacing the older Carboniferous types, spread gradually towards
the North, probably from a Southern Continent.
The recent valuable researches by Clement Reid and others on
Pleistocene plants indicative of a cold climate in Britain, as some of
Pliocene age witness low mean annual temperatures for Switzer-
land and elsewhere at that particular period, are referred to in
Chapter VIII. Further accurate research is required to approach
with certainty any conclusion as to climatal conditions in the several
62 Miscellaneous.
geological periods. Some indications have been offered in the
course of the essay, as shown above. The Mesozoic floras in par-
ticular have yet to be carefully reviewed as tests of climate. Sug-
gestive remarks on fossil Ferns and Conifers are then offered ; and
the Author says:—* We may expect that a closer study of the
Geological floras, not only from phylogenetic and anatomical, but
also from biological points of view may enable us to penetrate further
into the life-conditions of those forests of which the Earth’s crust
affords us such numerous, though often too fragmentary, relics.”
The list of works referred to in the text, occupying pages 134-
146, well arranged and made serviceable with reference to the
numerous footnotes, is a welcome bibliography for paleobotanists.
The Essay has also a useful Index. Like other scientific works
issuing from the University Press at Cambridge, this is well printed
on good paper.
MISCELLANEOUS.
Comparative Researches wpon the Organization of the Brain in the
principal Groups of Arthropods. By M. H. Viatranes.
I nave the honour of commupicating to the Société de Biologie
the principal results of researches which I have been conducting for
several years upon the organization of the nervous system of
Arthropods, and of which I have hitherto only published detached
fragments, some in the ‘ Annales des Sciences Naturellés’ and the
rest in the ‘ Comptes Rendus de l’Académie des Sciences.’
Organization of the Brain of Insects.—In the Insects the brain is
formed of three segments corresponding to the first three cephalic
zonites. The first segment, or protocerebron, innervates the eyes;
it is the seat of the visual perceptions, while the psychic centres also
reside in it. The second segment, or deutocercbron, imnervates the
antenne ; it is the seat of the olfactory perceptions. The third
segment, or tritocerebron, innervates the labrum and the initial
portions of the digestive canal; in it is situated the centre of the
gustatory sense.
Before entering further into detail as to the constitution of the
cerebral segments, it may be mentioned that the first two are
entirely precesophageal, that is to say that the commissures which
unite their symmetrical portions are situated in front of the
cesophagus. In the case of the third segment the conditions are
different; here all the commissural fibres pass behind the ceso-
phagus, where they constitute the commissure known under the
name of the transverse commissure of the esophageal ring.
The protocerebron is composed of a pair of lateral masses termed
optic ganglia and of an intermediate median mass. The constitu-
tion of the optic ganglia is most remarkable and most constant ;
Miscellaneous. 63
they are composed of a series of three ganglionic masses united to
one another by decussating fibres. The median protocerebral mass
is formed of a pair of lobes, which are intimately fused together and
contain in their interior—(1) the pedunculate bodies, which are the
seat of the psychic functions; (2) the central body, the organ
whither fibres coming from all points of the brain converge ; (3) the
pons of the protocerebral lobes, a portion discovered by myself, the
significance of which is as yet unknown.
The deutocerebron is composed of a dorsal portion, the structure
of which has nothing particularly noteworthy about it, and of two
olfactory lobes, which are highly differentiated in view of their
special functions, and are characterized especially by the presence
of organs known under the name of olfactory glomerules. The
olfactory lobes, whence the sensory fibres of the antennary nerve
originate, are united to the optic ganglia and to the pedunculate
bodies by fibres which decussate in the median line ; this connexion,
which is absolutely constant, seems to be bound up with physio-
logical necessities. Besides the antennary nerves, the deutocerebron
gives origin to a pair of tegumentary nerves and to a pair of roots
destined for the visceral nervous system.
The tritocerebron is represented in the Insect solely by a pair of
ganglionic masses, which we will designate cesophageal ganglia ;
these are separated from the median line and united to one another
behind the cesophagus by the transverse commissure of the cesopha-
geal ring. Each of the esophageal ganglia gives rise by means of
a common trunk to a nerve, which is destined for the labrum, and
to a root of the visceral nervous system.
In the Insects the visceral nervous system is composed—(1) of a
series of three median ganglia, which are connected with one
another, and of which the first, known by the name of the “ frontal
ganglion,” is united to the esophageal ganglia by a pair of roots,
which are often double ; (2) of a pair of lateral ganglia*. Each of
these latter is connected, on the one hand, with one of the median
ganglia, and on the other with the deutocerebron, by means of a
nerve-root which has already been mentioned.
The brain of the Myriapods is precisely similar to that of the
Insects in structure. But the visceral system of these animals
exhibits a remarkable condition, for it preserves throughout the
whole of life certain characters which in the Insect are only found
in the course of embryonic development.
For our knowledge of the cerebral structure of Per ipatus we ure
indebted to the researches of M. Saint-Rémy Tf ; it is connected in
the closest manner with that of Myriapods and Insects.
* The lateral ganglia are generally each divided into two little masses,
termed by M. Blanchard the. ganglia of the vessels and trachee (“gan-
glions angien et trachéen”).
+ M. Saint- Rémy has published (Arch. Zool. exp. vol. ili. bts) a most
conscientious paper upon the structure of the brain of Arachnids, Myria-
pods, and Pertpatus; this has been of great assistance to me.
64 Miscellaneous.
The Brain of the Crustacea.—Like the Insects and the Myriapods,
the Crustacea possess a brain formed of three segments—protocere-
bron, deutocerebron, and tritocerebron.
The protocerebron of the Crustacea is constructed upon the same
plan as that of the Insects ; we find in it an optic ganglion formed
of the same portions, as well as pedunculate bodies, a central body,
and a pons of the protocerebral lobes. In the same way their
deutocerebron is in every respect similar to that of the Insects and
the Myriapods.
In the Insects and the Myriapods the third cephalic zonite is
devoid of appendages and only bears the labrum; in the Crustacea,
on the contrary, the same zonite bears, in addition to the labrum,
the second pair of antenne. ‘This difference entails a slight modifi-
cation in the structure of the tritocerebron. While in the Insects
and Myriapods the tritocerebron is represented only by a pair of
cesophageal ganglia, in the Crustacea the same cerebral segment is
formed by a pair of cesophageal ganglia, and, in addition, by a pair
of antennary lobes intercalated between the latter and the deuto-
cerebron.
The cesophageal ganglia in the Crustacea, as in the Insects and
Myriapods, are united with one another behind the esophagus by
the transverse commissure of the cesophageal ring, and each of them
gives rise, by means of a common trunk, to the nerve of the labrum
and to a root of the visceral nervous system.
The antennary lobes, the commissural fibres of which pass with
those of the cesophageal ganglia behind the esophagus, give rise to
the nerves of the second antenne, to a pair of tegumentary nerves,
and to the motor nerves of the eye-stalk.
The visceral nervous system of the higher Crustacea differs from
that of the Insects in a single point, which is, however, of but slight
importance. In the Crustacea the unpaired and the lateral ganglia,
instead of being separated as in the Insects, are all fused into a
median mass applied to the wall of the stomach, and known by the
name of the stomatogastric ganglion. This mass is united to the
brain by roots, which are strictly homologous with those which we
find in the Insects. Like the frontal ganglion of the Insects, the
stomatogastric ganglion of the Crustacea is united to the cesophageal
ganglia by a pair of roots, which are generally double, and, like the
lateral ganglia of the same animals, it is connected with the deuto-
cerebron,
We therefore conclude, from what has been stated above, that,
from the point of view of cerebral structure, there exists the closest
relationship between the Crustacea, Insecta, Myriapoda, and
Peripatus.
Limulus and the Arachnids, of which in other respects many
zoologists recognize the affinities, constitute, as regards the organiza-
tion of their brain, a most homogeneous group, but one which
recedes considerably from the rest of the Arthropods.
In Limulus and the Arachnids the brain is composed of two
Miscellaneous. 65
segments only, the protocerebron and deutocerebron, both of which
are provided with precesophageal commissures. The protocerebron,
which innervates the eyes, is comparable to the protocerebron of the
Crustacea and Insects; it is nevertheless to be observed that in
Limulus the pedunculate body attains truly colossal proportions.
The same organ, although considerably modified, is still recognizable
in the Arachnids, in which M. Saint-Rémy has described it under
the name of the stratified organ (“ organe stratifié’’),
In Limulus and the Arachnids the deutocerebron, instead of inner-
vating olfactory antenne, as in the Crustacea and Insects, performs
the same function for the chelicerse, which are simply tactile appen-
dages, and so is not differentiated in view of special sensorial
perceptions. The tritocerebron is wanting in Limu/us and the
Arachnids, and the first ganglionic mass which follows the deuto-
cerebron is devoted exclusively to the innervation of the first
maxillipede or mandible *.
The visceral nervous system of Limulus and the Arachnids is repre-
sented only by lateral ganglia, which, as in the case of the Insects,
derive their roots from the deutocerebron ; the median ganglia are
wanting ; the absence of these centres is evidently correlated to that
of the tritocerebron.
Finally, we may express the differences and resemblances pre-
sented by the different types of Arthropods as regards the organiza-
tion of the brain by dividing these animals into two great groups.
The first of these, comprising the Arachnida and Limulus, is
characterized by the absence of the tritocerebron and the non-
differentiation of the deutocerebron into an olfactory centre.
The second, which embraces the Crustacea, Insecta, Myriapoda,
and Peripatus, is characterized by the presence of a tritocerebron
and the differentiation of the deutocerebron into an olfactory centre.
This group may be itself subdivided into two sections, the first
containing only the Crustacea, which are provided with two pairs of
antennee, the second embracing Myriapoda, Insecta, and Peripatus,
which possess a single pair of antennee.—Comptes Rendus Hebdo-
madaires des Séances de la Société de Biologie, n. s., t. iv. (May 6,
1892), pp. 354-357.
On the Circulation of the Blood in young Spiders.
By M. Marcer Cavsarp.
I have examined the circulation in young spiders belonging to
fifteen genera of Dipneumones, of which the following twelve have
been determined by M. Eugéne Simon:—Dictyna, Tentana, Theri-
dion, Epeira, Zilla, Micariosoma, Chiracanthium, Teatrix, Clotho,
* The rostrum of Arachnids is analogous to the labrum of Crustacea
and Insects, but it belongs to the second zonite und is innervated by the
deutocerebron.
Ann. & Mag. N. Hist. Ser. 6. Vol. xi. 5
66 Miscellaneous.
Pardosa, Phlegra, Heliophanus. Three other genera examined
came from eggs found under stones without the adult females.
My results differ in certain points from those obtained by Clapa-
réde for Lycosa. The ramitications of the aorta are indeed such as
were observed by this author, but the recurrent branch arising from
the cephalic arteries conducts the globules into a lacuna occupying
the median portion of the upper face of the cephalothorax, and
which is not, as was supposed, an actual gutter ; the globules which
circulate in it from the front towards the rear, returning from the
ophthalmic lacunee, form a sort of sheet spread out beneath the
integument ; then, overflowing at the sides without following any
definite route, they unite with the currents which skirt the lateral
portions of the cephalothorax. In the Saltigradw, in which the
latero-posterior eyes are very large and placed well behind the
others, the globules which have circulated round these eyes arrive
at the central lacuna by following real gutters. Between the
median anterior eyes globules are observed to penetrate between
these organs to giin the sternal face of the cephalothorax.
In young spiders which have already undergone the first ecdysis
and are still transparent other ramifications of the cephalic arteries,
not remarked by Claparéde, may be observed in the cephalothorax,
These canals, which communicate with the recurrent branch of the
cephalic artery, of which mention has been made above, carry the
globules back towards the posterior portion of the cephalothorax ;
at certain points of their course globules may even be observed as
they rise from the depth of the organs. The arrangement of these
canals is the same in all the genera studied, except in the Salti-
grade, where it is slightly different. The appearance of these
ramifications, which do not exist immediately after the animal is
hatched, is highly interesting, since it shows that if, in consequence
of the exclusive study of young spiders, the arterial system of these
creatures has been considered as being very slightly ramified, it is
nevertheless susceptible of complication and of attaining in the adults
the development which the researches of M. Blanchard, confirmed
by those of M. Schneider, have demonstrated.
In the appendages the globules of the arterial current follow one
another in narrow file. The venous current is broader and forms a
sort of sheet heneath the integument on the extensor side; it may
be temporarily subdivided by muscular masses,
In all the genera which I examined I found the sternal branches
of the pedal arteries, which open into the transverse sternal lacune.
In the median sternal lacuna the globules come from the deep
parts chiefly through a sort of notch, corresponding to the interval
between the second and third pairs of limbs. Of these globules
some go towards the rear, and the rest towards the front, to empty
themselves at last into the transverse lacune and rejoin the venous
currents returning from the appendages upon the sides of the thorax.
It seems, moreover, that the route followed by the globules of the
Miscellaneous. 67
blood in the various cephalothoracic lacune may be slightly
variable.
At the point where it divides to give off the pedal arteries each
trunk of the aorta emits a short branch towards the sternal face ;
the two sanguineous currents thus formed encircle the sucking
stomach, and, uniting forthwith, travel below the cesophagus and
come into view beneath the integument, pouring numerous globules
into the median sternal lacuna. This is probably the first of the
anastomoses described by M. Schneider, that which gives off the
subeesophageal vessel.
Though the abdomen is but slightly transparent the heart is
generally quite visible, as also the two anterior pairs of its orifices
(pylocardia of Schneider). The posterior pair is usually very diffi-
cult to observe. The blood which returns from the lungs into the
pericardium enters the heart partly by the anterior and partly by
the median orifices. In the posterior portion of the pericardium
the globules circulate from behind forwards, to arrive at the median
and posterior orifices. The blood which, penetrating into the ante-
rior portion of the heart, is not sent into the aorta, circulates in
this organ from in front backwards. I have nevertheless once seen
in the anterior region of the heart the sanguineous current produced
from behind forwards, because the greater portion of the blood
returning from the lung gained the median instead of the anterior
orifices of the heart.
I have not been able to directly observe globules issuing from the
heart by the lateral arteries of M. Schneider ; but in young speci-
mens of Heliophanus I have succeeded in seeing in the posterior
portion of the body a sanguineous current recede from the heart
and branch off to lose itself in the liver. In young individuals of
Dictyna and Chiracanthium I have perceived at the sides of the
abdomen a current of globules proceeding towards the ventral face,
which probably belonged to one of these arteries.
The blood which escapes from the posterior portion of the heart
passes into the pygidial lacuna, which surrounds the anus and the
spinnerets. It divides into two currents, which reach the ventral
face; a few globules separate from the rest to circulate in the
spinnerets before rejoining the common route. These two currents
flow forwards, following the longitudinal ventral muscles, and
mingle with those which have come from the cephalothorax, in the
interval which separates the two lungs. The globules disappear
behind the respiratory lamellae, to reappear on the outer side and
be swallowed up in eddies by the corresponding pulmonary vein,
which conducts them to the pericardium opposite the anterior
orifices.
The whole of the blood which reaches the heart has not previously
passed through the lungs. Asa matter of fact globules may be
observed at the sides of the abdomen, which, issuing probably from
the mass of the liver, appear beneath the integument and there
circulate until they reach the pericardium, whence they penetrate
68 Miscellaneous.
into the heart. In young specimens of Pardosa I have seen globules
leave the two abdominal currents, some near the spinnerets and
the others towards the middle of the ventral face, pass round the
sides of the abdomen and fall directly into the pericardium. Finally,
in some young individuals of Pardosa and Heliophanus globules,
instead of penetrating into the lung, skirted its external border, and
then, cireulating beneath the integument, passed directly into the.
pericardium.
In conclusion: the vascular system, which is very little ramified
in newly hatched spiders, becomes complicated later on ; the venous
blood circulates in a very extensive series of lacune. The whole of
the venous blood of the cephalothorax is arterialized before reaching
the heart: a portion of that of the abdomen returns directly to the
pericardium, and from thence to the heart, without passing through
the lungs.—Comptes Rendus, t. exiv. no. 18 (May 2, 1892),
pp. 1035-1038.
A Contribution to the Knowledge of the Anatomical Structure of the
Sexual Organs in the Galeodide. By A. Brrvta, of the Zooto-
mical Institute of the University of St. Petersburg. (Provisional
Communication.)
The chief results of my investigations into the anatomico-histo-
logical structure of the genital organs in the Galeodide are the
following.
My studies were conducted upon :—
a. Galeodes araneoides, Pall. (¢ & 2);
b. Galeodes ater, Bir. (9 ).
The male genital organs are constructed as follows :—
1. The external genital aperture is represented by a longitudinal
slit in the protuberance of the posterior margin of the first abdo-
minal segment ;
2. Aciniform (so-called accessory) glands, with a chitinized
intima, open into the uterus masculinus, which is clothed with
chitin ;
3. Each of the seminal ducts (vasa deferentia) divides in the third
abdominal segment into two rami, which, suddenly narrowing, pass
into the filiform testes ;
4. In the walls of each vas deferens, at their opening into the
uterus masculinus, lie aciniform accessory glands, with columnar
epithelium, but without an intima;
5. At the period of the maturity of the sexual products the end
of each ramus of the vasa deferentia, which is histologically indis-
tinguishable, swells up into a vesicle and functions as a vesicula
seminalis ;
6. The testes consist of four thin and very long coiled tubes, which,
Miscellancous. 69
before they open into the vesicula seminalis, lose their typical
epithelium and form
7. The special glandular portion, which serves to secrete the
chitinous substance for the membrane of the spermatophores ;
8. The semen enters the sexual organs of the female in the form
of oval and somewhat flattened spermatophores.
The female sexual organs are constructed in the following
manner :—
1, The external genital aperture has the same appearance as in
the male ;
2. The vagina is clothed with a thick chitinized intima ;
3. The receptacula seminis are represented by two vesicles with
chitinized intima, and open into the vagina in the neighbourhood of
the genital aperture ;
4, On its posterior wall the uterus is provided with two auricu-
late appendages, which do not differ in histological structure from
the remaining portions of the uterus, and, as it appears, play no
physiological part whatever ;
5. The oviducts pass immediately into the ovaries. The walls of
these two sections are longitudinally folded, in consequence of which
when the sexual organs become filled with ova or spermatophores
they are capable of considerable expansion, whereby the cavity of
the organs is increased ; they consist of (a) the external adipose
layer, (6) the circular musculature, (¢) the longitudinal musculature,
(d) the tunica propria, and (e) the columnar epithelium. In the
first three layers an abundant ramification of trachez is observable ;
6. The ova develop from a special epithelial layer, which clothes
the wall of the ovaries on the interior ;
7. The ripe ova, which already lie in the follicles which become
evaginated, have a so-called “ stylum ” * ;
8. In the cavity of the ovaries and of the oviducts there may be
observed a considerable number of free cells which bear a strong
resemblance to the bloed-corpuscles. The cells possess the power
of amcebvid movement and exhibit figures of karyokinetic division.
They demolish the envelopes of the spermatophores, thereby libera-
ting the spermatozoa, and at the same time destroy the superfluous
spermatozoa and the unfertilized ova 7;
9. The ripe ova fall into the cavity of the ovaries, where the
development of the embryo is completed ;
10. Even before the formation of the rudiments of the appen-
dages a great difference in form is noticeable between the thoracic
* Bertkau, “ Ueber den Generationsapparat der Araneiden,” Archiv f,
Naturgeschichte, 1875, p. 245.
+ Corpuscles of this kind have been described by Prof. A. Schneider in
Nephelis, Aulostomum, and Hirudo—A. Schneider, “ Ueber die Auflésung
der Hier und Spermatozoen in den Geschlechtsorganen,” Zool. Anz. 1880,
no, 46, p. 19.
70 Miscellaneous.
and abdominal segments of the germinal streak. The segment
which bears the chelicere is separated later than the remainder of
the thoracic segments, at a period when from three to four abdo-
minal segments have arisen from the caudal section ;
11. The segmentation of the appendages appears at a somewhat
early stage ;
12. There are no embryonic envelopes ;
13. A flexure of the embryo takes place, as in the Ara-
neina;
14. The lateral organs, which were described by Croneberg *,
are represented in younger stages by large elongate vesicular sacs,
which are connected with the body above the first pair of legs by
means of a thin stalk. In the young immediately after birth the
lateral organs are considerably diminished in size and shrunken.
In the adult animal, apparently, the linguiform triangular folds of
skin which are found beneath the mandibles must be regarded as a
remnant of the lateral organs.—Biologisches Centralblatt, xii. Bd.,
no. 22 (November 15, 1892), pp. 687-689.
On Two Species of Myzostoma parasitic upon Antedon phalangium,
Miller. By M. Henri Provuo.
Antedon phalangium is the host of two species of Myzostoma
described by von Graff under the names Myzostoma pulvinar and
M. alatum, and which were both discovered in the Minch during
the expedition of the ‘ Porcupine.’ I have met with these two
parasites on their usual host in the dredgings made in the course
of last summer by the boat belonging to the Arago Laboratory ;
and this enables me to communicate forthwith certain interesting
features in the history of these Myzostomidee, which are so little
known. I pass over the anatomical and histological details, which
will be dealt with elsewhere.
Myzostoma pulvinar.—Herr yon Graff, whose description of this
species was founded upon a unique specimen, has well characterized
its external form ; he has drawn attention to the dorsal position of
the mouth and the cloacal orifice, but he must have been led astray
by the bad state of preservation of the specimen which he studied,
for he states that the organs which are known in the other species of
Myzostoma under the name of suckers are absent in this form,
though these structures are really present, although not so well
developed as in the majority of the other species.
Contrary to the opinion of von Graff, Myzostoma pulvinar does
not live upon the disk of Antedon phalangium ; it inhabits the
* A. Croneberg, “Ueber ein Entwicklungsstadium von Galeodes,”
Zool. Anz., 10 Juhrg., 1887.
Miscellaneous. 71
alimentary canal of its host, in which it is sufficiently deeply
ensconced to be invisible from the exterior. The Myzostoma occupies
almost the whole of the first portion (esophagus and stomachal sac)
of the alimentary canal of the Comatula, and is situated in such a
way that its anterior extremity is turned towards the aboral pole.
It applies itself by its ventral face, which is very convex and bears
the ten parapodia, to the digestive epithelium of the Comatula, while
its concave dorsal surface, looking towards the buccal orifice of the
latter, forms a channel which affords a passage to the food-currents,
which nourish at once host and parasite. The gutter-like form of
the dorsal surface of Wyzostoma pulvinar explains how it is that the
latter, in spite of its considerable dimensions, does not completely
obstruct the alimentary canal of the Comatula.,
Myzostoma pulvinar, which is the first endoparasitic form
belonging to this genus with which we are acquainted, is a dicecious
species with well accentuated sexual dimorphism. In this, as in
other points of its organization, it approaches the cysticolous
members of the genus.
In linear dimensions the female is four and a half times larger
than the male; it measures 4°5 millim. in each direction. It
presents no trace of testes either in the adult or young state. Asin
the hermaphrodite species, its uterus communicates with the exte-
rior by three ducts—a median one, which is a direct prolongation
of the uterus itself, and two lateral ones, which open into the ante-
rior portion of the cloaca (rectum).
The male only measures 1 millim. in length by 0°8 millim. in
breadth ; it hooks itself on to the integument of the female, upon
which it is able to move pretty rapidly. In shape it is flattened
and elliptical, recalling by the form of its body the free species of
Myzostoma. Its alimentary canal is not branched, but exhibits on
each side indications of the three ramifications which are seen in all
the other species of the genus; its mouth, which is situated quite
close to the marginal border, is ventral in position. It possesses
two testes, one on each side of the alimentary canal, and each pro-
vided with a vas deferens which opens on the ventral surface.
The two sexes must become associated at a very early period, for
I have observed a young female measuring 1-7 millim. in length
which bore upon its back a male of 0-7 millim. The female in this
instance did not as yet present any trace of ovaries, but already had
the characteristic form of the adult.
Myzostoma alatum.—This species lives as a parasite upon the disk
of Antedon phalangium, and its relations to its host are the same as
those of Myzostoma glabrum to Antedon rosacea : its anatomy does not
essentially differ from that of Myzostoma glabrum. Like the latter
it is hermaphrodite, and my object in drawing attention to this
species is occasioned by the fact that the observations which I have
been able to make with regard to it raise a serious doubt as to the
existence of the so-called complemental males in the hermaphrodite
species. It is true that there are very frequently found upon the
72 Miscellaneous.
dorsal surface of Myzostoma alatum what are supposed to be com-
plemental males to the number of one or two, which exactly recall
those which were described by Beard in Myzostoma glabrum, and
which von Graff had probably ‘been right in considering as young
ones. Now a series of observations has convinced me, without any
possible doubt, that the individuals attached to the dorsum of the
hermaphrodite Myzostoma alutum are young ones of the same species,
which, while they are males in their youth, with well-developed
spermatozoa and vasa deferentia equal to those of the adult, increase
in size and acquire during growth ovaries which are identical with
those of the hermaphrodite form which bears them, and this without
abandoning the dorsal surface of the latter. These supposed comple-
mental males moreover possess, even in their youth and at a time
when they show no trace of ovaries, genital ducts corresponding to
the female genital ducts of the hermaphrodites (a fact which hus
already been observed by Nansen in Myzostoma giganteum and
M. Carpenteri).
The complemental male therefore has no existence, in the proper
sense of the word, in the case of Myzostoma alatum. This species is
a proterandrous hermaphrodite form, in which the two male and
female genital glands coexist in the adult. That the young male at
the outset of its existence plays the part of a complemental is
possible, but it is not proved; and in all cases its male condition is
only transitory, which indicates perhaps that hermaphroditism in
Muyzostoma is an acquired condition, and not a primitive one,
I do not as yet possess proof that the complemental males of
Myzostoma glabrum also acquire ovaries ; but the facts which I have
ohserved in the case of Myzosioma alatum lead me to believe that
they do.
In conclusion, I would remark that the four species of Mzyzostoma
with which we are at present acquainted as existing in the Mediter-
ranean present us with three, or perhaps even with four, types,
which are of interest from the point of view of the sexual organi-
zation of the genus Wyzostoma, viz. :—
Hermaphrodite type, Myzostoma cirriferum.
Proterandrous hermaphrodite type, M/yzostuma alatum.
Hermaphrodite type with complemental male(?), Wyzustoma
glabrum.
Dicecious type with dwarf male, Myzostoma pulvinar.
— Comptes Rendus, t. exy. no, 20 (November 14, 1892), pp. 846-849.
Ann.k Mag. Nat. Hist.8.6. Vol. AL Pl. 1.
Nintern bros. Iith.
Ann. & Mag. Nat Mist. &.6.Vol AT. PUL.
Mintern Bros. lith.
Anrw.& Mag. Nat. Hist Vol.5.6. Vol 11. PUL.
————4
eC
ll. 12. 1s.
; R Mmtern del. et lith Mintern Bros . mmp-
THE ANNALS
AND
MAGAZINE OF NATURAL HISTORY.
{SIXTH SERIES.]
No. 62. FEBRUARY 1893.
XV.—Natural History Notes from H.M. Indian Marine Survey
Steamer ‘ Investigator, Commander C. F. Oldham, R.N.,
commanding.—Series IL., No. 7, An Account of the Collec-
tion of Deep-sea Asteroidea. By A. Atcock, M.B.
Surgeon-Captain I.M.S., late Naturalist to the Survey.
[Plates IV.-VI.}
§ 1. Introduction.
THE Starfishes to be described in the sequel have been
dredged since the year 1885, all in water over 100 fathoms,
most in water over 200 fathoms, and very nearly half in
water over 1000 fathoms deep. A large collection of littoral
and shallow-water forms has also been made, but these are
not here considered. If it be thought objectionable to have
separated the deep-water from the shallow-water forms, it
may be urged in justification that within the limits of Indian
seas, so far as our experience at present goes, there is no
instance of the two sections overlapping, and, on another
ground, that almost nothing has been published, and nothing
else is promised, about the extremely interesting Asteroidea
of the deeper waters of India. Of the basins into which
these waters may conveniently be divided the Bay of Bengal
proper—the basin best explored by the dredge so far—gives
us the smallest number of unknown species. Beyond the
Ann. & Mag. N. Hist. Ser. 6. Vol. xi. 6
74 Mr. A. Alcock on
limits of the 380-fathom line it would seem as if the over-
whelmingly muddy bottom of the Bay presented conditions
specially unfavourable to the existence of starfishes ; and
after passing this limit we usually dredge nothing until we
reach true bathybial conditions in the middle of the Bay. At
561 fathoms on the Andaman side, however, we have met
with Brisinga, and at about 500-700 fathoms, opposite to
the Godavari and Kistna Deltas, on a bottom of hardening
clay that seems to bea special habitat for F/abellum (japont-
cum and laciniatum), Bathyactis, Phormosoma, and Spatan-
goids, Pentagonaster again appears. When we come to the
middle of the Bay, where Globigerina-ooze is accumulating,
we find forms that are familiar to deep-sea explorers all the
world over, namely Pararchaster, Dytaster, Porcellanaster,
Styracaster, Hyphalaster, Paragonaster, Zoroaster, Marsi-
paster, Hymenaster, and Freyella.
The enclosed basin of the Andaman Sea in its moderate
depths appears to be peculiarly favourable to starfish life,
though of its greatest depths—6V0 to 1200 fathoms—we
know nothing. The calcareous sand and ooze (coral, Glodi-
gerina, and Pteropod) accumulating between 200 and 300
fathoms seems to afford to Asteroids, as to Ophiuroids and
Echinoids, an optimum of development. Not only is there
abundance, but there is also variety. Though our dredgings
have been few, they have brought in twenty-one species, of
which sixteen (including three remarkable generic types) are
new toscience. The species belong to the following genera:—
Pseudarchaster, Porcellanaster, Astropecten, Pentagonaster,
Dorigona, Nymphaster (common), Anthenotdes, Palmipes,
Chetaster, Zoroaster (common), Cribrella, Pedicellaster,
Stolasterias, Brisinga, Persephonaster (uear Plutonaster),
Dipsacaster (an Astropectinoid), Milteliphaster (near Calli-
aster), and Dictyaster (near Plectaster).
Concerning the Laccadive Sea our limited zoological
knowledge is almost entirely confined to the depths (700 to
1200 fathoms) in the immediate vicinity of the atolls. The
Gulf of Manaar may here be very naturally included. The
calcareous bottoms of these basins seem to be very favourable
to starfishes, and we have in a few hauls found representa-
tives of eighteen species of the following genera :—Parar-
chaster, Pontaster (very common), Porcellanaster, Pentagon-
aster, Nymphaster, Mediaster, Zoroaster (common), Brisinga,
and Persephonaster.
Our observations have done but little to illustrate the
habits of life of the deep-sea Asteroidea. Like some of the
common reef-forms they must sometimes live in swarms, as,
a
ro
‘
Indian Deep-sea Asteroidea.
for instance, Zoroaster carinatus, of which over a score have
been taken at one haul, Pontaster hispidus, of which about
fifty have been dredged at the same time, and Nymphaster
Jlorifer, of which a hundred and fifty have come up on the
tangle-bar. Like their shallow-water relatives they seem to
subsist largely on mollusks, the shells of which, along with
the chitinous remains of prawns and amphipods, are often to
be found in their stomachs; but some of the characteristic
bathybial forms, as Pararchaster, Pontaster, Porcellanaster,
Styracaster, and Hyphalaster, appear to gorge themselves
with Globigerina-ooze. A curious case of symbiosis, which
has been observed too often to be a merely accidental associa-
tion, occurs between Dictyaster renophilus and an annelid.
Some of the species noticed in this paper have already been
described in joint papers by Professor Wood-Mason and
myself upon the recent dredging-operations of the ‘ Investi-
gator ;’ but owing to the discovery of further material in the
examination of the collections of former years, I have found
it necessary to amend the diagnoses of Persephonaster and
Dictyaster, and I have redescribed Dytaster anacanthus,
Persephonaster croceus, Persephonaster rhodopeplus, Dictyaster
zenophilus, and Asterias mazophorus, and have more fully
described three species of Brisinga, namely B. tnsularum,
B. andamanica, and B. bengalensis, which before were merely
named.
I have to express my great obligations to Mr. Sladen’s
classical volumes on the ‘ Challenger’ Asteroidea, to which I
owe guidance on every point. Mr. Sladen’s classification is
followed throughout, in the conviction that it is the expression
of natural relations.
§ 2. List of the Asteroidea, with Descriptions of the
Species believed to be new.
Family Archasteride.
PARARCHASTER, Sladen.
1. Pararchaster semisquamatus, Sladen.
Pararchaster semisquamatus, Sladen, ‘Challenger’ Asteroidea, p. 7,
pl. ii. figs. 1 and 2, pl. iv. figs, 7 and 8; and in Wood-Mason and
Alcock, “ Indian Deep-sea Dredging,” Ann. & Mag. Nat. Hist., Dec.
1891, p. 428.
Bay of Bengal, on a bottom of Globigerina-ooze, in 1664
fathoms.
Colour uniform salmon-red.
G2
76 Mr. A. Alcock on
2. Pararchaster Huddlestonii, sp. n.
Rays 5. R=117. R=about 170 millim. in the
type specimen.
Disk very small; rays depressed, very long, tapering.
Abactinal surface covered with small, rather distant,
unequal-sized nodular plates, each of which bears a single
erect acute spine or spinelet, the spines on the disk being
very stout and measuring in the type specimen 5 to 8
millim. in length, while the spinelets on the rays are small
and slender and decrease in length from about 2 millim. near
the base to less than a millim. at the periphery of the ray.
Supero-marginal plates more than 60, entirely lateral and
slightly oblique in position, elongate, oval, tumid ; each with
a central tubercle bearing a stout acute spine with a basal
circlet of spinelets, one of which is sometimes considerably
enlarged, especially in the interbrachial region; the odd
interbrachial plate is in the form of a salient eminence, upon
which stands a spike of preeminent size with a basal circlet
of spinelets larger than those on the other plates.
Infero-marginal plates alternate with the supero-marginals
and very oblique, elongate, pyriform, tumid; each bears
usually three stout sharp spines in an oblique series, the most
abactinal being the largest and often having at its base a pair
of spinelets; in the interbrachial are the spines, especially
those of the odd plate, are a good deal aborted. The intervals
between the infero-marginal plates throughout the ray are
occupied each by a large multivalve or pectinate pedicellaria.
Adambulacral plates large, broad, tumid, each with a
furrow-series of six to eight compressed obtuse spinelets in a
semicircular comb, and with three spinelets obliquely placed
on its actinal surface, the two of these nearest the furrow
being of conspicuous size.
Mouth-plates large, each with a furrow-series of six or
seven spinelets, increasing in size from periphery to centre,
and with a suture-series of three much larger spinelets.
Actinal interradial areas very small, not extending beyond
the second or third marginal plate; the intervals between the
plates occupied by large multivalve (pectinate) pedicellarie,
but the plates are otherwise quite unarmed usually.
Anus subcentral. Madreporiform plate large, subcircular,
flat, with fine radial striations.
Ambulacral furrow wide; tube-feet very large, with a
small sucker at point.
Papule confined to rudely stelliform areas at the very base
Indian Deep-sea Asteroidea. 77
of the rays, reaching to the level of the fourth marginal
plate.
Bay of Bengal, Carpenter’s Ridge, 1520 fathoms, Globi-
gerina-ooze.
3. Pararchaster violaceus, sp. n.
Rays 5. R=9-107. R=about 100 millim. in the
type specimen.
Disk very small ; rays depressed, very long, tapering.
Abactinal surface covered with close-set nodular plates of
unequal size, each of which bears from two to four minute
capillary spinelets, visible only with a lens on the rays, but
plain to the naked eye on the disk, where a few in the centre
become distinct spines.
Supero-marginal plates more than 35, entirely lateral and
slightly oblique, elongate, rudely oval, tumid; each with a
central tubercle, bearing a very stout and sharp spine
surrounded at its base by three to five spinelets, one or two ot
which are usually large; the odd interbrachial plate is a
salient eminence which supports a spike of preeminent size
rising out of a circle of large spinelets.
Infero-marginal plates alternate with the supero-marginals
and oblique, elongate, pyriform, tumid; each with two stout
sharp spines, of which the upper is nearly equal in size to
and the lower about half the size of the supero-marginal
spine, and with several distant spinelets ; in the interbrachial
ares the spaces between the plates to the number of four are
occupied each by a large pectinate pedicellaria.
Adambulacral plates large, broad, tumid; each with a
furrow-series of seven or eight obtuse spinelets in a semi-
circular comb, and actinally with two large spinelets. Mouth-
plates large, each with a furrow-series of seven or eight
spinelets, increasing in size from periphery to centre, and with
a suture-series of two much larger spinelets.
Actinal interradial areas extremely small, with apparently
in each area only two pairs of distantly spinate plates and a
large pectinate pedicellaria between each pair.
Anus subcentral and very distinct. Madreporiform plate
rather large, strongly convex, with close radial striations.
Ambulacral furrow narrow.
Papulz confined to oval areas quite at the base of the rays,
reaching only to the level of the second marginal plate.
Colours in the fresh state delicate pink, with the disk
violet owing to the mud contained in the stomach and its
ceca.
78 Mr. A. Alcoek on
Laceadive Sea, 1200 fathoms, coral and Globigerina-
00ze.
PoNTASTER, Sladen.
4. Pontaster mimicus, Sladen.
Pentaster mimicus, Sladen, ‘Challenger’ Asteroidea, p. 48, pl. vi. figs. 1
and 2, pl. vii. figs. 5 and 6,
Laccadive Sea, 1000 fathoms, olive mud.
5. Pontaster cribellum, sp. n.
Rays 5. R=5r. R=45 millim. in the
type specimen.
Very closely related to P. subtuberculatus, Sladen, from the
South Pacific Ocean.
No pedicellarize of any kind present.
Disk of moderate size; rays rather short.
Abactinal surface covered with small paxille formed of
from six to twelve uniformly minute spiniform granules, three
or four round the madreporiform body, and one in each inter-
radius having about twenty granules.
Marginal plates uniformly covered with similar granules.
Supero-marginal plates about 21, broad, tumid, almost
entirely abactinal in position, articulating rather obliquely
with one another; each with a central eminence supporting
a single stout sharp spine.
Infero-marginal plates broad, tumid, alternating with the
supero-marginals in the basal haif of the ray, coinciding with
them in the distal half; each with a stout sharp spine at its
upper (abactinal) end.
Adambulacral plates distant, broad, each with a furrow-
series of about twelve spinelets, standing in an almost circular
yalisade round a single large actinal spine. Mouth-plates
Jurge, each with a furrow-series of six or seven spinelets, of
which the most central (adoral) is many times larger than any
of the others and slightly larger even than the large actinal
spine of the plate, and with a suture-series of small spinelets,
Actinal interradial areas small, not extending beyond the
third adambulacral plate, and carrying only about six finely
spinate plates.
Anus central, distinct. Madreporiform plate small, situated
midway between the centre and the margin of the disk, and
outside a very big paxilla.
Indian Deep-sea Asteroidea. 79
Papularia small and compact, with only five or six papule
in each,
Colour in the fresh state delicate pink.
Laccadive Sea, 1200 fathoms, coral and Globiyerina-
o0ze.
6. Pontaster hispidus, Aleock and Wood-Mason.
Pontaster hispidus, Ann. & Mag. Nat. Hist., Dec. 1891, p. 428.
This species is very closely related both to P. forerpatus,
Sladen, and to P. mimicus, Sladen, but especially to P. fored-
patus, var. echinata, Sladen, from the southern limit of the
Indian Ocean.
Abundant on the green mud of the Laccadive Sea at about
1000 fathoms.
7. Pontaster pilosus, sp. n.
Also allied to P. mimicus.
Rays 5. R=about 67. R=70 millim. in the
type specimen.
Disk of moderate size; rays rather long.
Abactinal surface covered with densely crowded spinose
paxille of two kinds in nearly equal proportions, the one kind
crowned with about eight spinelets of uniform size, the other
kind bearing also a central spine—these, seen in mass, pro-
ducing a shaggy appearance.
Marginal plates extremely tumid, uniformly invested with
capillary spinelets.
Supero-marginal plates 30 to 35, forming a broad abactinal
border to the disk and rays, each with a large spine quite on
its inferior (actinad) margin. [In P. hispidus the spine is
quite on the superior (abactinad) margin of the plate. ]
Infero-marginal plates alternate with the supero-marginals
nearly to the tip of the ray, each with a large spine at its
superior (abactinad) margin, and with from one to three much
smaller and slenderer spines vertically beneath it.
The distant adambulacral plates are so convex marginally
as almost to meet from opposite sides across the furrow; each
has a furrow-series of about eight spinelets upstanding in a
semicircle around a large actinal spine, which is nearly as big
as the spines of the corresponding supero-marginal plate ;
outside the actinal spine an irregular row of capillary spinelets
completes the circle with the furrow spinelets. ‘The mouth-
plates are hardly modified from this type, but, like the first
80 Mr. A. Alcock or
two or three adambulacral plates, they bear two large actinal
spines, and the most central (adoral) furrow-spine is conspic-
uously enlarged.
The actinal interradial areas form small crescents, the horns
of which touch the proximal end of the third infero-marginal
plates ; each area carries 21 to 24 small plates in three con-
centric rows, the plates being closely covered with capillary
spinelets, which occasionally fall into the semblance of a
pectiniform pedicellaria, and sometimes bearing also a central
spine.
Anus subcentral. Madreporiform plate small, placed about
midway between the eentre and the margin of the disk.
Papularia small and compact, each with about 10 papule.
No true pedicellarie.
Gulf of Manaar, 597 fathoms, green mud.
Dyvaster, Sladen.
8. Dytaster exilis, Sladen.
Dytaster evilis, Sladen, ‘ Challenger’ Asteroidea, p. 65, pl. 11. figs. 3
and 4, pl. iv. figs. 9 and 10; and in Wood-Mason and Alcock,
“Indian Deep-sea Dredging?’ Ann. & Mag. Nat. Hist., Dec. 1891,
p. 429,
‘This species has several times been met with in the Bay
of Bengal between 1748 and 1924 fathoms on Globigerina-
OO0ZEe.
9. Dytaster anacanthus, Alcock and Wood-Mason.
Dytaster anacanthus, Ann. & Mag. Nat. Hist., Dec. 1891, p. 429,
Rays 5. R=about 67. R=82 millim. in the
type specimen.
Disk small, irregularly inflated ; rays long, narrow, rigid,
tapering ; interbrachial ares rather acute.
Abactinal surface densely crowded with small paxille
formed of narrow tabule capped with numerous clones
granules or papilliform spinelets, those in the centre of the
disk and in a narrow band along the middle of the rays being
smaller than those elsewhere, and those on the rays being
disposed in obliquely transverse rows on each side of this
median band,
The supero-marginal plates, 40 to 45 in number, are
entirely vertical and lateral in position, almost perfectly
square, except in the inte:brachial ares, where their vertical
diameter is greater, and are uniformly covered with papilli-
torm granules without any enlarged spines or tubercles,
Indian Deep-sea Asteroidea. SI
The infero-marginal plates correspond in number, size, and
shape with their exactly superposed fellows of the upper
marginal series ; they are uniformly covered with papilliform
granules, and, except in the apical third of the ray, each
bears near the middle of its distal (aboral) margin a long fine
accumbent spine, and often another, and occasionally two
other, similar but smaller spines vertically below the first.
Adambulacral plates long and large, fairly coinciding base
to base with the infero-marginals, each with a straight
furrow-comb of 6 to 8 equal obtuse spinelets, and actinally
with a dense mass of smaller spinelets which usually fall into
three fairly distinct longitudinal series. Mouth-plates very
prominent, with widely open sutures, and with the most
central (adoral) furrow spine enormously enlarged ; actinally
each plate is thickly covered with spinelets in a triserial
arrangement.
Actinal interradial areas narrow, but extending to the
eleventh or twelfth infero-marginal plate, and bearing thickly
spinulate plates, the spinelets of which have a tendency to
fall into pedicellaria-like clumps.
Anus distinct, central, forming a longish chink-like vent to
a short broad intestine. Madreporiform plate placed near the
margin of the disk and much concealed by paxille.
Colour in the fresh state uniform light rose-madder.
Bay of Bengal, 1748 fathoms, Globigerina-ooze and
pumice.
The determination of the relations of this species has always
given me great trouble. Of described and figured bathybial
starfishes it comes nearest to Psilaster gracilis, Sladen
(‘Challenger’ Asteroidea, p. 230, pl. xli. figs. 5 and 6,
pl. xli. figs. 9-11), which Mr. Sladen states to be so far
different from all its congeners that he only places it with
them with hesitation. As far as mere external characters go,
our species seems to agree with Psilaster gracilis in almost
every detail; but I have satisfied myself by dissection that
our species has an alimentary canal with both ends opening
to the exterior, and for the present I leave it with
Dytaster.
PERSEPHONASTER, Alcock and Wood-Mason.
Close akin to Plutonaster, Sladen.
Disk large, flat ; rays of moderate length, rigid.
: ’ y Tay Stn, rs Yad
Marginal plates more or less covered with papilliform
stars Paar i:
spinelets and bearing each one or many strong rigid spines :
the supero-marginals, which form a broad massive border on
82 Mr. A. Alcock on
the abactinal surface of the disk and rays, directly super-
posed to the infero-marginals, plate to plate.
Abactinal surface covered with close-set paxille, which on
the rays are arranged in curved transverse rows without any
enlarged median series; papule distributed everywhere
between the paxille. .
Actinal intermediate areas large, extending far along the
ray; the plates closely covered with coarse spinelets.
Adambulacral plates with a furrow-series of obtuse, com-
pressed, slightly radiating spinelets, and actinally with two
longitudinal series of coarse papilliform spinelets.
All the spinelets on the actinal surface of the animal are
covered with membrane and are grooved, somewhat resembling
pedicellariz to the casual view.
Anus distinct, subcentral. Madreporiform body small,
concealed, distant from the margin of the disk.
No true pedicellariz are present.
The generative glands have the usual position in the inter-
brachial ares.
‘The ambulacral furrow is remarkably wide.
This genus was first defined in the Ann. & Mag. Nat.
Hist., Dec. 1891, p. 430. The examination of further
material now permits more precision in the definition.
10. Persephonaster croceus, Alcock and Wood-Mason.
(PI. TY .tigs. 4, 2.)
Persephonaster croceus, Ann. & Mag. Nat. Hist., Dec. 1891, p. 430.
Rays 5. R=45r. R=95 millim. in the
type specimen.
Disk and rays quite rigid, their abactinal surface covered
with close-set spinose paxillee, which become small and very
crowded towards the subcentral anal aperture, and are arranged
in transverse series on the rays; a few incipient pedicellarie
adjoin the marginal plates on the disk; the whole abactinal
surface is perforated by close-set papule.
‘The very massive supero-marginal plates number 31, and
are directly superposed on the infero-marginals, plate to
plate; each is coarsely granular in the centre and finely
spinulate at the edges, and each bears two sharp rigid spines,
oue at the abactinal, the other near the actinal end, the former
being the smaller and often bifid.
The massive infero-marginal plates correspond, plate to
plate, with the supero-marginals; they are uniformly covered
with papilliferm granules, which are largest in the middle of
the plate, and each bears near its upper (abactinad) end a
Indian Deep-sea Asteroidea, 83
sharp rigid spine, beneath which is an obliquely vertical row
of three or four very slender accumbent spines.
Adambulacral plates with the furrow-margin slightly con-
vex, and armed with a comb of six or seven longish compressed
spinelets; actinally each carries two longitudinal series of
small inflated membrane-clad spinelets, which are grooved
down the middle like a barley-com, four in each series.
Mouth-plates small, tumid, with a close suture ; each with a
furrow-series of about seven spinelets, the most central (adoral)
of which is of enormous relative size, and with two longitu-
dinal series of close barleycorn spinelets.
Actinal interradial areas large, the intermediate plates
extending to the thirteenth or fourteenth infero-marginal ;
all are closely covered with barleycorn spinelets, which in
the plates adjoining the adambulacrals fall into clumps
resembling multivalve pedicellariz.
Madreporiform plate small and inconspicuous, placed about
two diameters from the margin of the disk.
Ambulacral groove extremely broad; tube-feet large,
conical. The spirally coiled sacculated intestine opens at a
distinct subcentral vent.
Colour in the fresh state olive-yellow, marginal plates
pink, tube-feet red.
Gulf of Manaar, 738 fathoms, green mud.
This species, like the rest, seems to resemble Psi/aster
patagiatus, Sladen (‘ Challenger’ Asteroidea, p. 232, pl. xli.
figs. 3 and 4, pl. vil. figs. 11 and 12), which Mr. Sladen
considers to be an abnormal Astropectinoid of Archasteroid
affinities, and quite unlike any other species of Psd/aster.
11. Persephonaster rhodopeplus, Alcock and Wood-Mason.
Persephonaster rhodopeplus, Ann. & Mag. Nat. Hist., Dec. 1891,
p- 431.
Rays 5. R=3:5r. R=about 65 millim. in the
type specimen.
Disk and rays quite rigid, their abactinal surface covered
with very close-set tabulate paxille surmounted by numerous
uniform flat-topped granules ; the paxilla: are very small and
crowded towards the subcentral anal aperture, and on the
rays fall into transverse curved rows; the whole abactinal
surface is perforated by close-set papule.
The supero-marginal plates number about 28, and are
directly superposed on the infero-marginals, plate to plate ;
they are covered with granules, which are coarse and distant
in the middle of the plate, and are armed with sharp rigid
spines—those in the interbrachia with one, those along the
84 Mr. A. Alcock on
rays with one, or with two or three in a vertical series; at
the base of the rays each plate usually bears on its extreme
abactinal edge one or two incipient pedicellariz.
The infero-marginal plates correspond, plate to plate, with
the supero-marginals ; they are almost smooth in the middle
and are covered with papilliform granules round the edge, and
are armed with from two to four stout accumbent spines in a
median vertical series, the most abactinad being the largest.
Adambulacral plates with a strongly convex furrow-margin
armed with six to eight short, truncated, longitudinally-
grooved spinelets, and actinally with two longitudinal series
of similar spinelets, about five in each series ; these spinelets
are sometimes almost clavate. Mouth-plates small, very
narrow, with the suture widely open; each with a furrow-
series of about ten spinelets, of which the most central (adoral)
is much enlarged, and with a single suture-series of eight or
nine barleycorn spinelets.
Actinal interradial areas large, the intermediate plates
extending to about the eighteenth infero-marginal; in the
interradial areas each plate carries a pedicellaria-like clump
of from six to eight truncate or clavate grooved spinelets,
and along the rays each usually carries two short longitudinal
series of similar spinelets.
Madreporiform plate small and inconspicuous or quite
concealed; placed about midway between margin and centre.
Ambulacral groove very broad; tube-feet large, conical.
The intestine is similar to that of P. croceus and opens by
a long narrow vent.
Colour in the fresh state “ crushed strawberry,” with some-
times a golden suffusion; marginal plates pink, tube-feet
blood-red.
Laccadive Sea and Gulf of Manaar, 738 to 902 fathoms,
green mud.
In a very young specimen a granular epiproctal tube is
observed.
12. Persephonaster celochiles, sp. n.
Rays 6. R=4:75r. R=117 millim. in the
type specimen.
Disk thin, flat, flexible; rays very broad at base, but
tapering to a fine point, thin and flat, with a remarkably
broad ambulacral furrow.
Abactinal surface with close-set paxille consisting of a
slender cylindrical column crowned by a thick brush of
capillary spinelets ; the paxille of the disk small and irregu-
—
Indian Deep-sea Asterotdca. 85
larly placed, those on the rays in beautifully regular trans-
verse rows; the whole abactinal surface closely perforated
with papule.
Supero-marginal plates 42, superposed on the infero-
marginals, plate to plate ; almost entirely abactinal in position ;
closely covered with membrane-clad granules, and bearing at
the actinad end each a stout procurrent spine.
Infero-marginal plates corresponding, plate to plate, with
the erased covered with similar granules, and
armed with an obliquely vertical series of, usually, three stout
procurrent spines.
The adambulacral plates are practically within the furrow,
lying far below (¢. e. above in the normal position of the
animal in life) the level of the infero-marginals; they are
distant inter se, and their slightly convex furrow-margin
bears a comb of, usually, six long needle-like spinelets, while
on the actinal surface there are two longitudinal rows of
inflated membrane-clad barleycorn spinelets. Mouth-plates
long and large, extremely tumid actinally; each with a
furrow-series of seven or eight spinelets, the most central
(adoral) of which is of enormous relative size, with a suture-
series.of ten or eleven spinelets, of which the most central is
enlarged, and with a few scattered granules between.
Actinal interradial areas narrow, but long, the intermediate
plates reaching in a discontinuous series to the twenty-third
infero-marginal, and those along the ray lying in a furrow
between the adambulacrals on the one side and the infero-
marginals on the other; each plate with a clump of barley-
corn spinelets.
Madreporiform plate small, somewhat concealed by paxille ;
placed midway between margin and centre.
Anus central, distinct, receiving the termination of a wide
intestine.
Ambulacral groove of great width, its width being two
fifths that of the ray ; tube-feet very large, conical, in two
rows.
Andaman Sea, 230 to 250 fathoms.
This species is readily distinguished by the actinal hollowing
of the rays and by the flexibility of the disk.
PSEUDARCHASTER, Sladen.
13. Pseudarchaster mosaicus, Alcock and Wood- Mason,
Pseudarchaster mosaicus, Ann. & Mag. Nat. Hist., Dec. 1891, p. 432.
Andaman Sea, 188 to 220 fathoms, green mud.
86 Mr. A. Alcock on
Family Porcellanasteridz.
PoRCELLANASTER, Wyville Thomson.
14. Porcellanaster ceruleus, Wyville Thomson.
Porcellanaster ceruleus, Wyville Thomson, Voy. Chall, Atlantic, vol. i.
p- 378, figs. 97 and 98; Sladen, ‘Challenger’ Asteroidea, p. 134,
]. xx.; and in Wood-Mason and Aleock, “Indian Deep-sea
Jredging,” Ann. & Mag. Nat. Hist., Dee. 1891, pp. 433 and e
Andaman Sea, 683 fathoms, blue mud; Laccadive Sea,
738 fathoms, green mud; and Bay of Bengal, 1664 to 1748
fathoms, Globigerina- ooze.
STYRACASTER, Sladen.
15. Styracaster horridus, Sladen.
Styracaster horridus, Sladen, ‘ Challenger’ Asteroidea, p. 150, pl. xxiii.
figs. 5-7, pl. xxvii. figs. 17-20; and in Wood-Mason and Alcock,
“Indian Deep-sea Dredging,” Ann. & Mag. Nat. Hist., Dec. 1891,
p. 434.
Bay of Bengal, 1748 and 1803 fathoms, Globigerina-ooze
and pumice.
16. Styracaster armatus, Sladen.
Styracaster armatus, Sladen, ‘ Challenger’ Asteroidea, p. 153, pl. xxiv.
figs. 1-4, pl. xxviii. figs. 1-4.
Bay of Bengal, 1840 and 1924 fathoms, Globigerina-ooze.
17. Styracaster clavipes, Alcock and Wood-Mason.
Styracaster clavipes, Ann. & Mag. Nat. Hist., Dec. 1891, p. 434.
Bay of Bengal, 1748 fathoms, Globigerina-ooze and
pumice.
HYPHALASTER, Sladen.
18. Hyphalaster tara, Aleock and Wood-Mason.
Hyphalaster tara, Ann. & Mag. Nat. Hist., Dee. 1891, p. 434.
Bay of Bengal, 1748 and 1997 fathoms, Globigerina-ooze
and pumice.
Indian Deep-sea Asteroidea. 87
Family Astropectinide.
ASTROPECTEN, Linck.
19. Astropecten, sp.
A small Astropecten, which I think is undescribed, occurs
in the Andaman Sea at about 250 fathoms.
DIPSACASTER, gen. nov.
Disk large, with flat rigid rays of moderate length.
Abactinal surface of disk and rays with compact definitely
arranged paxill and numerous intervening papule.
Marginal plates with highly developed ridges and fasciolar
channels ; the infero-marginals with enlarged spines.
Actinal interradial areas large, with numerous intermediate
plates extending far along the ray, and bearing compact
rosettes of spinelets.
Adambulacral plates with a palmate or pectinate furrow-
series of spinelets, and actinally with several series of
spinelets arranged in rosette-like groups.
Madreporiform plate large but concealed.
No anus exists, and though a minute pore is present in the
centre of the disk, it has no communication with the lumen of
the intestine. No pedicellaria,
20. Dipsacaster Sladeni, sp.n. (PI. V. figs. 3, 4.)
Rays 5. R=3 to 3:3 r. R=110 millim. in the
type specimen.
Disk large, quite flat; rays very broad, quite flat, bluntly
pointed ; interbrachial ares wide.
Abactinal surface densely covered with paxillee which have
broad bases, long slender cylindrical pedicles, and a glome-
rular crown of crowded capillary spinelets ; the paxille are
arranged with beautiful regularity in close series of chevrons,
the angles of the chevrons pointing to and converging at the
centre of the disk.
The marginal plates have the form of great salient cubical
blocks, separated from one another by deep trenches; they
are densely covered with papillary granules, which at the
edges become capillary spinelets that stretch across and com-
pletely fill up the trenches.
The supero-marginal plates number 32 (exclusive of the
terminal plate); they are without any enlarged spines or
tubercles ; they are entirely abactinal in position, forming a
88 Mr. A. Alcock on
broad border to the abactinal surface of the disk and rays.
Beyond them, when viewed abactinally, project the infero-
marginals for about one fifth of their extent, so that from this
view the disk and rays have a double-festooned border—an
inner broad border formed by the supero-marginal plates, and
an outer narrower fringe formed by the uncovered projecting
ends of the infero-marginal plates.
The infero-marginal plates correspond, plate to plate, with
the supero-marginals ; each carries on its projecting abactinad
end a tuft of enlarged spinelets, of which about three, usually
in an obliquely vertical row, are particularly large; in the
interbrachial ares, as far as the sixth plate, there are usually
from two to four additional enlarged spines in a vertical row
down the middle of each plate.
The adambulacral plates are large and distant, each bears
a palmate furrow-series of about eight long needle-like
spinelets, and actinally a radiating rosette-like or paxilla-like
group of about twelve very much shorter spinelets. Except
that the mouth-plates are a little larger and have slightly
larger spines, they are not much modified from the adambu-
lacral type.
The actinal interradial areas are very large; on the disk
each area carries about ninety strongly carinated plates
arranged in slightly oblique rows; and on the rays similar
plates extend in a double series to about the tenth, and in a
single series to about the twentieth infero-marginal. In the
undenuded state each plate has the appearance of a very large
compact and beautifully expanded glomerular paxilla, owing
to the fact that the central carina of the plate bears a multi-
tude of stiff radiating spinelets arranged like the florets of a
composite flower.
Madreporiform plate very large, entirely concealed by
enlarged paxille, placed a little nearer to the margin than to
the centre. There is a central pore, anal in position, but
having, as is seen by dissection, no connexion with the gut.
The tube-feet, which are in two rows, have a simple
papillary point.
Andaman Sea, 250 fathoms.
I beg to name this fine species after Mr. Percy Sladen.
Family Pentagonasteride.
PENTAGONASTER, Linck, Sladen.
21. Pentagonaster investigatoris, sp. n.
R=16r. R=82°5 millim.
Disk pentagonal, much inflated in the radial, very concave
Indian Deep-sea Asteroidea. 89
in the interradial areas, flexible ; rays very short and broad,
flat, pointed, rigid, upturned at the tip.
Abactinal surface covered with uniformly small round
tabular plates, which are distinctly isolated from one another
and are fringed with a single row of flat squamous membrane-
clad granules flush with the general surface, but are otherwise
naked, except that some of the plates (perhaps one fourth)
bear a very excentric or quite marginal broadly bilobed pedi-
cellaria. Papule long, crowded everywhere, except in a
narrow band down the middle of each interradial area, where
the plates are in close apposition.
Marginal plates 17 in each series from mid-interbrachium
to tip of ray, quadrangular, fringed with squamous granules
like the abactinal plates, and occasionally bearing a deciduous
bilobed pedicellaria; the supero-marginals with a_ small
central patch of deciduous granules, the infero-marginals with
a much larger similar patch ; at the very tip of the rays these
patches of granules are placed on distinct elevations.
Adambulacral plates with a furrow-series of 6 or 7 coarse
prism-shaped spinelets, and actinally with (i.) a longitudinal
series of three or four large coarse distant granules, one or two
of which in a series are often replaced by a bivalve pedi-
cellaria ; and (ii.) numerous distant conical granules, arranged
in two irregular rows, but sometimes only in one. Mouth-
plates large, with the furrow spinelets chisel-shaped, the most
central (adoral) one being enlarged.
Actinal interradial areas very large, continued to the tip
of the ray; the intermediate plates are rather closely covered
with small rounded granules which are more or less decidu-
ous; many of those nearest the actinostome carry one or two
rather broadly bilobed pedicellariz.
Anus excentric. Madreporiform plate of moderate size,
laced a third of the way from the centre to the margin, with
fine radial striations.
Colour in life brilliant orange.
Bay of Bengal, off the Kistna Delta, 678 fathoms, brown
mud,
22. Pentagonaster arcuatus, Sladen.
Pentagonaster arcuatus, Sladen, ‘ Challenger’ Asteroidea, p. 277, pl. li.
figs. 1 and 2, pl. xviii, figs, 5 and 6.
1 am not quite certain about this determination, although
our species conforms exactly to Mr, Sladen’s description, and
although the habitat supports it—the ‘ Challenger’ specimen
being dredged in the green mud of the Japanese Sea at
id
Ann. & Mag. N. Hist. Ser. 6. Vol. xi. i
90 Mr. A. Alcock on
345 fathoms, and our specimen being dredged in the green
mud of the Andaman Sea at 271 fathoms.
23. Pentagonaster intermedius, Perrier.
Pentagonaster intermedius, Perrier, Nouv. Archiv. du Mus. d’Hist. Nat.
sér, ji, tom. vi., 1883, p. 243, pl. v. fig. 5.
From the Laceadive Sea, off Minnikoy, 1200 fathoms,
coral and Globigerina-ooze.
Colour in the fresh state salmon-red.
24. Pentagonaster pulvinus, sp. n.
Rays 5. R=2:2 r. R=33 millim. in the
type specimen.
Near Pentagonaster mirabilis, Perrier (Arch. Zool. expér.
vol. v., 1876, p. 40).
Disk pentagonal, much inflated abactinally and hollowed
actinally ; rays relatively long and narrow, blunt-pointed,
strongly upeurved in the distal half. The strongly convex
abactinal surface of disk and rays is covered with hexagonal
or polygonal plates, so close-set that, although their boundaries
are quite definite, no papule are visible on denudation nor
any papular pores, and all closely covered with angular
granules which show a distinctly paxilliform arrangement ;
the basal interradial plates are more than twice the size of any
of the other abactinal plates.
Marginal plates 17 in the upper, 19 in the lower series,
measured from mid-interbrachium to tip of ray, all very
closely and uniformly covered with granules except the
terminal six to eight in the supero-marginal series, which
have a central smooth oval boss.
Adambulacral plates with a furrow-comb of about seven
nearly equal-sized lamellar spinelets, and actinally with two
longitudinal rows of granules, and between these and the
furrow a row of three coarse spinelets, the adcentral (adoral)
of which is often replaced by a pedicellaria with two spathu-
late valves,
Actinal surface deeply concave; the actinal interradial
areas are large ; the actinal intermediate plates extend to the
fourteenth infero-marginal, they are large and roughly quad-
rangular, and ‘are so closely covered with granules that their
limits are not easily discerned.
Anus very indistinct. Madreporiform plate small and also
very indistinct ; it lies outside a much larger basal interradial
plate, and is inconspicuous not merely because it is small,
Indian Deep-sea Asterotdea, 91
but also because its coarse discontinuous vermicular erosions
give it a granular appearance much like that of an ordinary
plate.
Colour in the fresh state salmon-red.
Laccadive Sea, off Minnikoy, 1200 fathoms, coral and
Globigerina-ooze.
MILTELIPHASTER, gen. nov.
Allied to Calliaster, Gray.
Rays 5, moderately long.
Abactinal surface of disk and rays covered with a close
mosaic of round or hexagonal naked plates, which (like the
marginal, adambulacral, and actinal interradial plates) have a
fringe of squamous membrane-clad granules lying flush with
the surface of the plate, and often carry a single large spike
or a large pedicellaria; papule conspicuous between the
abactinal plates.
Marginal plates large, naked, armed with one or two great
spines, which may be sharp or may end in swollen bifid or
multifid points ; sometimes with a pedicellaria also.
Adambulacral plates with a close comb of furrow-spines,
and actinally with (two) great spines which end in swollen
multifid points ; sometimes with a pedicellaria also.
Actinal interradial areas large, some of the plates with
great spines swollen and multifid at tip, or with a pedi-
cellaria.
Madreporiform plate small, nearer to the centre than to
the margin of the disk. Anus distinct.
The most characteristic feature of the species upon which
this genus is, not without hesitation, established is the curious
form of the large spines of the under surface; they end in
swollen bifid or multifid points, and, recalling as they do the
long spines of certain Cidaroids, may be called incipiently
florigerous.
25, Milteliphaster Wood-Masoni, sp. n. (PI. VI. figs. 5-7.)
Rays 5. R=36r. R=80 millim. in the
type specimen.
Disk rather large, very thin, rigid, flat, with slight central
and radial abactinal convexities ; rays rather long, broad at
base, thin, rigid, tapering and gently curving upwards from
base to tip, the tip being turned right over.
Abactinal surface closely covered with quite smooth,
naked, circular, oval, or hexagonal plates, which are fringed
7%
92 Mr. A. Alcock on
with a single close series of small squamous membrane-clad
granules lying nearly flush with the general surface; the
central plates and the mid-radial plates to a point about
halfway along the ray are much enlarged, and usually bear a
central large coarse erect spine from 5 to 8 millim. in length,
and often also a large excentric pedicellaria formed of two
short-stalked spoons or obovate leaves ; the plates.elsewhere
are small and occasionally carry a pedicellaria, but never a
spine ; papule emerge everywhere between the plates, but
much more numerously in the centre and along the margin of
the disk.
Marginal plates quite smooth, edged like the abactinal
plates with squamous granules.
The supero-marginal plates number 20, excluding the
globular terminal plate, and are entirely lateral and vertical
throughout; they are tumid, and each carries at its abactinad
end a large coarse sharp spine, from 5 to 9 millim. long,
standing rigidly outwards almost at right angles to the ray ;
at the tip of the ray the plates have two spines ; many of the
plates have also one of the large pedicellaria formed of two
spoons or obovate leaves.
The infero-marginals correspond se to plate with the
supero-marginals in the interbrachial ares, but not in the
distal half of the ray; each bears on a central eroded
eminence one or two, occasionally three, coarse spines, not
quite so long as those of the supero-marginal series, and
differing from them in having the points bifid or trifid; a few
of the plates carry also one of the large obovate pedi-
cellariz.
The adambulacral plates, which, in addition to the edging
of squamous membrane-clad granules, have a distantly
granular surface, bear a furrow-comb of eight to ten close
truncate spinelets, and actinally a transversely arranged pair
of large spines (Pl. VI. fig. 7) equal in length to those of the
infero-marginal series, and, somewhat like them, ending in
swollen multifid points not unlike the long spines of Doro-
cidaris bracteata or of Goniocidaris florigera; a few of the
plates have also one of the large pedicellarie. Mouth-plates
with the central (adoral) furrow-spine enlarged—4 millim.
long in the type specimen—and each with one large “ flori-
gerous ” spine actinally.
Actinal interradial areas large, extending to the eighth
infero-marginal plate; the actinal intermediate plates are
arranged in concentric chevrons or semicircles, in addition
to the edging of squamous granules they often have an
incomplete second marginal series of granules, and occa-
Indian Deep-Sea Asteroidea. 93
sionally either a central large “ florigerous”’ spine or a large
obovate pedicellaria.
Anus central. Madreporiform plate small, radially striated,
placed a little nearer to the centre than to the margin,
Colour in life a network of cinnabar-red on a yellow
ground abactinally.
Andaman Sea, 230 to 290 fathoms.
I have named this magnificent species after its discoverer.
Doricona, Gray.
26. Dorigona pentaphylla, sp. n.
A magnificent species, very near to Dorigona longimana,
Mobius.
Rays 5. R=3:257r. R=115 millim. in the
type specimen.
Disk large, flat, pentagonal; rays long, broad at base,
tapering. All the plates—actinal, marginal, and abactinal—
except the adambulacral perfectly smooth and edged with
foliaceous granules lying flush with the general surface.
Abactinal surface of the rays entirely occupied by the
supero-marginal plates, which from the seventh onward meet
across the ray; abactinal surface of the disk covered with
hexagonal plates, of which those in a broadly oval petal-
shaped area in each radius are distant regular hexagons—in
the mid-radial line of predominant size—with long tentacle-
like papule standing near the angles, while those in a trian-
gular area in each interradius are crowded irregular hexagons
without any papule intervening.
The marginal plates number about 30 in each series,
measured from mid-interbrachium to tip of ray; they are
broad massive squares and form a broad margin on both
surfaces; the supero-marginals from and including the
seventh meet across the ray in a zigzag suture; the infero-
marginals are smaller than the supero-marginals and alternate
with them, except in the interbrachia, where the two series
nearly correspond,
The adambulacral plates are small ; each has a semicircular
furrow-series of small foliaceous spinelets—ten or eleven in
number in the basal half, eight to six in number in the apical
half of the ray—of which those at the ends of the series are
thickened, while actinally there are three irregular longitudinal
rows of depressed granules; most of the plates near the
actinostomal end have a central bivalve or trivalve pedi-
94 Mr. A. Alcock on
cellaria. On the mouth-plates the furrow spines become
more prism-shaped.
Actinal interradial areas very large, the intermediate plates
extending to about the fourteenth infero-marginal ; the plates,
which number about 250 in each area, are arranged in about
nine chevrons, and decrease conspicuously in size from
actinostome to margin and from ambulacrum to mid-inter-
brachium. .
Madreporiform plate pentagonal, with fine radial striations,
placed not quite a third of the way from the centre to the
margin of the disk.
Tube-feet with a broad sucker.
Colour in alcohol ivory-white, the papular areas combining
to form a beautifully symmetrical, dark-coloured, five-petal
flower.
Andaman Sea, 271 fathoms.
NyYMPHASTER, Sladen.
27. Nymphaster florifer, sp. n.
Rays 5. R=3r. R=32 millim. in the
type specimen.
Disk pentagonal, large, very thin and flat; rays of mode-
rate length, very narrow and flat, quadrangular in section ;
interbrachial arcs wide.
Abactinal surface of the disk displaying in the centre and
in the radial areas a large and particularly beautiful and con-
spicuous rosette of hexagonal paxilliform plates, each inflated
petal-shaped segment of the rosette consisting of seven
longitudinal rows of plates, of which those in the middle row
—coinciding with the mid-radial axis—are of predominant
size; the interradial (interpetaloid) areas are small and
triangular and are clothed with small irregularly pentagonal
or subcircular plates.
The abactinal surface of the rays, between the plates of the
supero-marginal series, is occupied chiefly by a continuation
of the large plates, now become rather irregular in shape, of
the middle row of the petals, with on each side a discon-
tinuous row of platelets.
The hexagonal plates of the rosette are true paxille and
consist of a raised tabulum bearing in the centre from one to
six (usually two or three) and round the margin from twelve
to sixteen flat-topped bacillary granules; the irregular plates
of the interradial (interpetaloid) areas and of the rays are
merely covered with minute granules.
Indian Deep-sea Asteroidea. 95
The marginal plates are large and rectangular and are
uniformly covered with small granules, without any other
armature; many of them bear an entrenched pedicellaria,
found only on denudation ; they number about twenty-two in
each series, which correspond plate to plate; the supero-
marginals do not meet across the ray even at its very tip.
The adambulacral plates have a pectinate furrow-series of
seven to eight long fine spinelets of nearly equal size, the
proximal (adoral) one of the series alone being diminutive,
and actinally a row of from three to five papillary spinelets,
and outside these a row of four or five granules. Mouth-
plates small, the conjoint pair nearly circular in outline, the
armature hardly differing from the adambulacral type except
that the granules are more numerous.
Actinal interradial areas large, each area carrying about
sixty irregularly quadrangular plates arranged in chevron
series; all of these plates are granular like the marginal
plates, and many of those nearest the ambulacrum have an
entrenched pedicellaria.
Madreporiform plate small but conspicuous, placed very
much nearer to the centre than to the margin of the disk.
The papule are found only in the rosette of the abactinal
surface, where they stand with great regularity at the angles
of the hexagonal plates.
Andaman Sea, 130 to 250 fathoms.
This species much resembles Mymphaster bipunctus,
Sladen.
In young specimens R=about 2:2 r, the interbrachial ares
are not so wide as they are in the adult, the apical plates are
conspicuously large, and the marginal plates number eight or
nine in each series.
28. Nymphaster protentus, Sladen.
Nymphaster protentus, Sladen, ‘ Challenger’ Asteroidea, p. 303, pl. 1.
figs, 3 and 4, pl. hii. figs. 9 and 10.
Andaman Sea, 220 to 250 fathoms.
29. Nymphaster basilicus, Sladen.
Nymphaster basilicus, Sladen, ‘Challenger’ Asteroidea, p. 308, pl. lvil.
figs. 8 and 9.
Two fine specimens, one from the Laccadive Sea,
1370 fathoms, coral-mud, the other from the Gulf of Manaar,
597 fathoms, green mud,
This species, as Mr. Sladen observes, appears to be very
near Dorigona ternalis, K. Perrier.
96 | Mr. A. Alcock on
30. Nymphaster nora, sp. n.
Rays 5. R=63r. R=120 millim. in the
type specimen.
Disk small, flat, pentagonal; rays very long and slender
and tapering, quadrangular in section, but at the tip eylin-
drical ; interbrachial ares wide.
Abactinal area of the rays entirely occupied by the supero-
marginal plates, which from the fifth onwards meet across
the :ay ; abactinal area of the disk covered with hexagonal
paxilliform plates, which consist of a slightly raised tabulum
closely covered with angular many-facetted granules, of which
six to nine form a central group and twelve to sixteen a
marginal ring ; of these plates the basal interradials are the
largest, but those in the radial areas, especially those in the
mid-radial line, are also very Jarge; papule are found in the
radial areas only, emerging at the angles of the plates.
The marginal plates number about 40 in each series, they
are quadrangular and massive, and are closely covered with
angular granules without any other armature ; each supero-
marginal articulates with two infero-marginals unequally.
The supero-marginals of opposite sides, from the fifth
onward, meet across the ray, plate to plate, in a straight
suture; each plate presents a broad abactinal plane and a
narrow lateral plane, and at the junction of these two planes
an inflated angle; the cap-like terminal plate bears five spines
in a half hoop.
The infero-marginals are smaller than the supero-marginals ;
their lateral surface, except in the interbrachium, is twice the
breadth of the actinal surface, and the angle at which the
two surfaces meet is so inflated that the rays, whether viewed
actinally or abactinally, seem to have beaded or festooned
borders.
The large adambulacral plates completely close the furrow,
the distantly isolated pairs of tube-feet emerging from semi-
circular gaps which, by the close apposition of the plates of
opposite sides, become isolated circular holes. Each adambu-
lacral plate is, in fact, rudely L-shaped, the broad horizontal
limbs of the L’s meeting across the furrow, and the much
restricted concavities of the L’s forming the gaps which, by
the apposition of the plates, are converted into closed
chambers; actinally the plates are closely covered with
granules, while on the furrow-edge each plate has about
twelve long compressed spinelets arranged (except in the case
of a few plates close to the actinostome) in two divergent
Indian Deep-sea Asteroidea. 97
series so as to form a circular palisade round each isolated
pair of tube-feet.
Actinal interradial areas large, the irregularly quadrangular
and closely granular intermediate plates being arranged in
concentric chevrons and numbering about fifty in each area.
Madreporiform plate very small, with coarse radiating
striations ; placed close to the centre, encircled by three much
larger plates, the adcentral of which—the largest of all—is a
basal interradial.
Andaman Sea, 490 fathoms.
PARAGONASTER, Sladen.
31. Paragonaster tenuiradiis, sp. n.
Rays 5. R=4:57. R=40 millim. in the
type specimen.
Disk small, pentagonal ; rays long, extremely slender, oval
in section, rigid.
The abactinal area of the small disk bears large crowded
paxilliform plates arrauged in regular chevrons and elosel
covered with angular flat-topped granules, the mid-radial
plates being the largest and the mid-interradial plates the
smallest of all; the narrow abactinal surface of the rays bears
a single series of narrow granular plates intervening between
the supero-marginal plates of opposite sides.
The marginal plates, which are large and closely covered
with angular granules, number over twenty-five in each series ;
the supero-marginals, which are a good deal larger than the
infero-marginals, are almost entirely lateral on the disk, but
Jargely abactinal on the rays, where those of opposite sides
are separated only by a narrow series of abactinal plates ; the
infero-marginals alternate (more markedly in the distal half
of the ray, less markedly in the interbrachia) with the supero-
marginals, and those in the interbrachia bear a median vertical
series of two to four small spines or spinelets.
The adambulacral plates bear a semicircular marginal
series of nine or ten radiating spinelets encircling a few
distant actinally-placed spinelets, of which one, standing
close to the middle of the marginal series, is more conspicuous
than the rest. Mouth-plates prominent actinally, with a close
furrow-series of ten or eleven spinelets and a single suture-
series of nine or ten spinelets.
Actinal interradial areas small, barely reaching to the third
infero-marginal plate; in each area there are about thirty
minute scale-like plates bearing distant papilliform granules.
98 Mr. A. Alcock on
Madreporiform plate completely hidden. Anus central,
Colour in the fresh state pale yellowish pink.
Bay of Bengal, 1748 fathoms, Globigertna-ooze and
pumice,
32. Paragonaster, sp.
Paragonaster, sp., Ann. & Mag. Nat. Hist., Dec. 1891, p, 436.
A specimen too much mutilated for description was taken
with the above ; it is characterized by having the papule in
distinctly circumscribed papularia, over which the paxille are
singularly large and prominent.
MEDIASTER, Stimpson.
33. Mediaster roseus, sp. n.
Mediaster, sp., Ann. & Mag. Nat. Hist., Jan. 1891, p. 13.
Rays 5. R=4r. R=42 millim., in the
type specimen,
Disk small, flat; rays flat, tapering, rigid.
Abactinal surface of the disk covered with paxilliform
plates which in the centre and interradii are small, irregular
in shape, and crowded together, but which in the radial areas
are large and regularly hexagonal and are arranged in
regular longitudinal parallel rows; the plates are surmounted
by large flat-topped hexagonal granules of unequal size, which
fit close together to form a mosaic for each separate plate ;
abactinal surface of the rays formed principally by the
supero-marginal plates, which, however, are separated
throughout the ray by at least three rows of small irregularly
hexagonal paxilliform plates.
Marginal plates massive, closely covered with hexagonal
flat-topped granules, and forming on both aspects a very
broad border to the disk and rays; they number thirty-five
in each series, and the two series correspond plate to plate;
the infero-marginals of the interbrachia have a median
vertical series of distant papilliform spinelets or pedicellaria.
The large adambulacral plates have each a radiating
furrow-series of seven or eight flattened prismatic spinelets,
slightly decreasing in size in each series from the ends to the
middle, and actinally three unequal longitudinal series of
close prismatic granules. Mouth-plates narrow and elongate,
each with a furrow-series and a suture-series of coarse papilli-
form spinelets, which increase in size from periphery to
centre.
Indian Deep-sea Asteroidea. 99
Actinal interradial areas small, semielliptical in shape,
reaching to the third infero-marginal plate; about thirty
plates in each area, of which those in the row nearest the
ambulacrum are much the largest; all the plates are closely
covered with prismatic granules, grouped, except in the
particularly large adoral pair of plates, in paxilla-form.
Anus central but indistinct. Madreporiform plate almost
completely hidden, rather nearer to the marginal plates than
to the centre.
Ambulacral furrow narrow,
Colour in the fresh state light pink.
Laccadive Sea, 740 fathoms, coral-mud.
‘This species appears to be closely related to Astrogontum
fallax, Perrier (Ann. Sci. Nat. tom. xix., 1885). Guided by
the work of Mr. Sladen and by the description and figures in
the ‘ Boston Journal of Natural History,’ vol. vi., 1857,
p- 5380, pl. xxi., I have placed it, though not without
misgiving, in Stimpson’s Pacific genus.
ANTUENOIDES, Perrier.
34. Anthenoides sarissa, sp. n.
Rays 5. R=2:°5r. R=46 millim. in the
type specimen.
Disk large, flexible, inflatable, especially in the interradial
areas ; rays moderately long, rigid, broad at base, but quickly
becoming narrow and then tapering.
The abactinal surface of the disk and of the rays, except
at the extreme tip, where the supero-marginal plates meet
across, is cased by flat plates which are covered with a finely
and distantly granulose membrane; these plates are of two
kinds and of two modes of disposition :—(1.) large stellate or
somewhat polygonal plates arranged in series radiating from
the centre of the disk, and (i1.) small or minute platelets inlaid
everywhere between the large plates. Papule emerge in the
seams between the plates everywhere except in a narrow band
in each interradius.
The marginal plates, which number about 20 in each
series, are large and finely and distantly granular. ‘The
supero-marginals form a very broad bevelled margin to the
disk in the interradii and a narrower margin to the rays,
except in the distal third of the latter, where they nearly meet
across the ray ; except for one, two, or three small inconstant
papilliform pedicellariw occurring on the extreme distal edge
100 Mr. A. Aleock on
of the plates of the apical half of the ray they are unarmed,
but the cap-like terminal plate has a fringe of five spinelets.
The infero-marginals, which correspond plate to plate with
the supero-marginals, have each a strong lateral bulge on
which stands a horizontal series of three (sometimes only
two) spines, but in the apical half of the rays there is only one
spine, placed on the extreme distal edge of the plate.
The small adambulacral plates have each a palmate furrow-
series of six or seven spinelets, and almost in serial continuity
with these, and to the central (adoral) side of them, a pedi-
cellaria with two long spoon-shaped valves; actinally each
plate has near its distal end an eminence on which stands a
stout spine longer than the plate itself and usually longer
than any of the infero-marginal spines, in addition to two or
three pustules; on the mouth-plates, which are extremely
prominent actinally, the spine and pedicellaria are usually
absent, and only the pustular granules present, but the central
(adoral) furrow-spines are much enlarged.
The actinal interradial areas are very large, reaching to the
sixth or seventh infero-marginal (thirteenth or fourteenth
adambulacral) plates; each area bears eighty to ninety large
platesarranged inconcentricchevrons; all the plates are covered
with a distantly granular membrane, and some or all of them
(but most constantly those of the series next the ambulacrum)
bear one or more papilliform pedicellariz ; the plates of the
second series from the ambulacrum are of superior size.
Anus central, distinct. Madreporiform plate large, with
fine radial striations ; placed rather nearer to the centre than
to the margin.
Ambulacral furrow wide; tube-feet with a terminal sucker.
Andaman Sea, 130 to 250 fathoms.
Family Asterinide.
PaumiPEs, Linck.
35. Palmipes pellucidus, sp. n.
Rays 5. R=1:57r. R=48 millim. in the
type specimen.
Form « curvilinear pentagon with bluntly rounded angles ;
flat, very thin and leaf-like, membranous, semitransparent.
Abactinal surface with regular longitudinal and oblique
rows of scale-like rhomboid or subhexagonal plates, which
gradually diminish in size from the centre to the margin; all
the plates are covered with distant erect spicules and bear
Indian Deep-sea Asteroidea, 101
each a central tuft of capillary spinelets, the tufts and their
constituent spinelets being enlarged in a mid-radial row.
The papulx are in four rows, two on each side of the mid-
radial line, the inner row on each side extending from the
centre of the disk to the tip of the ray, and the outer extending
continuously only about halfway along the ray, and thence
onward in a broken series. In young specimens there are
only two rows of papul, one on each side of the mid-radial
line.
The papular pores, like the papule themselves, are of two
kinds; all are encircled by a close palisade of spinelets, but
from three to six of the pores of the inner two rows are much
enlarged, to give exit each to a singularly long papula, and
these are encircled by spinelets of superior length; the small
pores give exit each to a small short papula.
The marginal plates, which lie in alternation with the
oblique rows of abactinal plates, are minute, and each is
edged with a tuft of spinelets and each has a very minute
scale-like platelet at its base.
The adambulacral plates have each a palmate furrow-series
of six spinelets and actinally an oblique or curved series of
three or four capillary spinelets; the mouth-plates have a
furrow-series of eight or nine spinelets which increase in size
adorally and a suture-series of five or six capillary spinelets.
The actinal plates are in regular longitudinal and oblique
rows, and diminish in size from the actinostome to the
margin; each plate carries a radiating marginal series of
capillary spinelets, four in a series near the actinostome, three
elsewhere.
Colour in life pellucid hyaline grey.
Andaman Sea, 112 fathoms, blue mud.
Family Linckiide.
Cuzraster, M. & T.
36. Chetaster, sp.
Jn our collection there is a small broken specimen of an
undoubted Chetaster which I do not at present venture to
describe. In appearance it corresponds with the figure of
Chetaster munitus, Mobius (‘ Neue Seesterne des Hamburger
und Kieler Museums,’ pl. i. figs. 1 and 2), and, so far as the
description (op. cit. p. 3) of that species goes (a description
which, however, is certainly incomplete), it corresponds with
it exactly. With Professor Mobius’s form I should have
identified our species, had not Chetaster munitus, Mobius,
102 Mr. A. Alcock on
been pronounced by Mr. Sladen to be merely a synonym of
Nectria ocellifera (Lamarck).
The single specimen is from the Andaman Sea, 238 to 290
fathoms.
Family Zoroasteride.
ZOROASTER, Wyville Thomson.
37. Zoroaster Alfredi, sp. n.
Rays 5. R=9r. R=about 190 millim. in the
type specimen.
Disk small, hemispherical, tumid above the tumid rays;
rays long, narrow, tapering, subcylindrical to cylindrical.
Abactinal surface of disk with large distant, subhexagonal,
or substellate primary radials and interradials surrounding a
large dorso-central plate, and with numerous small intervening
plates; all the plates are closely covered with simple or
grooved or bifid membrane-clad spinelets, and the large
hollows between neighbouring plates contain each a group ‘of
papulz and a group of pedicellaria, one of which is of con-
spicuous size, being about as big as a grape-stone.
The rays have a longitudinal mid-radial row of large
subhexagonal plates co- serial with the large primary plate of
the apical system, and on each side of it and parallel with it
six (in the interbrachia seven, at the end of the rays five)
rows of smaller plates, which also form transversely parallel
series, the lowest row articulating with the cramped adambu-
lacrals ; all these plates are closely covered with simple or
grooved membrane-clad spinelets and pedicellarix, and beara
centrally-placed spine, which is small, erect, and often obso-
lescent in the plates of the abactinal rows, but large and
acumbent in the four lower rows on each side, gradually
increasing in size from above downwards; the large hollow
intervals between neighbouring plates, which, in consequence
of the symmetry of the plates themselves, also fall into longi-
tudinally and transversely parallel series, contain groups of
pedicellaria, one of which in each group is conspicuously
enlarged, and (except between the lowermost two rows of
plates) from one to three papule.
The adambulacral plates extend vertically far upwards into
the furrow, and the whole system is so cramped that the two
middle tube-feet of each row are quite perceptibly atrophied
by pressure; every alternate plate has a strongly salient
intra-ambulacral ridge, upon which stands a row of three or
Indian Deep-sea Asteroidea. 103
four large spinelets, each with a large pedicellaria and often
also a bunch of small pedicellariz attached by ligament to its
base, and which, deep in the roof of the furrow, “bifurcates to
give origin to a pair of large bunches of pedicellarie, twelve
to twenty in each bunch; all the adambulacral plates are
covered with spinelets and small pedicellariz on their narrow
actinal edge.
Madreporiform plate large, with coarsish radial striations.
Colour in the fresh state “ deep salmon-colour throughout,
spines a little paler” (G. MZ. Giles).
Bay of Bengal, 13800 to 1380 fathoms, Globigerina-ooze.
This large and very beautiful species is readily distinguished
by the extraordinary development of its pedicellariw, espe-
cially by the two large ceespitose masses of these organs which
are borne on every alternate adambulacral plate between the
bases of the tube-feet, and by the large pedicellarize (much
more conspicuous than any of the spines except those on the
plates of the two most actinal rows) arranged in regular
longitudinal and transverse parallel series along the rays.
It was named by Messrs. Wood-Mason and Giles after
Commander Alfred Carpenter, R.N.
38. Zoroaster Barathri, sp. n.
Rays 5. R=16r. R=about 180 millim. in the
type specimen.
Disk extremely small, not differentiated from the bases of
the rays, tumid; rays extremely long and finely tapering,
semicy lindrical.
Abactinal surface of disk with large, close-set, subhex: agonal
primary radials and interradials surrounding an apical mass
formed of a dorso-central and radial underbasal plates, all of
equal size; all the plates are very closely covered with
‘apillary spines, and the small depressions which intervene
between the plates are perforated for one or two papule and
bear from one to three pedicellariz, of which one is sometimes
slightly enlarged.
The rays have a longitudinal mid-radial row of large tumid
hexagonal plates coserial with the large primary radial, and
on each side of it six (in the interbrachia seven, at the end of
the rays five) parallel rows of smaller plates, very close-set,
the lowest row abutting on the adambulacrals; these plates
also fall into close-set transversely parallel series, and all are
densely covered with capillary spines that become gradually
longer and more slender in each successive row from the
104 Mr. A. Alcock on
abactinal to the actinal surface ; the plates of the lowest (most
actinad) two or three rows bear each one or two centrally-
placed spines which are of slightly superior size. The small
intervals between the angles of the closely apposed plates give
exit, except between the two lowermost (most actinad) rows,
each to a papula, and (on the plates) between every two
papule is a pedicellaria,
‘The adambulacral plates are short and do not extend far
up into the furrow; each plate bears on its actinal edge a
transverse row of two or three large sabre-shaped spinelets, of
which the innermost often carries a large pedicellaria, and
every alternate plate has also a prominent intra-ambulacral
ridge, on which stand two more spines, of which the outer
carries a pedicellaria almost as large and conspicuous as a
tube-foot and the inner a small cluster of small pedicellariz.
The mouth-plates bear each two series of long needle-like
spines.
Tube-feet quadriserial.
Madreporiform plate very small and much hidden, with a
deep, coarse, vermicular sculpture like a peach-stone.
Bay of Bengal, 1520 fathoms, Globigerina-ooze.
This fine species is recognized (i.) by its extremely small
disk not differentiated from the bases of the rays, and (ii.) by
the close capillary spinulation of the plates, which gives the
animal a uniformly hairy appearance when dried. It appears
to be more nearly related to Zoroaster longicauda, Perrier,
than to any other described species.
39. Zoroaster planus, sp. n.
Rays 5. R=167. R=210 millim. in the
type specimen.
Disk extremely small, depressed ; rays extremely long and
finely tapering, semicylindrical, depressed.
Abactinal surface of disk with slightly enlarged, close-set,
subhexagonal, primary radials and interradials surrounding a
mass composed of a dorso-central and radial under-basal plates
all of equal size; all the plates are rather closely covered
with capillary spinelets, and the dorso-central and the radial
plates carry in addition a stout conical fluted spine; the
narrow intervals between neighbouring’ plates each with one
apula and often with a single large pedicellaria.
The rays have a longitudinal mid-radial row of large tumid
plates, each of which bears, besides the capillary spinelets, a
central stout, conical, fluted spine much like a pedicellaria ;
Indian Deep-sea Asterotdea. 105
on each side of this row are six (seven in the interbrachia, five
at the end of the rays) very close parallel rows of smaller
plates, the lowest row articulating with the adambulacrals ;
these plates, which also fall into transversely parallel series,
are rather distantly covered with capillary spinelets, the central
one of which in each plate (except in the row immediately
adjoining the large mid-radial series) becomes a long slender
spine that gradually increases in size in each successive row
from the abactinal to the actinal surface. ‘The narrow
intervals between the angles of neighbouring plates give
passage (except between the two lowermost rows of plates)
each to one papula, and (on the plates) between the papule
is a rather large pedicellaria.
The adambulacral plates are short and do not extend far
upwards within the furrow; each plate bears on its actinal
edge two transversely placed spinelets and occasionally a
pedicellaria, and every alternate plate has a prominent intra-
ambulacral ridge on which is borne a row of three spinelets,
the innermost being furnished with a cluster of eight small
pedicellariz, and the one next the innermost with a single
large pedicellaria. The mouth-plates are armed with large
needle-like spines.
Tube-feet quadriserial.
Madreporiform plate large, tumid, and conspicuous, with a
coarse peach-stone sculpture.
Colour in life salmon-red.
Laccadive Sea, 1200 fathoms, coral and Globigerina-ooze.
This species is very like Z. Barathri, from which it is easily
distinguished, (i.) by the flattened disk very definitely
delimited from the bases of the rays; (il.) by the large salient
madreporiform plate ; and (iil.) by the large spinelet borne
centrally on every plate except the basal interradials and the
plates of the row immediately adjoining the mid-radial row
on each side.
40. Zoroaster angulaius, sp. n.
Rays 5. R=12to137r. R=148 millim. in the
type specimen.
Disk small, high, flat-topped; rays long and tapering,
high, compressed, and strongly keeled abactinally.
Abactinal surface of disk with close-set, large, tumid,
stellate primary radials and basals surrounding a very large
tumid dorso-central plate and a ring of small depressed
radial under-basals ; all the plates bearing distant capillary
Ann. & Mag. N. Hist. Ser. 6. Vol. xi. 8
106 Mr. A. Alcock on
spinelets and small marginal pedicellaria, and the crevices
between the plates giving exit to scattered distant papule.
The rays with a longitudinal mid-radial row of very large
tumid vertebra-shaped plates co-serial with the primary radial,
each bearing a central globular or squatly conical spine; and
on each side of this mid-radial row seven (in the distal half
of the ray six) parallel rows of much smaller tumid octagonal
plates, the lowermost of which articulates with the adambu-
lacrals ; these plates, which also fall into transversely parallel
series, all bear distant capillary spinelets and small marginal
pedicellaria, and all but those in the row immediately
adjoining the mid-radial row have a central long needle-like
spine, which is largest in the plates of the row adjoining the
adambulacrals. ‘The pin-hole intervals between the angles of
neighbouring plates (except between the two lowermost rows
of plates) are just large enough to give exit to a single small
papula.
Adambulacral plates small, almost cylindrical ; each plate
bears on its actinal surface either three or four long sharp
spinelets in two transverse series, one of the spinelets which
adjoins the furrow often furnished with one or two small
pedicellariz ; every alternate plate has a prominent intra-
ambulacral ridge, on which stands a row of two or three long
spinelets, of which the innermost bears a cluster of from six
to ten small pedicellariz and the outermost a single large
pedicellaria. The mouth-plates are armed with needle-like
spines similar to those of the other adambulacral plates.
Tube-feet quadriserial, but the two middle feet of each row
reduced in size.
Madreporiform plate very small, not half the size of the
basal plate, outside of which it lies almost hidden by over-
hanging spinelets.
Gult of Manaar, 597 fathoms, green mud; Laccadive Sea,
705 fathoms, coarse coral shingle.
This species is distinguished, (i.) by the high but flat-
topped disk, with its tumid stellate primary plates sparsely
covered with spinelets; (ii.) by the strongly carinated rays,
pentagonal in section, with their median row of large
vertebra-shaped plates; and (iii.) by the broad cylindrical
adambulacral plates with their two transverse series of actinal
spinelets.
In a young specimen the disk, with its relatively enormous
tumid apical plates, recalls the appearance of the Ophiuroid
Ophiomastus ; the relatively short rays (R=8 7) end in a
large inflated, helmet-like, two-horned plate, and the tube-
feet are biserial.
Indian Deep-sea Asterowdea. 107
41. Zoroaster carinatus, sp. n.
Rays 5. R=about 107, R=about 90 millim. in
the type specimen.
Disk small, hemispherical, higher than the rays; rays
semicylindrical, strongly carinated, tapering in the distal half,
but not much diminishing in the basal half.
Abactinal surface of disk with close-set, large, tumid,
stellate, primary radials and basals surrounding a very promi-
nent dorso-central plate and a ring of small depressed under-
basals ; all the plates are densely covered with thorn-like
spinelets, which are enlarged in the middle of the plate;
isolated papule surround the plates, and a few small scattered
pedicellariz: occur.
Rays with a median longitudinal row of very large and
tumid stellate plates, closely covered with spinelets, of which
those in a tuft in the middle of each plate are enlarged. On
each side of this median row is a depressed discontinuous
series of very small irregular platelets, below which are five
parallel rows of close-set tumid octagonal plates ; all these are
closely covered with thorn-like spinelets, and those of the
four lower rows have also a long slender median spine. The
minute apertures between the angles of neighbouring plates
(except between those of the two lowermost rows) give exit
to a single papula, and close to every papula is a small
pedicellaria,
Adambulacral plates small, cylindrical, each with a trans-
verse series of two (rarely three) long slender spinelets ; every
alternate plate has a prominent intra-ambulacral ridge bearing
a row of two or (usually) three similar but larger spinelets,
one of which, within the furrow, has occasionally a small
pedicellaria, or sometimes two. Mouth-plates with long
spinelets like those of the ordinary adambulacrals,
Tube-feet quadriserial.
Madreporitorm plate very small, hardly one third the size
of the basal plate outside of which it lies.
Colour in life brick-red.
Andaman Sea, 130 to 250 fathoms.
This species, which is one of the commonest starfishes met
with in the Andaman Sea between 200 and 300 fathoms, is
characterized, (i.) by the dense thorny armature of the very
tumid apical plates, each of which has a central tuft of
enlarged spinelets; (ii.) by the strong carination of the rays,
which is due partly to the tufts of enlarged spinelets of the
tumid mid-radial 1ow of plates and partly to the sinking of
gt
108 Mr. A. Alcock on
the small plates on each side of this prominent mid-radial
row; and (iii.) by the paucity of esinosilteres even the spines
of the adambulacral plates being oftener without these organs
than with them.
In the young stage (R=6°5 r) the apical and abactinal
mid-radial plates are extremely prominent, the tip of the rays
is capped by a large, inflated, spiny terminal plate, and the
tube-feet are but indistinctly quadriserial.
42. Zoroaster Gilesit, sp. n.
Rays 5. R=about 87. R=70 millim.
Disk comparatively large, low, flat or very gently convex,
very distinctly delimited from the rays (above which it is not
much elevated) by a complete ring of massive semiglobular
plates; abactinally it is encased in concentric series of these
great semiglobular plates (with or without small sunken
plates intervening) in the following order :—a dorso-central,
five basal interradials, five primary radials (the largest of all),
and twenty-five marginal plates, of which one succeeds each
primary radial and four close in each interradial area. All
the plates are covered with capillary spinelets with a few
coarser spinelets interspersed, and the intervals between the
plates are occupied by papule and rather numerous large
pedicellariz.
The rays are rigid, broadish, and flat, with a gentle abac-
tinal convexity, and are encased in thirteen longitudinal
parallel series of small and very close-set tumid plates, which
also stand in obliquely transverse parallel series. These
plates have the following arrangement :—(i.) a mid-radial
row of suboctagonal slightly imbricating plates lying in a
furrow between two rows of distant papule (which two rows
of papule are the only papule present on the rays), flanked
on each side by (ii.) six rows of very close-set bead-like
plates, of which the upper (abactinad) three rows are much
the larger, the lower (actinad) three rows consisting of minute
plates so crowded together as to appear at first sight like a
single row. All these plates are covered with coarse spinelets,
which are larger and denser in the three lower (actinad) rows,
and those of the mid-radial and three upper (abactinad) rows
on each side have also a small coarse spine centrally, and on
each suture margin a largish pedicellaria, while those of the
three lower rows have each a long lanceolate spine.
Adambulacral plates small and short, but extending high
up into the furrow; each plate has two transversely placed
Indian Deep-sea Asterotdea. 109
spinelets, and every alternate plate has a prominent intra-
ambulacral ridge bearing a row of three spines, the inner-
most of which carries a single large pedicellaria. Mouth-
plates armed with spines similar to those of the ordinary
adambulacrals.
Tube-feet quadriserial.
Madreporiform plate small and much hidden by neigh-
bouring spinelets, with obsolescent sculpture.
Colour in life “ red ochre” (G. M. Giles).
Andaman Sea, 490 to 500 fathoms.
43. Zoroaster squameus, sp. N.
Rays 5. R=about 117. R=135 millim. in the
type specimen.
Disk pentagonal, flat-topped, distinctly delimited from the
rays by a circumferential series of large tumid plates ; its
abactinal surface bears concentric series of massive tumid
stellate plates—a dorso-central, five basal interradials, five
primary radials (the largest of all), and twenty-five marginals
arranged as in Z. Gulesi/; all these plates bear distant
deciduous granules, and the intervals between them display
papulz and numerous small pedicellariz.
Rays long, tapering, compressed, strongly convex abac-
tinally, with thirteen to fifteen parallel longitudinal rows of
plates (which also fall into transversely parallel series), each
row, with the exception of the mid-radial row, strongly imbri-
cate over the row below. The following is their arrange-
ment :—(i.) a mid-radial row of non-imbricate plates, hardly
enlarged, and flush with the surface of the ray; flanked on
each side by (ii.) six or seven rows of scale-like plates which
overlap one another from above like tiles on a roof, those of
the upper four rows being the more massive and conspicuous,
those of the lower two or three rows being thin and crushed
up beneath one another. The mid-radial row of plates
appears to be quite smooth and unarmed; the plates of the
three or four upper (abactinad) rows on each side are quite
smooth, but bear each a more or less deciduous procumbent
median spine, and in the distal half of the ray some mar-
ginal pedicellariz; the plates of the three lower (actinad)
rows on each side are very closely covered with acumbent
seale-like spinelets, with a long procumbent, needle-like,
deciduous median spine. Papule occur, accompanied each
by a pedicellaria, in an incompletely double row on each side
of the mid-radial row of plates ; and between the second and
110 Mr. A. Aleock on
third row of plates throughout the ray, and between the third
and fourth row of plates in the basal fourth of the ray, there
are minute perforations (seen only in the dried specimen)
which appear to be too small for the passage of papule.
Adambulacral plates very small, but extending high up
into the furrow; each plate has two short transverse series of
spinelets, and every alternate plate has a prominent intra-
ambulacral ridge bearing three stout spinelets, of which the
innermost is furnished with a large pedicellaria emerging
from a cluster of small pedicellarize. Mouth-plates with long
needle-like spines and large clusters of pedicellariz.
Tube-feet quadriserial.
Madreporiform plate very small, with coarse peach-stone
sculpturing.
Colour in life orange-pink.
Laccadive Sea, 1043 fathoms, green mud.
44, Zoroaster zea, sp. n.
Rays 5. R=about 127, R=144 millim. in
the type specimen.
Disk semicircular, flat-topped, well raised above the rays,
from which it is delimited by a circumferential series of
massive oval or substellate plates, arranged exactly as in
Z. Gilesti and Z. squameus, with small plates intervening.
All the plates are quite smooth and membrane-clad, but the
small intervening plates bear each a small coarse spine; the
intervals between the plates show distant papule and pedi-
cellaria, the last often in pairs.
Rays long, rigid, subcylindrical, tapering, with thirteen
longitudinal parallel series of Indian-corn-like or bead-like
membrane-clad plates, which also fall into transversely
parallel series. A. single ray, viewed abactinally, has much
the appearance of a seed-spike of maize. The plates are
disposed as follows:—(i.) a mid-radial row of slightly
enlarged plates, flanked on each side by a deep furrow, in
which lies (ii.) a discontinuous row of minute platelets con-
cealed by membrane, and only revealed either by a small
pedicellaria or by a coarse spikelet which they sometimes
bear ; and outside these (iii.) six rows of plates, decreasing in
size and inclining to imbricate actinally, of which the three
abactinad rows are, like the mid-radial row, quite naked and
unarmed, while the three actinad rows are thickly covered
with membrane-clad squamous spinelets and bear a median
spine and sometimes a marginal pedicellaria. Long papule
Indian Deep-sea Asterotdeua. 111
are found in a close double row in the furrow on each side of
the mid-radial series of plates; minute perforations occur
between some of the other rows of plates, but they are only
visible with a lens, and seem to be far too small to give exit
to papule.
The adambulacral plates are covered with flat foliaceous
spinelets ; every alternate plate has a prominent intra-ambu-
lacral ridge bearing a row of three sabre-shaped spines, and
deep within the furrow a slender spinelet furnished with
several pedicellaria, one of which is large. Mouth-plates
with stout spinelets and clusters of pedicellariz.
Tube-feet quadriserial.
Madreporiform plate small—half the size of a basal inter-
radial plate—but tumid and conspicuous, with coarse radial
striations.
Gulf of Manaar, 597 fathoms, green mud; Laccadive Sea,
1200 fathoms, coral and G'lobigerina-ooze.
In a youngish specimen (R=65 millim.) R=about 8 7;
the mid-radial plates are relatively larger; the rays terminate
in a rather inflated bilobed plate fringed with spines; and
the papule, which are in a double row on each side of the
mid-radial series of plates, extend only about halfway along
the rays.
Zoroaster Gilesit, Z. squameus, and Z. zea form a very
well-defined group within the genus Zoroaster, and, along
with Z. Sigsbeet, Perrier (Nouv. Archiv. du Mus. sér. i.
vol. vi., 1883, p. 195, pl. ui. fig. 2), may perhaps be con-
veniently recognized asasubgeuus. ‘The following characters
are common to the first three species, and, as far as can be
judged from the rather brief though very succinct description
of Professor Perrier and from the phototype figure, appear
to be shared by the last:—(i.) the disk is very distinctly
demarcated from the rays abactinally by massive plates
which form a circumferential series—one plate at the base of
each ray and four in each interradius; (ii.) the rays are
particularly ngid, with the longitudinal mid-radial row of
plates neither conspicuously large nor very prominent, and
the papule of the rays are more or less restricted to two single
or double rows, one on each side of the mid-radial row of
plates {in 7, Sigsbeei the papular orifices seem to have been
only doubtfully recognized (tom. et loc. cit.)]; (iii.) the three
actinad rows of plates are much crushed together (?7. Sigsbeet),
and are always more densely covered with spinelets and more
conspicuously armed than the plates of the abactinad rows,
112 Mr. A. Alcock on
which last are often quite smooth and unarmed; (iv.) the
anal aperture is very distinct.
This subgenus shows an interesting series of gradations
between Zoroaster, Wyville Thomson, and Cnemidaster,
Sladen. At the true Zoroaster extreme is Zoroaster Gilesii,
with all the plates spinate and armed, with numerous pedi-
cellarize, and with a broad ambulacrum and four rows of tube-
feet. Near the Cnemidaster extreme comes, as Mr. Sladen
has already pointed out, Z. Sigsbeet, with the abactinad plates
smooth and unarmed and only the actinad plates spinate, with
few pedicellariz, and with crowded tube-feet which only in
the basal part of the ray are quadriserial. While between
the two come (i.) Z. squameus, with the abactinad plates
granular or quite smooth and their armature reduced to one
deciduous spinelet, and (ii.) Z. zea, in which the abactinad
plates are quite smooth and unarmed and the pedicellarize
reduced in size and number.
Family Pterasteride.
MARSIPASTER, Sladen.
45. Marsipaster hirsutus, Sladen.
Marsipaster hirsutus, Sladen, ‘Challenger’ Asteroidea, p. 487,
pl. Ixxviil. figs. 3 end 4, pl. Ixxix. figs. 4 and 6; and in Wood-Mason
and Alcock, “ Indian Deep-sea Dredging,” Ann. & Mag. Nat. Hist.,
Dec. 1891, p. 437.
Bay of Bengal, 1997 fathoms, Globigerina-ooze and
pumice.
HYMENASTER, Wyville Thomson.
46. Hymenaster nobilis, Wy ville Thomson.
Hymenaster nobilis, Wyville Thomson, Journ. Linn. Soe., Zool.
vol. xiii. p. 73, fig. 11; Sladen, ‘Challenger’ Asteroidea, p. 495,
pl. lxxxvii. figs. 1-3; and Wood-Mason and Alcock, “ Indian Deep-
sea Dredging,” Ann. & Mag. Nat. Hist., Dec. 1891, p. 438.
Bay of Bengal, 1748 fathoms, Globigerina-ooze and
pumice.
Family Echinasteride.
CRIBRELLA, Agassiz, Forbes.
47. Cribrella prestans, Sladen.
Cribrella prestans, Sladen, ‘Challenger’ Asteroidea, p. 545, pl. xevi.
fig. 7, pl. xeviii. figs. 7 and 8.
Four specimens from the Andaman Sea, 240 to 480
Indian Deep-sea Asteroidea. 113
fathoms, on bottoms of green mud or of coral and Globige-
rina-ooze, only differing from the described type, which was
dredged by the ‘Challenger’ in the Indian Ocean off the
Crozet Islands, in having the disk relatively smaller. In
one specimen the rays are most remarkably inflated in their
basal fourth.
Dicryaster, Alcock and Wood-Mason.
Rays 5.
Disk large, quite flat actinally, but slightly inflated abac-
tinally, as are the broad rays; both disk and rays are invested
throughout by a thick, smooth, coriaceous membrane, beneath
which is a reticulum of plates.
Abactinal surface with coarse irregular plates, some or all
of which bear coarse spinelets, and forming an irregular wide-
meshed network, the meshes being occupied by large groups
of papule.
Supero-marginal plates absent or inconspicuous; infero-
marginals conspicuous and bearing one or more stout spines ;
the ao of papule extend downwards as far as the infero-
marginal plates.
Actinal interradial areas large, the smooth intermediate
plates extending to the tip of the rays.
Adambulacral armature in the form of a double furrow-
palisade.
Tube-feet in a double row, their tip ending in a sucker.
Madreporiform plate small, single. Anus subcentral. No
pedicellaria.
This genus appears to be nearest related to Plectaster ; it
is instituted for the reception of two Andaman forms, Dicty-
aster xenophilus, from deep water, and Dictyaster Wood-
Masont, a littoral form recently discovered in Prof. Wood-
Mason’s rich Andaman collection.
48. Dictyaster xenophilus, Alcock and Wood-Mason.
(Pl. V. figs. 8, 9.)
Dictyaster xenophilus, Ann. & Mag. Nat. Hist., Dec. 1891, p. 438.
Rays 5. R=2°5r. '
Disk large and, like the short broad rays, flat actinally,
and often a little inflated abactinally ; both disk and rays are
everywhere invested in a thick leathery membrane which
covers all the plates and their spines—abactinal, actinal, and
adambulacral.
114 Mr. A. Aleock on
The narrow salient abactinal plates form a very wide-
meshed network, the sunken meshes of which are occupied by
large grouped masses of papule ; all the plates carry coarse
spines, either solitary or in rows of two or three.
The supero-marginal plates hardly differ from the ordinary
abactinal plates in form and armature, but they constitute a
fairly well-defined border to the rays.
The infero-marginals are separated from one another by an
interval equal in length to the plates themselves; they are
distinct lamine, with their surface horizontal and with a
sharp edge that projects laterally and bears a horizontal comb
of three to five (usually three) coarse spines. The space
between the two series of plates is occupied by papule in
large isolated groups.
Adambulacral plates small, each with two, but near the
actinostome often three, narrow ligulate spines along and
almost within the furrow, and on the margin of the furrow
with one much larger broadly ligulate spine, the truncated
end of which is usually bilobed or trilobed; so that the
narrow ambulacral groove is bounded on each side by a
double palisade of ligulate spines, those of the outer series
being about half as numerous but about twice as big as those
of the inner series. Mouth-plates with a furrow-series of
from four to six ligulate spines, increasing in size from the
periphery to the centre, and actinally with two of the broadly
ligulate spines, placed transversely.
Actinal interradial areas large ; the plates, which are quite
smooth and which are separated from one another by
intervals equal in breadth to the plates themselves, extend
nearly to the tip of the ray.
Madreporiform plate situated almost in an interbrachial are,
very finely and closely striated, Anus subceutral, surrounded
by blunt spinelets.
Tube-feet in a double row, ending in a sucker.
Colour in the fresh state chestnut-brown.
A symbiotic Chetopod is often found on the actinal inter-
radial areas, on which also it often lays its eggs.
Andaman Sea, 170 to 290 fathoms.
Family Pedicellasteride.
PEDICELLASTER, Sars.
49. Pedicellaster atratus, sp. n.
Rays 5. R=117. R=about 100 millim. in
the type specimen.
Disk extremely small, circular (button-like), abactinally
5
Indian Deep-sea Asteroidea. 11
delimited from the rays, with which it is flush, by a groove
that passes across the base of each ray. Rays long, semi-
cylindrical, constricted laterally, as well as somewhat
depressed, near their junction with the disk.
Abactinal surface closely covered with partly imbricate
cruciform plates which have no particular arrangement on
the disk, but which on the rays stand in beautifully regular
longitudinally and transversely parallel rows forming a
quadrangular network of great symmetry. ‘There are from
twelve to fourteen longitudinal rows of plates on each ray
between the adambulacrals. All the plates are embedded in
a continuous membrane, and each plate carries at the decus-
sation of its cross-pieces a stout erect spinelet. The mem-
brane that closes over the quadrangular meshes between the
plates is thickly studded with large pedicellaria and is
perforated by papulee—one to three in each mesh.
Marginal piates quite undifferentiated.
There are no actinal plates, the actinal surface of the
rays being almost entirely taken up by the broad ambulacral
furrow.
The adambulacral plates are very small; each plate carries
two long cylindrical truncated spines placed transversely, the
outer spine being about twice the length and four times the
thickness of the inner one, and usually having external to its
base a large pedicellaria.
Although the ambulacral furrow is so broad the tube-feet
are biserial.
Madreporiform plate small, radially striated, placed near
the margin of the disk.
Colour uniform jet-black.
Andaman Sea, 240 to 220 fathoms, coral-sand and forami-
niferal ooze; Bay of Bengal, 290 fathoms.
Family Asteriide.
Astertas, Linn.
Subgen, SroLastreRtiAs, Sladen.
50. Asterias mazophorus, Alcock and Wood-Mason.
Asterias mazophorus, Ann. & Mag. Nat. Hist., Dec. 1891, p. 436.
Rays 5. R=11r. R=115 millim. in the
type specimen.
Disk extremely small, circular (button-like), delimited
abactinally from the rays, beneath the level of whicia it is
116 Mr. A. Alcock on
sunk, by a deep groove passing across the base of every ray.
Rays long, semicylindrical, much constricted laterally at the
junction with the disk, and very deciduous.
Abactinal surface of the disk with a reticulum of plates
embedded in tough membrane ; each plate bears a large spine
with its base sheathed in a globular fleshy mass closely
covered with minute pedicellariz ; the meshes between the
plates are perforated by papule in groups; numerous large
pedicellarie fill the interbrachial ares and extend nearly to
the actinostome.
Abactinal surface of the rays with five beautifully regular
parallel rows of partly imbricating plates that may best be
described as compositely cruciform—that it to say, each com-
posite plate is made up of two cross series of tightly imbri-
cating platelets. The outermost row of plates on each side
forms a distinct supero-marginal series. All the plates are
embedded in a tough membrane, and each plate of the mid-
radial and supero-marginal rows, and in large specimens each
plate of the other two rows also, bears at the decussation of
its composite cross-pieces a large spine with its base en-
sheathed in a globular fleshy mass closely covered with
pedicellarie ; other large pedicellariz lie scattered over the
plates. The quadrangular meshes between the plates are
filled with papule in oval plots of from five to nine.
Marginal plates distinct; the supero-marginals, which
otherwise resemble the abactinal plates, are distinguished by
their mainly vertical direction, their long outer limbs forming
a high vertical palisade along the sides of the ray; the broad
intervals of the palisade are filled by papule in groups. The
infero-marginals are simply oblong plates of small size lying
at right angles to the long vertical limb of the supero-
marginals, and having a considerable lateral horizontal
bulge; each carries two strong horizontally-directed spines,
one above and behind the other, the upper being the fates
and exactly resembling the abactinal spines in form and in
its ensheathing fleshy pedicellaria-covered base, the lower
being quite simple.
Adambulacral plates very small, but extending up into the
furrow ; each carries a pair of spines placed transversely, and
almost every alternate plate has inside the furrow a large
pedicellaria.
The adambulacral plates are separated from the infero-
marginals by a single row of small narrow plates distant
almost their whole length apart, the long intervals between
the plates being filled each with a large papula encircled by
Indian Deep-sea Asteroidea. 117
pedicellarix. Mouth-plates ambulacral, each with two large
spines.
Madreporiform body placed in the middle of a large basal
(interradial) plate.
Anal aperture indistinct.
Tube-feet quadriserial, ending in a sucker.
Colour in the fresh state deep orange-yellow, with large
chestnut-brown blotches.
Andaman Sea, 120 to 250 fathoms.
This is a very remarkable and interesting form, and has
repeatedly led me on to difficult and uncertain ground. Its
affinities, unlike those of some of its fellows of the Stolasterian
alliance, appear to be more Stichasteroid and Zoroasteroid
than Brisingoid.
Family Brisingide.
BristnGA, Asbjornsen.
51. Brisinga insularum, Alcock and Wood-Mason.
Brisinga insularum, Ann, & Mag. Nat. Hist., Dec. 1891, p. 439.
Allied to B. coronata, Sars.
Rays 13.
Disk comparatively large (its diameter being 44 millim. in
the type specimen), with vertical edge; rays not extremely
deciduous (six of them still being firmly attached to the disk
in the type specimen), but fragile, inflated and broadened in
the basal (ovarian) region.
Abactinal surface of the disk invested by a tough opaque
membrane, closely covered with coarse granules which form
a base each for a tuft of minute spinelets.
Abactinal surface of the rays covered with a very delicate
transparent membrane, which in the basal part of the rays is
strengthened by from thirteen to seventeen sinuous, very
salient, transverse ridges, armed with strongly spinate plates
and covered with microscopic pedicellaria. In the intervals
between these ridges, as throughout the ray at regular close
intervals, occur broad, slightly raised, felt-like, transverse
bands of microscopic pedicellariz.
The vertebra-like adambulacral plates have the following
armature :—(1.) abactino-marginally, on a distinct but closely
fused platelet, a needle-like spine ten to twelve times as long
as the plate itself; (ii.) actino-marginally, a similar spine
four to five times as long as the plate; (iii.) on the furrow-
118 Mr. A. Alcock on
margin, one at each end of the plate, two spinelets as long as
the plate; (iv.) deep within the furrow, on a distal epiphysis,
a spinelet which stretches across the furrow to overlap with
its fellow of the opposite plate and separate the pairs of tube-
feet from one another. All these spines are hyaline, fluted,
and invested in membranous sacs which are closely felted
with pedicellariz.
Actinostome large (25 millim. in diameter in the type
specimen). Mouth-plates of moderate size, each plate dis-
tinctly formed of two fused adambulacrals, of which they
bear the furrow and cross-furrow spinelets unchanged and the
actino-marginal spines but little modified.
Madreporiform plate rather large, excavated centrally and
divided by a deep groove into two halves, with fine radial
striations, placed close to the margin of the disk.
Colour in the fresh state bright cinnabar-red.
Laccadive Sea, 1043 fathoms, green mud with foraminifera.
This species is characterized by its large disk with tough
membrane and coarse tufts of spinelets, and by the great
length and coarseness of its spinature.
52. Brisinga andamanica, Alcock and Wood- Mason.
Brisinga andamanica, Aun. & Mag. Nat. Hist., Dec. 1891, p. 439.
Rays 15. R=27r.
Disk of moderate size (its diameter being 38 millim, in the
type specimen), with vertical edge ; rays very deciduous, but
not fragile, long, slender, a little depressed, but not inflated
in the ovarian region.
Abactinal surface of the disk much as in B. énsularum, but
the membrane is not so tough, and the tufts of spinelets and
the tubercles on which they stand are not so coarse.
Abactinal surface of the rays covered with a very delicate
transparent membrane, which in the proximal third of the
ray is strengthened by about forty-five rather distant cross
bands or half-bands ot feebly developed spiny plates. In the
intervals between these bands, as throughout the whole ray at
regular intervals, occur very narrow felted bands of micro-
scopic pedicellariz.
‘he vertebra-like adambulucral plates have the following
armature :—(i.) abactino-marginally a spine about six times
as long as the plate; (ii.) actino-marginally a spine about
four times as long as the plate ; (iil.) a single furrow spinelet,
placed at the adoral end, about equal in length to the plate;
(iv.) deep within the furrow, on a distal epiphysis, a cross-
Indian Deep-sea Asteroidea. 119
furrow spine separating the pairs of tube-feet. All these
spines are hyaline, fluted, and invested in sacs formed of
membrane and felted microscopic pedicellariz.
Actinostome large (25 millim. in diameter in the type
specimen). Mouth-plates small, each plate formed of two
adambulacrals, with the furrow and cross-furrow spinelets
unchanged and the actinal spines not much modified; the
most adoral actinal spine is enlarged and is often united with
its fellow of the same pair in a common membranous
investment.
Madreporiform plate rather small, excavated centrally, with
fine radial striations, marginal in position.
Colour in the fresh state bright cinnabar-red.
Andaman Sea, 405 fathoms, green mud.
This species stands intermediate between B. ctnsularum
and the next species, B. bengalensis ; it is well characterized
by the extensive development along the rays of feeble calca-
reous plates.
53. Brisinga bengalensis, Alcock and Wood-Mason.
Brisinga bengalensis, Ann. & Mag. Nat. Hist., Dec. 1891, p. 439.
Rays 14. . R=25 r.
Disk small (its diameter being 30 millim. in the type
specimen), depressed centrally, with a strongly bevelled
abactinal margin; rays very deciduous, but not otherwise
very fragile, long, slender, hardly inflated in the ovarian
region.
Abactinal surface of the disk much as in B. andamanica ;
abactinal surface of the rays covered with a very delicate
transparent membrane, which in the basal ninth of the ray is
strengthened by about twenty close cross bands or half-bands
of spiny plates that are not very much more salient than the
cross bands of felted pedicellariz that occur throughout the
ray.
The vertebra-like adambulacral plates have the following
armature :—(i.) abactino-marginally a spine about five or six
times as jong as the plate; (ii.) actino-marginally a spine
twice to two and a half times as long as the plate ; {iii.) two
furrow-spinelets not quite half as long as the plate; (iv.)
deep within the furrow, on a distal epiphysis, a cross-furrow
spine not quite as long as the plate, separating the tube-feet.
All these are hyaline, fluted, and clothed with membrane and
felted pedicellarie.
Actinostome very large (21 millim. in diameter in the type
120 On Indian Deep-sea Asteroidea.
specimen). Mouth-plates very small, each being distinctly
formed of two fused adambulacrals, of which it carries the
furrow-spinelets and cross-furrow spinelets unchanged ; of the
two actinal spines the adoral one is remarkable in being
united with the corresponding spine of the fellow mouth-
plate in a common membranous sac, so that each pair of
mouth-plates appears to possess in this situation but one large
ligulate spine between them.
Madreporiform plate large, salient, marginal, marked with
fine but deep radial striations.
Colour in the fresh state bright cinnabar-red.
Bay of Bengal, 561 fathoms, grey mud.
This species is well characterized by the small disk with
its bevelled edge ; by the slenderness of the rays and by their
short and very delicate spinature; and by the union in a
common investing membrane of the two large actinal spines
of each pair of mouth-plates.
54. Brisinga Gunnii, sp. n.
Rays 14-15.
Disk remarkably thin, its abactinal margin slightly
bevelled; rays both very deciduous and very fragile, slender,
not inflated in the ovarian region.
Disk covered abactinally with a very thin semitransparent
membrane bearing small tufts of spinelets, of which the central
in each tuft is sufficiently elongated to give en masse a downy
appearance to the disk. In one of these specimens midway
between the centre and the margin is a pair of large spines
covered with microscopic pedicellariz.
Rays covered with a membrane of extreme delicacy, which
in the basal part of the ray is strengthened by from twenty
to thirty very contorted calcareous ridges, these standing far
out on each side like hoops, but becoming inconspicuous or
quite obsolete abactinally. In the intervals between the
ridges are narrow felted bands of pedicellariez, which also
occur in the region beyond the ridges, though, owing to
excessive denudation, their exact disposition is not deter-
minable.
The vertebra-like adambulacral plates have the following
armature :—(i.) abactino-marginally a spine about six times
as long as the plate; (ii.) actino-marginally a spine about
two and a half times as long as the plate; (iii.) a single
furrow-spinelet about as long as the plate; (iv.) deep within
the furrow, on a distal epiphysis, a cross-furrow spine about
On the Myriopoda of the ‘Challenger’? Expedition, 121
as long as the plate, separating the pairs of tube-feet. All
these are hyaline, fluted, and invested in a felt of membrane
and microscopic pedicellariz.
Actinostome large. Mouth-plates very small; each plate
is made up of two adambulacrals so incompletely fused that
in place of the usual groove there is a ligamentous symphysis
between the two plates, and each bears the usual furrow and
cross-furrow spinelets unchanged and the actinal spine not
meh changed—only diminished in size in the distal (aboral)
ate.
Madreporiform plate marginal, small, deeply cross-fissured,
radially striated. A membranous (epiproctal?) appendage is
found on the disk excentrically.
Colours in the fresh state dull reddish ochre.
Off the Konkan Coast, 559 fathoms, green sand.
This species is well characterized by the strong contortion
of the caleareous ridges at the base of the rays and their hoop-
like lateral elevation ; by the incomplete fusion and partially
ligamentous union between the two adambulacral elements
that make up a mouth-plate ; and by the curious membranous
epiproctal (?) appendage.
FREYELLA, Perrier.
5d. Freyella tuberculata, Sladen.
Freyella tuberculata, Sladen, ‘ Challenger’ Asteroidea, p. 658, pl. exvii.
figs. 1-3.
Bay of Bengal, 1840 fathoms, Globigerina-ooze.
56. Lreyella benthophila, Sladen.
Freyella benthophila, Sladen, ‘ Challenger’ Asteroidea, p. 641, pl. exi.
figs. 5-8; and in Wood-Mason and Alcock, “Indian Deep-sea
Dredging,” Ann. & Mag. Nat. Hist., Dec. 1891, p. 440.
Bay of Bengal, 1520 to 1997 fathoms, usually on Globi-
gertna-00ze.
XVI.—Report upon the Myriopoda of the ‘Challenger’ Expe-
dition, with Remarks upon the Fauna of Bermuda. By
R. I. Pocock, of the British Museum (Natural History).
[Plate IX.]
SINCE marine biological research was the main object of the
cruise of the ‘ Challenger,’ and the capture and preservation
of terrestrial forms but a pastime, so to speak, of the zoologists
Ann. & Mag. N. Hist. Ser.6. Vol. xi. i)
122 Mr. R. I. Pocock on the
on board, it is not difficult to understand why the species
collected from time to time inland of the different ports that
were touched at, were for the most part colisiderel by those
into whose hands they ultimately fell to be, if of interest at
all, certainly not worthy of special reports. However sey
fiable this opinion was touching the majority of the land-
forms of life, it can scarcely be said to hold good when we
take the Myriopoda into consideration. Nor is this a sur-
prising fact, seeing that many of the localities visited were
isolated islands or localities rarely, if ever, explored by
collectors ; and consequently a large percentage of new forms
was obtained—large, that is, considering that those who
discovered them were without experience of the group, and
merely picked up such specimens as presented themselves
during the search after land-forms in general. But, apart
from the new species, of which the value can only be appre-
ciated by a specialist, the interest of the collection touching
geographical distribution is in some respects very great.
This is notably the case with regard to Bermuda.
Prior to the small report published by Mr. Bollman upon
the Myriopoda obtained by Dr. Angelo Heilprin in Bermuda,
I am not aware that any species of the group—authentically
named or otherwise—have been recorded from this island.
Mr. Bollman mentioned five species in his paper; in the
‘Challenger’ collection there are in all seven, five of them
being new to the locality and two of them doubtfully new to
science,
The following is a complete list of the known species, with
notes respecting their extra-Bermudan distribution :—
1. Scutigera coleoptrata (Linn.).
Common in South and West Europe, Madeira, and Azores,
and probably introduced into Bermuda from the east. Not
Antillean; but possibly Nearctic, if Sc. forceps is the same
species.
2. Lithobius provocator, Pocock.
Certainly closely allied to, if indeed it be truly distinct
from, the common European and N.-American ZL. forficatus.
Not Antillean.
3. Lithobius bermudensis, Pocock.
Also a doubtful species. No doubt of European or Nearctic
origin. Almost certainly not Antillean, the genus Lithobius
being unknown in the West Indies.
Myriopoda of the ‘Challenger’ Expedition. 123
4. Lithobius lapidicola, Meinert.
This species was identified with some hesitation by Boll-
man, to whom authentic examples of L. lapidicola were
unknown. I seeno reason, however, for doubting the correct-
ness of the determination, seeing that ZL. lapidicola is a
tolerably common European form and occurs also in Madeira.
5. Scolopendra subspinipes, Leach.
A widely distributed tropical species. Occurring commonly
in the West Indies, whence it has doubtless made its way to
Bermuda.
6. Mecistocephalus Guildingii, Newp.
Recorded by Bollman. A West-Indian species; occurs
also in Demerara.
7. Strongylosoma coarctatum (Sauss.).
Like Scolopendra subspinipes, this is a widely distributed
tropical species. Abundant in Guyanaand the West Indies ;
occurs also in Europe, but only in conservatories or places to
which tropical plants have been imported.
8. Strongylosoma Guerinii, Gervais.
Common in Teneriffe, Madeira, Algeria, &c., but not
known upon the western side of the Atlantic.
9. Iulus Moreleti, Lucas.
Common in the Azores and Madeira, and closely allied to
several species of the South-European fauna.
10. Spirobolus monilicornis, Porath.
Described from Bermuda under the name Sp. Heilprini by
Bollman ; but Bermudan specimens are not specifically distin-
guishable from others occurring in many of the West-Indian
Islands and in Demerara, which are almost certainly monili-
cornis of Porath.
From the above list it will be seen that out of the ten
known species four have been doubtless introduced from the
West Indies, three are either of Palearctic or Nearctie origin
(7. e. nos. 1, 2, and 3), while the remaining three belong
#
124 Mr. R. I. Pocock on the
unquestionably to the Mediterranean fauna of the Palearctic
region.
Thus the western Palearctic element is almost equal to
the Antillean—a circumstance which is both interesting and
surprising, seeing that the plants, land-mollusca, insects, and
spiders appear to be almost wholly West-Indian or Kast-
American. ;
Also of considerable interest was the small series of species
obtained from Teneriffe, showing the distinctly Mediterranean
character of the mainland fauna of this island.
Class CHILOPODA.
Fam, Scutigeride.
Scutigera coleoptrata (Linn.).
Loc. Bermuda.
Scutigera planiceps, sp. n.
Scutigera rugosa, Porath, “ Myriopoda Afric australis in Museo Regis
Holmiensi asservata,” C2fvy. Vet-Akad. Forhandlingar, 1871, no. 9,
pp. 1158, 1189 (not Seutigera rugosa, Newport, Linn. Trans, xix.
p. 353, 1845).
Loc. Simon’s Bay (Cape Town). A single specimen.
Porath’s description of this species is sufficiently accurate
to leave no doubt as to the specific identity of his specimens
with the one mentioned above. But since these specimens
are certainly totally distinct from rugosa of Newport, the type
ot which is preserved in the British Museum, it is necessary
that they receive a new name. It is proposed therefore to
call them planiceps, in commemoration of perhaps their most
marked characteristic, namely the flatness of the upper surface
of the head.
Porath’s specimens were evidently somewhat faded, for he
merely describes them as being ‘ supra nigrescens, subtus
pallidior.” In reality the general tint of the upper surface is
a deep black-brown; but when more critically examined the
tergites are seen to be adorned with a median black longitu-
dinal band and to be black at the sides, the intermediate area
being of a reddish-brown tint; the legs are a pale olivaceo-
testaceous colour and the tarsi are ochraceous. ‘The stomata
are small and the stoma-saddles very low,
Se. rugosa of Newport, from East Africa, which Porath
mistook for this species, is very different. It is a very beautiful
form, the tergites being a deep black, this sombre colouring
being relieved by an orange-yellow median dorsal band which
Myriopoda of the ‘Challenger’ Expedition. 125
runs without interruption from the fore part of the head to
the hinder end of the body ; moreover the lower surface and
legs are also orange-yellow, these last being beautifully ringed
with black. The head, too, is deeply excavated, the stoma-
saddles are elevated, and the generative forceps has its distal
segment smooth and not serrate internally.
Sc. planiceps seems to have a wide distribution in South
Africa. Porath has recorded it from the Cape of Good Hope
and Caffraria, and the British Museum has a single example
which was collected by Gueinzius at Port Natal.
Fam. Lithobiide.
Henicops maculatus, Newport.
Henicops maculata, Newport, Trans, Linn. Soc., Zool. xix. p. 372,
pl. xxxiii. fig. 27, and pl. xl. fig. 3; id. Catalogue Myr. Brit. Mus.
22
Henicops impressus, Hutton, Ann. & Mag. Nat. Hist. (4) xx. pp. 114,
115 (1877); id. Tr. N. Z. Inst. x. p. 288 (1877).
Henicops maculata, Haase, “ Die Indisch-Austral. Chilopoden,” Ab-
handlungen Dresden Museum, 1887, no. 5, p. 36; Pocock, Ann, &
Mag. Nat. Hist. (6) viii. p. 154 (1891).
Loc. Wellington (New Zealand).
Henicops insignis, Pocock.
THenicops insignis, Pocock, Ann. & Mag. Nat. Hist. (6) viii. p. 154.
Loc. Juan Fernandez.
Henicops tristant, sp. nu.
Colour fusco-ochraceous ; antenne entirely flavous, legs
flavous distally.
Head convex from side to side, the anterior median portion
angularly notched. Antenne short, composed of 23-24
segments.
Coxe of the maxillipeds produced forwards mesially and
armed with 2+2 sharp teeth. Tergites smooth, with all
their posterior angles rounded and their hinder borders not
emarginate. Coxal pores rounded, 3, 3, 3, 2. Generative
forceps of the female with two short spurs rising just below
the apex of the internal posterior angle of the segment ; claw
simple.
Length up to 10 millim.
Loc. Tristan Island (Tristan d’Acunha).
This species is closely related to LH. africanus of Porath,
from South Africa. It appears to differ, however, in having
126 Mr. R. I. Pocock on the
fewer antennal segments. In Porath’s species, judging from
a number of examples sent to the British Museum from Cape
‘own and Port Elizabeth, the usual number of segments to
these appendages is 30, but it varies from 26 to 35, whereas
in HZ. tristant the number is 23 to 24 in four specimens.
Lithobius pilicornis, Newp.
Lithobius pilicornis, hay ee Ann, & Mag. Nat. Hist. xiii. p. 96. no. 5
(1844) ; id. Tr. Linn. Soc. xix. p. 369; Pocock, Ann, & Mag. Nat.
Hist. (6) vii. p. 872.
Lithobius Sloanei, Newport, Ann. & Mag. Nat. Hist. xiii. p. 96. no. 6
(1844); id. Tr. Linn. Soe. xix. p. 569.
Lithobius longipes, von Porath, (ify. Vet.-Akad. Forh. xxvii. p. 816
(1870) ; Meinert, Nat. Tidskr. (3) viii. p. 823.
Lithobius galatheez, Meinert, Vid. Medd. Foren, 1884-86, p. 109.
Loc. Teneriffe (Canary Islands).
Lithobius provocator, Pocock.
Lithobius provoeator, Poeock, Ann. & Mag. Nat. Hist. (6) viii. p. 152.
Loc. Bermuda.
Lithobius bermudensis, sp. ti. (?).
Colour uniform fusco-ochraceous ; legs flavous, head ochra-
ceous.
Head smooth, cordate; eyes composed of about 10 ocelli,
1+4, 3,2. Antenne broken off.
Coxe of the maxillipedes armed with 2+2 strong acute
teeth.
Tergites at the posterior end of the body lightly rugose ;
the posterior angles of 9, 11, and 13 strongly dentate.
Coxal pores rounded, in a single series, 4, 5, 5, 4.
Anal legs long but moderately thick, armed beneath with
1, 3, 2, 1 spurs ; claw spurred.
Generative forceps of the female with two longish, subequal,
subparallel spurs and trilobate claw.
Length 14°5 millim., with anal leg 19 millim.
Loc. Bermuda.
I am rather doubtful as to the distinctness of this species,
partly owing to the imperfection of the antenne of the type
specimen and partly to my ignorance of the North-American
species of the genus. I cannot, however, match it with any
European species with which I am acquainted.
Myriopoda of the ‘Challenger’ Expedition. 127
Fam. Scolopendridz.
Scolopendra morsitans (Linn.).
Loc. St. Iago (Cape Verde) ; Zebu (Philippine Islands).
Scolopendra mutilans, L. Koch.
Scolopendra mutilans, L. Koch, Verh. z.-b. Ges, Wien, 1878, p. 791 ;
Haase, op, cit. pp. 47, 48.
Loc. Yokohama.
Scolopendra chilensis, Gervais.
Scolopendra chilensis, Gervais, Ins. Apt. iv. p. 285 (1847); id. in Gay’s
Hlist. de Chile, Zool. iv. p. 67, fig. 7 (1849); Meinert, Proc. Am.
Phil. Soc. 1886, pp. 199, 200.
Loc. Valparaiso.
Pithopus calcaratus, Pocock.
Pithopus calearatus, Pocock, Ann. & Mag. Nat. Hist. (6) vii. pp. 224,
225.
Loc. Bahia.
Cormocephalus violaceus (Fabr.).
Scolopendra violacea, Fabr. Suppl. Ent. Syst. p. 289; Guérin-Ménéville,
Icon. Régne Anim. ii., Ins. pl. i. fig. 8.
Cormocephalus calcaratus, Porath, CEfv. Vet.-Akad. Férhandl. 1871,
no. 9, pp. 1159, 1160,
Loc. Simon’s Bay (Cape of Good Hope).
There can be very little doubt, I think, as to the correct-
ness of the above synonymy. Fabricius’s description and
Guérin’s figure are accurate enough to preclude the likelihood
of error in the identification of the species. Newport never
saw an example of it; he consequently retained the name
violacea under the genus Scolopendra, an error which gave
rise to two others. For, in the first place, Newport him-
self described a New Zealand species of Cormocephalus as
violaceus, and Porath, following Newport’s opinion respecting
the generic position of Fabricius’s violaceus, recharacterized
it as a Cormocephalus which he called calcaratus. C. calca-
ratus is consequently a synonym of C. violaceus (Fabr.), and
the New Zealand violaceus of Newport, which is different
from the African, must be renamed. I propose to call it
pu rp ureéus.
12 Mr. R. I. Pocock on the
io 2)
Cormocephalus rubriceps, Newp.
Cormocephalus rubriceps, Newport, Tr. Linn, Soc. xix. p. 419; Haase,
op. cit. p. 57.
Loc. Maua Island (New Zealand). een
On the parchment label accompanying this species is written
“ Centipede now confined to the island of Maua.”
Cormocephalus Huttont, sp. u.
Cormocephatlus violaceus, Hutton, Tr. N. Z. Inst. x. p. 289, 1877 (not
violaceus of Newport).
Colour (in aleohol and certainly faded) olivaceous or ochra-
ceous, often with metallic lustre, the head, first segment, anal
segment, legs, and antenne pale ochraceous.
Head usually a trifle longer than wide, distinctly punctured.
Antenne composed of 17 segments, whereof about the basal
7 or 8 are naked.
Prosternal plates of the coxe tolerably long, in contact,
each furnished with 4 acute teeth. Tergites punctured, from
the second bisulcate, from the eighth marginate.
Sternites bisulcate, not otherwise impressed.
Anal somite.—Tergite not mesially suleate, nearly parallel-
sided, a little wider than the head, but narrower than the first
and twentieth; pleure densely porous, the process long,
slender, tipped with two spines, another minute spine on the
posterior border of the pleur ; sternite posteriorly narrowed.
Legs short and moderately robust, the femur incrassate
posteriorly, a little more than twice as long as wide; the
process moderately large, tipped with two spines, the inner
surface usually armed with 6 small spines, 2 above, 2 in the
middle, and 2 below, the lower surface only very shallowly
excavated, the external carina being low and armed with 2, 2
spines in two series ; claw without basal spurs.
Rest of the legs with unspined tarsi.
Length up to 70 millim., of antenna 10, of anal leg 11;
width of anal tergite 4°2, of head 4+.
Loc. New rostani
‘These specimens are specifically identical with one of Prof.
Hutton’s examples of his violaceus, Newp., which the British
Museum obtained from the Otago University Museum. This
specimen is ticketed Wellington; so in all probability the
‘Challenger’ examples came from the same locality. The
Museum also has an example from Waikouaito.
The species differs from C. purpureus, Pocock (=violaceus,
Newp., cf. supra), in having no spurs at the base of the claw
on the anal leg and in having 4 spines in two series on the
Myriopeda of the ‘Challenger’ Expedition. 129
lower surface of the femur of the anal leg. In purpureus
there is a single series of 3 spines in this position and the claw
of the anal leg is spurred ; the anal pleura, moreover, have a
shorter and more angular process. C. Huttoni is most nearly
allied to C. fecundus of Newport, from Van Diemen’s Land
and South-east Australia. It differs, however, at least in
having no median sulcus on the anal tergite.
Heterostoma rubripes, Brandt.
Heterostoma rubripes, Brandt, Recueil &c. p. 65; Haase, op. cit. p. 89,
pl. v. fig. 93.
Loc. Cape York (N. Australia).
Heterostoma viridipes, Pocock.
Heterostoma viridipes, Pocock, Ann, & Mag. Nat. Hist. (6) vii. pp. 56,
57, pl. iv. fig. 2.
Loc. Ternate.
? Cryptops hortensis, Leach.
Loc. Teneriffe.
The identification of this species is doubtful on account of
the absence of the anal legs.
Cryptops australis, Newport.
Cryptops australis, Newport, Tr. Linn. Soc. xix. pp. 293 and 408.
Loc. Wellington (New Zealand).
Geophilus challengeri, Pocock.
Geophilus challengeri, Pocock, Ann, & Mag. Nat. Hist. (6) viii. pp. 217,
218, pl. xii. fig. 3.
Loc. St. Iago (Cape Verde).
Fam. Geophilide.
Himantarium dimidiatum, Meinert.
Himantarium dimidiatum, Meinert, Nat. Tidskr. Kroyer, (3) vii. pp. 30,,
31 (1870-71).
Loc. ‘Teneriffe (Canary Islands).
Recorded originally from Seville and Madeira.
Geophilus antipodum, Pocock.
Geophilus antipodum, Pocock, loc. cit. pp. 222, 223, pl. xii. fig. &
Loc, Wellington (New Zealand).
130 Mr. R. I. Pocock on the
Geophilus provocator, Pocock.
Geophilus provocator, Pocock, loc. cit. p. 225, pl. xii. fig. 10.
Loc. Wellington (New Zealand).
Class DIPLOPODA.
Order ONISCOMORPHA.
Fam. Glomeride.
Spherotherium angulatum, Butler.
Spherotherium angulatum, Butler, Tr. Ent. Soc. 1878, p. 299, 9.
Spherotherium walesianum, Karsch, Arch. Nat. 1881, p. 31, pl. ii.
fig. 1, gd.
Loc. Queensland.
Butler’s specimens were from Rockhampton, the type of
walesianum trom Sydney.
Order HELMINTHOMORPHA.
Fam. Polydesmide.
Strongylosoma gracile (C. Koch).
Fontaria gracilis, C. Koch, Syst. d. Myr. p. 142 (1847) ; id. Die Myr. 11.
p- 51, tig. 175.
Paradesmus gracilis, Latzel, Tomésvary, &e.
Loc. Sandwich Islands (Hilo and Honolulu); Cape of
Good Hope.
Strongylosoma coarctatum, Sauss.
Paradesmus coarctatus, Sauss. Mém. Mex. Myr. p. 39, fig. 18 (from the
Mém. Soc. Phys. Genéve for 1860).
? Paradesmus coarctatus, Humb. & Sauss. Verh. z.-b. Wien, xix. pp. 670,
571 (1869).
Paradesmus vicarius, Karsch, Arch. f. Nat. 1881, p. 38, pl. iii. fig. 8.
Strongylosoma Poeyi, Bollman, Ent. Amer. iii. p. 81.
Loc, Bermuda ; Mactan and Zebu (Philippines).
Very common in the West Indies and in Guyana, Burma.
Strongylosoma Guérinii, Gervais.
(PI. IX. fig. 2.)
Polydesmus Guérinii, P. Gervais, Ann, Soc. Ent. Fr. v. p, 686 (1836) ;
id. Ins. Apt. iv. ri 116, pl. xlv. fig. 3 (1847); Humbert and Saussure,
Verh. z.-b. Ges. Wien, xix. pp. 685, 686 (1860).
Polydesmus cylindraceus, P. Gervais, Ins, Apt. iv. p. 117, pl. xlv. fig. 3 ¢.
Loc. eneriffe ; Bermuda.
Myriopoda of the ‘Challenger’ Expedition. 131
This species has hitherto been recorded from Barbary,
Marocco, Algeria, and Madeira. The British Museum has
many from the two latter localities.
Strongylosoma Gervaisii (Lucas).
(Pl. IX. figs. 1, 1a.)
Polydesmus Gervaisti, Lucas, Hist. Nat. Anim. Articul. Apt. p. 525
(1840) ; Gervais, Ins. a iv. p. 118.
Strongylosoma trilineatum, Newport, Ann. Nat. Hist. xiii. p. 266 (1844).
Strongylosoma Petersii, L. Koch, Verh. z.-b. Ges. Wien, 1865, p. 882;
Karsch, Arch, Nat. 1881, p. 44.
Loc. Paramatta.
Stenonia tuberosa, sp. n.
(Pl. IX. figs. 3-34.)
9. Colour deep café-au-lait, with the antenne, legs, and
margins of the keels brunneo-fulvous.
Antenne short.
Collum convex from side to side, the anterior border evenly
rounded from the apex of one keel to that of the other, the
keel triangular, depressed, its posterior border oblique and not
continuing the curve of the hinder border of the tergite; a
transverse groove sometimes running along the anterior border,
closely covered with rounded tubercles, those along the two
borders larger than the others. The rest of the segments
granular or tubercular like the first, all of them marked with
three distinct rows of larger tubercles; the keels slightly de-
pressed, rising a little above the middle of the side, not large,
squared, the anterior border smooth, with a distinct basal
shoulder, the posterior border smooth, the lateral border
furnished with from 4 to 6 smooth rounded tubercles (that is
including the anterior and posterior angle). Pores close to
the lateral margin, separated from it by a space about equal
to their own diameter.
Anal tergite evenly rounded, at most lobulate. Lateral
surface of the segments closely granular and rugose.
Sterna not spined.
g. A little narrower and flatter than the female, the keels
being a trifle larger. The copulatory feet terminating distally
in two processes, the distal simple, long, slender, and lightly
curved, the proximal itself dividing into two—an upper
strongly curved and dilated at its distal end, an under simple
and pointed.
Length, g 40 millim., width 65; ¢, length 44, width 7:3.
Loe. Ki Dulau (Ki. Islands).
132 Mr. R. I. Pocock on the
Fam. Iulide.
Iulus Moreleti, Lucas.
Iulus Moreleti, Lucas, in Arthur Morelet’s ‘Notice sur l'Histoire
Naturelle des Azores,’ Paris, 1860, p. 96 ; Porath, Bih. Sy. Vet.-Akad, ~
Handl. no. 7, p. 821 (1870).
Loc. Bermuda.
Described originally from the Azores by Lucas and subse-
quently recorded by Bollman from Bermuda. The British
Museum has examples of what is apparently the same species
from Madeira.
Iulus canariensis, sp. n.
Colour black, in spirit greyish blue banded with black ;
legs pale.
Body very long and slender.
Head smooth, with a frontal sulcus and a transverse stria
between the eyes; without apparent frontal sete. Eyes large,
with ocelli distinctly defined, arranged in six transverse rows.
Antenne absent.
Collum smooth above, triangular laterally, striate just above
the angle. The rest of the segments with complete transverse
suleus and a median dorsal longitudinal sulcus, all of them,
including the second, longitudinally striated behind the
sulcus, the strie fine, complete, and very close-set, the area in
front of the sulcus smooth. Pores large, situated about one
third of the distance behind the sulcus, which is at this spot
lightly sinuate forwards, the space between the sulcus and the
pore not striate.
Anal tergite not surpassing the valves; valves convex, not
compressed or marginate.
Anterior legs of the male hook-like.
Number of segments about 60.
Length up to about 26 millim,
Loc. ‘Teneriffe.
In its black colouring, slender build, position of pores, &e.
this species at first sight seems to fall into the Ophiulus section
of Berlese; but the absence of a spine on the terminal seg-
ment serves at once to distinguish it.
lulus tristant, sp. n.
(Pl. 1X. figs. 5, 5a.)
Colour fuscous or flavous and fusco-annulate ; head yellower,
with fuscous fascia between the eyes; legs and antenne
flavous ; anal somite fuscous.
Myriopoda of the ‘Challenger’ Expedition. 133
Head smooth, without frontal sete. Hyes obscurely mani-
fested, represented by a patch of black pigment. Antenne
short, scarcely surpassing the collum.
Collum laterally angular and marked with a few strie.
The first five segments smooth above, striate at the sides ;
the rest striate above and at the sides, the striz not close-set
and not reaching the hinder border of the segments. ‘The
sulcus complete and deep, crenulate, the area in front of it
smooth. The pores conspicuous, situated just behind the
sulcus, and, except at the hinder end of the body, touching it,
the sulcus being posteriorly angled to meet the pore.
Anal tergite covering the summit of the valves, sometimes
just surpassing them; valves convex, not compressed, and
scarcely hairy ; sternite triangular.
Legs short.
d- Smaller and thinner than female. The first pair of
legs small and hook-like; the second pair without coxal pro-
cesses ; the second segment of the mandible produced back-
wards into a large rounded prominence.
Number of segments up to 44.
Length up to 15 millim.
Loc. Inaccessible Island and Tristan Island (Tristan
d’Acunha).
Iulus solitarius, sp. n.
(Pl. 1X. fig. 4.)
Colour very like that of J. puszllus of Leach ; two longitu-
dinal flavons dorsal bands, the median dorsal line and the
sides of the body black, the lower surface and legs flavous ;
antenne and space between the eyes fuscous.
Head lightly punctulate, without distinct frontal pores.
Eyes well developed, subcircular, composed of at least five
distinct rows of well-defined ocelli. Antenne a little longer
than in the preceding species.
The collum laterally acutely triangular, margined, its ante-
rior border lightly sinuate ; the first three or four segments
smooth above, striate at the sides, the rest striate above and
below, the strize deep but not close-set and not reaching the
hinder border of the segments. ‘The sulcus deep, the area
behind it raised, that in front of it smooth. Pores scarcely at
all conspicuous, situated just behind the sulcus but touching
it, the sulcus lightly anteriorly angled.
Anal tergite surpassing the valves a little, the process short
and down-curled ; valves convex, not compressed, and smooth ;
sternite acutely triangular and very long.
134 Mr. R. I. Pocock on the
Legs longer than in J, tristant.
@. Smaller and thinner than female. The first pair of
legs small and hook-like, the second pair with simple cox ;
mandible angularly produced.
Number of segments 33-34.
Length up to about 10 millim.
Loc. Tristan d’Acunha (Inaccessible Island and Tristan
Island).
Iulomorpha Porathi, sp. n.
Colour black, with two flavous or testaceous spots on the
dorsal area of the segments, constituting two parallel flavous
bands; anal tergite black and polished, anal valves, legs, and
antenne testaceous; head fusco-ferruginous, with a deeper
fascia between the eyes.
Head smooth, with weak frontal sulcus and a fine stria
between the eyes ; labral pores 4+4. Eyes widely separated,
small and triangular, composed of about three transverse
rows of ocelli. Antenne longish, reaching past the second
segment.
Collum laterally elongate, triangular, with a marginal
sulcus and two longitudinal sulci, smooth above. ‘The rest of
the segments without trace of a transverse sulcus, merely
marked by a shallow transverse groove, the area behind this
groove smooth above, inferiorly finely ridged longitudinally,
the covered area of the anterior portion closely and finely
transversely striate in front. The pores large, above the
middle of the side at some little distance behind the trans-
verse groove. Sterna finely striolate.
Anal tergite covering but not surpassing the valves; valves
convex, not compressed and not marginate.
Sternum obtusely rounded.
Legs sparsely setose beneath.
Number of segments 47.
Length about 36 millim., width 2-5.
Loc. Malamaui (Philippine Islands), Two female ex-
at a
take great pleasure in dedicating this striking species,
the second known of the genus, to Dr. C. O. von Porath, who
was almost the first zoologist to describe exotic Myriopoda
with accuracy.
This new form J. Porathi agrees tolerably closely with the
South-African J. Kinbergi in most of its structural features.
It may at once be recognized, however, by its marked colouring
and smaller number of segments.
The second example, measuring only about 12 millim., with
Myriopoda of the ‘Challenger’ Expedition. 135
35 segments, and apparently the young of the large one
described, has the yellow dorsal spots much smaller and
visible just above the pore, the collum almost entirely flavous,
and a faint though distinct transverse stria on the segments.
Spirostreptus chilensis, Gervais.
Tulus chilensis, Gervais, Ins. Apt. iv. p. 193; id. in Gay’s Fist. de
Chile, Zool. iv. p. 61, fig. 3 (1849).
Spirostreptus chilensis, Porath, Bih. Sv. Vet.-Akad. Handl. iv. no. 7,
p- 41.
Loc. Valparaiso.
Spirostreptus erythropareius, Brandt.
Spirostreptus erythropareius, Brandt, Bull. Ac. Sci. St.-Pétersb. viii.
nos. 175 and 176, p. 111 (1841); Gervais, Ins. Apt. iv. p. 155;
Porath, Géfy. Vet.-Akad. Forh. 1872, no. 5, p. 56. a J
Spirostreptus rubripes, Newport, Ann. & Mag. Nat. Hist. xiii. p. 270
(1844).
Loc. Cape of Good Hope.
Sptrostreptus Moseley?, sp. n.
(Pl. IX. figs. 6, 6a.)
Colour brunneo-fuscous ; legs and antennz ferruginous.
Body moderately robust, not attenuated at its posterior
end, narrowed at about the fourth segment.
Head sculptured below, nearly smooth and faintly sulcate
above. yes acutely angled internally, separated by a space
that is greater than a diameter. Antenne about as long as
the head.
First tergite narrowed laterally, both the anterior and pos-
terior border lightly emarginate, the anterior angle roundedly
rectangular, the posterior acutely rectangular; a strong
marginal sulcus and two abbreviated sulci. The rest of the
somites with conspicuous and complete transverse sulcus, the
anterior covered area finely transversely striolate, the median
and postericr portions finely sculptured, the sculpturing con-
sisting of fine close-set longitudinal striole; the posterior
portion longitudinally striate about a third of the way up to
the pore; pores behind the sulcus. Ventral grooves long, but
not quite as long as the two basal segments of the leg.
Anal somite large, punctulate; tergite posteriorly angled,
lightly constricted, the process not projecting beyond the apex
of the valves ; valves convex, prominent, with their margins
strongly compressed ; sternite triangular, without trace of
suture.
136 Mr. R. I. Pocock on the
Legs short, mostly with a single seta on the lower surface
of each segment.
Number of segments 70.
Length about 115 millim.
A single female example from Malamaui (Philippines).
This species differs from all the Indo- and Austro-Malayan
forms that are characterized by long ventral grooves in that
there is no caudal process overlapping the valves.
ACANTHIULUS, Gervais.
Acanthiulus, Gervais, Ann. Sci. Nat. (3) 1, p. 70; Ins. Apt. iv. p. 173.
The species of Acanthiulus can scarcely be regarded as
more than extremely well-marked forms of Spirobolus. The
constitution of the mouth-parts and the number and dispo-
sition of the legs of the anterior segments are the same in the
two groups. In fact the only character by which Acanthiulus
is to be distinguished is the presence of a transverse row of
(8) spiniform tubercles on most of the segments. Nevertheless
the collum is undoubtedly much longer than is usually the
case in Spirobolus. This last character no doubt misled
Gervais iuto stating that his species belonged to the genus
Sptrostreptus of Brandt, and probably this statement, coupled
with the presence of the spines on the segments, will account
for Latzel’s erroneous surmise that Acanthiulus and T’rachy-
tulus were synonyms.
Acanthiulus Murrayt, sp.n. (PI. IX. figs. 7-7 6.)
Colour black or olivaceous, with a series of red spots on
each side marking the pore; legs and antenne olivaceous,
with the distal ends of the segments flavo-annulate.
Face entirely smooth, the frontal sulcus mesially interrupted ;
labral border somewhat deeply excised; pores 2+2. yes
well developed, subcircular, separated by a space equal to
about twice a diameter. Antenne long and slender, much
longer than the face.
Collum smooth above, projecting laterally below the level
of the second segment, the anterior border of the lateral portion
widely emarginate and defined by a deep sulcus, which curves
round the inferior border, the posterior border deeply and
abruptly emarginate; the inferior surface of the second
segment flat, with carinate anterior border, the posterior
margin of this segment longitudinally grooved below, irregu-
larly wrinkled above, with a squamiform tubercle in the
middle of the lateral surface; the third, fourth, and fifth
segments sculptured like the second, but more coarsely, the
Myriopoda of the ‘Challenger’ Expedition. 137
squamiform tubercles that are so characteristic of the succeed -
ing segments becoming gradually more and more manifested.
The rest of the segments with their posterior portions lon-
gitudinally grooved or wrinkled from base to summit and
furnished with 8 equidistant, spiniform, squamiform tuber-
cles, forming 8 longitudinal rows of spines: the lowest of
these series is situated just below the middle of the side
and begin on the seventh segment; the second series,
beginning at the sixth segment, is situated just below the
level of the pores, the spines are larger than those of the
lowest series; the upper series on each side is composed
of flatter tubercles, which, nearly obsolete on the anterior
segments, become spiniform only at the hinder end of the
body ; in the middle line of the dorsum there is a faint indi-
cation of a similar series of tubercles. The tubercular area
of the segments is lightly elevated and separated from the
anterior area, which is rugose and finely transversely striolate
in front, by a shallow depression representing the transverse
sulcus. Pores conspicuous, situated in front of the sulcus
and just above and in front of the large tubercle of the median
lateral series ; the anterior pore far below (1°5 millim.) the
level of the rest, and situated immediately in front of the
tubercle, which is itself lower than those on the rest of the
segments. Sterna finely striolate transversely.
Anal somite large ; tergite without spines but rugulose, the
posterior angle elongate but not surpassing the summit of the
valves; valves with strongly compressed margins, finely
rugulose in front, coarsely rugose and punctured behind ;
sternite large and angular.
Legs longish and slender, with a single seta below each
segment, except the tarsus, which has about three; the ante-
rior legs, at least in female, shorter, thicker, and more setose
beneath. :
Number of segments 50.
Length 111 millim., width 11.
Loc. Wokan Dobbo (Aru Islands).
Acanthiulus Blainvillet, Le Guillou*, from New Guinea, the
only other known species of the genus, differs from this new
form, if we may trust Gervais’s figure, in having the antennz
shorter than the face, the antero-lateral border of the collum
less widely emarginate, and the posterior angle more strongly
produced. Moreover the spiniform tubercles are well deve-
loped on the anterior segments.
* Bull. Soc. Phil. Paris, 1841, p. 86; Gervais, Ins. Apt. iv. p. 173,
pl. xliv. fig. 8.
Ann. & Mag. N. Hist. Ser. 6. Vol. xi. 10
138 Mr. R. I. Pocock on the
I have great igre in dedicating this interesting new
species to Dr. John Murray.
Sptrobolus monilicornis, Porath.
Spirobolus monilicornis, Porath, Bih. Sv. Vet.-Akad Handl, iy. no. 7,
p. 31 (1876).
Spirobolus Heilprini, Bollman, Proc. Ac. Philad. 1889, p. 127.
? Spirobolus virescens, Daday, Term. fiizetek, xiv. p. 140, pl. vii.
figs. 8-10.
Loc. Bermuda.
This species occurs commonly in many of the West-Indian
Islands and also abundantly in Demerara. Dr. Porath’s
examples were ticketed Brazil. S. Heilprini was described
from Bermuda, and virescens, which appears to be the same
species, from Trinidad.
Sptrobolus digrammus, sp. 0.
(Pl. IX. figs. 9-9 d.)
? Sptrobolus tessellatus, Porath, op. cit. p. 21 (in part only, ze. the
example from the Cape).
Colour.—Head fuscous above, flavous below the eyes; legs
and antenne bright red, concolorous; first tergite black,
with its anterior border blood-red, the rest of the segments
blood-red in their lower half, black in the upper, with two
complete parallel blood-red bands running from the posterior
margin of the first tergite to the anal somite; anal valves red.
Head punctulate, with a median sulcus above and below,
with four labral pores. yes separated by a distance about
equal to twice a diameter, subcircular, each composed of
about 44 ocelli.
First tergite punctulate or striolate, the lateral portior.
narrowed to an angle, with only an anterior marginal sulcus.
The rest of the tergites punctulate and striolate throughout,
longitudinally striate laterally behind the transverse sulcus,
but the striz not extending as high as the pore; the trans-
verse sulcus nearly obsolete above. The pores large, above
the middle of the side, the anterior the lowest, close to each
is a posterior longitudinal stria; scobina present in most of
the somites, but small.
Anal somite punctulate, the tergite with a somewhat sharp
but short median posterior angle ; valves convex, with margins
neither suleate nor compressed ; sternite posteriorly angled.
Legs short, with a single seta above the claw and each
segment furnished distally beneath with one seta; the male
with the distal segment padded beneath.
Myriopoda of the ‘Challenger’? Expedition, 139
Copulatory feet of the male as in fiz. 9 4.
Number of somites 45-46.
Length about 40 millim.
Loc. Cape Town (Simon’s Bay).
So far as colouring is concerned this species approaches
Sp. formosus of Porath; but [ infer from the description of
this last that the red dorsal bands are not continuous. More-
over, formosus is said to be “ leve,” and the anal valves are
compressed. In colouring again this species very much
resembles Sp. litoralis; but in litoralis the whole head is
red and the dorsal spots are not continuous from segment to
segment. Porath mentions one specimen of (itoralis (his
tessellatus) as coming from Cape Town and having a fuscous
band between the eyes. This example, [ strongly suspect, is
in reality to be referred to this species.
Spirobolus cupultfer, Voges.
Spirobulus cupulifer, Voges, Zeitschr. wiss. Zool. xxxi. p. 188.
Loc. Zamboanga and Zebu (Philippine Islands).
Recorded originally from Bohol.
Spirobolus challengeri, sp. n.
(Pl. IX. figs. 10-10 c.)
Colour olivaceo-fuscous, most of the segments adorned
posteriorly and laterally with an inferiorly narrowed orange-
yellow stripe, which runs along the hinder margin, beginning
on a level with the pore and extending halfway towards the
legs; legs and antennz narrowly testaceo-annulate.
Head convex, shining, smooth or only very finely striolate,
with two pores on each side of the labrum, with a longitudinal
mesially nearly obsolete sulcus. yes subcircular, consisting
of about seven transverse rows of ocelli, separated by a space
equal to twice a diameter. Antenne short, shorter than the
head, segments subequal.
First tergite smooth, its anterior edge lightly emarginate
on a level with the eye, the inferior angle wide, subacute,
with a faint anterior sulcus. The rest of the somites finely
striate laterally, the strie falling far short of the pore, the
posterior portion smooth above, the anterior portion only very
finely striolate, the transverse sulcus visible only at the sides.
The pores conspicuous, situated just behind the position of
the sulcus, above the middle of the side. Scobina large,
erescentic, extending from about the tenth to about the
fortieth segment, but very small towards the end of the body.
Anal tergite smooth above and scarcely constricted, pro-
*
140 Mr. R. I. Pocock on the
duced behind into a short caudal process which scarcely covers
the tops of the valves; valves with their margins widely
compressed ; sternite large, rounded, obtuse-angled.
Legs with a single seta on the lower surface of each
segment; none of the segments padded; the coxew of the
third, fourth, and fifth legs produced, and the inferior surface
of these legs, as well as of the sixth and seventh, inflated
beneath.
Copulatory feet with the anterior median lamina wide, half-
moon-like above, bearing a median, downwardly directed,
slender, pointed linguiform process, which projects far below
the level of the anterior lateral lamina and also below that of
the posterior lateral lamina.
Number of somites 46-48.
Length 83 millim.
Two male examples from Ki Dulau, one of the islands of
the Ki or Kei group, in the Banda Sea, off the coast of New
Guinea.
Spiroholus flavo-collaris, sp. n.
(Pl. IX. fig. 11, 11 a).
Closely allied to the preceding species.
Colour dark greenish black, with the legs, antenne, labial
border, and posterior border of the somites ochraceous ; the
collum entirely bordered with ochraceous, the anterior stripe
wider than the posterior. The somites smooth, striate below,
the sulcus very feeble, obsolete above; scobina large and
extending to about the thirty-sixth segment.
The anal somite small, the hinder end of the body being
narrowed from above downwards and from side to side; the
posterior border of the tergite produced into an angle of about
90°; the valves prominent, very lightly compressed behind.
Number of somites 52.
Length about 50 millim., width 6°5.
A single example from Wokan Dobbo, Aru Island.
Spirobolus dorsalis (Le Guillou).
(Pl. IX. fig. 8.)
Iulus dorsalis, Le Guillou, Bull. Soc. Phil. Paris, 1841, p. 86; Gervais,
Ins. Apt. iy. p. 175.
Colour.—The posterior borders of the segments fusco-
ferruginous, the anterior part testaceous; a large anteriorly
narrowed black spot in the middle line of the back, the series
of spots constituting a continuous longitudinal dorsal band
from the collum to the end of the anal tergite; a fuscous
patch marking the pore; the collum widely flavous along its
Myrtopoda of the ‘Challenger’ Expedition. 141
anterior border and lateral angles; upper half of head fus-
cous, lower flavous ; antenne distally fuscous; legs flavous.
Face smooth, indistinctly sulcate. Pores 2+2. LHyes
large, separated by a space that is about equal to a diameter.
Antenne slender and longish, extending to the end of the
second segment.
Collum laterally acutely angled, the posterior border
vertical, the anterior obliquely cut away and lightly emar-
ginate, the apex not reaching the inferior border of the
second. The rest of the segments lightly punctulate and
striolate above, the longitudinal striz of the lateral portion
extending almost up to the pore and passing on to the area
in front of the transverse sulcus; this sulcus represented
dorsally by a shallow depression, sinuate opposite the pore,
which is situated in front of it above the middle of the side.
Sterna very finely striolate. Scobina absent.
Anal tergite covering the valves but not surpassing them,
roundly angled; valves lightly compressed, not marginate ;
sternite acutely triangular.
Legs \ong and slender, with a single seta on the lower edge
of each of the first four segments, and a series of three or four
on the tarsus, not padded.
Number of segments 50.
Length 51 millim., width 4.
A single male from Wokan Dobbo (Aru Island).
This species falls into the same category with Sp. Goési of
Porath, but the sculpturing is much finer and the colouring
very different.
Spirobolus hemorrhantus, sp. n.
(PL. IX. figs. 12, 12a.)
Colour (in alcohol) bright blood-red throughout.
Face smooth ; sulcus mesially obsolete. Pores2+2. Eyes
large, with upper inner border angular, separated by a space
slightly greater than a diameter. Antenne long and slender,
much longer than the face, surpassing the collum. -
Collum polished, acutely angled below, and projecting
nearly as far as the inferior portion of the second segment.
The rest of the segments smooth and polished behind dorsally,
at most lightly wrinkled longitudinally, striolate in front. No
scobina. ‘lhe lateral striz extending in front of the transverse
sulcus and reaching almost up to the pore; the transverse
sulcus obsolete or nearly so, represented by a shallow depres-
sion. ‘The pores minute above the middle of the side, in front
of the transverse sulcus ; sterna striate.
The posterior end of the body noticeably compressed; the
142 M. A.-E. Malard on the Influence of Liyht on
anal tergite scarcely covering the valves, which have their
borders prominent and deeply compressed ; sternite large and
angular.
Legs long and slender.
Number of segments 55.
Length about 70 millim., width 5°5.
Loc. Ki Dulau (Ki Islands).
This species, of which a single female specimen was
obtained, in colouring resembles Sp. sanguineus of C. Koch
(Die Myriop. i. p. 16, fig. 15). It, however, at least differs
in its long legs and antenne and prominent anal valves.
EXPLANATION OF PLATE IX.
Fig. 1. Strongylosoma Gervaisti (Lucas). Dorsal view, X 1}. In this
figure the anterior angles of the keels are too squared and the
legs are wrongly placed and wrong in number.
Fig. 1 a. Ditto. Left copulatory foot from below.
Fig. 2. Strongylosoma Guérini, Gery. Left copulatory foot from
below.
Fig. 8. Stenonia tuberosa, sp. n. Keel of the twelfth segment from
above.
Fig. 3a. Ditto. Anal segment from above.
Fig. 36. Ditto. Copulatory foot from below.
Fig. 4. Iulus solitarius, sp. n. Anal segment from the side.
ig. 5. lulus tristani, sp.n. Anal segment from the side.
Fig. 5a. Ditto. Part of segment to show position of pore.
Fig. 6. Spirostreptus Moseley, sp.n. Lateral view of head and collum, x.
Fy. 6a. Ditto. Lateral view of anal segment.
Fig. 7. Acanthiulus Murray, sp. n. Lateral view of anterior end of
body, x.
Fig. 7a, Ditto. Lateral view of two of the middle segments.
Fig.7 6. Ditto. Lateral view of posterior end of body.
Fig. 8. Spirobolus dorsalis (Le Guillou). Lateral view of head, x.
Fig. 9. Spirobolus diyrammus, sp.n. Lateral view of head and collum, x .
Fig. 9a. Ditte. Lateral view of anal segment, x.
Fig.9b. Ditto. Anterior view of copulatory feet.
Fig. 10. Spirobolus challenger, sp.n. Lateral view of head and collum, x.
Fig. 10a, Ditto, Lateral view of anal somite, x.
Fig. 106, Ditto. Third leg of male.
Fig. 10 e, Ditto. Anterior view of copulatory feet.
Fig. \1. Spirobolus flavo-collaris, sp. n. Lateral view of anal segment, x.
Fig. lla. Ditto. Anterior view of copulatory feet.
Fig. 12. Spirobolus hemorrhantus, sp. n. Lateral view of head and
eollum, X.
Fig. 12 a, Ditto. Lateral view of anal segment, x.
XVIT.— The Influence of Light on the Coloration of
Crustaceans. By A.-E. MALarp*.
M. Martin has ‘yecently drawn the attention of our Society
to a peculiar ease Of albinism which he had studied in a lobster
* Translated from the ‘ Bulletin de la Société Philomathique de Paris,’
Bieme série, t. iv. no. 1 (Paris, 1802); pp. 24-30,
the Coloration of Crustaceans. 143
observed at St.-Vaast-la-Hougue. In this lobster the absence of
pigment did not extend to the eyes, which had preserved their
ordinary colour. This phenomenon is not confined to the
lobster, but seems to be even of very frequent occurrence in
other crustaceans, such as the common edible crab (Cancer
pagurus) *, in which I have met with very numerous examples
of it in the young individuals living beneath the stones of the
old oyster-beds near the Isle of Tatihou.
This more or less complete absence of pigment seems to be
related to the nature of the habitat of these animals; the
fossorial crustaceans, such as Gebia, Aaia, and Callianassa,
are almost entirely white, and the greater portion of those
which live in grottos or crannies of rocks likewise assume
similar tints. In the lobster especially considerable diffe-
rences of hue have been observed by Mr. E. Lovett F in the
neighbourhood of the Channel Islands; he has described one
specimen in particular, which was “ of a pale lavender,” with
a mauve patch on the cephalothorax, and the chele of a bright
pale blue. This lobster, which had been sent to him alive by
M. Sinel, of Jersey, was still living when he examined it: it
was a female with eggs. Mr. Lovett also mentions other
colours in the lobster—white spotted with blue, “ dappled
blue and grey,” uniform grey, and finally a curious variety of
a pale reddish colour with the antenne ‘‘ of a decided bright
red.” It was maintained at a certain period by Bell that the
variations in colour are purely local, each race of lobster thus
having a peculiar facies, so much so that a Jersey lobster
would be distinguished at once from a French one; the fact
in itself is certainly true, but only arises from the different
conditions of medium which the animals meet with at diffe-
rent places. As is remarked by Lovett, the coloration must
be related to the depth of the water, which allows more or less
light to penetrate, and also to the number of the grottos and
natural crannies, which, as we know, are very numerous at
certain points of the coast of the Channel Islands, as, for
instance, in Sark.
I therefore think that these cases of pseudo-albinism in
lobsters, such as that of the roseate specimen of Mr. Lovett,
the greenish-white individual which has been reported to us
by M. Martin, and finally the one which M. Biétrix has told
us is under observation at Concarneau, are only normal indi-
viduals which have lost their pigment of the cyanic series as
* Cases of melanism in the same species, due to iujury to the derm,
are also met with ; the crab becomes entirely black, or “charbonné” as
the French sailors say.
T ‘Zoologist,’ ser. 3, vol. vili., 1884, p. 491; and ser. 3, vol. ix., 1885,
p. 102.
144 M.A.-E. Malard on the Influence of Light on
an actual consequence of the insufficiency of light in the
medium in which they have lived.
Far from stopping here, indeed, the influence of light on
the coloration of crustaceans is enormous, and we may say
that, as an almost general rule, the animals belonging to this
class have a tendency to vary the coloration of their bodies
according to the coloration of the surrounding medium.
Moreover this chromatic adaptation of the animal to the
ambient medium seems to be effected in two different
fashions :-—
1. By chemical means, that is to say, by the modification of
a pignient under the direct influence of light.
2. By physiological means, that is, by the action of pig-
ment-cells or chromatoblasts working under the influence of
light, but indirectly, and by the intervention of a sort of reflex
process which actually originates from the eyes of the
animal.
Chromatic adaptation is met with in many Copepods,
Amphipods, Isopods, and Macrura; it appears to be of less
common occurrence in the Brachyura, where it is often re-
placed by another form of mimicry.
As long ago as 1863 Claus mentioned the changes of colour
which are undergone by Copepods belonging to the genus
Sapphivina. According to this author, in the Copepods of
this genus the shield can assume changing colours, which the
animal varies according to the media in which it happens to
be. One species alone forms an exception to the rule and
seems to be devoid of this power of mimicry. In the greater
portion of the others the male alone is endowed with this
power *. In 1867 an analogous statement was made by Sars
with regard to Copepods of the Norwegian coasts. And,
lastly, at a more recent date the same phenomenon was
obseived by Herrick in the case of Diaptomus castor f. _
The changes of colour in Sguilla and Mysis, under the
influence of light, have been known for a very long time;
they have formed the subject of recent papers by Weber §
and Schmidlein||, and 1 merely mention them here in passing.
Among the Isopods the phenomenon is most interesting
and easiest to determine in the genus Jdotea.
In the line of foam and floating sea-weed which distin-
* Claus, ‘ Die freilebenden Copepoden,’ 1863, p. 35.
+ G. O. Sars, ‘ Histoire Naturelle des Crustacés d’EKau douce de Nor-
vege,’ 1867, p. 23.
} Herrick, ‘American Naturalist,’ vol. xvii. p. 381].
§ Weber, Archiv f. mikr. Anat. Bd. xix. pp. 591, 597.
|| Schmidlein, Mittheilungen zool, Stat. Neapel, Bd. i., 1879, p. 513.
the Coloration of Crustaceans. 145
guishes the flood-tide from the ebb, it is not unusual to find a
somewhat large quantity of a floating weed, Halidrys sili-
quosa, and nearly always there are to be found in this alga
considerable numbers of an Isopod Crustacean, Idotea marina,
which in the general shape of its body, and especially in
colour, bears a deceptive resemblance to the brownish elongate
elliptical floats of the weed, which have gained the latter its
name. ‘This property of chromatic mimicry which is
possessed by Idotea marina seems to be shared by several
other species of the genus. In the case of Idotea tricuspi-
data, a species which is very common in the vicinity of
Saint-Vaast, and especially on the tower which serves as a
beacon at the Dranguet reefs, it is not difficult to ascertain
that the individuals which live in the acorn-barnacle zone
are usually of quite a different colour from those living amidst
the mussels which cover the base of the tower, and that the
latter likewise generally differ in tint from those which are
to be met with among the Ulva.
P. Mayer has observed these changes of colour in specimens
of Idotea tricuspidata in one of the tanks at the Naples Zoolo-
gical Station *, and has found that the same animals change
colour according to the objects which surround them, and that
in the space of half an hour.
If in some way or other the animal is rendered blind, as by
extirpating the eyes for example, this curious adaptation does
not take place. Carl Matzdorfft, who has studied very
thoroughly the colour-variations of Jdotea tricuspidata, has
shown that these changes are due to chromatoblasts which
possess the power of expansion and contraction. The indi-
viduals which live upon alge or hydroids are, according to his
observations, in the majority of cases less high!y coloured than
those which run about on the bottom. Matzdorff even
succeeded in causing specimens of Jdotea to change colour by
the experiment of placing them in differently coloured glasses.
Perhaps this adaptation to the general colour of the ambient
medium is to be regarded as the cause of the coloration
observed in that singular blue pelagic species /dotea annulata ;
at any rate its commensalism with a Physalia which is like-
wise blue would explain this mimicry}. Several Lemodi-
poda, including species of Proto, Protella, and Capreila§,
* P. Mayer, Mittheilungen zool. Stat. Neapel, Bd. i. pp. 520, 521.
+ Matzdortf, “Ueber die Fiarbung von Idotea tricuspidata” (Dissert.
inaug., Jena, 1882), Jenaische Zeitschrift, Bd. xvi. p. 158.
} Spence Bate, Ann. & Mag. Nat. Hist. ser. 4, vol. i., 1868, pp. 443,
447, i xxi. fig. 1.
§ Proto pedata, Flem., Protella phasma, Latr., Caprella equilibra,
Spence Bate and Westwood; Haller, Zeitschr. f. wiss. Zool. Bd. xxxiii.
1879, p. 391.
146 = M. A.-E. Malard on the Influence of Light on
likewise mimic to a deceptive degree, as I have very often
been enabled to observe, the ramifications of the alge on
which they are found. M. E. Chevreux * has already men-
tioned the singular adaptation which is exhibited in particular
by Caprella acutifrons to Cystocira granulata, the sea-weed
which is most often green with yellow spots (as at Les
Jonchéres, near the Croisic) ; the Caprella is seen to assume
the same colour and to cover itself with yellow spots of the
same shade and arranged in the same fashion; while at other
points of the coast, where the alge are red, it assumes a
uniform brownish-red tint, identical with that of the sea-weed
upon which it lives (as at Baie d’Eslandes, near the Croisic).
For a long time it was believed that the food of the animal
was responsible for these changes in coloration ; but in a case
like the present it is very evident that, as M. Chevreux
remarks, the colour of the Caprella cannot be attributed to the
nature of its food, since it is an exclusively predaceous
animal, It may be that we ought rather to regard this simi-
litude of form and colour as indicating a sort of rational act
on the part of the animal, which seeks a favourable place to
hide itself, and actually chooses the alga on account of its
shape, not adapting itself as regards colour until afterwards.
In Hippolyte, a genus allied to Palemon but of a slightly
smaller size, the adaptation of colour seems likewise to be
related to the choice of habitat; thus Hippolyte varians,
which owes its name to these changes of coloration, will be
green in Zostera, brown in Fucus, red in Floridea, and trans-
parent or almost transparent when it is found amidst Antennu-
laria and Sertularia.
Now what is the cause of these changes of colour ?
It seems to be the same as that of the coloration of the
alge. Asa matter of fact experiments which have been made
upon this subject have proved that the same specimen of
Hippolyte assumes a red tint in complete darkness, while in
bright light it becomes a vivid emerald-green colour, and
semi-obscurity renders it brown. ‘These experiments, which
I have myself repeated, would appear decisive. I am bound,
however, to state that I have met with a case which strangely
complicates the question.
Thanks to the courtesy of the Board of Bridges and High-
ways (Buoying Service of the Arrondissement of Cherbourg),
and especially to the extreme kindness of M. Rouland, who is
er Chevreux, ‘Les Plages du Croisic,’ par Adrien Dolfuss, pp. 9
an .
+ This remark had already been made in the case of Idotea by Mobius,
Bericht Exped. ‘ Pommerania,’ p. 121.
the Coloration of Crustaceans. 147
especially intrusted with this service, [ have been enabled for
several years past to be present at the picking up of the buoys
included in the district of La Hougue: upon one of these it
was my fortune to observe an immense quantity of Comatula
(Antedon rosacea), with which the chain of the buoy was
literally covered.
These specimens of Comatula were of three very distinct
colours—more or less deep violaceous red, orange-yellow
inclining towards saturn-red, and, lastly, alternately white and
red with whitish pinnules. Now I was not a little surprised
at observing along the chain of the buoy specimens of /Hippo-
lyte apparently living side by side with the feather-stars, which
they in many cases clasped with their limbs, and agreeing, at
least in the majority of instances, so closely with their neigh-
bour in colour that it became difficult to perceive them.
The fact, strange as it is, isnot unique. Lucien Joliet has
recorded a similar faculty in a Mediterranean Pontonia living
as a commensal with Diazona; this Pontonia, which is allied
to P. tyrrhena and which Joliet has described as a new species
under the name P. diazone*, also bears a deceptive resem-
blance to the Diazona ; the transparency of its body blends
with the hyaline jelly ef the colony, and the yellow spots with
which its thorax, abdomen, and chelz are marked harmonize
so perfectly with those of the Ascidian itself, that it becomes
impossible to perceive its presence so long as it remains upon
its host.
Specimens of Palemon also exhibit variations in colour
according to the nature of the bottom on which they are
found, becoming green when the bottom is covered with
Zostera and grey or reddish yellow when the bottom is of
sand.
Some years ago M. Georges Pouchet made some very
interesting observations upon this subject ft. Taking some
earthenware vessels coloured black and white inside, he placed
in them for the purpose of observation some specimens of
Lalemon of medium size (3 to 4 centim. in length), which
experience had taught him to be most readily subject to
variations of colour. These prawns, which on leaving the
fishermen’s nets are usually of a roseate or faint lilac tint,
become colourless, or at the most faintly yellowish, in the
vessels with a white bottom ; while in the black vessels they
become, on the contrary, dark brown.
* L. Joliet, “ Observations sur quelques Crustacés de la Mediterranée,”
Arch. Zool. expér. t. x. p. 118.
+ G. Pouchet, ‘Journal d’Anatomie et de Physiclogie,’ 1872, t. iy.
pp. 401 -407; C. Rt. Acad. Se. Paris, 20 mai, 1872,
148 Influence of Light*on the Coloration of Crustaceans.
The change of colour from the pale to the dark condition
occupies but a few minutes ; it is different with regard to the
opposite process. During the disappearance of the deep
reddish-brown tint to become pale yellow and almost trans-
arent the observer noticed that the animal passed through an
intense deep blue stage.
M. Pouchet has furnished the explanation of these pheno-
mena* by showing how the action of two kinds of pigments
took place in them; on the one hand the pigments of the
xanthic series (red, orange, and yellow) by the action of the
chromatoblasts, on the other the pigments of the cyanic series
generally free and in solution. The removal of the eyes
produces in Palemon the same effect as a black bottom f.
In certain Brachyurous Crustaceans we again meet with
facts of the same kind; but here they are less general and
not so numerous. Fritz Miller t mentions an instance in a
Brazilian species of Gelasimus which is of a uniform greyish-
brown colour in the female. In the male of this Gelasimus at
the breeding-season the posterior portion of the cephalothorax
is of a pure white, while the anterior region assumes a rich
green colour, passing into dark brown ; in the event of danger
and on the animal being alarmed its colours are subject to
modification in a few minutes, the white becoming dirty grey
or even black and the green losing all its brilliancy. Not
until we come to Carcinus menas do we find that similar facts
have been reported; I have myself often observed that the
crabs living upon a bottom clothed with Ulva, as at the mouth
of the Serre, near St.-Vaast, for example, when angry have a
more decidedly green tint on the dorsum of the cephalothorax
than those which are met with among the stony bottoms of
the old oyster-beds and in the Laminaria-zone, where they
assume olivaceous hues, passing into dirty yellow and brownisa
red, a shade which is in perfect agreement with the general
tone of the bottom. Messrs. Carrington and Lovett §, in
recording analogous observations, state that they have been
able to study the mechanism of the phenomena in the tanks
of the Westminster aquarium; here again the facts observed
are probably due to the action of chromatoblasts.
I do not know whether the chromatoblasts also play a
similar part in the Mediterranean Lambrus, the curious
* C. R. 1878, t. Ixxxvii. pp. 802-303.
+ M.S. Jourdain has since shown that by removing the eyes and
leaving the animal in the dark a red coloration is always obtained.
} According to Darwin, ‘The Descent of Man,’ French edition, p. 361
[2nd English ed., 1883, p. 271.
~ § Carrington and Lovett, ‘ Zoologist,’ 1832, pp. 12 and 14,
Mr. J. E. 8. Moore— Observations upon Amceba. 149
mimicry of which has been described by M. de Varigny in
the ‘Revue Scientifique’ *; it appears to me that this is
probably the case, as also in Portumnus varitegatus, which
disappears almost completely upon coarse granitic sand, as [
have myself been able to observe. Be that as it may, we see
from these examples that concealment by isochromatic adap-
tation seems to be a very widely spread fact in the different
orders of Crustacea, and that albinism in these animals
appears to be only a particular case of a very much more
general phenomenon of chromatic adaptation to the medium.
XVIII.— Observations upon Ameeba, with especial reference to
the existence of an apparent Micro-nucleus in that Organism.
(Preliminary Communication.) By Joun E. S. Moore,
A.R.C.S. (from the Huxley Research Laboratory, R. Coll.
Sci. Lond.).
[Plate XII. }
THROUGH the laborious investigations of Maupas, Biitschli,
Hertwig, and others we are to-day pretty well acquainted with
the minute structural peculiarities and life-history of the
ciliate Protozoa. Regarded in the light of single cells these
little beings present points of structure at once both strikingly
different and similar to those apparent in the cells that build
up the Metazoan tissues.
The karyokinetic division of the micro-nucleus (‘ Neben-
kern,” “ endoplastule,” ‘‘nucleolus”’) in the ciliate Infusoria
is undoubtedly strictly comparable, step by step, with the
similar process apparent in the cells of higher forms; but the
coexistence of this structure itself with the macro-nucleus,
which divides akinetically, is something totally unlike those
conditions which ordinarily present themselves in Metazoan
cells.
Our knowledge of the multitude of structures included under
the somewhat comprehensive title of ‘ Nebenkern” in the
Metazoa is still in a sufficiently unedifying condition to render
it impossible to say whether those remarkable bodies met with
in the gland-cells of many animals, ¢. e. in the cells actively
secreting, are normal or parasitic, or whether in such cells we
may not have to deal with a third structure besides the
* de Varigny, ‘ Revue Scientifique,’ 3° série, t. x. p. 92, 2* sem., 1885,
150 Mr. J. &. S. Moore— Observations upon Amooba.
nucleus and attraction sphere, which may well retain the
name of ‘* Nebenkern ”*,
In May last I succeeded in adducing evidence in support of
the view, first initiated by Flemming I believe, that the
karyokinetic division of a cell is in reality an expression of
two metamorphoses, each to a certain extent independent of
the other f: one affects the attraction-sphere and the dividing
central bodies, the other the nucleus and the dividing chromo-
somes. And it is probable that this apparent duality in the
metamorphosis may have had much to do with the modern
conception that the protozoan micro-nucleus is more or less
equivalent to the metazoan central body, and the metazoan
chromatic element to the protozoan macro-nucleus f.
1 am not of this opinion, for I fail to see how a structure
presenting all the features of a karyokinetic division, such as
is seen in the micro-nucleus of a Paramecium or a Colpidium,
can be compared with the central body of a cell, metazoan or
other.
That the great macro-nucleus, where it exists in the Infu-
soria, 1s intimately related in some way or other to the
digestive activity of the animal seems highly probable.
Indeed it is difficult to see what other functions could be
assigned to it; and the Ciliate as a single cell will have to
peiform all those secretive operations which in more highly
specialized organisms would be told off to particular glandular
organs §.
‘Thus it becomes possible that the existence of the “ Neben-
kern”’ in many gland-cells of the Metazoa (if that structure
can be shown to be there normal) may offer an analogy to
that differentiation of the nuclear elements apparent in the
ciliate Infusoria. Whatever be the ultimate solution of
* See “ Contributions to Morphol. and Physiol, of the Cell,” Macallum,
Trans. Canadian Institute, vol. i. pt. 2; Steinhaus, ‘ Ueber parasitiire
Einschliisse in den Pancreaszellen der Amphibien,” Ziegler’s Beitrige
zur Path. Anat. und zur Allgem. Path. Bd. vii. p. 367; Nussbaum,
“ Ueber den Bau und die Thatigkeit der Driisen,” Arch. fiir mikr. Anat.
Bd. xxi. p. 296.
+ Unpublished paper in the hands of the editor Quart. Journ, Mier.
Sci.
{ Cf.II. E. Ziegler, ‘ Biologisches Centralblatt,’ Bd. xi. nos. 12 and 13,
p- 372-889 ; and Biitschli, “ Ueber die sog. Centralkorper der Zelle und
ihre Bedeutung,” Verhandl, d. Naturhist.-med. Vereins zu Heidelberg,
Bd. iv.
§ This view receives some support from a consideration of the multi-
nucleate parasite Opalina. Here, where there is no need for digestive
activity, a8 the animal is immersed in already digested food-material, we
find a corresponding simplitication of the nuclear elements, all of which
present the micro-nuclear karyokinetically dividing structure.
Mr. J. E. S. Moore— Observations upon Amoeba. 151
these problems, it would be interesting to find anything com-
parable to such nuclear differentiation in the other Protozoa.
Turning to the Rhizopods, we find a great variety of
nuclear conditions; for example, those described by Gruber
in a compendious summary published in the ‘ Zeitschrift
fiir wissenschaftliche Zoologie’ *, and, again, in his ‘ Studien
iiber Amében’ +, where there is an admirable description of
the mono- and multinucleate forms of Amcebe, of the great
diffused nucleus of A. proteus, and of the numerous vesiculate
nuclei of Pelomyxa.
The most interesting thing, however, about these nuclei
seems to be the fact that whereas the multinuclei of Pelomyca
divide by a process equivalent to karyokinesis, the great
diffused nucleus of A. proteus, according to the beautiful
figures of Schulze and others, divides akinetically in the
strictest sense of the term. So also the primary division of
the nuclear element in Arcella, when nearing the spore-
forming stage, is typically karyokinetic; but whether this
process of multiplication is maintained is not apparent.
The numerous nuclear elements in the Heliozoa appear,
according to Gruber, to increase in a mitotic fashion, while
the duplication of the nuclei in Huglypha alveolata is repre-
sented as proceeding by more than one method.
In an interesting paper in the ‘ Zeitschritt fiir wissenschaft-
liche Zoologie’ { Verworn describes in Digfugia lobostoma the
conjugation process and the existence of corpuscles answer-
ing to the micro-nuclei of the Ciliata. Of these structures he
says :—‘‘ Neben den Individuen, welche die obenbeschrie-
benen normalen Kernverhialtnisse zeigten, fanden sich seir
hiiufig auch einzelne lebende Exemplare mit ganz abweich-
endem Verhalten. Diese Individuen besassen zwar auch den
grossen runden, blassen Kern, mit ganz normalem Ausselhen
. . . ausser ihm noch einen zweiten, der eine vollig verschie-
dene Beschaffenheit aufwies ;”’ and further, “ Ofter fand ich
Konjugationen, in denen, ausser dem normalen grossen Kern,
keine kleine Kerne vorhanden waren.” He sums up thus:
“Dass der Kern eine bedeutsame Rolle bei der Konjugation
spielt, indem die Konjugation charakterisirt ist durch das Auf-
treten je eines kleinen eigenthiimlich gestalteten Kerns neben
dem gewodhnlichen, der méglicherweise dem Nebenkern der
Ciliata Infusorien entspricht, und ferner dadurch, dass diese
kleine Kerne der beiden Individuen wiihrend der Konju-
gation in nahe Bezichung zu einander treten.”’
* Bd. xl. 1884, pp. 121-152.
+ “Eine Mittheilung tiber Kernvermehrung und Schwarmerbildung
bei Siisswasser-Rhizopoden,” tdid. Bd. liii. 1592, pp. 114-118.
t Ba. 1. 1891, p. 443, Taf. 18,
152. Mr. J. E. 8S. Moore— Observations upon Ameeba.
From these observations it becomes probable that we shall
have to deal, as Verworn himself says, with such bodies related
to conjugation in other Rhizopods. At the same time it is
orobable that the further study of the apparent discrepancies
in the nuclear division of these animals may hereafter lead up
to very interesting conclusions, with respect to a possible
periodicity in the occurrence of direct and indirect division in
such forms of life.
Mr. Lister has kindly sent me, through Prof. Howes, the
MS. of a paper read before the Linnean Society of London*, in
which he demonstrates the very important fact that in the
Mycetozoa the numerous nuclei in the streaming plasmodia
divide directly till just before the formation of the spores,
when the division passes into typical karyokinesis.
Of the life-history of the Rhizopods we know in reality
very little, and it seems to me highly probable that much of
the apparent confusion with respect to their direct and indirect
division may be due to a possible periodicity in their recur-
rence.
During the examination of some Amcebe last autumn I
became struck with the regularity in appearance and persis-
tence of a small granular body, usually near the nuclear
element, and in its general relationships very like a micro-
corpuscle during certain phases of its evolution (Pl. AIL.
fig. 1).
~ Although loath to enter into a description of a new structure
where we have already descriptions and structures enough to
last for half a century, I would state that this body appeared
to those who saw it a definite entity, and that it showed a
remarkable tendency, when displaced in the living animal by
the pressure of the cover-glass or other means, to return
eventually to its position beside the nucleus.
It did not stain with picro-carmine, but remained perfectly
visible after fixing with either gold or platinum chloride
(fig. 1) ; and it could be stained slightly with orange, as was
the case with Verworn’s corpuscle in Difflugia. Flemming’s
fluid rendered it nearly invisible, on account of the induced
contraction ; while no better results were obtained with either
mercuric chloride or acetic acid. Indeed, it was much best
seen when the animal was in full activity (figs. 2, 3).
Of both the significance and metamorphosis of this body
I am absolutely ignorant ; and my knowledge is summed up
in the assertion that there is a small body of fairly constant
appearance near the nuclei of some Amcebx, and (as those
* Meeting held December lst. Cf. reports in scientific journals.
Mr. J. E. S. Moore— Observations upon Ameeba. 153
originally observed were of large size, and as I have not found
it in smaller broods since examined) that it possibly only
occurs in certain conditions of activity of the same *.
Respecting the other bodies and granules apparent in
Amcebe I have a few remarks to offer.
Exclusive of matter obviously indigestible, which is rapidly
extruded, it appears that the remaining granules may be
grouped within three categories: one, containing all those
fragments of undigested matter, some of whose original
characters remain, and two others of a totally different appear-
ance, viz. a second, containing all those spherical homogeneous
globules, usually supposed to be fat (but, as the vast majority
of them never blacken with osmic acid, this can hardly be the
case), and a third, including the so-called crystalline bodies
(concretions of Ray Lankester) t.
The origin and significance of the two latter remain still
somewhat enigmatical. Considerable light, however, may
be thrown on the inter-relations of the bodies in question if an
active Amceba be ruptured under the weight of the cover-
glass (which, if the animal be surrounded by clear water,
usually occurs in from ten to fifteen minutes). If the issuing
matter as well as that remaining, now relieved of pressure, be
watched under a high power, not only are the more conspicuous
masses of little-digested material seen to be surrounded
with the usual vacuoles of ingestion, but a vast number of
other vacuoles make their appearance, of all sizes, down to one
just sufficient to include the smallest apparent concretions
(fig. 9). Both the spherical globules and the so-called
crystalline bodies are seen to occupy small vacuoles and to
issue with them into the surrounding water.
During the later stages of digestion many fragments of
nutritious matter bear a closer and closer resemblance to these
globules, and it is possible to arrange complete series extend-
ing (i.) from the undigested food on the one hand to the
homogeneous spheroids on the other {figs. 4-8), and (ii.) from
these to the so-called crystalline bodies, which are not doubly
refractive (fig. 10). As Miss Greenwood has pointed out f,
the vacuoles about the food-material decrease up to a certain
point; and they are, as she thought possible, secondarily
acquired.
From these observations it seems that the food, after what
* It is probable that a body figured by Leidy in his monograph on the
Rhizopods (plate v. fig. 8) is identical with that herein referred to.
Tt Quart. : a Micr. Sci. vol. xix. p. 484.
} ‘Journal of Physiology,’ vol. viii. pp. 264, 283.
Ann. & Mag. N. Hist. Ser. 6. Vol. xi. 11
154 Mr. ©. J. Gahan on the Sensory Nature of the
we may call the primary process of digestion, enters into the
condition of the spherical globules, each surrounded by its
vacuole; and that these spheroids, gradually decreasing in
size, are ultimately worked up into the so-called crystals *.
T’o the further consideration of this and other questions
raised I intend to return at length.
EXPLANATION OF PLATE XIL
Fig. 1. Amceba after treatment with gold chloride. a, refractive body
(? micro-nucleus).
Fig. 2. Living Ameeba. a, refractive body.
Fig. 3. Nucleus of living Amceba, showing its relation to the refractive
body (a) when at rest.
Figs. 4-6. Digesting matter in successive stages.
Figs. 7, 8. Homogeneous spheroids contained in vacuoles.
Fig. 9. Concretionary matters in relation to the small vacuoles.
Fig. 10. Relation between the spheroids and the so-called crystalline
bodies,
Figs. 1] and 2 drawn under Zeiss’s apochromatic system, oc. 8 compens.
obj. 1-4 homog. immers. Figs, 3-10 drawn under same objective, with
substitution of oc. 18.
XIX.—On the probable Sensory Nature of the “Appendix”?
of the Antenne of Coleopterous Larve. By CHARLES J.
Gauan, M.A., of the British Museum (Natural History).
Many Coleopterous larve are provided with a remarkable
structure which is situated upon the distal surface of the
penultimate segment of the antenne. Though this structure
has been noticed by more than one writer on Coleopterous
larvee and has been described as an ‘‘ appendix,” an ‘‘ appen-
dicular joint,” a ‘ blunt tubercle,” and in other terms, it does
not seem to have attracted much attention. At least, no
author, so far as I am aware, has attempted to describe its
microscopical characters in detail.
Some observations that I have recently made upon the
antenne of the larva of Pterostichus—a genus of Carabidee—
have led me to believe that the so-called appendix is in reality
a eae | organ. When the antenne of this larva are
examined under the microscope the appendix is seen as a
tolerably conspicuous object projecting from the oblique outer
(or posterior) surface of the distal extremity of the third
segment, its transverse diameter being very little less than
* Le Dantec has shown the vacuolar fluid of several Protozoa to be
acid (‘ Annales de l'Institut Pasteur,’ 1890, pp. 776-791).
“Appendix” of the Antenne of Coleopterous Larve. 155
that of the fourth or terminal segment. It consists of a short
chitinous or semichitinous collar or stalk, supporting a cap
composed of a thin transparent cuticular membrane, which
appears to be of the same thickness throughout and to be
lined by very small cells. The cap is in the form of a short
cone with curved sides, and is strengthened at the base where
it joins the collar by a narrow and thickened chitinous ring.
Lying within the laterally expanded distal portion of the
third segment, at a short distance from the base of the collar,
I was able to recognize what seemed to me to be a ganglionic
swelling of the antennary nerve, containing a number of
nerve-cells from which fibres or rods were seen to extend into
the collar. These fibres did not seem to pass beyond the
chitinous ring, and were only visible when focusing between
the upper and lower portions of the ring. In view of the
probable auditory nature of the organ I tried to detect the
presence of the characteristic auditory rods which are found
to be associated with the auditory apparatus in other insects.
My failure in this endeavour was perhaps due to the want of
a sufficiently high microscopic power. I am inclined, never-
theless, to suspect that within the collar, and just below the
ring, rods of this kind may be found. I was also unable to
satisfy myself as to the nature of the contents of the cap,
though it seemed quite evident that it contained no otoliths.
Whether the ring supports a transverse partition is another
point I could not satisfactorily determine, though the ap-
pearances were rather in favour of believing that it does.
These, however, are points that will have to be settled by
further investigation. ‘The presence of a nerve-ganglion with
fibres passing out to the organ will also, I admit, need con-
firmation. In the few preparations I was able to make
nothing more was attempted than to clear the integument
slightly before finally mounting the antenne in balsam. [
regret that, owing to lack of sufficient material, I could not
proceed to examine the structure of the organ more thoroughly
by means of staining and sectioning. ‘This I hope to be able
to do later on.
Should the organ eventually be definitely proved to possess
a sensory function little doubt would, I think, arise as to its
auditory character. From its position and the way in which
it is guarded by some long stiff sete it would be more or less
prevented from coming in contact with external bodies; so
that it would be almost impossible for it to function as a tactile
process ; while, on the other hand, its general structure seems
to preclude the idea of its being an olfactory organ. There
are, moreover, on the same distal surface of the er segment,
1
156 Mr. B. B. Woodward on the
as well as at the apex of the antenna, a few very much
smaller hair-like structures somewhat similar to those to
which an olfactory function has been usually ascribed. One
of these, in close proximity to the organ in question, and two
at the apex have a shape somewhat like that of a hand-bell.
The remaining two at the apex are more or less cylindrical,
with rather blunt extremities. Another on the third segment
is shorter and broader and seems to have the form of a trun-
cated cone.
It may be mentioned, in conclusion, that the presence of
the ‘‘appendix”’ is not confined to the larve of a few
genera of beetles. According to the observations of Schiédte
and of Chapuis and Candéze it seems to be characteristic
of the larve of Carabidex, of Chrysomelids, and of certain
other large families of Coleoptera, and that, though usually
placed towards the outer or posterior side of the distal surface
of the penultimate segment, it is, in the Staphylinide, situated
on the anterior side.
The only other species in which I have had an opportu-
nity of examining it was one of Telephorus, in which it had
the same relation and pretty much the same general structure
asin Pterostichus, the chief difference being that the chitinous
ring, instead of having a uniform width throughout, is narrow
at one side and extends upwards, to reach its greatest width
on the opposite side. It was first noticed in this larva by the
late G. R. Waterhouse, who referred to it as a “‘ process.”
Considering, therefore, the widespread existence of this
appendix amongst Coleopterous larvee, and of its very definite
localization on the penultimate segment of the antennae,
together with its rather peculiar structure, it seemed to me
that some elucidation of its precise nature and meaning was
desirable. And even should it ultimately be proved that I
have been mistaken in my first observations upon it, there
will be some justification for having directed attention to a
structure which seems to have been hitherto almost entirely
neglected.
XX.— Classification of the Pelecypoda: Fischer's Families
rearranged tn accordance with Pelseneer’s Scheme. By B. B.
Woopwarp, F.G.S., F.R.M.S., of the British Museum
(Natural History).
THE classification of the Pelecypoda has been from the first a
matter of great difficulty owing to their comparatively uniform
structure.
The characters hitherto employed have been more or less
Classification of the Pelecypoda. 157
superficial in their nature. Thus the muscular impressions
were for a long time solely relied upon; but an arrangement
based on the existence of only one or of two adductor
muscles, and, in the latter instance, on their relative size,
proved unsatisfactory. Nor did the change in favour of the
presence or absence of siphons and of a pallial sinus yield any
better result ; whilst classification based on the form of the
foot is now no longer even thought of.
Neumayr, regarding the matter from a_ paleontological
point of view, suggested a system based on the hinge cha-
racters *, which, however, does not satisfy the student of
living forms.
Fischer, going deeper, made use of the respiratory organs f,
but, misled by their external appearance, divided the class
into two-gilled and four-gilled groups, ruthlessly sundering
closely related families in the process.
Pelseneer it was who first, in 1889 f, pointed out the true
reading of the Pelecypod gill-structure and founded on the
morphology of that organ a classification which, thouzh
doubtless not final in minor details, promises to be by far the
best of any.
It is extremely interesting to note that Pelseneer fore-
stalled by but a very short time an independent worker,
Ménégaux, who had arrived at almost precisely similar con-
clusions §.
Yet another classification has recently been proposed by
Prof. C. Grobben ||. In this the external form of the gill is
taken as a primary character, whilst the hinge is relied upon
as the secondary basis of the arrangement. The result,
* “ Zur Morphologie des Bivalvenschlosses,” Sitzb. k. Akad. Wissensch.
Wien, Bd. lxxxviii. Abth. i. (1884), pp. 885-419. ‘The amplification of
which, so far as completed at the time of his death, appeared in the
Denkschr. vol. lviii. (1891), pp. 701-801.
+ ‘ Manuel de Conchyliologie’ (1886-87).
¢ “ Classification phylogénétique des Pélécypodes (communication
préliminaire),” Bull. Scient. France et Belgique, tom. xx. (1889), pp. 27-
52; the complete memoir was published as “ Contribution 4 l’étude des
Lamellibranches,” Arch. Biol. (Liége), xi. (1891), pp. 147-312, 18 pls.
§ In fact his first note, “Sur la Branchie des Lamellibranches et sur
la comparison avec celle des Scutibranches,” Bull. Soc. Philom. sér, viii,
tom. i. (1889), pe. 137-144, was elicited by Pelseneer’s “ preliminar
communication,” the full text having been sent in asa Thesis, whic
was afterwards published under the title ‘ Recherches sur la circulation
des Lamellibranches Marins’ (4to, Besangon, 1890).
|| “ Beitrage zur Kenntniss des Baues von Cuspidaria.... nebst Be-
trachtungen uber das System der Lamellibranchiaten,” Arb. Zool. Inst.
Univ. Wien, x.; the latter section also appeared in Zool. Anz. xv.
(1892), pp. 371-375, whilst diverse translations are given in Journ. R.
Mier. Soc. 1892, p. 772, and Nat. Sci. ii. p. 9.
158 On the Classification of the Pelecypoda.
however, as set forth by the author will hardly commend it
to practical malacologists.
Pelseneer’s scheme appears so decidedly in advance of the
others that after consultation with my friend and colleague
Mr. E. A. Smith it has been decided to adopt it m future
volumes of the ‘ Zoological Record,’ retaining Fischer's
families as far as possible for the present for the sake of
eonvenienee in reference.
Since, however, the ‘ Zoologieal Record’ does not admit of
the new arrangement being shown in its entirety, it seems
fairer that it should im the first instanee be published where
it will be available to all in a handy form.
Hitherto only the relationship of the main groups has been
published, and that in a “tree” which should be studied in
conjunction with the linear arrangement that here follows ;
for in the latter of course the essential features of the “‘ tree”
eannot be reproduced, though some help is sought to be given
by “indenting” those families which lie on the divergent
branches and placing them between “ rules,”’ after the method
familiar in a railway time-table.
The three families not in Fischer are printed in italics.
Class PELECYPODA. Subord. iii. Myacra.
Order I. SEPTIBRANCHIATA. | Fam. 17. Solenide.
18. Glycymeride.
Fam. l. Poromyide (Poromya and 19. Myide.
Silenia). 20. Lutrariide.
2. Cuspidaride.
Order I. EULAMELLIBRAN- Subord. iv. CaRDIACEA.
ee Fam. 21. Radiolitide.
Subord. i, ANATINACEA. 22. Hippuritidz.
Zs. asa ee 23, Caprinide.
Fam. 3. Clavigellide. 24. Monopleuride.
4, Pholadomyide. 25. Chamide.
5. Preecardiide. 26. 'Tridacnide.
6. Grammysiide. ist! 27. Lunulicardiidee.
7. Anatinidie (excepting 28. Cardiide.
Poromya).
8. Arcomyide,
9. Ceromyide. 2. .
10. Livondiidss (exeepting | Subord. v. VENERACEA.
Silenia). _ Fam. 29. Psammobiide.
11. Verticordiide. 30. Cyrenellide.
12, Chamostreide. | 51. Rangiide.
15. Pandoride. 32. Cyrenide.
35. Glaucomyide.
Subord. ii. PHOLADACEA, 34. Petricolide.
* a: 35, Veneride.
Kom. = a 36. Pachydomide.
15. Pholadide. 37. Cyornid
16, Gastrocheenide. i. prinide.
38. Megalodontida.
Mr. H. O. Forbes on Apteryx Haasti. 159
Subord. vi. TELLINACEA. Order III. PSEUDOLA-
MELLIBRANCHIATA.
Fam. 39. Serobiculariide. ; :
40. Tellinidee. Fam. 60. Ostreide.
41. Donacide.
42. Cardiliide. 61. Aviculide.
43. Mesodesmatide. 62. Prasinide.
44. Mactridie (excepting | Ee EWS i Bee
Lutraria). 63. Pectinidee.
64, Limide.
65, Spondylide.
66. Dimyidee.
Subord. vii. SUBMyTILACEA. |
Fam. 45. Galeommide. | Order IV. FILIBRANCHIATA.
46. Erycinide (excepting | Fam, 67, Mytilide (excepting
Montacuta). Dreissensia and Dreis-
47, Kellyellide. sensiomya).
48. Crassatellide., |
49, Tancrediide.
50. Unicardiide.
51. Ungulinide.
52. Lucinidée (including |
Montacuta).
53. Astartidee.
54. Carditide.
55. Cardiniide. | Order V. PROTOBRANCHIATA.
56. AEtheriide.
57. Unionide. Fam. 71. Solenomyide.
58. Dreissensiide.
59. Modiolopside. 72. Nuculide.
68. Trigoniide.
69. Arcide.
70. Anomiidee.
XXI.—Notes on Apteryx Haasti.
To the Editors of the ‘Annals and Magazine of
Natural History.’
GENTLEMEN,—In the last number of the ‘ Annals,’ page 43,
I observe an article on Apteryx Haasti, Potts, by the Hon.
Walter Rothschild. “ Mr. Forbes,” it affirms, “has ex-
pressed it as his opinion that Apteryr Haastii is a natural
hybrid between A. australis and A. Owenii. 1 have paid
much attention to this question as well as to the study of this
genus generally,” &e.
Iam not sure whether or not I am the “ Mr. Forbes”
referred to; but as I am led to believe that I am, I hope you
will grant me the privilege of making a few observations on
Mr. Rothschild’s note. I am surprised, in the first place, to
have opinions that I privately entertain publicly criticized; for,
so far as I can charge my memory, I am not aware of having
160 Mr. Hl. O. Forbes on Apteryx Haast.
ever committed to print my opinions upon the species of
Apteryx. 1 should be glad, therefore, if Mr. Rothschild will
be so kind as to give mea reference to the publication in
which I have made the statement that Apterya Haasti “is a
natural hybrid of A. australis and A. Owenti.” Having had
charge, however, of the type specimens of A. Haasti for some
years, I naturally made a somewhat careful examination of
them; and the opinion I formed then I still retain, that
A. Haasti is most probably a natural hybrid between the
two above-named species. This, however, is, I believe, the
first occasionon which I have expressed ¢n writing this opinion.
The type specimens of this bird in the Canterbury Museum
were obtained on the west coast of the South Island, and
from the region in which A. australis and A. Owent both
abound. I am therefore again surprised to read Mr. Roth-
schild’s affirmation that Apteryx Haast? is known only from
the North Island, as on page 330 of his ‘ Birds of New Zealand’
(2nd ed. vol. ii.) Sir Walter Buller has the following remark
in reference to these very specimens :—“‘ These [specimens of
A. Haasti] were obtained on the high ranges above Okarita,
on the west coast of the South Island, where, according to
the resident natives, the large Grey Kiwi is_tolerably
common.” Mr. Rothschild bases his belief in the distinctness
of A. Haasti as a good and pure species on its geographical
distribution, on the size of the bird and the length of its beak,
and on variations in itscecum. As I have shown, the infer-
ence from its distribution signally fails. It is well known,
too, that the various species of Apteryx vary greatly in size.
In the Canterbury Museum, N. Z., there is a very large
specimen of A. australis, nearly equalling the Apteryx
maxima belonging to Mr. Rothschild, as I remarked to one
or two of my friends at the recent meeting of the Zoological
Society at which it was exhibited. Before leaving New
Zealand J had also, through the kindness of Mr. A. Hamil-
ton, of the Otago University, Dunedin, an opportunity of
examining a large collection of Aptery« crania (both australis
and Owent) discovered in one of the limestone caves of
Otago, and was particularly struck by the great variability in
length and strength of the beaks, some being remarkably long.
I hope Mr. Hamilton will ere long favour ornithologists with
a description of these remains. In regard to the variations
observable in the caecum, does not their divergence in the two
sexes of A. Haasti afford a presumption in favour of hybridity ?
If it were a true species, should we not look for a close agree~
ment between the internal anatomy of the male and female ?
Further, Mr. Rothschild states that the native name for
On Indian Deep-sea Dredying. 161
’
Apteryx Haasti is ‘* Roa-Roa” and it is not known by the
name of “ Kiwi.” The name ‘ Roa-Roa,” however, is
applied to several species of Apteryx, and specially to Apteryx
maxima of Stewart Island. I would, however, again refer
Mr. Rothschild to Sir Walter Buller’s second volume, p. 330,
where he will find that the native names for the South-Island
types in Christchurch, N. Z., are both ‘f Roa” and “ Roa-
Roa ” as well as “ Kiwi Karuai.”
To the best of my knowledge Apteryx Haast? has hitherto
been found on/y in the South Island, and in that district alone
in which Apteryx australis and A. Owent both occur, which
so far seemed very significant. Its occurrence in the North
Island is to me a new and most interesting fact, and, if sub-
stantiated, would certainly go far to upsetting the opinion
I at present hold that Apteryx Haasti is a hybrid. Is
Mr. Rothschild quite certain of the localities of his specimens
and of the accuracy and bona fides of his collector? I shall
look forward with much interest to learn the actual localities
whence have come his specimens, also if the young specimens
of A. Haasti which Mr. Rothschild has had alive at Tring
were reared by him, or if he has had eggs from a pair of
these birds when confined by themselves. Does Apteryx
Hlaasti occur in the localities in which A. Bulleri (=A. Man-
tell’) is found? Or can Mr. Rothschild himself have mistaken
Jarge west-coast specimens of A. Oweni tor A. Haasti?
Henry O. ForsBEs.
1 Philbeach Gardens,
Jan. 6, 1893.
XXII.—WNatural History Notes from H.M. Indian Marine
Survey Steamer ‘ Investigator, Commander Rh. F. Hoskyn,
LR.N., commanding.—Series L., No. 1. On the Results of
Deep-sea Dredging during the Season 1890-91. By J.
Woop-Mason, Superintendent of the Indian Museum, and
Professor of Comparative Anatomy in the Medical College
of Bengal, and A. Atcocx, M.B., Surgeon I.M.S., Sur-
geon-Naturalist to the Survey.
[Continued from vol. ix. p. 370. ]
[Plates X. & XI]
Family Pasiphaida.
PASIPHAE, Savigny.
52. Pasiphaé sivado (Risso).
Pasiphaé sivado, Savignyi, et brevirostris, Milne-Edwards, Hist. nat.
des Crust. vol. ii, 1857, p. 426, et Atlas du Régne Anim. de Cuvier,
162 Messrs. J. Wood-Mason and A. Alcock on
Crust, pl. xxii. fig. 8; Bell, British Stalk-eyed Crust. 1853, p. 312,
woodcut; Heller, Crust. Siidl. Europ. 1868, p. 243, pl. viii. figs. 4-6;
Wood-Mason, Ill. Zool. ‘ Investigator,’ pt. i., Crust. pl. iii. fig. 6,3*.
Two males were taken on November 29th, 1888, north of
Port Blair, Andaman Sea, in 200 fathoms.
They differ from a Mediterranean specimen of the female
— RAY A
TAS \ \\ NS
Fasiphaé sivado (Risso), ¢ —Basal portion of endopodite of second abdo-
minal limb of the right side from in front. en, endopodite; at,
appendix interna ; am, appendix masculina; the spines of the hinder
row are shaded to distinguish them more clearly from those of the
front row. Magnified.
* ‘Tllustrations of the Zoology of H.M. Indian Marine Surveying
Steamer ‘ Investigator,’ under the command of Commander A. Carpenter,
R.N., D.S.O., and of Commander R. F. Hoskyn, R.N.’—Part I., Fishes,
Plates I.-VIL., under the direction of A. Alcock ; Crustaceans, Plates I.-
V., under the direction of J. Wood-Mason. Published under the autho-
rity of the Director of the Royal Indian Marine. Calcutta: printed and
sold by the Superintendent of Government Printing. 1892.
Indian Deep-sea Dredging. 163
in the collection of the Indian Museum in their slenderer form,
in the minuteness of the postfrontal spine, which is only about
as large as the apical half of that of the female, and in their
shorter abdominal pleura. In the female these are longer
and squarer, and those of opposite sides are abruptly bent
inwards near the lateral margins so as to overlap one another
ventrally, completely closing the subabdominal cavity at all
events when the abdomen is partially flexed, and entirely con-
cealing from view the four intermediate pairs of appendages,
which are laid forwards upon the sternal region, thus forming,
there is little doubt, an incubatory cavity for the eggs. In
our specimens of the male, which are preserved with the
abdomen and its appendages fully extended, the pleura are
not bent inwards, being kept straight by the extended limbs,
but at each end of some of them a short longitudinal crease
is distinctly to be made out, indicating that the male, in
common with the female, possesses the power of closing the
subabdominal cavity.
The appendix masculina is armed at the extremity with
two curvilinear rows of slender and moderately curved spine-
like sete, one row slightly in front of the other; the front
row, consisting of six spines, commences about the middle of
the inner margin and extends to the inner apex of the part ;
the hinder row, consisting of four, commences opposite to the
interval between the third and fourth spines of the front row,
extending to the same level.
Total length, from anterior end of carapace to tip of telson,
48 millim.; of carapace, from middle of anterior to middle of
posterior margin, 15 millim. ; of antennal scale 6°25 millim. ;
of abdomen, from base to tip of telson, 32 millim.; of its
sixth tergum 7°75 millim.; of telson 5 millim.; breadth of
thorax across branchial regions 3°5 millim.; of abdomen
across hump 3 millim.
53. Pasiphaé unispinosa, sp. n.
Pasiphaé unispinosa, W.-M., Ill. Zool. ‘ Investigator,’ pt.i., 1892, Crust.
pl. iii. fig. 7, 2.
Differs from P. s¢vado in the following points :—The body
is not quite so strongly compressed. ‘The carapace is longi-
tudinally convex in the mid-dorsal line and is furnished on
each side with a blunt lateral carina, which commences just
behind the eye and extends downwards and backwards to the
hepatic region, whence, after giving off a branch obliquely
downwards and backwards towards the inferior margin, it is
164 Messrs. J. Wood-Mason and A. Alcock on
continued in a nearly straight line along the branchial region
almost to the posterior margin ; its sides bulge so as to throw
the narrow dorsal region into relief as a blunt carina, and its
postfrontal spine is rather larger. The abdominal hump
appears to be more strongly developed; the sixth abdominal
tergum is deeper, more strongly arched both above and below,
and is not produced to a spine in the middle of its posterior
margin, which, when viewed from above, appears quite
straight. In the first pair of legs the basipodite is furnished
at the apex with one spine, but the meropodite is unarmed ;
in the second pair there is also a spine at the apex of the .
basipodite, but only one on the lower margin of the meropo-
dite. Both eye-peduncles and corne are more elongated.
Length, from anterior end of carapace to tip of telson, 57
millim.; of carapace, from middle of anterior to middle of
posterior margin, 16°5 millim. ; of antennal scale 7-0 millim. ;
of abdomen, from base to tip of telson, 40-0 millim. ; of its
sixth tergum 9°2 millim.; of telson 7-0 millim. ; breadth of
thorax across branchial regions 6 millim. ; of abdomen across
hump 4 millim.
‘wo females were taken on April 11th, 1888, 7 miles S.E.
by 8. of Ross Island, Andaman Sea, in 265 fathoms.
PHYE™*, gen. nov.
Differs from Pasiphaé in the carapace and abdomen being
more or less extensively and distinctly carinated dorsally, in
the former being armed in front with a pair of branchiostegal
spines, and in the telson being forked at the extremity.
Includes P. princeps, 8. I. Smith (‘ Albatross’ Crust. 1884,
p- 37, pl. v. fig. 2, 9, et 1886, p. 78, g 9, Western Atlantic,
444-1342 fathoms); P. acutifrons, Sp. Bate (‘ Challenger’
Macrura, 1888, p. 871, pl. exli. fig. 3, South of Japan, 775
fathoms, and Coast of Patagonia, 245 fathoms) ; P. forceps,
A. M.-Edw. (Miss. Se. du Cap Horn, Crust., 1891, p. 51,
pl. vi. fig. 2, Straits of Magellan, 326 metres) ; and the
tollowing :—
54. Phye Alcocki, W.-M.
Parapasiphaé Alcocki, W.-M., Aun. & Mag. Nat. Hist. (6) vii., 1891,
p- 196, ¢; et Ill. Zool. ‘Investigator,’ pt. i., 1892, Crust. pl. ili. fig. 5.
From P. princeps and P. acutifrons it differs in the form of
the postfrontal spine (which is thin and foliaceous, terminates
abruptly in front in a strongly sinuous and almost vertical
edge, and extends nearly to the posterior slope of the carapace
* Opyrray 7 dvopa bin.—Arist., Ath, Pol. 14.
Indian Deep-sea Dredying. 165
as a backwardly diminishing carina, which is sharp and
foliaceous in its anterior half and blunt in its posterior half) ;
in the pigment of the eyes being brown instead of black; in
the sixth abdominal tergum alone being distinctly carinated ;
and probably in the relative proportions of its different parts.
From P. princeps it differs in the telson being shorter than
the sixth abdominal somite and in the meropodites of the first
pair of legs being armed with three spines upon their lower
margin.
From P. acutifrons it differs very markedly in the form of
the postfrontal spine, and, according to Spence Bate’s figure,
in the strongly arched outer margin of the antennal scale.
From P. forceps it differs no less markedly than from
P. acutifrons in the form of the postfrontal spine, and, besides,
in its more unequal and absolutely much longer antennulary
flagella; in the notch at the extremity of the telson being
acutangular and fringed to the bottom on each side with
minute spinules; in the point of the fixed arm of the claws of
the first pair of legs being simple; and probably in other
details.
Of the three, P. princeps is the one to which it is most
nearly related.
Length, from middle of frontal margin to tip of telson,
59 millim.; of carapace, from middle of frontal to middle of
posterior margin, 18 millim. ; of abdomen, from base to tip
of telson, 41 millim.; of telson 8 millim.; of sixth abdominal
somite 9°75 millim.; of antennal scale 8 millim.
PARAPASIPHAE, S. I. Smith.
a, Antennal and branchiostegal spines absent...... Sect. Parapasiphaé.
The following species come into this section :—
1. Parapasiphaé sulcatifrons, S. 1. Smith, ‘ Albatross’ Crust.,
1884, p. 40, pl. v. fig. 4, pl. vi. figs. 1-7,¢ 9. Western
Atlantic, 515 to 2949 fathoms.
2. Parapasiphaé cristata, id. ibid. p. 44, pl. v. fig. 3, ?.
Western Atlantic, 1628 fathoms.
6. Antennal and branchiostegal spines present ...... Sect. Zupasiphaé.
To this section belong the following :—
55. Parapasiphaé (Hupasiphaé) latirostris, W.-M.
Parapasiphaé latirostris, Wood-Mason, Ann. & Mag. Nat. Hist. (6) vii.
1891, p. 196, 2.
We give a figure of this fine species the size of nature
(fig. 2, p. 166).
166 Messrs. J. Wood-Mason and A. Alcock on
56. Parapasiphaé (Eupasiphaté) Gilesii, sp. n.
Parapasiphaé Gilesii, Wood-Mason, Ill. Zool. ‘ Investigator,’ Crust.
pl. iii. fig. 8, 9, x 2.
Rostrum slender, acute, awl-shaped, slightly compressed,
slightly curved, ascendant, extending by rather more than
Parapasiphaé latirostris, nat. size.
one third of its length beyond the extremities of the eye-
peduncles. Postfrontal ridge cristiform, armed throughout
from the base of the rostrum to the posterior slope of the
carapace with minute forwardly-inclined denticles, the first of
which is placed well on the base of the rostrum and is more
than double the size of any of the rest, which are subequal ;
Indian Deep-sea Dredging. 167
it is divided by a distinct cervical groove into two lobes, the
anterior of which is somewhat the higher and longitudinally
somewhat the more convex of the two.
The anterior margin of the carapace bears on each side two
spines—the one smaller, situated just behind the edge of the
orbital sinus, and answering in all probability to the antennal
spine of the Penzidea; the other larger, which arises from
the margin itself, opposite to the insertion of the antenna, and
is, there is little doubt, a true branchiostegal spine. About
midway between the branchiostegal spine and the obtuse
extra-orbital angle on each side the anterior margin projects
a process of about the same size and shape as the latter, and
hence presents an angularly sinuous or zigzag outline between
the two spines. The branchiostegal spine is continued back-
wards along the side of the carapace as a well-defined slightly
upwardly concave ridge ; this is subacute nearly as far as the
hepatic region, and thence blunt to its abruptly upcurved
extremity near the hinder end of the carapace; it curves
upwards for a short distance from its origin, then descends
almost imperceptibly to about the middle of its length, whence
it rises by a no less gentle ascent to its upturned extremity ;
it emits three branches—one, faint, from the hepatic region,
upwards and forwards towards a point in the anterior margin
which is just internal to the antennal spine ; a second, as well-
defined as itself, from the point at which it is intersected by
the cervical groove, straight downwards and backwards
towards the inferior margin; and a third, equally well-defined,
from a point a little to the rear of that from which the second
is given off, upwards, inwards, and backwards, so as to mark
out the upper boundary of the subjacent branchial chamber.
The terga of the abdominal somites are increasingly carinate
from the second to the fourth; the carina of the fourth is
faintly notched, as in Acanthephyra, and terminates _poste-
riorly in astrong spine. ‘Those of the remaining somites are
transversely rounded.
The telson, which wants its extreme tip, agrees, as far as it
is preserved, with the description of that of Parapasiphaé
sulcatifrons, S. 1. Smith.
The eye-peduncles are compressed from above downwards
and bear on the inner and upper margin, at a short distance
from the edge of the cornea, a small conical tubercle, which is
directed inwards and slightly forwards; the compressed
hemispherical and distinctly faceted cornea occupies the whole
width of their apex, upon which it is somewhat obliquely set ;
and their pigment is bright brown in spirit.
The peduncle of the antennules agrees with Prof. S. I.
168 Messrs. J. Wood-Mason and A. Alcock on
Smith’s description of that of P. sulcatifrons; but the inner
flagellum of these organs is only about one half the length
of the carapace measured from the apex of the rostrum to the
middle of the hinder margin in a straight line, and less than
one third the length of the outer flagellum; this is much
stouter than the inner and expanded at the base in the usual
manner.
The thoracic limbs differ in the relative proportions of their
parts and in armature from those of P. sulcatifrons to an
extent and ina manner which will be best appreciated by
comparison of the figures of the two species.
Length, from apex of rostrum to end of caudal appendages,
49 millim.; of carapace, from apex of rostrum to middle of
hinder margin, 19°5 millim.; of outer flagellum of antennules
34-5 millim., of inner 9°5 millim.; of antennal scale 7°25
millim.; length of abdomen from base to end of telson 27°5
millim., of its sixth somite 5°5 millim., of telson (tip wanting)
65 millim.
A single female was taken on Jan. 15th, 1888, off Cinque
Island, Andaman Sea, in 650 fathoms.
PSATHYROCARIS, gen, nov.
Psathyrocaris, Wood-Mason, Admin, Rep. Marine Survey of India,
1890-91, p. 19.
Integament thin and delicate; all appendages and processes
of extraordinary fragility.
Body moderately compressed.
Carapace shortly rostrate, longitudinally carinate for a
longer or shorter distance in backward continuation of the
dorsal crest of the rostrum; its anterior margin unarmed—
antennal and branchiostegal spines being absent,—at first
broadly arched, then narrowly emarginate between the blunt
triangular extraorbital angle and the antero-lateral angle on
each side ; its sides distinctly ridged, the ridging varying with
the species; the efferent portion of the branchial chamber
relatively spacious.
Antennules and antennz broken off in all the specimens
near the base.
Eye-peduncles and cornee depressed, the latter hemi-
spherical, occupying the whole width of the extremities of the
former, on which they are set quite square. ‘The usual papilla
is present close to the edge of the corne on each side.
Mandibles (Pl. XI. fig. 1) of typical Pasiphaidean form,
with a very large two-jointed palp.
First maxille (Pl. XJ. fig. 2) with the inncr lacinia (ez)
Indian Deep-sea Dredging. 169
pointed and recurved towards the outer (4p), and with a small
triangular process on the inner margin of the endopodite
(en) as in Parapastphaé.
Second maxille (Pl. XI. fig. 3) with well-developed inner
(ca) and middle (4p) laciniz, the latter of which is bilobed,
thus differing from those of all* previously described genera,
in which both laciniz are reduced to a quite rudimentary
condition; and with a relatively very large and powerful
scaphognathite, in correlation to the spacious efferent branchial
channel.
First maxillipedes (PI. XI. fig. 4) with well-developed endo-
podite (en) and middle (4p) and inner (cz) laciniz, the latter of
the two last of which, though decidedly weaker, is nevertheless
produced inwards to the same level as the former, and is fairly
well fringed, thus also differing from all previously described
genera, in which indeed little besides the exopodite of these
jaws remains, the lacinia being both reduced to a more or
less scanty fringe of weak sete and the endopodite to a
minute projection of the inner margin of the exopodite, whilst
the epipodite when present is smaller (Parapasiphaé) or alto-
gether wanting (Pastphaé). ‘The exopodite gives off from its
Inner margin just below the true apex a pedunculated oval
plate, which serves as the occlusor apparatus of the efferent
branchial aperture and, in correlation with the large size of
that aperture, is much larger than in any other genus.
Second maxillipedes (Pl. XI. tig. 5) almost exactly as in the
Peneidea, seven-jointed, the division between the second and
third joints being still quite distinct at the edges ; furnished
with a short tapering exarticulateexopodite (ex) , whichscarcely
exceeds the third joint in length ; and with a complete podo-
branchia (ep + dr), thus contrasting strongly with those of previ-
ously described genera, in which the corresponding appendages
are weaker and but six-jointed, thesecond + third being indistin-
guishably fused into one, there is never any trace of an exo-
podite, and the podobranchia is represented at most by a small
epipodite and may be entirely wanting.
‘Third maxillipedes(Pl. XI. fig. 6) four-jointedf, asin Pasiphaé,
Phye, and Parapasiphaé, turnished at the base with a rudimen-
tary epipodite, and, quite close to the base of the second joint
(2+3+4), with a minute tapering exarticulate exopodite,
similar to, but much smaller than, that of the second maxilli-
pedes, and lodged in a groove in the side of the joint.
* Except Leptochelu, which would appear to have fully developed
second maxille.
+ Leptochela has five, aud is so far less modified than any other genus.
Ann. & Mag. N. Hist. Ser. 6. Vol. xi. 12
170 Messrs. J. Wood-Mason and A. Alcock on
The first and second pairs of legs are dissimilar both in
form and structure.
The first pair is the shorter, and their claws are shaped
much as in Pasiphaé. ‘The inner edge of the dactylopaalite
is unarmed, but is raised into a sharp lamellar cutting-edge ;
that of the prolonged part of the propodite, on the other hand,
is armed throughout with minute acicular spines of tolerably
uniform size and all slanting towards the apex of the joint.
The second pair is much the longer, and their claws differ
in form in the different species and would appear to be
unequal on the two sides. ‘The inner edge of both dactylo-
poditic and propoditic elements of these claws is armed with
minute spines similar to those of the first pair, and, in addi-
tion, at intervals with much longer ones (three or four times
as long) of the same form.
The third and fourth pairs of legs are alike. They are
greatly reduced in thickness, but little if anything in length,
forming long setaceous filaments of excessive tenuity and
fragility.
The fifth pair of legs is the shortest of all and is much
stouter than the third and fourth and much slenderer than the
first and second; it is, in fact, in point of thickness about
intermediate between the two sets of legs. They are set on
and directed in the manner which seems characteristic of the
family. Their propodite bears at the distal end of the lower
surface a conspicuous whisp of longish sete, which is directed
towards the apex of the joint, while the dactylopodite is
covered below with a dense brush of short spiny sete, and is
evidently intended to fold back against the propoditic whisp,
so as to form therewith a sort of prehensile subchela.
All the legs possess the full number of joints and all are
furnished with exopodites. The exopodites of the first pair
of legs are small and inconspicuous, but those of the second
to the fifth pairs are long and excessively fine articulated
setaceous filaments, which form a gradually increasing series
to the fourth, the fifth being suddenly much longer and fully
equalling in length the third or fourth pairs of legs. They
are all very sparsely clothed with long, lax, obsoletely plu-
mose sete, and are all produced at the base into a little
tongue-shaped spur.
The first to fifth pairs of abdominal appendages are remark-
able for the enormous inequality of their two branches, as
well as for the excessive tenuity of the outer branch, which,
in the case of the second pair, is in one species no less than
twenty-four times the length of the inner, which is quite
minute ; the exopodites of the abdominal appendages, in fact,
Indian Deep-sea Dredging. 171
closely resemble those of the legs, differing therefrom only in
being somewhat stouter, more distinctly articulated, and more
richly provided with more strongly plumose sete.
Abdomen transversely rounded, not carinate, dorsally.
Telson quadrangular, tapering gradually from base to apex,
terminating in a minute fixed median spine and two unequal
airs of articulated lateral spines; its dorsal spines obso-
Tpaient
There are twelve functional branchie and two epipodites
on each side distributed as follows in Psathyrocaris fragilis :—
Somites and
their Podo- Arthro- Pleuro-
appendages. branchiz. branchive. branchie.
WEE, nese. Olen.) 0 O = O+ep
js a Oe ! 0 Oe
fp os | a 0 = 2+ep
ee ne 6 1 | ae ea
BLS cheorD 1 nee — ae
» 2 1 Lb, 2.2
BE ts am O 1 iL. 2
Pe a 0 je — ae
lt2ep.+ 6 + 6 = 12+42ep.
Psathyrocaris has hence one more gill—and that a complete
podobranchia consisting of plume and epipod attached to the
second maxillipedes—than in Parapasiphaé, the least modified
of the genera so far described, in which the corresponding gill
is reduced to a rudimentary epipod,
57. Psathyrocaris fragilis, sp.n. (Pls. X., XL.)
Colour in spirit very dark reddish purple or wine-red above,
the ventral surface and legs lighter, the thoracic and abdo-
minal exopodites and antenne colourless.
Thirteen specimens, all more or less broken, though admi-
rably preserved as to their soft tissues, were taken at Station
120, 240 fathoms.
In dealing with a single species it is impossible satisfac-
torily to define the specific characters, and, as the results of
the dredgings carried out during the season 1891-92 contain
several additional species, including a fine ovigerous female
of one of them, we have thought it better to defer the specific
description of the present species until we shall have
thoroughly examined our new material. Meantime the
characters of the genus will suffice for those of the species,
especially as they are accompanied by detailed figures.
12
172 Mr. A. 8. Packard on Aglia tau.
EXPLANATION OF THE PLATES.
PuatTE X.
Fig. 1. Psathyrocaris fragilis, 2, from the left side. Nat. size. Both
the legs of the second pair are figured, to show their inequality.
Fig. 2. Rostrum of the same. Enlarged.
Fig. 3. The caudal swimmeret of another specimen, from above. X 3.
Fig. 4. A leg of the fourth pair of another specimen. X 4.
Fig. 5. An abdominal appendage of the second pair of another specimen.
x 5.
PuaTE XI.
Fig. 1. Psathyrocaris fragilis, Mandible. x 9.
Fig. 2. First maxilla. x 9
Fig. 8. Second maxilla. x 9.
Fig. 4. First Spee x 9.
Fig. 5. Second maxillipede. x 9.
Fig. 6. Third maxillipede. xX 4°.
[To he continued. ]
XXIIJ.—Aglia tau, a connecting-link between the Cerato-
campide and Saturniide, and the Type of a new Subfamily,
Agliine. By ALpHeus 8. PACKARD.
In this European Bombycine moth we have a most interesting
form surviving side by side with Saturnia, which seems to
be the most generalized form of its family. Aglia appears
to be a Ceratocampid in its earlier larval stages, the cater-
pillar in its final stage, however, and the moth being closely
related to the Saturnians. It seems quite reasonable to
suppose that the Saturnians have directly descended from a
form like Aglia, and we could scarcely expect a clearer
demonstration of the origin of one family from another by
direct genetic descent.
‘The transformations of this form, originally figured in
Duponchel and Guenée’s ‘ Iconographie’ * (tom. i1.), have
been more fully elaborated by Mr. Poulton +.
Having received, through the kindness of Dr. Heylaerts, a
young larva of Aglia tau in its third stage, I have been able
* Guenée states that, after attaining its full size, “elle se retire 4 la
surface de la terre, entre des mousses et des débris de végétaux qu’elle
attache avec de la soie, et elle s’y change en une chrysalide grosse, courte,
d’un brun foncé saupoudré de grisitre, et dont l’anus est terminé par un
faisceau de pointes recourbées.”
+ Trans. Ent. Soc. London, 1888, pp. 555-568, pl. xvii. figs. 1-7.
Mr. A. 8. Packard on Aglia tau. 173
to compare it with Hacles imperialis in its third stage, which
Mr. Poulton could not do for want of specimens. The resem-
blance between the two genera at this stage is most striking,
although the fully-fed larve are so different, Aglia passing at
a single moult (the third and last, this larva only having four
stages) from one family to another !
We know of no parallel case, or, at least, of one so very
striking and conclusive*. ‘Thus the ontogenetic development
of this caterpillar epitomizes that of the two families, whereas
that of most Bombyces is simply usually only an epitome of
that of a subdivision of a family or of a small group of
genera.
Aglia tau in its third stage differs from Hacles imperialis
in its third stage in having a pair of dorsal “ horns”’ on the
first and third thoracic segments, 4. ¢mperialis having only
minute ones on the prothoracic segment, while those on the
second thoracic segment are as well developed as those on the
third segment ; those on the second segment are minute; all
the “ horns” are forked, as in Hac/es. ‘The dorsal spines on
the abdominal segments are simple and minute, like those on
the second thoracic segment.
The shape of the head and of the anal legs is much as in
Eacles, but the suranal plate differs strikingly in being pro-
duced into a rather large spinulated spine, a feature not known
to exist in any Ceratocampids.
It should be observed in regard to the large size of the
prothoracic horns of Aglia that those of Citheronia regalis are
quite well developed, being about two thirds as long as those
on the two succeeding segments.
Upon examining the adult of Aglia I find that its head and
antenne are closely similar to those of Hyperchiria to and
the Hemileucide f in general; the antenne form a close
approach to those of H. zo, as on careful examination with a
good lens a second branch of the pectinations of the male
antenne can be perceived ; it forms a long separate branch,
but is in the dead and dry specimens very closely appressed
to the anterior main pectination. In the venation of both
wings Aglia shows a most unexpected resemblance to Hacles
* Nearly twenty years ago, in 1863, when first beginning my studies
on the Bombyces, my attention was attracted to the singular changes of
Aglia tau, and I compared the young larva to the full-grown larva of
Citheronia regalis, and pointed out that the latter genus was an “ embry-
onic form, and therefore inferior in rank to the tau moth ” (‘ Amer. Natu-
ralist,’ June 1870, and ‘ Our Common Insects,’ p. 52).
+ For the sake of clearness, and on account of the differences presented
by the larve as well as the imagines, I am inclined to provisionally regard
this group as of family rank and equivalent to the Ceratocampidz.
174 Mr. A. 8. Packard on Aglia tau.
imperialis ; like that and other Ceratocampide and _ the
Hemileucide having five subcostal branches, while in the
Saturniide there are only four, the first one wanting.
Thus the moth belongs to the Ceratocampide, while the
larva after the last moult loses all its spines and becomes very
much like a Saturnian, perhaps of the type of Telea, and
especially resembling a smooth form like the larva of Attacus
betes, Walker, figured by Burmeister in his ‘ Atlas of the
Lepidoptera of the Argentine Republic,’ though it is without
tubercles or spines. I therefore suggest that Aglia tau
should be regarded as the type of a distinct subfamily of
Ceratocampide, and thus the latter group may be divided
into the two subfamilies, Ceratocampine and Agliine.
At present, both from their larval and their imaginal
characters and from their spinning a cocoon, I am disposed
to consider the Hemileucide as a family closely allied to the
Ceratocampide.
On examining the European genus Endromis we are
disposed to think that the family Endromidz is a natural one.
It would, however, be a violation of the principle of classifi-
cation to include Aglia with it. The two genera, both as
regards their larval and their adult characters, are quite
distinct. I find that Endromis versicolor has the head, palpi,
and antenne, and the hairy abdomen very closely like those
of Hemileuca maia; but the median vein of both wings
divides into four branches, and the subcostal vein of the fore
wings divides into five branches, as in H. maza and the other
Hemileucide. Judging by the coloured figure of the larva in
European works the larva of Hndromis is smooth, with a
small retractile head, oblique bars, and a conical caudal horn.
The group Endromidz is a branch of the Bombycian tree
parallel to but distinct from the Hemileucide, and stands
above the latter, connecting this group and the Cerato-
campide and Saturniide with the higher families of the
Bombyces in which there are four branches of the median
vein, all the families mentioned agreeing with the Notodon-
tide in having but three.
In its general shape, the small retractile head, the mode of
coloration, and the caudal horn the larva of Endromis appears
to be remarkably near the Sphinges. Buckley describes the
cocoon as ‘‘ composed of an open-worked reticulation of coarse
black or black-brown silk threads, with round or broad oval
interstices ; the fabric is extremely strong, tough, and elastic,
covered externally with moss and birch-leaves; fringe adhe-
rent” (iii, 65).
It is interesting that in the transformations of Rhescynthis
A General History of the Marine Polyzoa. 175
erythrine, as figured by Burmeister, we have a parallel to
the case of Aglia tau. The fully grown larva is smooth-
bodied and without the four long large thoracic spines and
the caudal horns on the eighth and ninth abdominal segments
of the previous stage. ‘The genus appears to belong to the
Ceratocampide.
Although we are not yet acquainted with the early larval
stages of Endromis, we do not see why the Sphingide may
not have sprung from a form like this as much as from Aglia,
as the shape and markings of the full-grown caterpillar are
much nearer a typical Sphinx than those of Aglia. More-
over, taxonomically Agia is by no means so “closely ”
allied to the Sphingidw as Mr. Poulton in his able papers
would lead us to infer. In its venation Hndromis is much
nearer, and the latter is a more generalized or synthetic form
than Aglia. From the Ceratocampide the families of Satur-
niide and also of Hemileucide may have originated, and,
indeed, all the Bombyces (unless we except the Arctians and
Lithosiide) may have evolved before the Sphingide appeared.
Judging by the characters of the head, the antenne, thorax,
and especially the venation, the Sphingide are far removed
from the Ceratocampide, and their origin from the latter
family was at least remote, and there must be some lost,
extinct, annectant forms which originally connected them.
XXIV.— Contributions towards a General History of the
Marine Polyzoa, 1880-91.— Appendix. By the Rev.
Tuomas Hincxs, B.A., F.R.S.
[Continued from vol. ix, p. 334.]
‘ Annals,’ February 1882 (p. 82 sep.).
Steganoporella (Vincularia) Neozelanica, Busk.
In a note on p. 85 (sep.) the last clause of the first para-
graph should read thus :—‘‘ The latter is a Membranipora,
the former belongs to a different family.” The Vincularia
abyssicola mentioned in this paragraph is the Smittipora
abyssicola of Jullien, which he ranks in the family group of
the Onychocellide *.
The passage relating to the ocecium on the same page has
been criticized by Dr. Jullien, who challenges the interpre-
tation which I have given of the upper chamber in the zocecia
* Bulletin de la Société Zool. de France, t. vi. (1881).
176 Rev. T. Hincks’s Contributions towards a
of Steganoporella on the ground that no one has demonstrated
the presence of reproductive bodies within it. This may be
true, and may be easily explained. All the probabilities point
to this interpretation as the true one. The Sheiletarase are
generally furnished with external ocecia in which the embryos
complete their development. These are wanting in Stegano-
porella, and it is reasonable to suppose that the internal
chamber provided by the peculiar structure of the zocecium
may be the equivalent. Amongst the ordinary Cheilostomata
species occur which are destitute of ocecia of the ordinary
type; their function is discharged by specially modified
zocecia, which are commonly distinguished by the larger size
and sometimes the altered shape of the orifice. In the case of
Steganoporella, as Busk has pointed out*, the upper or ocecial
compartment is more developed in some of the zocecia than in
others, and the difference is marked by a difference in the size
and pattern of the chitinous framework of the operculum.
These structural variations throw light one on the other.
On p. 86 (sep.) alist has been given of the species belonging
to the genus Steganoporella; but when it was prepared the
structure of the group to which this genus belongs had not
been thoroughly investigated, and important changes have
since been made in the classification. The following table
shows these changes:—Genus STEGANOPORELLA: Species
S. magnilabris, S. Neozelanica. Genus 'THALAMOPORELLA,
Hincks: Species 7. Rozier’, Audouin, and its forms gothica,
Indica, and falcifera; T. Smittii, Hincks; T. Jervoisii,
Hincks; 7’. steganoporoides, Goldstein.
Micropora elongata, Hincks, and M. perforata, MacGillivray,
were wrongly referred to Steganoporella. Vincularia Nove-
Hlollandiew, Goldstein, which was doubtfully included in
Steganoporella, is Thalamoporella Roziert, formindica, Hincks.
Ibid. (p. 86 sep.).
Monoporella albicans, sp. n.
Ibid. (p. 89 sep.).
Schizoporella aperta, sp. n.
The forms described under the above names agree in general
character and in most of the details of structure; but as I was
unable to detect an oral sinus in my specimen of the former,
it seemed necessary to refer it to Monoporella, noting the
remarkable similarity in other points between the two forms.
* ‘Challenger’ Report, part i. p. 74.
General History of the Marine Polyzoa. 177
I now believe that in the specimen (or specimens) on which
Monoporella albicans was founded the sinus was not absent,
but was entirely concealed by the suboral umbo, as it is, I
find on reexamination, in many of the cells of the other form.
Monoporella albicans therefore must merge in Schizoporella
aperta,
‘ Annals,’ August 1882 (p. 96 sep.).
Euthyris obtecta, sp. n.
The larger cells with modified orifice are now known to
occur on many species and are no doubt subservient to repro-
duction. MacGillivray rightly refers Carbasea (Flustra)
episcopalis to this genus; but he does not mention whether
it possesses the two classes of cell.
‘ Annals,’ March 1883 (p. 104 sep.).
StirpPariA, Goldstein.
It may be a question, I think, whether this genus can be
maintained. The erect segmented stem seems to be the one
peculiarity which separates it from Bicellarta *, with which
it entirely agrees so far as the zocecial characters are con-
cerned; and the morphology of this structural element has
hardly been determined as yet with certainty. There is a
close resemblance between the stem of Stirparia and that of
the genus Kinetoskias of Koren and Danielssen, also a
Bicellarian form,
Ibid. (p. 108 sep.).
Family Cellariide.
FarcrmiA, Pourtales.
Farcimia appendiculata, sp. n.
In a paper on “ Tertiary Chilostomatous Bryozoa from New
Zealand” t— Mr. Waters has identified his Membranipora
articulata t with the present species, and expresses an opinion
that if I had “ decalcified’ my specimens I should have taken
the same view. Circumstances have prevented me from
* Busk refers Stirparia glabra (mihi) to the genus Bicellaria (‘ Chal-
lenger’ Report, part i. p. 35, pl. vi. fig. 1).
+ Quart. Journ. Geol. Soc. for February 1887,
¢ “Fossil Chilostomatous Bryozoa from South Australia,” Quart.
Journ. Geol. Soc., August 1882,
178 Rey. T. Hincks’s Contributions towards a
recurring to the subject before ; but after a careful reexami-
nation, recently made, I still find myself quite unable to
accept his conclusion. Fareimia appendiculata, when deprived
of all its chitinous appendages and reduced, as far as possible,
to the condition of the fossil, differs markedly from Memb.
articulata, and in my judgment the differences are specific.
If we compare the individual zocecia we can hardly fail to be
struck by the points of contrast which they present and which
are clearly shown in Mr. Waters’s figures and in my own.
In the first place there is a remarkable difference in the shape
of the cells as well as in much of their detail. That of
M. articulata, which is inclosed by a conspicuous raised
boundary line (the “‘ band” of Waters), is elongate and hexa-
gonal in form. The zocecia are ranged in single file along
each face of the quadrilateral stem, and are quite distinct, but
are linked together by an extension of the marginal line given
off from the summit of each cell. The aperture is compara-
tively small, occupying the upper two thirds of the area,
narrow, with straight sides, rounded above and slightly incurved
below. A calcareous lamina surrounds the aperture, closing
in the space between it and the outer wall.
In Farcimia appendiculata the zocecia are elongate-oval,
contracted above, so as to form a framework for the operculum,
the extremities in contact or nearly so, the margin rather
thin, sloping slightly outward and usually expanded at the
base of the cell; the aperture occupying the whole of the area
and closed in by a membranous covering ; the orifice semi-
circular, at the very top of the zocecium, ‘There is some
slight diversity in the shape of the cell, the regular oval
giving place in many cases to a form which narrows off
towards the top and expands considerably below ; but in both
conditions it is equally removed from that which characterizes
the other species. he dissimilarity of the apertures in size and
shape is very marked and significant, whilst the total absence
of the raised boundary-line which surrounds the cells and links
them together, and which in M/. articulata gives its distinctive
aspect to the colony, is strong evidence against specific identity.
In the presence ot such important zocecial differences I am
fully justified in regarding the two forms under consideration
as distinct species. 1 may add that a more searching method
of investigation has only made these more apparent.
It is difficult to compare the avicularian appendages in the
recent and fossil forms. ‘Those of J. appendiculata are
remarkable for their size and structure, and almost fill in the
space between the lines of cells. Mr. Waters represents in
his figure two very small pointed avicularia of the ordinary
General History of the Marine Polyzoa. 179
form near the top of each zocecium, placed one on each side
and pointing downwards. There are no traces of anything
more. The appendages of the Farciméa are remarkable for
their size and are of a peculiar type.
In my previous description of this species I have referred
to a structural similarity between its avicularia and those of
Scrupocellaria ; but more accurate observation has convinced
me that there are most important differences between them.
They consist of an elongate avicularian chamber, tapering
slightly downward, convex in front, and adnate to the side of
the cell a little below the oral extremity. The top (which is
flattened) is occupied by the mandibular apparatus and carried
out in front into a small beak-like process; the mandible is
rounded. A raised calcareous margin surrounds the chamber
(which is also calcareous), and the front is closed in by a
chitinous covering. As I have mentioned, the space between
the rows of cells is almost entirely occupied by the avicularian
appendages.
The differences between Farcimia appendiculata and
Memb. articulata as figured by Mr. Waters, which I have
just described, can hardly be due to the changes which have
taken place in the fossil. Traces of the large avicularian
chamber, which is formed of calcareous material, must have
survived, for this structure in the recent form, with the excep-
tion of its chitinous envelope, was not materially affected by
incineration.
The ocecium in F. appendiculata is immersed, as it is in
many of the Flustre. In ovicelligerous cells the margin is
not carried round the top, its place being taken by the oral
arch of the ocecium ; the latter occupies a small intercellular
space. ‘The ocecium is somewhat shallow, subglobular, the
front occupied wholly by a large circular orifice facing the
interior of the cell and probably closed by a membrane.
Above the operculum of the cell, immediately under the oral
arch of the ocecium, is a narrow slit-like opening through
which the embryos escape.
In a previous description of this species* I have referred to
certain remarkable appendages which are distributed in large
numbers over the zoarium. ‘These consist of tall, erect, strap-
like bodies, formed of shining membrane of a light brown
colour, broad below and for a great part of their length, but
tapering off abruptly within a short distance of the tip and
terminating in a sharp point. They seem to be always
connected in some way with the avicularium—sometimes
* Ann. & Mag, Nat. Hist. for March 1883.
180 Rev. T. Hincks’s Contributions towards a
attached to the side of it, sometimes to the top also, as if to
shield it from some danger. But we must have more definite
knowledge than we now possess of the function of the avicu-
laria themselves before we are likely to determine the precise
office of the subsidiary appendages. It may be noted that
the extremity of these curious organs is commonly bent
downwards from a definite point.
I do not propose to discuss at any length the generic posi-
tion of this very interesting form. It possesses characters
which, I think, should separate it from Membranipora, and
may probably be referred to the genus /arcimia of Pourtales*,
of which Smitt, in his ‘ Floridan Bryozoa’ (part il. p. 3), has
given an interesting account, pointing out its distinctive
peculiarities. The generic name, as he has remarked, was
originally given by Fleming to a Cellaria, and is therefore a
mere synonym available for further use.
Ibid. p. 109 (sep.).
Schizoporella cinctipora, sp. n.
Mr. Waters has described and figured a variety (personata)
of this species + from New Zealand Tertiary beds. The
differences, however, between the supposed varietal form and
the recent S. cinctipora as figured seem to be so striking and
important that I venture to question their specific identity,
Ibid. (p. 109 sep.).
Lepralia foraminigera, sp. n,
This species has also occurred in the New Zealand Tertiaries.
Mr. Waters notes that only the two upper openings in the
cell-wall occur in fossil specimens.
Ibid. (p. 110 sep.).
Lepralia rectilineata, sp. n.
This has also been found fossil in New Zealand. Waters
mentions that there is ‘often a small ridge or boss at each
side of the aperture, just below which there are two small
avicularia.” The occium, which I had not met with, is
described as “raised, globular, about half as wide as a
zocecium.”’
* Bull. Mus. Comp. Zool. Haivard Coll, Cambridge, no. 6, p. 110,
+ Quart. Journ, Geol. Soe., Feb. 1887,
General History of the Marine Polyzoa. 181
Ibid. (p. 110 sep.).
Mucronella bicuspis, sp. n.
This species is ranked by Waters as a variety of his
Smittia biincisa*, I have not had the opportunity of
examining specimens of the fossil form, which was procured
from Tertiary deposits in South Australia and New Zealand ;
but, judging from the figure, I can hardly think that there is
sufficient ground for identifying it with MZ. dicuspis. The
general character of the orifice seems to me to be very different
in the two; the lateral avicularia, a very constant feature of
the recent species, are wanting in the fossil, while the tubular
avicularia which are so conspicuous a characteristic in the
fossil are wanting in J. bicuspis. The oral denticle of the
latter is peculiar; but Mr. Waters has noticed some variability
in this portion of the structure.
The difference between the characters of the cell-wall in the
two forms is remarkable. Not only are there more of the
large pores in S. biincisa, but they differ in shape and
arrangement from those of the recent species. They form a
reticulate covering over the whole surface of the cell, whilst
in MM, bicuspis they are disposed in a single line running
across the front of the cell. At the same time it must be
admitted that the superficial character of the cell-wall is liable
to much variation. Mr. Waters’s experience as a student of
fossil Polyzoa entitles his judgment to much respect in a case
of this kind, and on the whole I prefer to leave the question
an open one.
‘ Annals,’ May 1884 (p. 358) T.
Membranipora marginella, sp. n.
Two of the four cells figured bear ocecia; the cells men-
tioned in the description as furnished with a large dark-
coloured operculum, “‘ occupying nearly half the area,” are
probably avicularian.
Ibid. (p. 358).
Smittipora abyssicola, Smitt.
The genus Smittipora is, in my judgment, a synonym of
Onychocella, Jullien, the differences between the two being
quite immaterial.
* Quart. Journ. Geol. Soc., August 1882, p. 272; wed. February 1887,
p. 58. we uy
+ From this point to the close of the “ Appendix” the paging is that
of the ‘ Annals’ and not of the separate copies.
182 Mr. B. T. Lowne on the
Ibid. (p. 360).
Microporella Fuegensis, Busk (sp.).
This is not a Microporella, as it wants the suboral pore
characteristic of this genus. It is furnished with the peristo-
mial pore, which is a leading character of Busk’s Adeonells ;
but this has a totally different structural significance.
As there is considerable doubt about the latter genus, [
shall postpone the discussion of the systematic place of the
present form.
[To be continued. ]
XXV.—A Reply to some Observations on the Mouth-organs of
the Diptera. By B. THompson Lowne, F.L.S.
Mr. Cuartes O, WATERHOUSE in the January number of
this Journal appears to invite me to reply to what, for want of
a better term, | may designate a “ quip courteous,” in which
he has availed himself of the saving qualities of an ‘ if.”
My critic has, curiously enough, seen more in my book than
I ever wrote or intended, and has failed to see what I did
write ; therefore I avail myself of an “ if.”
If Mr. Waterhouse had used no more acumen in the inter-
pretation of the mouth-parts of the Diptera than he has
brought to bear on the interpretation of what I have said I
should not have been surprised that he still holds the old and
time-honoured opinions regarding the mandible of the dipte-
rous mouth. I do not, however, for a moment suppose that
he reads “‘ Nature” as carelessly as he reads my work ; but
I think he might have rewarded the “ skill and care” which
he credits me with by a little more attention before he con-
signed me to oblivion in the pit of error in some unknown
region ; for if I have fallen into “some error,” the nature of
which is not even indicated, my position is no better, and
there is small chance that a passing friend may draw me out.
Therefore it behoves me to make an effort to save myself.
The main argument I use in favour of the views I have
adopted is the manner in which the parts in question are
developed. If I have falsely interpreted the appearances
relating to their development | am as likely to be wrong as
another ; therefore the question at issue is: Are the mouth-
parts ot Musca developed as Mr. Lowne states or are they
not? ‘There are no side issues to the question.
Mouth-organs of the Diptera. 183
Now with regard to my sins of omission. Mr. Waterhouse
is doubtful as to what I mean by parts of the maxilla; yet on
page 154 of my book I have indicated that I regard the lancets
as homologous with the palpiger and the lacina respectively.
My critic then makes it appear that ‘‘ I blow hot and cold”
according to my necessities. It is true I said that the position
of the pseudo-labium is no evidence from a morphological point
of view. In this Mr. Waterhouse concurs. But I have
nowhere stated that position is never of value in establishing
the morphology of a part.
It is possible that two pairs of appendages are fused in the
dipterous proboscis ; it is conceivable that three are so united,
and it is further possible that these limbs are so entangled
and crossed, that position would afford no evidence of the
morphology of their distal extremities. Moreover the mere
similarity of a terminal joint to that of an homologous appen-
dage, such as Mr. Waterhouse appeals to, is not evidence that
this joint belongs to a mandible or a maxilla. The hoofs of
a horse’s feet are similar; but it would be hazardous to con-
clude that a limb with two hoofs consists of a fore and hind
limb united. I see no reason at all why the terminal joint of
one of the divisions of the maxilla should not resemble a
mandible when it has similar functions to perform.
As regards the simple eye of the flea the case is very
different. Its relation to the antenna is such that it cannot
be explained on the supposition that the sternal plates of the
cephalic segments are dorsal, as the position of the antenne
is explained in 7ruxalis and Fulgora. Moreover Mr. Water-
house supposes an imaginary case, which does not exist so
far as we know. It will, I think, be time to consider its
bearings on the view I have adopted after its discovery. In
the meantime I assume that it does not happen that the
compound eye ever bears the relation to the antenne which
the simple eye of the flea exhibits.
Mr. Waterhouse credits me with a consistency which I do not
deserve. When I published my book on the Blow-fly in 1870 I
never said of the proboscis that “ it is mainly formed from the
maxille.” I then regarded it as a complex of several meta-
meres. I was wrong; but until I discovered the manner in
which it is developed no one had done so. The contradictory
statements of various writers on the subject will themselves
speak for the difficulty which exists in making the parts of
the proboscis conform to the received theory ; and the very fact
.that some have regarded it as composed of several meta-
meres whilst others deny it metameral characters shows that
the theory does not fit with the facts.
184 Mr. O. Thomas on a new Species of Sminthus.
Mr. Waterhouse is unfortunate in having ascribed a_pre-
conceived opinion to me, as my early writings show that I
formerly held very different views—views which I have since
given up; and I first published the idea that the proboscis is
developed mainly from the maxille in a short paper in the
‘Quekett Club Journal’ of 1887.
Although when my present work was first contemplated I
was asked to bring out a second edition of my former book, I
soon found that it would be entirely new, and Mr. Waterhouse
has committed a slight inaccuracy in speaking of my present
book as a second edition of one published in 1870. I should
have thought that the mere fact of its containing four times
the matter already, and a prospect of its containing 700 pages
when complete, might have indicated this; moreover it has
a different title. But perhaps it is too much to expect in
these days of high pressure that those who quote a work
should look at the title.
XXVI.— Description of a new Species of Sminthus from
Kashmir. By Ouprietp THOMAS.
Tue remarkable genus Sminthus, which, nota bene, is a
member of the Dipodide *, not of the Muride, has been until
recently considered to consist of only a single species, S. sub-
tilis, Pall. (S. vagus, auct. plurim.), ranging from Denmark
to Central Asia. Quite recently Dr. Biichner ¢ has described
a second species, S. concolor, from Ganssu, China, a species
which has not the characteristic black dorsal stripe of S. sub-
tilis ; and I have now the opportunity of describing a third
one of the same most interesting group.
The type specimen is a skin with skull, obtained in
Kashmir, at an altitude of 10,000 feet, by Major G. H.
Leathem, of the East Surrey Regiment, in whose honour I
propose to name the species
Sminthus Leathemi, sp. n.
Extremely similar in size, colour, and general appearance to
Mus sylvaticus, from a British specimen of which it is at first
sight hardly distinguishable, except that the ears are decidedly
smaller.
* This view of the true affinities of Sminthus, first published by Winge
in 1887 (‘ Gnavere fra Lagoa Santa—Ii Museo Lundui,’ p, 109), had been
held by me long previously, and I still think it is unquestionably correct.
¥ Bull, Ac. Sci. St. Pétersb. xiti. p. 267 (1892).
Mr. O. Thomas on a new Species of Sminthus. 185
Fur long and soft. General colour rufous grey, becoming
clearer rufous on sides; no trace of a darker dorsal stripe.
Belly white from chin to anus, but the hairs slate-coloured
for their basal three fourths. Ears short, rounded, their
visible parts when folded (i. e. the posterior two thirds of
their inner and the anterior third of their outer surfaces)
thickly clothed with short chocolate-brown hairs. Arms and
legs whitish ; hands and feet silvery white on their upper
surfaces ; palms and soles naked ; hallux reaching to the base
of the second toe, fifth toe to the middle of the second joint of
the fourth. Tail elongated, conspicuously bicolor, brown
above, both hairs and scales, white below.
Skull decidedly larger than that of S. subtilis, narrower
and more elongated anteriorly, and with a longer and
more oval brain-case ; interparietal bone decidedly broader
than that of S. subtilis, its antero-posterior only about a third
of its transverse diameter. Anterior palatine foramina
shorter, extending backwards only to the level of the middle
of the premolar; palate posteriorly ending close behind the
posterior molars, while in the allied species it is continued
backwards for a distance equal to the combined lengths of ®-*
and ™-1,
Teeth apparently quite as in S. subtilis, except that "3 is
longer, as long as broad, instead of being transversely oval ;
m.3 18 also correspondingly slightly longer in proportion to the
other teeth.
Dimensions of the type (a male) :—
Head and body (measured in the flesh by Major Leathem)
66 millim.; tail (ditto) 112; hind foot (from skin) 19; ear
(from skin, approximate) 8-3.
Skull: greatest length (occiput to nasal tip) 20°7; nasals,
length 7°7, greatest breadth 2°7; interorbital breadth 42 ;
greatest breadth across brain-case 9°6; interparietal, length 2°3,
breadth 7:0; palate, length 8-2; diastema 5°0; length of
palatine foramina 4:0; length of upper tooth-series 3°1.
Hab. Krishnye Valley, Wardwan, Kashmir. Altitude
10,000 feet. Type obtained June 24, 1892.
As may be seen from the above description, this new species
is widely removed from S. subtilis and approaches in many
characters the Chinese S. concolor, From this it is distin-
guished externally by its conspicuously bicolor tail, by its
brighter coloration, and shorter ears. In the skull it agrees
closely with the same animal as regards the general measure-
ments and in the “stirker verschmilert und schlanker aus-
gezogen vordere Partie” (both being compared with S. subtilis) ;
but it is impossible to believe that so’careful an observer as
Ann. & Mag. N. Hist. Ser. 6. Vol. xi. 13
186 Mr. O. Thomas on the Genus Chiroderma.
Dr. Biichner could have overlooked the marked differences
in the palate and interparietal had they also been present in
his species; so that I suppose S. concolor agrees with S. sub-
ti/is in these respects.
As to the structure of the teeth, all the members of the genus
seem to have the small fifth cusp on ™1, which is stated by
Dr. Biichner not to be present in S. subtilis; for in three
specimens of that species in the Museum, from widely different
localities, I find it clearly visible, although smaller than in
S. Leathemi. I presume, therefore, that Dr. Biichner had
under examination only specimens with worn dentition.
Major Leathem is to be congratulated on his discovery of
this interesting little animal, the first representative of its
genus found within British Indian territory.
XXVII.—Further Notes on the Genus Chiroderma.
By OLpDFIELD THOMAS.
TuE following points with regard to the Chiropterous genus
Chirederma have arisen out of the publication of Dr. H.
Winge’s ‘ Bats of Lagoa Santa’* and of Dr. H. Allen’s
remarks on the genus in connexion with the description of
his Vampyrops zarhinus +.
In the first place, my own remarks on the genus itself seem
to have been unaccountably misunderstood by the latter
author, for, far from “claiming that Chiroderma is not
distinct from Vampyrops,” I maintained, in the paper criticized
by him f, that it was a perfectly distinct genus, and gave it a
place in the synopsis of genera on p. 170, although at the
same time two species referred by Dr. Dobson to Chiroderma
were transferred to Vampyrops. Nor do I by any means
“‘ acknowledge that the nasal cleft disappears in old indivi-
duals,” as Peters’s statement to this effect was not accepted in
my paper and was explained away in the last paragraph of
. 169.
P One mistake in my paper, however, I should like to correct
more prominently than I have hitherto done, namely as to the
spelling of the new species there described. This should be
Vampyrops Caraccioli, instead of V. Caracciole, the mistake
having been due to a misconception as to the name of the
discoverer, Mr. Caracciolo, whose proper name is now well
* ‘Jordfundne og nuleyende Flagermus fra Lagoa Santa, Minas Geraes
—E Mus. Lundii,’ 1892.
+ Proc. Ac. Philad. 1891, p. 400.
t Ann. & Mag. Nat. Hist. (6) iv. p. 167 (1889).
Mr. O. Thomas on the Genus Chiroderma. 187
known to zoologists in connexion with the foundation of the
Trinidad Field Naturalists’ Club, of which he is president.
Again, in 1891 I had the opportunity of describing* a
Stenodermatous bat, this time of the genus Chiroderma itself.
That description was based on the Minas Geraes specimen
called in Dobson’s Catalogue Chiroderma villosum, Peters ;
and reasons were shown why it should be looked upon as
distinct, and the name Ch. Dorie was given to it. In the
contrasted descriptions of the teeth of the two forms on p. 882
two important misprints have crept in, which, although they
might be detected on a careful reading of the whole paper,
yet, placed in so prominent a position, might easily deceive
any one working on the subject. The mistake consists of the
transposition of the dimensions of the posterior teeth of
Ch. Dorie and Ch, villosum, so that each is made out to have
the dimensions of the other. The erroneous lines should
therefore read :—
Ch. Dorie. Ch. villosum.
Posterior upper premolar} Larger, combined Smaller, combined
and two molars ...... length 7-4 mm. length 60 mm.
Posterior lower premolar| Combined length Combined length
and two molars ...... 82 mm. 63 mm.
Thanks to the kindness of Dr. H. Winge, the National
Collection has now obtained from the Copenhagen Museum a
number of the specimens from Lagoa Santa described by him
in the valuable paper quoted above. Among these there is a
specimen called by him Chiroderma villosum, apparently in
ignorance of my paper, for on examination this proves, as
might be expected trom its locality, to be an example of
Ch. Dorie, agreeing in every respect with the type, except
that its outer incisors are rather shorter, perhaps owing to
greater wear. This very species, according to Dr. Winge,
was the one termed by Lund “ Phyllostoma dorsale,”’ but
without description ; so that my own much later name will still
have to stand for it.
The full synonymy of the Minas Geraes species will
therefore be :—
Chiroderma Dorie.
Phyjllostoma dorsale, Lund, Blik Bras. Dyrey., Dansk. Afh., ix. p. 200
(1842) (nom. nudum),
Chiroderma villosum, Dobs. Cat. Chir. B. M. p. 534, pl. xxix. fig. 2
(dentition) (1878); Winge, E Mus. Lundii, 1892, p. 9, pl. 1. fig. 12
(skull and teeth) (nec Peters).
Chiroderma Dorie, Thos. Ann. Mus. Genoy. (2) x. p. 831 (1891).
* Ann. Mus, Genoy. (2) x. p. 881.
188 R. 8S. Bergh on the Development of the
XXVIII.—On the Development of the Germinal Streak of
Mysis. By R. 5. Bande, of Copenhagen *.
My investigations commence approximately at the stage at
which the segmentation has concluded and the formation of
the germinal layers begins. On the termination of the process
of segmentation the blastoderm has extended round the yolk
in every direction, and there now arises a thickening of the
blastoderm in the form of a transverse streak (this streak
occupies a transverse position with reference both to the longi-
tudinal axis of the ovum as also to the subsequent longitudinal
axis of the embryo). In this thickened streak the cells are at
first arranged in only a single layer, and the thickening is
consequently occasioned merely by the greater height of the
cells of this regidn. Soon, however, the streak becomes
bilaminate at a certain spot in the neighbourhood of the
median line, since some few cells push their way inwards and
shortly afterwards undergo active multiplication within the
outer Jayer. I have observed stages in which only two, four,
or six inner cells are present, but the number soon becomes
much larger; figures of nuclear division are frequently met
with. The inner mass of cells which has thus been formed
now speedily separates into three different rudiments :—
(1) certain cells wander about and develop into vitellophaga
(“ Vitellophagen ”’) ; (2) other cells (which adjoin the median
line) become more firmly united together into a plate and
constitute the true (intestinal) endoderm; (3) towards the
sides certain others develop as primitive cells of the muscle-
plates (mesoderm of authors). The final number of these
primitive cells is four on each side, though in earlier stages
not so many of these larger lateral cells are to be seen. As
soon as the definitive number is reached these cells commence
to produce smaller ones in front by the process of budding
(“ Knospung”’); thus four longitudinal rows of cells are
formed on each side within the ectoderm, so that in certain
stages any transverse section from the region in question
contains four muscle-plate cells on either side. With further
growth the muscle-plates become very distinctly segmented,
and ] see no reason to doubt that their divisions correspond to
actual proto-segments. ‘The latter soon become separated
from one another, since the growth of the muscle-plates does
* Translated from the ‘Zoologischer Anzeiger,’ xv. Jahrg., no, 406,
November 28, 1802, pp. 436-440,
Germinal Streak of Mysis. 189
not keep pace with that of the ectoderm, so that the divisions
of the germinal streak which contain proto-segments alter-
nate with others in which no elements of the muscle-plates
are to be found: by this means this “ primitive segmentation”
can be recognized with peculiar distinctness. Hach proto-
segment at first consists of a simple transverse row of cells :
it is not until later on that these multiply, so that the rudi-
ment gradually comes to consist of several rows and several
layers, when, owing to the fact that the proto-segments then
fuse together, the muscle-plates develop into a continuous
layer within the ectoderm.
The ingrowth alluded to above, through which the deeper
cell-layers of the embryo are formed, without doubt corre-
sponds to the gastrula-invagination, from the lateral margins
of which the formation of the muscle-plates consequently
proceeds in this case also. Whether these muscle-plates
belong genetically to the ectoderm or to the endoderm it was
impossible to decide, and the question is one of those which
in many cases are most difficult of all to determine, but never-
theless are often “decided” with the utmost arbitrariness.
The blastopore has no relation whatever either to the mouth
or anus; its situation is in the neighbourhood of the future
anus: this, however, does not arise until much later, long
after the blastopore has become completely unrecognizable.
Before the formation of the gastrula-ingrowth no yolk-cells
are to be found.
At the anterior margin of the blastopore a very peculiar
differentiation of certain ectoderm-cells takes place: these
develop into primitive cells of the ectodermal portion of the
germinal streak, 'The definitive number of these cells is
seventeen or nineteen (1 find sometimes the one, sometimes
the other) ; they form a transverse arcuate streak in front of
the blastopore. ‘The first stages of the differentiation and
grouping of these cells seem to last only a very short time ;
for, in spite of the fact that I examined a very large number
of germinal disks at such stages, I am only able to assert that
I have found stages with nine, eleven, thirteen, and fifteen
primitive cells; less than nine primitive cells were not found,
and consequently no transition between this stage and the
earlier phase, at which such a grouping of the cells is alto-
gether indistinguishable ; it was likewise impossible to deter-
mine whether the original nine cells multiply into the seven-
teen or nineteen by means of fission or whether their number
is augmented by accessions from other neighbouring cells.
190 R. 8. Bergh on the Development of the
So soon, however, as the definitive number * is attained, these
cells commence precisely the same process as the primitive
cells of the muscle-plates (the cells lying nearest to the median
line are the first to begin, and are followed by those occupying
a more lateral position) : they produce smaller cells in front
by budding (“ Knospung’’). In this manner there conse-
quently arises an ectodermal germinal streak, formed of seven-
teen or nineteen longitudinal rows of cells. ‘The cells of this
germinal streak are also seen to be arranged very regularly
in transverse rows, and the cell-divisions take place precisely
in the same manner and with the same regularity as I recently
described in the case of Gammarus. A median row of cells
is always found, which is derived from a median primitive
cell; therefore the number of the primitive cells and of the
rows of cells is always an uneven one.
This ectodermal primitive streak extends in front as far as
a line which connects tbe points of insertion of the right and
left mandible. In front of this line we find in all stages under
consideration a mosaic of ordinary polygonal ectoderm-cells,
which are not arranged in rows and are not derived from the
above-mentioned primitive cells. It seems to me that this
fact, that the ventral ectoderm is differentiated, so to speak, into
a nauplial and (sit venia verbo) metanauplial rudiment, is not
entirely devoid of interest. The Nauplius appendages grow
out from the anterior mosaic of cells; but the whole of the
appendages which are situated behind the mandibles owe their
origin to the germinal streak which is derived from the primi-
tive cells. Behind the primitive cells there is formed at an
early period an embryonic (provisional) forked caudal fin
(“ Schwanzflosse ”), which is very distinct in the Nauplius
stage. ‘The epidermis withdraws by degrees from the chitinous
covering of this caudal fin, and the definitive caudal fin is
formed considerably further forward from the material of the
germinal streak. It is probable that the only other structure
which arises from the cell-material lying behind the primitive
cells is the telson, though this is difficult to prove.
The position of the endoderm-plate in different stages is
worthy of notice. Situated at first behind the ectodermal
primitive cells, it travels by degrees on the inside of these and
of the germinal streak very far forwards, until it enters the
region of the mandibles. The primitive cells of the muscle-
* The numerical theorists of Prague are here furnished with interesting
subjects for study. A comparison of the conditions of Mysis and Gam-
marus With reference to the validity of their laws would certainly have a
brilliant result.
Germinal Streak of Mysis. 191
plates, too, lie at first close behind, but in later stages close in
front of the ectodermal primitive cells. At last all the primi-
tive cells split up into smaller cells,
When the organs (nervous system and appendages) begin
to develop from the germinal streak the regular arrangement
of the ats in rows gradually disappears, since they commence
to divide in different planes. I am entirely unable to state
positively how many of the original longitudinal rows enter
into the formation of the ventral nerve-chain; it seems to me
most probable that it is only the median row and the one lying
next to it on each side which take part therein. Another
process, however, is very distinctly recognizable in the forma-
tion of the ventral chain; for the ectoderm-cells which are
destined for the production of the ganglion-cells become
developed as primitive cells, which, by means of budding,
give rise to rows of smaller cells towards the interior in a
manner precisely similar to that which Wheeler* has
described in the case of Insects. Yet in Mysis the “ neuro-
blasts”’ (as Wheeler terms them) are not covered by the
epidermis, but actually represent the most superficial layer of
cells of the region in question, and, so far as I am able to
observe, persist throughout as epidermis-cells, while in the
case described by Wheeler they are situated within the
epidermis.
A torsion of the germinal streak, such as I recently
described as occurring in Gammarus T, does not take place in
Mysis.
As is evident from what has been stated above, besides
many points showing great agreement with the conditions
which are found in Gammarus, several noticeable deviations
from what is seen in the last-named form also occur. In this
connexion the existence of the larger primitive cells at the
posterior end of the germinal streak of Mysis is especially
worthy of mention, since these are wanting in Gammarus.
On the whole Mysis is a more convenient and more easily
manipulated object for the study of the processes which are
here alluded to. The above results were derived partly from
the study of series of sections, but chiefly from the examina-
tion of transparent surface-preparations. J. Nusbaum ft, the
most recent monographer of the development of Mysis, has
* Wheeler, “ Neuroblasts in the Arthropod Embryo,” Journal of Mor-
phology, vol. iv., 1891, p. 337 et seg.
t+ Zool. Anzeiger, 1892, no. 396.
¢ J. Nusbaum, “ L’embryologie de Mysts chameleo (Thompson),” Arch,
de Zool. exp. et gén., sér, 2, t. v., 1887, p. 123 et seg.
192 Bibliographical Notices.
entirely neglected the study of surface-preparations of this
kind, and consequently the conditions which are here
described were almost completely ignored by him. In the
present brief communication I am unable to make further
reference to the literature of the subject.
BIBLIOGRAPHICAL NOTICES.
A Catalogue of British Jurassic Gasteropoda. By W. H. Hupteston,
M.A., F.R.S., P.G.S., and Epwarp Witson, F.G.S. 8vo.
Pp. xxxiiiand 147. Dulau and Co.: London, 1892 (November).
Tue very aspect of the pages of this book reminds a working Geolo-
gist of the well-known ‘ Morris’s Catalogue of British Fossils,’ even
without the allusion in the Preface to the latter still useful book,
though it has long been out of print. This new Catalogue, however,
is limited (as its Title intimates) to one Molluscan group of one
Formation, and has very good additional features in its lists of
localities, bibliographic catalogues, table of genera, notes on some
genera and on doubtful and rejected species. Moreover, the syno-
nymy and references are far more liberally represented than in the
former work.
The plan of this Catalogue is clearly laid down in the “ Explana-
tory Note,” pp. xi-xvi; and the paleontological bearings both of
the whole group and of its divisions are treated of in the Preface,
. V-x.
PPThe long experience and the accurate knowledge of the two ener-
getic Authors may be well trusted for the satisfactory fulfilment of
the task they have undertaken and brought to publication in this
work; and their nomenclatorial and general literary style and
method are decidedly good. Great pains have evidently been taken
to have quite correct printing; and severe judgment has evidently
been exercised in coming to a conclusion in cases of doubtful priority,
in choosing the best and most necessary references, and in deter-
mining the synonymy.
Sixty-five genera and subgenera of British Jurassic Gasteropoda,
with their frequently numerous species (122 in Cerithium and 78 in
Pleurotomaria, for instance), constitute the chief material here
reduced to zoological order, as far as the often imperfect preservation
of the shells permits. The six genera from the Rhetic Beds are
similarly treated (pp. 137-139).
Thus all the evidences that the relics of Gasteropods in the
British and, in many associated instances, foreign Jurassic strata can
yield to the experience of experts, as to zoological and geological
conditions and changements, are here brought to our convenient
Bibliographical Notices. 193
notice in a well-printed Catalogue. This is worthy of the attention
and well fitted to the use of geologists, whether working earnestly
in the details of the science or taking up the pleasures of ‘“ col-
lecting ” at a given locality or in a given formation.
The abundant fruits of research among the British Jurassic Gaste-
ropoda during the last forty years, largely due to the energy and
acumen of W. H. Hudleston, and incorporated in this work, thus
occupy 120 pages in the new Catalogue, whilst 55 pages of ‘ Morris’s
Catalogue’ served for all the known fossil Gasteropoda of Britain ;
the very limited and condensed references, however, in the Jatter
somewhat affect the comparison.
It is to be hoped that the scientific public will liberally support
the publication of such excellent catalogues of the British Fossils as
the book under notice and Woodward and Sherborn’s ‘ Catalogue of
British Vertebrates,’ brought out by the same publisher, and re-
viewed in the Ann. & Mag. Nat. Hist. ser. 6, vol. v., 1890, pp. 337
&c. Such trustworthy exegetical catalogues of fossils as these are
much wanted. They clear the way for students and others; they
do much for the avoidance of error; and they save loss of time and
atience in looking for the history of known species and for the
probable relationships of newly-found fossils.
The Jurassic Rocks of the Neighbourhood of Cambridge. By the
late THomas Rosrrts, M.A., F.G.S. 8vo. Pp. vii and 96. C.J.
Clay and Sons. London, 1892.
Tuts memoir was the “Sedgwick Prize Essay” for 1886. The
Author, who had collected and discussed so many useful points in
the distribution and natural history of these Jurassic strata, unfor-
tunately died, at an early age, whilst adding new facts and perfecting
the views which he had advanced with care and _ perspicuity.
Lamenting his death and desirous that his good work should not
be lost sight of, some of his colleagues in the Woodwardian Museum
and ctker friends have put together the notes that he left and have
brought out this Prize Essay, so enriched, as a lasting memorial of
a geologist whom his many friends highly respected and wish to
honour.
The Oolites of Cambridgeshire and northwards differ from those
of the south in several particulars, on account of the two series
having been laid down on and against a ridge or ridges of Paleozoic
rocks, making shoals in the sea of the Jurassic period, and trending
north-easterly and then northerly. Hence not only does the strike
of the Oolitic strata vary in the East-Anglian district, but their
constituent deposits vary in character, both according to the local
depths of the sea and the kinds of material supplied by the organic
remains, and ty the sediments brought from the shores. ‘hus
“throughout the greater part of the period the deposits were laid
Ann. & Mag. N Bist. Ser. 6. Vol. xi. 14
194 Bibliographical Notices.
down under locally shifting geographical conditions, so that the
district was from time to time divided into different and changing
hydrographical areas, the sediments varying as barriers disappeared
or were introduced, and the forms of life more or less readily
yielding to the influence of external circumstances.”
Moreover the higher members of the Oolitic group, seen in the
south-west, are wanting in East Anglia, either not having reached
so far as deposits along the old sea-bottom or haying been removed
by denudation.
Particular points in the inquiry carried out by Mr. Roberts were
as to the exact geological value of the several bands in the Kimeridge
and Oxford Clays, and as to strata representing the Corallian series
of neighbouring districts. Here the careful collection and exact
determination of the fossils were most important; and chiefly by
means thereof the author came to the conclusion that the Oolites of
the district under consideration might be tabulated as follows :—
1, Kimeridge Clay pe
Upware Section.
2. Amptill Clay 4 Coral-rag.
Coralline Oolite.
3. Lower Calcareous Grit. (The Elsworth and St.-Ives
Rocks.)
4. Oxford Clay.
This last great formation is here represented by three palonto-
logical zones :—
1. Zone of Ammonites perarmatus (rare), A. crenatus, A. ocu-
latus, and the Cordati group of Ammonites of the
St.-Ives clay-pit.
. Zone of Waldheimia impressa, at the base of the St.-Ives
clay-pit.
3. Zone of Ammonites Duncani and A. Jason (the Ornati
group of Ammonites) of St. Neots. [Lowest zone. |
bo
The valuable, because trustworthy, lists of fossils from the several
bands of clay and stone and the full stratigraphical description of
other strata,—the careful references to foregoing observations and
descriptions,—and the correlation of the Oolites of Cambridgeshire
with those of other English districts (pp. 77-86) and with foreign
equivalents (pp. 87-94), render this Memoir of very great value to
all concerned with the physical geography, the geology, and the
paleontology of this classic ground and of the corresponding regions
in France and Germany.
Miscellaneous. 195
The Tertiary Fauna of Markusevee in Croatia. [Fauna fossile
Terziaria §c.| By S. Brustva. 8vo. I8 pp. Zagreb (Agram),
1892. [Reprint from the ‘Glasnika Hevats. Naravosl. Druztva,’
vii. Godina. |
Turs memoir, published by the Natural-History Society of Croatia,
is a continuation of the results of the researches of Spiridion
Brusina on the paleontology of Dalmatia, Croatia, and Slavonia,
more especially the Tertiary Mollusks (largely of estuarine or
brackish character). ‘The Director of the National Museum has
devoted much time and energy to this work, some results of which
were noticed in the Ann. & Mag. Nat. Hist. for June 1875; and
other results have been since published in Austria-Hungary.
Some fossils collected by him from asandstone of the ‘“‘ Congerian
Formation” at the ‘ Kelekovo polje,” not far from MarkuSevec,
were found to comprise some vertebre, teeth, and otoliths of fishes,
fragments of Polyzoa, numerous valves of Ostracods, tubes of Anne-
lides, spines of Echinoderms, spicules of Sponges, and many Forami-
nifera. This collection is so important to local and other geologists
that a full list is given of its species of Limaw, Helix, and Succinea
(few), 2 of Limnea, 8 of Planorbis, 1 of Melania, 17 and more of
Melanopsis, 2 of Melanoptychia, 26 and more of the Hydrobiide, 7 of
Valvatidee, of Orygoceras 4, Cyclostoma 1, Neritona 1, Neritodonta 6,
Congeria 7, Limnocardium 5, Pisidium 1, Of these several are new
species, most of which are described in this memoir, and some are
species not yet determined sufficiently to be named.
This group of fossil] mollusks is carefully compared in detail with
those of other localities in Eastern Europe and with the fauna of the
Caspian and of Lake Baikal; and the Author acknowledges his
indebtedness for help an1 cooperation to his several friends and
fellow-workers.
Mollusks from the “Sarmatian Formation” at Markusevec are
enumerated at pages 80-82; and an Appendix gives a list (with
localities) of the fossil Dreissensidee of Dalmatia, Croatia, and
Slavonia, namely 20 species of Congeria and 7 of Dreissensia.
An Index of the fossil species contained in Parts I. and III. of
S. Brusina’s work then follows, pp. 89-96, and of the special recent
representatives at pp. 97 and 98.
MISCELLAN¥OUS.
On a Sporozoon parasitic in the Muscles of the Crayfish.
By MM. F. Hennuevy and P. Tuéronay.
We have recently * reported the existeuce of a Sporozoon living
as a parasite in the muscles of Crangon wulyaris and very closely
* Henneguy and Thélohan, “Sur un Sporosoiire parasite des muscles
196 Miscellaneous.
allied to that which one of us * had already described as occurring
in Palemon serratus and P. rectirostris.
We have since had occasion to observe in the muscles of the
Crayfish a parasite which likewise belongs to the group of the
Sporozoa, but which exhibits somewhat remarkable differences from
the foregoing forms.
Thanks to the kindness of M. Contejean we have keen able to
examine fragments of Crayfish muscles from the Department of the
Doubs, which in the fresh state exhibited that remarkable opacity
to which we have already drawn attention in Crangon and Palemon.
In sections of these muscles we have found that the fibres are
attacked by a parasite in different stages of development.
At certain points the muscle-tibre is found to be crammed with
little ovoid spores having a clear vacuole at their large extremity.
The appearance of these spores reminds us of those of the Micro-
sporidia, of Glugea, and of the parasites of Crangon and Palemon.
They are much smaller than the spores in Crangon and in size
approach those in Palwmon.
By the side of these spores younger stages of the parasite are
found, represented by nucleated masses of protoplasm. Our obser-
vations, though they are as yet very incomplete, have nevertheless
enabled us to determine that this organism differs from the parasite
of the Crangon and of the two species of Palemon in the mode of
the development of the spores. These, instead of being formed in
agroup of eight in each sporogenous vesicle, are produced in variable
but always larger numbers. In this character the parasite of the
Crayfish approaches the Microsporidia and certain forms of the
Myxosporidia.
As regards the group to which this parasite ought to be assigned,
since we have not had fresh material at our disposal we have not
been able to study the structure of the spores sufficiently thoroughly
in order to pronounce an opinion upon the point.
The presence of this sporozoon in the muscles of the Crayfish
offers a double feature of interest: it causes our first observations
to extend to a larger number of species of Crustacea, and, moreover,
it seems that it ought to be of a certain practical importance from
the point of view of the etiology of the disease which for several
years past has destroyed the Crayfish of our watercourses in the
east.—Comptes rendus des Séunces de la Société de Biologie (Séance
du 30 juillet, 1892): from a separate impression communicated by
the Authors.
des crustacés décapodes,” Comptes rendus de la Société de Biologie,
June 25, 1892 (Ann. & Mag. Nat. Hist. ser. 6, vol. x. pp. 342-344),
* Henneguy, “Note sur un parasite des muscles du Palemon recti-
rostris,’ Mémoires publiés par la Société philomathique & Voccasion du
centenaire de sa fondation (1888).
PERSEPHONASTER CROCEUS. ABACTINAL VIEW.
Ann. & Mag. Nat. Hist. S. 6. Vol. Bly Fb. 496
VIEW.
ACTINAL
CROCEUS.
PERSEPHONASTER
Ann. & Mag. Nat. Hist. S. 6. Vol. XI, Pl. V.
DICTYASTER: XENOPHILUS.
1 ABACTINAL SURFACE
wr ae
AD 5
NS
77
Vj
WMA UA.
py hla
,
i}
i eh
4
ig
as
: ae
oy D\
Li aan a Z)
eee 4
TMG, SB
ta AD i AEA yl ES
Zs 2PUNAGS
5 2 eS
Bi oT xi r
Sy ahh. ie
SO. SWE Ra Gp
«Se %
oS pe
Li ye La £
ces
IPSACASTER Sllpacp, DICTYASTER XENOPHILUS.
-ACTINAL- SURFACE.
bs
&
G
Mig
ee
a gx
s TER n ‘ 4 ad i
4 )
B
Cc I
i Al
IPS
A AS
SLA
. ACT
Ann. & Mag
IPSACASTER SLADENI. ACTINAL SURFACE,
See etest, S. 0. Vol. XI, Fl. V.
CTYASTER XENOPHILUS.
ABACTINAL SURFACE
DITYASTER XENOPHILUS.
ACTINAL- SURFACE.
MI
\i \ Sree: M IRR Ks SOS
os oie RNG
seca tasmte eae any tes BOER RTO
ae DE 4 » / ja ae rats ,
5 < eS. HS Ape Ae sa aay Sere) Vets
<2 ating es Cah,
POS aN SSS <) ‘Ai Sark B
N
agar ieee > td A Lea SALTY
Y EEA PANS Pe AS NN.
SABRES Rag IE NL VTL ety Seta SN
Mer AL ae TES,
ae Pe Seso-774 SS
=a) 2 -
— ‘Val canle es 2S SSS a —
ay ‘a WE ila dees) =
ait iss as << pein
oe EN ee <3 SODA, oe
Rae ee es Aoi
sieee ee 32 ae Lapis; oa
eaneee a
vs
MILTELIPHASTER WOOD-MASONI.
ABACTINAL VIEW.
Ann. & Mag. Nat. Hist. S. 6. Vol. XI, Pl. VI.
Fig. 7
Oey
¥ ™~ awa wr, D.
Salinas ae ae
ee With Gi
if Mi,
sy
IPHASTER WOOD-MASONI.
OF THE ADAMBULACRAL SPINES. (x5)
UL
* yal? \s
2}
¢
‘
3
a
=
+.)
7 \e
b hee
=~
TAN
‘)
4
4 ee
i
u
h
yb
sae @
et
-
YN
oe,
—s
4ik 3
WES
“~
ae
tr; ~
Pe
x,
A
ly
SUA yi
Ne i Zire aS Bo OO
a Lv aly we an? es
: PON? OY
SEW a
Yee
\
?
I
\:
ys
(i
x
aN
p>
WOOD-MASONI.
VIEW.
MILTELIPHASTER
ACTINAL
3. G6, Vola. PU.I. .
f MEK MMM Mii:
! a
= 3 x ) ema pear om Y nett} E
nt emf nied hr ik il i as ik i ek es i id ss
Je a) et E91 ry ro On ert one
Sat gad a
we
CUCL Te UT UT Ie
SARA RRA AR :
i Loma |
Annick Magy.. Val Hist.
Ann. & Mag. Nat. Hist. S. 6. Vol. XI, Pl. X.
PSATHYROCARIS FRAGILIS.
ons = 2h .
~*~
Af,
heen ehels
Vol.
ri eal
IVGEs “L11Sbe
cw Mag.
Ann.
\
\\\
——"
—
——
SEB
———
\ \ \\
MSS AN INNIS
\
ZI
tpyfpLe:
LEY) YY
i
R
y
5
ee
6x4.5
PSATHYROCARIS FRAGILIS.
rn.d May. Nat. Hist. 8.6. Vol AT. PL. XL
J3.E.SM. dei. M-P Parker lth. Mintern Bros. imp.
THE ANNALS
AND
MAGAZINE OF NATURAL HISTORY.
{SIXTH SERIES.]
No. 63. MARCH. 1893.
XXIX.—The Affinities and Origin of the Tardigrada.
By Prof. J. von KENNEL *,
LiKE the majority of the so-called aberrant groups in the
Animal Kingdom the Tardigrada have met with the most
diversified experiences in systems of classification. At one
time they were ranked among the ‘‘ Worms,” at another
among the Arthropods, and they have been assigned at diffe-
rent periods both to Crustacea and Tracheata. They were
treated as an “appendage” to these groups or else as
“ancestors” of them, and were regarded as degenerate or
greatly reduced or as highly primitive forms. A comparison
of the various views upon this question has been furnished by
Plate t, who also discusses in a subsequent paper ¢ a theory
by Biitschli § which had previously escaped his notice.
Now I am in entire agreement with Plate’s view that the
Tardigrada can be brought into relationship only with the
tracheate Arthropods, and do not consider it necessary to
support or amplify the arguments which he advances against
* Translated from the ‘Sitzungsberichte der Naturforscher-Gesellschaft
bei der Universitat Dorpat,’ ix. Bd. Heft iii. (Dorpat, 1892), pp. 504-512.
+ Plate, “ Zur Naturgeschichte der Tardigraden,” Zool. Jahrb. Bad. iii.
¢ Plate, “ Die Rotatorienfauna des bottnischen Meerbusens &c.,”
Zeitschr. f. wiss. Zool. Bd. xlix.
§ O. Biitschli, “ Untersuchungen iiber freilebende Nematoden und die
Gattung Chetonotus,” Zeitschr. f. wiss. Zool. Bd. xxvi.
Ann. & Mag. N. Hist. Ser. 6. Vol. xi. 15
198 Prof. J. von Kennel,on the
their being allied to the Crustacea. Noten this,
however, with regard to the systematic position of these
animals I have arrived at quite a different conclusion, which
more nearly resembles the older ideas, and regards the Tardi-
rada as degenerate forms of higher Tracheates. Plate
Hecke that “ the Bear-animalcules are the lowest of all the
air-breathing Arthropods with which we are at present
acquainted, and must be placed at the bottom of the Tra-
cheata, even below the Onychophora.” Nevertheless, as may
well be imagined, he does not derive Peripatus from them,
but states that “they form an offshoot of the great Tracheate
stem, which, however, lies much nearer the root of the latter
than any other branch of that stock. They are the group
in which the transition from the Annelids to the air-breathing
Arthropods is most clearly expressed and most distinctly
recognizable.”
To this opinion I am unable to assent. In the species of
Peripatus we have animals which furnish the best transition
between Annelids and Tracheates, and I have elsewhere *
explained at length the reasons why it is probable that forms
resembling Pertpatus were the ancestors of the Myriapod-
like Tracheata, trom which again the rest of the Tracheata
are descended. The bodily form, the organs of locomotion,
the commencing formation of head and jaws, the nervous
system, the eyes, the sexual organs, and the trachez of this
group of animals may without difficulty be brought into agree-
ment both with the conditions of the Annelids as also with
those of the Tracheates, and, in addition, we have the seg-
mental organs, which have been transmitted from the
Annelids.
Conditions are different among the Tardigrada, which
present far less resemblance to the Annelids. In the first
place we are acquainted with no Annelids whose bodies con-
sist of so few segments as is the case among the Bear-
animalcules. It is true that such might have existed, or the
number of the segments might have been subsequently
reduced. But the Tardigrada do not possess even a trace of
true segmental organs, but are provided with Malpighian
vessels in their stead, and consequently in this respect they
must have receded far more from the Annelidan ancestors
than even the existing species of Peripatus ; they are in this
point much more Arthropods than the latter. The muscu-
* Kennel, “Die Verwandtschaftsverhaltnisse der Arthropoden,”
Schriften herausg. yon der Naturforscher-Gesellschaft bei der Universitat
Dorpat, vi.
Affinities and Origin of the Tardigrada, 199
lature of the Bear-animalcules, too, diverges from the dermal
muscle-sheath of the Annelids much further than does the
musculature of Peripatus ; they have isolated muscle-strands,
which run freely through the body-cavity, decussate, and are
inserted at different points of the integument; in accordance
with their small size these are of very simple structure and
consist of few fibres. This arrangement reminds us forcibly
of the distribution of the muscles in many highly developed
Tracheates, which will be discussed later on. The nervous
system of the Tardigrada can, it is true, be easily compared
with that of the Annelids, but it also differs in no way from
that of higher Tracheates, in which the sense-organs of the
head are slightly or not at all developed. At any rate in its
configuration as a chain of ganglia it surpasses that of Peri-
patus, which has a very irregular rope-ladder-like nervous
system. On the other hand, however, the eyes of the Tardi-
grada are such insignificant dots of pigment that we can
scarcely find more simple ones among the Annelids; they can
only be regarded as rudimentary structures, but not as engaged
in progressive development.
While the sexual organs of Peripatus can at once and by
means of embryological proofs be traced back to those of the
Annelids, and their ducts to segmental organs, this is not
possible in the case of those of the Tardigrada, since in the
first place they are unpaired, and, secondly, open dorsally into
the posterior portion of the intestine. This condition also is
more readily intelligible on the theory of reduction and dege-
neration, as I shall subsequently show.
If we regard the Tardigrada as an offshoot from the root of
the Tracheata, we certainly need make no attempt to think
out the conditions of their organization in their further deve-
lopment into Arthropods. If, however, they “most clearly
express the transition from the Annelids to the air-breathing
Arthropods,” the ancestors which were common to them and
the Tracheata would yet have to be supposed to have
resembled them fairly closely. But here great difficulties
crop up in relation to the appendages. The four pairs of
appendages of the Tardigrada are post-oral, and are inner-
vated from the ventral chain of ganglia, while the Tracheata
nevertheless possess a pre-oral pair of appendages in the shape
of the antenne.
Now this pair of appendages must either have disappeared
in the Tardigrada or it must be regarded as a new develop-
ment in the rest of the Tracheata; for the first pair of leg-
stumps of the Bear-animalcules is innervated from the first
ventral ganglion. Should we wish to interpret the absence
15*
200 Prof. J. von Kennel on the
of the antenne as due to reduction, which may well be the
only possible explanation, it would follow that the first three
pairs of appendages of the Tardigrada are homologous with
the mouth. parts of the Tracheata, and only a single segment
would be left for the body. This would entail the conclusion
that the rest of the trunk-segments of the Tracheates have
arisen in consequence of continued segmentation on the part
of the common ancestors of Tardigrada and Tracheata.
Although this would be an absolutely permissible suppo-
sition, we must nevertheless consider that in the Peripatus-
like Protracheata we have animals which, with a larger number
of segments, combine Annelidan characters even in a far
greater degree, and consequently fulfil all demands that can
be made upon transitional forms. Moreover all Myriapods,
from which the rest of the Tracheata can be derived by the
process of concentration of segments &c., possess altogether a
larger number of segments and a fully developed head, the
rudiments of which are already in process of formation in
Peripatus in the shape of antenne, jaws, and slime papille.
Let us now consider whether the peculiar conditions of the
Tardigrada cannot after all be derived more easily and simply
from higher Tracheates by the metlod of reduction and
simplification. Let us just take the Tardigrade body as we
see it—a segmented animal which, according to its nervous
system, consists of five segments. It must certainly be
admitted that we are acquainted with no Tracheate Arthropod
which is composed of so few segments. But we nevertheless
find in the entire Tracheate stem a reduction of the number of
segments from the Myriapods upwards, which is brought
about in part by the actual absence of posterior abdominal
segments, and in part by fusions and intimate union of
several segments. As a general rule among the higher
Tracheata four cephalic, three thoracic, and a somewhat
fluctuating number of abdominal segments are present; the
latter, however, in the Mites and the true Spiders are fused
into one piece, which is greatly reduced in the case of the
former. A reduction such as this may have occurred in other
Tracheata also. Moreover, we are acquainted with stages of
Tracheates, namely their larve, in which the divisions of head,
thorax, and abdomen are not differentiated; Jarvee in which
the appendages have disappeared in adaptation to certain con-
ditions of existence (maggots) ; and, finally, larvee in which
new stumps of appendages, in the form of simple dermal
protuberances provided with claws of different shapes, have
appeared secondarily upon all the segments of the body or
upon a portion of them. I merely recall the caterpillars with
Affinities and Origin of the Tardigrada. 201
their “ pro-legs ” and many dipterous larve with leg-stumps.
Most important of all is the fact that in dipterous larva of this
kind the entire head is wanting, since the four foremost seg-
ments of the body are invaginated and constitute the esophagus
of the larva.
In the metamorphosis of the larva all portions of the head
of the imago are formed in a reversed position out of rudi-
ments in the wall of the larval cesophagus, and are everted in
the pupa. That which is known to us as a maggot is merely
the trunk, externally wholly unsegmented, enveloped in a
resisting cuticle, and apodous or with a very variable number
of secondary leg-stumps. If we imagine the trunk of such a
dipterous larva reduced to four segments we have in essentials
precisely the body of a Tardigrade.
In the secondary cesophageal tube formed by invagination
of the four cephalic segments there lie in the case of the
maggots of Diptera the two stylets of chitin as masticatory
organs of local origin, not homologous with any appendages,
precisely like the chitinous rods in the cesophagus of the
Tardigrada. The musculature of a dipterous maggot is
certainly more complicated than that of a Tardigrade, but
exhibits so surprising a similarity in arrangement and distri-
bution that we are involuntarily impelled to institute a
comparison. It is true that the muscles of the Tardigrada
are not transversely striated, but this, in the light of present
views, will surely not carry much weight. In the larve of
many Diptera (Stratiomys, Mycetophilide, &c.) leg-stumps
occur in different numbers as independent new formations. I
myself when a student, some seventeen years ago, found
beneath a stone in the Black Forest a number of very peculiar
dipterous larvee, which I have never yet been able to deter-
mine and which bore upon the flat ventral surface four pairs
of such leg-stumps provided with claws. Salivary glands
and Malpighian vessels are present in the larve of Diptera.
How greatly the condition of the alimentary canal can alter
according to the mode of life is shown with the greatest
distinctness precisely by Insect larva. Thus we next come to
the trachee. These are, indeed, present in larve of Diptera,
but the number of the stigmata is considerably reduced; that
trachee, however, may disappear, especially in very small
Arthropods which live in damp surroundings, is shown among
others by the Acarina.
We have now only to deal with the nervous system and
sexual organs.
The nervous system varies exceedingly in dipterous larve,
even in those which we may designate as maggots. The
202 Prof. J. von Kennel on the
supra-cesophageal ganglion is, in accordance with the invagi-
nation of the first segments, placed far back, while the
ganglion-chain is sometimes exceedingly concentrated, forming
a short band-shaped mass, from the lateral margins and
hinder end of which the nerves radiate into the segments after
the manner of a cauda equina—and sometimes also a well-
developed chain. If we retain the last case and assume a
great reduction in the number of body-segmenis, the con-
ditions in the Tardigrades present no further difficulty.
Now, however, it is precisely among dipterous larvae that
we observe in a series of forms that they are capable of repro-
duction as larve (Cecidomyia). In these larve a head
is never developed: it remains after a fashion latent for
generations. The larval sexual organs, which are in this case
only female ones, are situated, in the shape of small paired
sacs, upon the dorsal side of the intestine and have no ers
the differentiation of the germ-cells into egg- and nutritive
cells appears to be abolished or at any rate not sharply
expressed. Now if we were to suppose that larve of this
kind had ceased to undergo their metamorphosis, that they
always reproduced their species as larve, and that, in conse-
quence of adaptation to the very peculiar conditions of
existence in damp moss and water, they had become modified
in one direction and had then undergone further development,
we might regard them as constituting a transition to forms
which we now know as Tardigrades—Arthropods in the larval.
stage, without a head, and with a body consisting of a reduced
number of segments, and bearing a few (secondary) leg-stumps.
In the course of this process a few peculiarities must naturally
have received especial emphasis. In the first place, not only
female but also male larve must have remained in this stage.
Now we actually find, as, for instance, in the case of Cecido-
myia, that after a series of peedogenetic and parthenogenetic
larval generations the spontaneous production of male larve
occurs, since finally both males and females appear as
imagines. The idea that under certain circumstances male
larvee of this kind also failed to undergo metamorphosis must
not be rejected offhand. We need simply and solely suppose
that the sexual organs gradually redeveloped a duct—in the
present case a short canal in communication with the rectum.
And this supposition is certainly not outside the limits of
what we otherwise concede to the capacity for modification
shown by the animal body. It is likewise conceivable that
independent efferent ducts, which were previously present,
came into communication with the rectum! through invagina-
tion of the posterior body-segments, while in this way the
Affinities and Origin of the Tardigrada. 203
reduction in the number of the segments would also become
more intelligible. The enigmatical “ gland,” which occurs
in both sexes of Tardigrades (cf Plate, loc. c7t.), is perhaps
nothing else than the degenerate second ovary or testis, just
as in the case of birds also, at least in the female sex, only
one half of the genital apparatus arrives at maturity.
Now it is by no means my intention to put forward dipterous
larvee as actually the ancestors of the Tardigrada; on the
contrary, I merely selected these larvee in particular because
they combine in themselves a series of peculiarities which
show how great the capacity for modification may be in such
animals, and because these peculiarities occur in a precisely
similar manner in the Tardigrada. Just as in the case of
Diptera such very far-reaching secondary changes were possible
in the larval stage in adaptation to certain conditions of
existence, so this might equally well have happened in the
case of the larve of other insects also of which we have lost
all knowledge. I merely mean that, of all tracheate Arthro-
pods with which we are at present acquainted, no single form
so simply and so readily enables us to interpret the Tardi-
grade body as these very dipterous larve. I do not believe
that the 'l'ardigrades can be placed at the root of the tracheate
stem or in the neighbourhood of it; for the conditions of their
organization diverge more from those of the Annelids than
do those of indisputable Tracheates of much higher rank. If
my memory serves me it was once declared by Ray Lankester
that in the case of animals of very small size but of relatively
complicated structure we must first direct our thoughts towards
degeneration and reduction from higher forms. If we derive
the Tardigrada in the manner indicated above from pedo-
genetic and greatly modified Tracheate larve we can regard
the entire body of these animals as an Arthropod trunk of
four segments, of which the head ceased to be developed, and
of the cephalic organs of which the supra-cesophageal ganglion
is the sole remnant. The first three ganglia of the ventral
cord, which in higher Arthropods are fused together to form
the subcesophageal ganglion and innervate the mouth-parts,
may very well in the course of time have degenerated and
disappeared, since the organs which they had to supply were
no longer developed. ‘The four ganglionic centres of the
Tardigrades may then correspond to three thoracic and one
abdominal ganglion. If we consider that the young of many
Myriapods leave the egg at a very early stage with quite a
small number of segments, and only develop the remainder
during free existence, we can also conceive that this may have
been possible several times, and that such immature larve
204 Mr. G. A. Boulenger on some Jurassic and
having ceased to develop at an early stage, and being re-
modelled atter the fashion of the dipterous larve which we
have been discussing, might have acquired the power of
pedogenetic reproduction. I readily admit that our hypo-
theses are somewhat many in number; but there is not a
single one among them which has not been actually observed
in the Arthropod phylum itself, and more frequently in combi-
nations. Whether the leg-stumps of the Tardigrades have
arisen by degeneration from Arthropod appendages of their
ancestors, or whether they may be new formations like the
pro-legs and claspers of the caterpillars, is a question which
is difficult to decide. Its solution, however, be it as it may,
needs to alter nothing in the whole conception.
If we once more briefly sum up the points of agreement
between the Tardigrades and greatly modified ‘Tracheate
Jarve, somewhat of the type of the maggots of Cecidomyia,
we find :—absence of a head, chitinous stylets in the cesopha-
geal tube, absence of any ciliated epithelium and of a dermal
muscle-sheath, musculature broken up into isolated cords,
supra-cesophageal ganglion and ventral ganglion-chain,
simple structure of the sexual organs, and, lastly, Malpighian
vessels. The differences depend upon further advanced
degeneration of the Tardigrades, and include :—small number
of the ganglia (disappearance of the parts of the subcesophageal
ganglion), smooth musculature, absence of trachez and circu-
latory organs, and the probable reduction of the one germ-
gland. As new formations we may perhaps regard the
efferent duct of the sexual organs and, at any rate, the leg-
stumps, if these are not an ancestral character,
Embryology as yet affords us no explanation ; besides the
development need no longer be of the typical Arthropod type,
but may have secondarily undergone great modifications.
XXX.—On some newly-described Jurassic and Cretaceous
Lizards and Ithynchoccphalians. By G. A. BOULENGER.
In a paper published two years ago (2), whilst dealing with
afew points in the osteology of He/oderma and the systematic
position of that genus of lizards, I ventured to express some
views on the probable phylogeny of the order Squamata,
which comprises the existing group of true lizards, chame-
leons, and snakes. I pointed out that the Cretaceous lizard
Hydrosaurus lesinensis, regarded by some authors as a member
Cretaceous Lizards and Rhynchocephalians. 205
of the family Varanide, agreed, so far as could be judged from
the figures published by Kornhuber, with Owen’s Dolicho-
saurus, and that the suborder Dolichosauria might prove to
be the ancestral group from which the Lacertilia, Pythono-
morpha, and Ophidia evolved. ‘This opinion was founded on
the archaic condition of the hind limbs and the number of
cervical vertebra ; the presence of the zygosphenal articula-
tion of the vertebra, present in all Ophidia and several
Lacertilia and Pythonomorpha, lent additional support to
this hypothesis.
As I expected, my views have not had the approval of
Dr. Baur, who, in a lately published paper on the skull of
Mosasaurs (1), adheres to his previously expressed opinion
that the Varanide, Mosasauride, and Helodermatide should
be grouped together as a suborder “ Platynota.”
With regard to the structure of the foot, he denies any
considerable difference between Kornhuber’s Hydrosaurus
lesinensis, which I referred to the Dolichosauria, and a true
Varanus. But unless he contests the correctness of Korn-
huber’s restoration of the metatarsals and propodials, his
statement does not refute my interpretation; the figures
which I have reproduced (after Marsh and Kornhuber) speak
for themselves. On the other hand, when he says that he
has “no hesitation to assume that unguiculated limbs can be
transformed into paddles with numerous phalanges,” I entirely
agree with him, and do not know that | have ever expressed
any opinion to the contrary.
His other argument is that there is no evidence for the
supposition that the number of cervical vertebre after having
increased in the Dolichosauria can have become gradually
reduced again until the Rhiptoglossan number five was
reached. If my critic admits, as I believe he does, that the
Rhynchocephalia are descended from the Stegocephala, which
have fewer than eight cervical vertebre, and that the Rhipto-
glossa are only an ultra-specialized branch of the typical
Lacertilia, he cannot well argue against the probability of
such a process of increase followed again by a reduction. In
fact, if he will refer to one of his previous contributions to the
phylogeny of the Reptilia, he will find that he has no diffi-
culty in assuming that the Chelonians, with eight cervicals,
may have been descended from Plesiosaurians with very
numerous cervicals, the latter having been, as he himself
admits, derived from short-necked forms. That he now holds
“All forms which show a greater or smaller number of
eervicals [to] have with very little doubt developed from forms
with eight cervicals”” shows that his views have uadergone a
206 Mr. G. A. Boulenger on some Jurassic and
considerable change since 1887, when, commenting on Parker’s
discovery of at least fifteen somatomes in the cervical region
of the embryo of Chelone, he regards the latter author’s
statement, that ‘“ This free suppression of segments suggests
a great secular modification by shortening of a form not
unlike a Plesiosaur,” as a “ proof of the affinity of the Testu-
dinata and Sauropterygia.”” What Dr. Baur proves with so
much assurance on one occasion he himself pretends to dis-
prove on the next, without even referring to the position he
has previously taken up.
Two recently published contributions throw fresh light on
the Jurassic and Cretaceous Squamata, and suggest some
further remarks on the subject.
The first of these contributions is a paper by Gorjanovié-
Kramberger (5), who, ignoring my previously published note
and reasoning from a different point of view, arrives at results
very similar to mine in dealing with the systematic position
of some Cretaceous lizards from Dalmatia.
He describes a new form, Adgialosaurus, which shows
points of affinity to the Dolichosauria, the Pythonomorpha,
and the Varanoid Lacertilia, and proposes to establish a group
named QOphiosauria to comprise the Aigialosauride and
Dolichosauride. It is needless to observe that the term
Ophiosauria must be superseded by that of Dolichosauria,
which is of older standing, although Kramberger appears to
be ignorant of its existence. His definition of the group is,
besides, deficient in truly diagnostic characters.
The Hydrosaurus lesinensis of Kornhuber is incidentally
dealt with, and the genus Pontosaurus is established for it in
the family Aigialosauride, which is stated to be distinguished
from the Dolichosauride by the number, 7 to 9, of cervical
vertebre. However, it seems clear to me, after reexami-
nation of the figure given by Kornhuber, that H. lestinensis
possessed about 15 cervical vertebrae, and I am still at a loss
to find how it is to be generically distinguished from Dolicho-
saurus. But this is a matter which cannot well be dealt with
without comparing the specimens themselves ; therefore the
genus Pontosaurus may be accepted provisionally, provided
it be not identical with Acteosaurus of H. v. Meyer or Adrio-
saurus of Seeley.
Aigialosaurus, of which the figure of a nearly perfect
specimen is given, is a remarkable lizard, with somewhat the
physiognomy of a Monitor or Varanus, but with the jugal in
contact with the postfrontal and closing the orbit behind,
shorter and stouter ribs, and limbs much of the same type as
in Pontosaurus, although more developed. The quadrate is
Cretaceous Lizards and Rhynchocephalians. 207
shown to differ considerably from that of the Varanide and
to agree very closely with that of Mosasaurs. Kramberger
is therefore fully justified in regarding this type as one of the
original stock from which the Varanoids and the Mosasaurs
were derived,
There are a few points in Kramberger’s description which
need criticism. First, as regards the number of cervical
vertebre : whilst admitting that, owing to the sternum not
being preserved, it is difficult to decide which is the first
dorsal vertebra (taking as such that which bears the first
sternal rib), the author assumes that only seven vertebre are
to be reckoned as cervicals, his reason being that the scapula
in his specimen is situated on a line with the fifth to seventh
vertebrae. In a specimen of Varanus niloticus which I have
before me I find that the scapula corresponds to the sixth and
seventh vertebre, and yet nine cervicals exist ; besides, the
last cervical is a little shorter than the first dorsal, the differ-
ence between the two being about the same as represented in
d.2 and d.3 of Kramberger’s figure. I would therefore say
that Azgialosaurus had nine cervical vertebre, or even ten in
the event of the atlas having been overlooked.
A second criticism I have to make is with respect to the
importance attached by Kramberger to the great development
of the cervical autogenous hypapophyses of his reptile as
differentiating it from existing lizards ; for on the five anterior
vertebre of the Agamoid Physignathus Lesueurii I find them
quite as long as in Adgialosaurus, and other recent lizards
approach this condition.
An interesting point in the specimen figured is the presence,
to which, however, no allusion is made in the text, of double
parapophyses to the second sacral and the first two caudal
vertebree, thus representing the well-known “lymphapo-
physes”” of snakes. On this occasion I would remark that
Dollo, in a recent contribution, is entirely mistaken when he
thinks that the lymphapophyses of snakes and apodal lizards
represent the combined ribs and hemapophyses. A glance
at the skeleton of a viper, to mention no other examples,
shows that the lymphapophyses may coexist with the paired
hypapophyses on one and the same vertebra. His statement,
“les lymphapophyses ne coexistent jamais sur la méme
vertébre, soit avec les cétes, soit avec les hemapophyses,” is
therefore erroneous ; and his deductions, so far as this point
is concerned, consequently fall to the ground.
We have so long been ignorant of any undoubted pre-
Tertiary Lacertilian in the restricted sense, that much
importance attaches to the description of the Upper Jurassic
208 Mr. G. A. Boulenger on some Jurassic and
Euposaurus Thiollierit, Lortet, which has just appeared in
Dr. Lortet’s splendidly illustrated memoir on the fossil
reptiles of the Rhone Basin (6). Although the fossil is unac-
countably referred to the Rhynchocephalia, and even to
the family Sphenodontide, which, in the: French author’s
classification, includes //oma@osaurus, there can be no doubt
that we have here to do with a true lizard, as is evidenced
by the absence of a quadrato-jugal arch and of a plastron.
The pleurodont dentition, the absence of supra-temporal
fosse, the non-dilatation of the clavicles, are characters which
approximate Huposaurus to the Anguide. ‘The interclavicle
(‘sternum ”’ of Lortet) is unfortunately not preserved. A
curious oversight is noticeable in the description of this lizard,
the fifth toe being described as the hallux, which is thus
stated to be opposable to the other digits, whereas in reality
the pes does not differ from that of an ordinary lizard.
Of still greater interest is Lortet’s account and figure of
Pleurosaurus Goldfussti, H. v. Meyer, likewise referred to
the Sphenodontide. It is, however, quite clear that the
cranial characters are not Rhynchocephalian. ‘he temporal
arch appears to be essentially of a Lacertilian type and to
correspond with what is found in the Agamide. But the
structure of the limbs is primitive, agreeing in the tibia and
ulna and the metatarsals with the Dolichosauria and Protero-
sauria; and as the specimens described by H. v. Meyer show
a plastron in the form of fine riblets, which are, however, not
preserved in Lortet’s specimen, Pleurosaurus should be
regarded as the type of a distinct order of reptiles, combining
characters of the Proterosaurian Rhynchocephalia and Squa-
mata, for which the name Acrosauria, proposed by H. v.
Meyer in 1860, may be used.
The number of cervical vertebra: in Pleurosaurus is stated
by Lortet to be only five. I have to repeat the criticism
made above respecting Kramberger’s Azgialosaurus, and to
add that the first rib-bearing vertebra does not represent the
atlas ; this vertebra is not even entirely concealed in Lortet’s
specimen. Two small bones visible behind the occiput are,
in my opinion, the neuroids of the atlas. By further adding
to the neck the two vertebra named by Lortet first and second
dorsal we have eight cervicals instead of five.
Dr. Lortet’s memoir is also rich in information respecting
the Rhynchocephalian genera Hommosaurus and Sauranodon
(which name must yield to the prior Saphaosaurus). The
latter genus was very imperfectly known; but the beautiful
figures and the detailed description now published leave little
to desire, although some important characters shown by the
Cretaceous Lizards and Rhynchocephalians. 209
figures are not alluded to in the text. A new family, Saura-
nodontes, is established by Dr. Lortet for its reception, and
is chiefly founded on the total absence of teeth and the pro-
ccelous vertebre. In dealing with the latter character the
author curiously contradicts himself, for in the definition of
the family (p. 29) the vertebral centra are stated to be con-
cave behind, whilst further on (p. 53) the reverse is described.
That the latter statement is the correct one is shown by the
figures on pl. iii. The skull, as in the Rhynchosauride, has
no parietal foramen, and the bones described as the posterior
portions of the parietals appear to be the supra-temporals,
distinct from the squamosals.
The position of this new family in the system is indicated
in the following revised scheme of the classification proposed
by me in 1891 (3). The Champsosauride, first included in
the Rhynchocephalia vera, are now shifted to the Protero-
sauria, in accordance with the recent researches of Dollo (4),
who has shown these reptiles to be related to Proterosaurus.
Order RHYNCHOCEPHALIA.
Suborder I. PROTEROSAURIA.
Each transverse segment of the plastron composed of numerous paired
pieces. Pubis and ischium plate-like. Fifth metatarsal not modified.
A. Nasal openings distinct.
Vertebree conically excavated at either end,
with persistent notochord, all with inter-
vertebral hypapophyses; limb-bones with-
out condyles; humerus with entepicondylar
TE on An a Oe cere phe ae 1, PALZOHATTERID £.
Vertebre fully ossified, cervicals opisthoccelous,
dorsals biconcave ; no hypapophyses be-
tween the dorsal vertebre ; fot Hondd
with condyles; humerus with ectepicon-
dylar foramen or groove .........56 seee 2, PROTEROSAURID.
B. Nasal opening single; vertebre fully
ossified, feebly biconcave; no hypapo-
hyses between the dorsal Seba
umerus with ectepicondylar groove .. 3. CHAMPSOSAURID™.
Suborder I. RHYNCHOCEPHALIA VERA.
Each transverse segment of the plastron composed of three pieces, a
median angulate and a pair of lateral. Pubis and ischium elongate and
fifth metatarsal modified, as in the’ Lacertilia.
A. Jaws toothed ; vertebree amphiccelous.
a. Nasal openings distinct ; mandible with
coronoid process, the rami not united
by suture. Vertebrae deeply biconcave.
210 Messrs. T. and A. Scott on some
Humerus with ectepicondylar and entepicon-
dylar foramen; ribs with uncinate pro-
cesses ; all the vertebrae with intercentral
hypApophySes :sa:< 6G ens iah hea wees 6089 4. Harreriip2.
Humerus with entepicondylar foramen; ribs
without uncinate processes; no hypapo-
physes between the dorsal vertebree...... 5. Homaosaurip™.
6. Nasal opening single. Mandible with-
out coronoid process, the rami united
in a solid symphysis ; vertebrae feebly
biconcave ; no hypapophyses between
the dorsal vertebrae. Humerus with
ectepicondylar foramen or groove.... 6, RHYNCHOSAURID.
B. Jaws toothless. Vertebre procelous.
Mandible without coronoid process, the
rami united in a solid symphysis. Hu-
merus with ectepicondylar foramen .... 7. SAURANODONTID2®.
References.
(1) Baur, G.— On the Morphology of the Skull in the Mosasauride,”
Journ. of Morphol. vii., 1892, pp. 1-22, pls. i. and ii.
(2) BouLtencer, G, A.—“ Notes on the Osteology of Heloderma horri-
dum and H., suspectum, with Remarks on the Systematic Position
of the Helodermatide,” Proc. Zool. Soc. 1891, pp. 109-118.
(3) BouLtencrrR, G. A.—“ On British Remains of Homeosaurus, with
Remarks on the Classification of the Rhynchocephalia,” ¢. e.
pp. 167-172.
(4) Dotxo, L.— Nouvelle Note sur le Champsosaure,” Bull. Soc. Géol.
Belg. v., 1892, pp. 5-53, pls. vi.—viii.
(5) Gorsanovr¢-KRraMBERGER, C.—“ Aigialosaurus, eine neue Eidechse
aus den Kreideschiefern der Insel Lesina, mit Riicksicht auf die
bereits beschriebenen Lacertiden von Comen und Lesina,” Glasnik
Soe. Hist.-Nat. Croat. vii., 1892, pp. 74-106, pls. iii. and iv.
(6) Lorret, L.—* Les Reptiles fossiles du Bassin du Rhéne,’’ Arch.
Mus. Hist. Nat. Lyon, v., 1892, 139 pp., 16 pls.
XXXI.—On some new or rare Scottish Entomostraca. B
Tuomas Scorr, F.L.S., Naturalist to the Fishery Board
for Scotland, and ANDREW Scort.
| Plates VI. & VIII.)
PARARTOTROGUS, gen. nov. (provisional name),
Anterior and posterior antenne and mouth-organs as in
Artotrogus, Boeck, except that the siphon is rudimentary.
First pair of swimming-feet with both branches two-jointed ;
new or rare Scottish Entomostraca. 211
third pair with both branches three-jointed ; fourth pair with
outer branch three-, inner branch two-jointed; fifth pair
rudimentary.
This genus has been provisionally instituted to include a
species closely related to Artotrogus, but from which it is at
once distinguished by the structure of the swimming-feet and
the rudimentary siphon.
Parartotrogus Richardi, sp.n. (Pl. VII. figs. 1-11.)
Length exclusive of tail-setee ‘5 millim. Seen from above
the first body-segment is subrhomboid, its greatest breadth
being equal to about one and an eighth times the length ;
forehead slightly produced, truncate. Anterior antenne
short, moderately stout, nine-jointed, the second joint con-
siderably longer than any of the others; the formula shows
the relative length of the joints—
1-6 — 6 10_ 13-4
1 —2 —3—4_5— 6 —7 —8— 9
The antenne when bent back upon the body are scarcely half
the length of the first segment. Posterior antenne four-
jointed, the third joint nearly twice the length of the preceding
one and three times longer than the next; armed with a strong
terminal claw, hooked at the apex ; a rudimentary one-jointed
secondary branch springs from the middle of the second joint.
Siphon very small. Mandibles small, stylet-shaped. Maxille
small, with two one-jointed spiniferous branches, one branch
much smaller than the other. Anterior and posterior foot-
jaws nearly as in Artotrogus magniceps, Brady. Both
branches of first pair of swimming-feet two-jointed, the last
joint of both branches (but especially of the inner branches)
dilated and bearing several plain sete; (?) third and fourth
pairs as in Lichomolgus fucicolus, Brady, but more slender and
with the marginal and terminal spines broadly dagger-shaped ;
fifth pair small, bilobed, furnished with a few apical sete.
Ovisacs two, large. No males were observed.
Remarks. Only three pairs of swimming-feet could be
observed, even after the most careful dissection of several
specimens; it appeared to be the second pair that was
wanting, as a considerable hiatus existed between the first
pair and the next, much greater than between the fifth and
the preceding pair.
This species was first known to us in 1889, having been
obtained in material dredged near Fidra Island, Firth of
Forth. Since that time it has been occasionally observed
212 Messrs. I’. and A. Scott on some
not only in material from the locality named, but also from
the “Fluke Hole,” off St. Monans. Towards the end of
June 1892 a quantity of dredged material from the vicinity
of Fidra was being examined, when a specimen carrying
two large ovisacs turned up—the only specimen with ovisacs
that has been obtained. The discovery of this specimen
set at rest to a great extent some doubts entertained by us
regarding the maturity of those previously observed.
We have named this species in compliment to M. Jules
Richard, of Paris, the eminent zoologist and student of the
Entomostraca.
Lichomolgus concinnus, Scott. (Pl. VIL. figs. 12-15.)
1892. Lichomolgus concinnus, Scott, Tenth Annual Report Fishery
Board for Scotland, part iii. p. 261, pl. xi. figs. 25-33,
This species was described (op. e/t.) froma single specimen,
a female, obtained in material dredged off St. Monans, Firth
of Forth, in the early part of last year.
Some time ago a specimen of Doris (?) tuberculatus was
taken in the neighbourhood of Granton; and while it was
being examined several copepod parasites were accidentally
observed in the vicinity of the branchial appendages ; the
thorax of the copepods was of pale whitish colour, but they
were otherwise nearly transparent and were almost undistin-
guishable from their surroundings.
A careful examination of some of the specimens showed
that the Dords parasite agreed with Lichomolgus concinnus in
all respects except that the distal angles of the fifth feet were
acute instead of being rounded, and that the abdomen was
slightly longer than is shown in the figure in the Fishery
Board’s Report ; but these differences, which might be due to
local causes, are comparatively unimportant.
No other specimen of the same species of Doris has since
been obtained; we are therefore unable to say whether this
Lichomolgus is commonly associated with the Doris or not,
but its strongly clawed posterior antenne and foot-jaws seem
to indicate that it is at least semiparasitic in its habits, and it
would be of some interest to know if it was confined to any
particular species of Doris.
The St. Monans specimen, from which the species was
described, may have become detached from some Doris during
the operation of dredging.
new or rare Scottish Entomostraca. 213
Morar *, gen. nov. (provisional name).
Somewhat like Cylindropsyllus, Brady, in general form and
structure, but the posterior foot-jaws are three-jointed; the
outer branches of the fourth pair of swimming-feet in the
female are similar to those of the second and third pairs, and
the fifth pair are two-branched and nearly as in Attheyella
eryptorum, Brady.
This genus is instituted, provisionally, to include an inter-
esting Harpactid from Loch Morar, Argyleshire, having
characters connecting the freshwater species Attheyella cryp-
torum, Brady, with the marine Cylindropsyllus levis, Brady.
Moraria Anderson-Smithi, sp.n. (Pl. VIII. figs. 1-14.)
Female.—Length exclusive of tail-sete *62 millim. (75 inch).
Body elongate-cylindrical. Anterior antennz short, mode-
rately stout, seven-jointed; the upper distal angle of the
fourth joint is strongly produced, and forms the base of a
stout olfactory appendage ; the last joint is rather longer than
any of the others. The relative length of the joints is shown
in the formula—
9 go 5710
Posterior antenne (fig. 5) small, three-jointed; a small one-
jointed secondary branch, with a few small apical set, springs
from the middle of the second joint. Mandibles with a mode-
rately broad biting part, armed with five cylindrical teeth and
a small seta; palp small, composed of a single two-jointed
branch. Maxille simple; primary branch broadly truncate,
with five moderately large apical spines ; secondary appen-
dage bilobed; the exterior and larger lobe bears a stout
terminal spine, plumose on the distal half; the smaller Icbe
terminates in a plain spiniform seta. Anterior foot-jaws
(fig. 8) short, stout, furnished with a terminal claw-like spine
and two marginal processes, each process terminating in a
moderately stout spine, and a curved spine-like seta plumose
on the inner edge. Posterior foot-jaws three-jointed, the last
joint being very short, and armed with a long terminal claw;
the proximal half of the inner margin of the second joint is
furnished with several short sete; a stout, setiferous, spini-
form appendage springs from the inner distal angle of the
first joint. Outer branches of the first four pairs of swimming-
* From Loch Morar, Argyleshire.
Ann. & Mag. N. Hist. Ser. 6. Vol. xi. 16
214 Messrs. T. and A. Scott on some
feet three-jointed, inner branches two-jointed ; both branches
of the first pair are of nearly equal length ; the other three
pairs, which are nearly alike, have the inner branches con-
siderably shorter than the outer ; all the branches are sparingly
setiferous, but the terminal and lateral spines are elongate and
taper gradually from a moderately broad base to the sharp-
pointed apex ; the inner branches of the first pair are fur-
nished with a very long subapical seta in addition to the
terminal spine. Fifth pair small, two-branched, nearly as in
Aittheyella cryptorum, Brady, but the terminal and marginal
sete are shorter and spiniform (fig. 13). The abdomen con-
sists of four segments, but the first is composed of two
segments coalesced. Caudal stylets about as long as the last
abdominal segment. Operculum small, subconical, apex
acuminate.
The male differs little from the female except in the form
of the anterior antenne, which are hinged and somewhat
dilated ; the upper margin of the third joint is produced near
the middle into a small lobe-like process, which forms the base
of acurved spiniform seta; a sensory filament springs from the
upper distal angle of the same joint. A stout conical process
with a slightly hooked extremity arises from the proximal
half and extends somewhat beyond the end of the first joint
of the inner branches of the second pair of swimming-feet.
The principal branch of the fifth pair is broadly truncate at
the apex and provided with two short, stout, terminal spines ;
the small secondary branch bears an elongate setiferous
terminal spine and two sete on the inner and outer margins.
The male abdomen consists of five segments.
Hab. Loch Morar (a freshwater loch in Argyleshire), in
material dredged in shallow water at the head of the loch and
also to the west of South Tarbet; specimens were more
frequent in material from the latter place than from the former.
‘This species is named in compliment to Mr. W. Anderson-
Smith, one of the directors of the Fishery Board for Scotland,
who, by his pen and otherwise, has done much to encourage
the a A of natural history in Scotland.
Note.—Besides Moraria Anderson-Smithi, now described,
several other interesting Entomostraca were observed in the
material from Loch Morar, two of which may be specially
mentioned here, viz. :— Cyclops Ewarti, Brady, first described
by Dr. Brady in the Sixth Annual Report of the Fishery
Board for Scotland from specimens obtained in the upper
reaches of the Forth in November 1887: this species has
not since been observed till now; and its occurrence in Loch
new or rare Scottish Entomostraca. 215
Morar confirms the opinion expressed by Brady, that it had
been carried down by some stream into the Forth. Adtthey-
ella cryptorum, Brady, a species described by Dr. Brady in
the ‘ Journal of Microscopical Science,’ 1868, from specimens
obtained among the gelatinous algz on the damp roof of the
pit-workings of the low main, West Cramlington Colliery,
near Newcastle. No record of any further occurrence of this
species has been observed ; and it is of some interest to find
Attheyella cryptorum in the waters of Loch Morar.
Specimens of these two species were sent to Dr. Brady, who
confirmed our identification of them.
EXPLANATION OF THE PLATES.
PLATE VII.
Parartotrogus Richardi, gen. et sp. 0.
Fig. 1. Female, dorsal view. x 126.
Fig. 2. Anterior antenna. X 253.
Fig. 3. Posterior antenna. X 253.
Fig. 4. Mandible. x 460.
Fig. 5. Maxilla. x 253.
Fig. 6. Anterior foot-jaw. xX 253.
Fig. 7. Posterior foot-jaw. x 253.
Fig. 8. Foot of first pair. x 253.
Fig. 9. Foot of ? third pair. x 190.
Fig. 10. Foot of ? fourth pair. x 190.
Fig. 11. Abdomen and last thoracic segment. xX 170.
Lichomolgus concinnus, Scott.
. Female, dorsal view. xX 43.
. Posterior foot-jaw, male. x 190.
. Foot of fifth pair. x 380.
. Abdomen and last thoracic segment. x 190.
y
Ss's'¢
7
OU Co bo
PuaTe VIII.
Moraria Anderson-Smithi, gen, et sp. n.
Fig. 1. Female, dorsal view. X 80.
Fig. 2. Male, lateral view. x 80.
Fig. 3. Anterior antenna, female. x 380.
Fig. 4. Anterior antenna, male. x 380.
Fig. 5. Posterior antenna. x 6500.
Fig. 6. Mandible and palp. x 760,
Fig. 7. Maxilla. x 760.
Fig. 8. Anterior foot-jaw. x 760.
Fig. 9. Posterior foot-jaw. xX 760.
Fig. 10. Foot of first pair. x 760.
Fig. 11. Foot of second pair, male. x 760.
Fig. 12. Foot of fourth pair. x 760.
Fig. 13. Foot of fifth pair, female. x 760.
Fig. 14. Foot of fifth pair, male. x 760.
16*
216 Mr. H. G. Smith on Four new
XXXII.—Descriptions of Four new Species of Butterflies from
Omet-shan, North-west China, in the Collection of H. Grose
Smith. By H. Grose SMITH.
Euthalia strephon.
Upperside. Olivaceous green. Anterior wings crossed
beyond the middle from the costal to the submedian nervures
by a pale greenish-yellow band, widest on the costa,
narrowest between the two upper median nervules ; a small
yellowish somewhat elongate spot near the costal margin and
a larger oval spot of same colour at the outer edge of the
band beneath it ; the space between and on each side of the
dark bars which cross the cell is also pale greenish yellow.
Posterior wings with a pale greenish-yellow curved band
following the contour of the outer margin from the middle of
the costa, gradually narrowing and becoming obsolete towards
the lowest median nervule; an oval dark ring crossing the
cell and an indistinct submarginal row of dark green hastate
spots.
P Underside, Olivaceous yellow. On the anterior wings the
yellowish-green band is more clearly defined and edged on
each side with black, narrowly towards the costa, gradually
and irregularly becoming broader towards the submedian
nervure, where it ceases. Posterior wings with the pale
greenish-yellow curved band as above, bordered on each side
with olivaceous green, the space round the spots in the cell
and on the outer margin being the same colour as the central
band.
Expanse of wings 23 inches.
Nearest to E. omeia, Leech, which it resembles in colour,
but the posterior wings are very different.
Five specimens were sent, apparently all males.
Euthalia trrubescens.
Male.— Upperside. Anterior wings with the basal half
dark green, almost black, the outer half paler and slightly
metallic, the veins, with streaks between them on the paler
portion of the wings; being the same colour as the basal half ;
the cell is crossed in the middle by an irregular crimson
bar, and there is another crimson bar, narrower and somewhat
indistinct, at the end of the cell. Posterior wings the same
colour as the basal half of the anterior wings, being paler
Species of Butterflies from North-west China. 217
across the disk and traversed by the dark veins with streaks
between, as on the outer half of the anterior wings; two dark
bars cross the cell, and there is a crimson elongate spot
parallel with the outer margin between the submedian nervure
and the lowest median nervule; the costal margin is pale and
tinged with a bluish shade.
Underside similar to the upperside, but paler. Anterior
wings with the crimson bars across and at the end of the cell
wider and more distinct, and a small black spot below the
median nervure at its junction with the lowest median ner-
vule. Posterior wings with two crimson bars crossing the
cell, two crimson spots below the costal nervure and upper
median nervule respectively, another crimson spot on the
costal margin near the precostal nervure, and another at the
base; on the outer margin at the ends of the dark streaks
between the veins is a row of crimson spots, those nearest the
anal angle the most distinct and those in the middle nearly
obsolete ; the inner margin from the base to the anal angle
is broadly edged with crimson. Antenna black, the collar
and palpi crimson.
Expanse of wings 2} inchies.
One specimen only.
Allied to 2. lubentina, Cramer, var. ludonia, Staudinger,
and Whiteheadi, Grose Smith.
Dichorragia nesseus.
Male.— Upperside. Dark bluish green. Auterior wings
with the cell crossed by three black bars, that nearest the base
nearly obsolete ; beyond the cell, across the disk between the
veins, is a curved row of indistinct greyish spots, the second
and third of which are elongate; a submarginal row of
sagittate grey markings, elongated inwardly to the extent of
about one third of the wings, those towards the apex being
more elongate and narrower than the others; there are no
spots on the outer margin. Posterior wings more bluish green
than the anterior wings and without grey markings, except
three small hastate lines near the apex, with grey lines at
their base, the lowest almost obsolete ; beneath these, on the
outer margin between the veins, is a series of black hastate
spots.
Underside. Anterior wings bluish black, with the spots
beyond the cell more distinct and the bars crossing the cell
and a small spot beyond violaceous ; the sagittate markings
as on the upperside, but more distinct. Posterior wings
olivaceous, the hastate spots at the apex as on the upperside,
218 Mr. T. D. A. Cockerell on a new
and the series of marginal black spots faintly and very narrowly
bordered with white.
Expanse of wings 2% inches.
One example.
Near to D. nesimachus, Boisduval, but considerably less
maculate.
Lethe sicelides.
Male.—Upperside brown. Anterior wings with a broad
darker brown indistinct band on the outer margin, and a large
triangular indistinct sericeous patch, eve its base on the
inner margin and towards its apex extending beyond the
end of the cell. Posterior wings without any tufts of hair
within the cell; a submarginal row of four dark brown spots
without any white centre, and surrounded by pale brown
rings, the spot nearest the apex and the next but one being
the largest.
Underside. Anterior wings with the cell crossed in the
middle by two dark bars, the inner bar thicker than the outer
and the space between being pale; a dark line at the end of
the cell; two indistinct spots underneath each other towards
the apex, with white centres and surrounded by pale rings;
the other markings on the anterior and those on the posterior
wings very nearly the same ason ZL. sicelis, Hewitson, but on
the posterior wings the spot (in the submarginal row of spots)
between the middle and lowest median nervules is much
smaller than in L. sicel’s, and the outer of the two narrow bands
which cross the wings is bifid at its lower end and further
apart from the inner band.
Expanse of wings 2} inches.
Three specimens.
Very like L. sicel’s in general appearance, but belonging to a
different section of the genus. The large sericeous patch on
the anterior wings and the absence of the tufts on the poste-
rior wings distinguish it from that species.
XXXIII.—On a New Species of Aplysiide from Jamaica.
By T. D. A. CocKERELL, F.Z.S., Curator of the Museum
of the Institute of Jamaica.
On January 19th Mr. Verona Carter brought to the museum
a remarkable Aplysiid which he had caught at Kingston.
The specimen was alive in a bowl of water, so I madea
description of it before putting it in spirit. It was well that
Species of Aplysiide from Jumaica. 219
I did so, as in alcohol (which was not too strong) it presents
a shapeless mass which would defy description. Probably it
is for this reason that the creature has not hitherto been
described.
Owing to the absence of a shell and other characters the
animal must be referred to the genus Acles’a, Rang, which
has been considered peculiar to the East Indian region.
Aclesia intrapicta, sp. 0.
Length about 4} inches. Body swollen, subglobose ; foot
flattened, posteriorly broad, terminally acute. Neck sub-
cylindrical, moderately thick. Anterior pair of tentacles
large, branched, antler-like, retractile. Posterior pair large,
cylindrical, somewhat tapering, hollow, with open truncate
ends, and with two whorls of spine-like soft lateral branches ;
these and the other tentacle-like processes on the body are
also retractile. On the middle line of the neck, between the
atk of tentacles, is a short but broad branched filament.
pipodia contiguous in the middle line, but with the anterior
and posterior parts separating alternately, forming wide
cavities, in respiration. ‘The anterior of these cavities serves
for inspiration, the posterior for expiration, and the whole
respiratory cycle takes about five seconds. Quite a jet of
water can be thrown from the posterior orifice. Sides of
epipodia and body with many branched processes, some short,
others long, the largest resembling the anterior cephalic
tentacles. On the sides of the epipodia are three longitudinal
series of these processes—one dorsal, one subdorsal, one latzral
or subpedal. Each row numbers four processes, and the rows
are so placed that, as a general rule, the processes of the
dorsal row are more posterior than the equivalent ones of the
lateral row. Sides of foot with many processes.
Colour.—Prettily marbled with black and pale grey,
dorsal portions of epipodia and sides of neck with most black.
Most of the tentacles or processes tinged reddish, the larger
ones mottled with white. Inside of epipodia grey with white
dots. Sole finely speckled all over grey and white.
Described from a living specimen.
The anatomy, so far as examined, agrees in all important
points with that of Aplysia. The narrow white fore-gat
enlarges rapidly to form the big gizzard, which is pale red in
colour. In this gizzard 1 found four (and a fifth rudimen-
tary) little bodies, more or less triangular in outline, about
5 millim. diam., colour pale yellowish brown. These, like
those described by Prof. Ray Lankester in Aplysia, are no
220 M. Julius Wagner on the
doubt for crushing the food. Posteriorly to the gizzard the
gut is grey and rather broad, winding round the large brown
liver. The genitalia are somewhat ordinary, but rather
curious for their bright colour, which suggested the specific
name I haveadopted. The albuminiparous gland and herma-
phrodite duct are pale ochreous yellow, as is usual, but the
gland has on one surface a large elongated patch of bright
red, which does not remain well in alcohol. ‘The ovotestis is
large and irregularly globular, yellow-green in colour, with
two blackish broad sulci. A strong ligament has its origin
Pi the ovotestis, close to the beginning of the hermaphrodite
uct.
It is perhaps this species that has sometimes been observed
here and taken for a true Ap/ysta. Gosse, in his ‘ Naturalists’
Sojourn in Jamaica,’ p. 55, and Mr. E. A. Andrews, in the
‘ Johns Hopkins University Circular,’ April 1892, both refer
to the occurrence of an Aplysia (species not identified) at
Jamaica.
Institute of Jamaica,
Kingstown, Jamaica,
Jan, 21, 1893.
XXXI1IV.—On the Embryology of the Mites : Segmentation of
the Ovum, Origin of the Germinal Layers, and Development
of the Appendages in Ixodes. By JuLIuS WAGNER, of St.
Petersburg *.
Our knowledge of the processes which take place in theovum of
the Mites during its development is very limited. ‘The causes
of this are to be found on the one hand in the small size of
the object, and on the other in the properties of the ovum,
such as the thickness of the chorion and the brittleness of the
yolk when hardened in alcohol, owing to the largeness of the
yolk-spheres.
Ixodes, upon which my investigations were conducted, is a
comparatively convenient subject—in the first place since its
development proceeds somewhat slowly, and, secondly, be-
cause its ova do not require any great amount of attention
and develop very well without especial precautions. When
an Jvodes has ouce ‘begun to deposit its ova it no longer stirs
from the spot, and takes no notice if disturbed, as is often the
* Translated from the ‘ Zoologischer Anzeiger, xv. Jahrg., no. 899
(August 29, 1892), pp. 316-320.
Embryology of the Mites. 221
case when clusters of the ova are removed for the purpose of
preservation. The deposition of the ova is a somewhat
lengthy process, and continues almost to the time when the
young emerge from the first batches ; one and a half months,
however, are requisite for development (in the northern
Caucasus). Towards the end of the period of oviposition the
tick is surrounded by a mass of eggs, which is twice or even
three times the size of its body.
This at first sight apparently paradoxical phenomenon is
due to the fact that the ova are heaped together in a loose and
not in a compact mass.
1. Segmentation of the Ovum *.—The segmentation of the
ovum of Jxodes differs from what was previously observed in
the case of the Mites, in that the cells in process of division
do not at first pass to the surface of the ovum (as is the case
in Tetranychus, according to Claparéde), though the yolk
does not divide at the same time (as was observed by Robin
and Megnin in Sarcoptide), since I found in the yolk four
and six cells in course of fission, and on one occasion nine
with resting nuclei. The process of segmentation conse-
quently belongs to the partially interlecithal type. In the
end the cells pass to the surface and form the blastoderm ;
no cells remain behind in the yolk.
2. Formation of the Endoderm.—A|though in the first stages
of development differences are noticeable between the indi-
vidual cells of the blastoderm, they are at all events not
pronounced and are connected by transitions in such a way
that it is impossible to describe them precisely: the con-
ditions, however, soon change. Certain cells are distinguished
by their large nuclei, which usually occupy an oblique posi-
tion with reference to the surface of the ovum; these nuclei
take a fainter stain from carmine than those of the ordinary
blastoderm cells, and contain in their interior from one to two
nucleoli, which stand out sharply and are not present in the
nuclei of the rest of the cells of the blastoderm. In one and
the same series of sections, but still better in series which
follow one another in succession, we observe how cells of this
kind recede from the surface into the interior of the yolk
and how the surrounding cells of the blastoderm close up above
them. ‘These cells constitute the so-called yolk-cells. As
* Before the segmentation I also observed the extrusion of two direc-
tive corpuscles ; in Zvodes this takes place prior to the deposition of the
ova.
222 M. Julius Wagner on the
regards the region where the yolk-cells are differentiated, no
regularity whatever can be detected; they are formed over
the entire surface of the ovum. At the same time, however,
certain of these cells form a cluster on the dorsal side of the
ovum nearer the posterior end. They appear not to partici-
pate in the formation of the yolk-cells, but represent the
remaining portion of the endoderm.
3. Formation of the Mesoderm.—At the stage which imme-
diately precedes the appearance of the appendages mesoderm
cells become visible at the sides of the above-mentioned
endodermal cluster. From this we see that although this
group of cells, which in its position corresponds with the
cumulus primitivus of the spiders, does not represent the sole
spot where the endoderm cells are formed, on the other hand
it is just here that the mesoderm cells arise, which extend
hence over the posterior end of the ovum on to the ventral
surface, and along this towards the front. As regards the
nature of the origin of the mesoderm cells, I find by a careful
examination of the sections that at this spot no division in a
tangential direction takes place; that is to say, that we here
get an immigration of cells. On each side of the cluster of
endodermal cells we find upon the surface of the ovum in the
early stages of the development of the mesoderm a furrow-
like depression, at the bottom of which an active immigration
of the cells takes place. ‘These depressions appear to me to
correspond to the lateral margins of the germinal streak of
Insects (Heider, Hydrophilus).
4, Segmentation of the Yolk.—As I have already indicated,
no division of the yolk is observable during the segmentation
of the ovum. Nevertheless when fresh ova are examined the
yolk in the stages of the development of the appendages does
not present a homogeneous structure; on the contrary, it is
differentiated into separate polygonal portions of large size,
which are divided from one another by clefts. In later stages,
when the appendages attain a considerable length, this appear-
ance stands out with especial distinctness. I was able to
detect it later on in sections also. ‘Thus we see that the yolk,
subsequently to the migration of the yolk-cells, which are
present at first in inconsiderable numbers (about thirty), splits
up according to the number of these into separate portions,
which are still preserved in the intestine of the larva shortly
before its escape from the egg.
5. Development of the Legs.—The germinal streak of the
Mites, like that of the Spiders, at the stage of the appearance
of the appendages takes up the greater portion of the cireum-
ference of the ovum, passing over on to the dorsal surface
Embryology of the Mites. 223
with its caudal and cephalic lobes. At this stage it already
consists of two ectodermal ridges, which are separated by a
band of flat cells and meet together at the ends of the ellip-
tical ovum. When examined from the side the appendages
appear very sharply defined. In addition to the usual three
pairs of legs, which are characteristic of the larvae of most of
the Acarina and especially of [xodide, the fourth and last
pair is also developed in the embryo *. In the course of the
development of the anterior legs this pair elongates and even
exhibits an indistinct segmentation. Shortly before the larva
is hatched it begins to degenerate rapidly ; it decreases in
length and becomes, so to speak, drawn in, so that soon no
trace of it is left on the outside. But sections of the larva
after its escape from the egg show that beneath the integu-
ment of the body on both sides, in the region of the transition
of the mid-gut into the hind-gut (the excretory vesicle), there
lies a cluster of cells, which represents the remains of the
fourth pair of legs. From these clusters of cells also the
fourth pair of appendages appears to develop on the transition
of the larva into the nymph.
6. Segmentation of the Abdomen and its Appendages.—In
the early stages of development the mesoderm consists of
isolated groups of cells corresponding to the appendages ; the
mesoderm cells completely clothe the cavity of the appen-
dages, but this cavity remains open to the yolk; the meso-
dermal elements are also localized in a similar manner behind
the appendages, that is in the abdomen, since the fourth and
last pair of legs must be regarded as appendages of the last
thoracic segment. Behind the legs lie from five to six meso-
dermal groups. I did not determine their number precisely,
because the sixth group passes quite gradually into the meso-
derm of the caudal lobes. The internal segmentation in the
region of the abdomen is consequently expressed through at
least five segments. In the first segment, as in that of the
thorax, the mesoderm lies in a layer; in the segments next
following it forms a closed half-somite. Most developed are
the second, third, and fourth segments, on each of which
there is a projection on each side in the shape of a scarcely
perceptible tubercle. These tubercles can only be observed
in sectional preparations, and although they apparently do
not develop into more noticeable protuberances, they yet
without doubt represent structures homologous with the
abdominal appendages of spiders.
7. The Cephalic Appendages.—I can positively assert that,
* Cf. Winkler, Gamasus, in no, vii. of Claus’s ‘ Arbeiten.’
224 On a new Buprestid from Madagascar.
in front of the chelicere (cf Jaworowsky, Trochosa) in the
case of Jzxodes no rudiments whatever of appendages are
present. If nevertheless we observe the early stages of the
development of the appendages, it is not difficult to discover
that between the cheliceree and pedipalpi there lies yet another
pair of tubercles; consequently there are altogether three
pairs of tubercles—a pair of pfelicarte, a pair of pedipalpi, and
a pair which is situated between the two former. This latter
pair arises later than all the rest of the appendages, and,
judging from its position, bears a close relation to the pedi-
palpi. This is proved also by sections, since they show that
the mesodermal elements of this pair of tubercles form a direct
process of the mesoderm of the pedipalpi. In subsequent
stages no trace of the tubercles is to be found: I have not
yet elucidated their significance.
XXXV.—Description of anew Buprestid from Madagascar in
the Collection of the Hon. Walter Rothschild. By C. J.
GAHAN, M.A.
Chalcophoropsis Rothschild, sp. n.
Viridi-metallica; capitis fronte subplana, valde inequaliterque
punctata, supra inter oculos canaliculato-depressa ; pronoto for-
titer ineequaliterque rugoso-punctato, medio longitudinaliter sat
profundeque canaliculato, utrinque ante medium plaga levi, atro-
violacea, paullo elevata, in depressione magna flayo-pollinosa,
posita ; scutello inviso; elytris cupreo-purpurascentibus, nitidis ;
costis octo, paullo elevatis (quarum interna prope basin utrinque
brevissima) et punctis foveolatis numerosis albo-fasciculatis,
obtectis, lateribus prope apicem breviter denticulatis; corpore
subtus dense foveolato-punctato et albido-fasciculato; tarsis
posticis articulo primo quam secundo manifeste longiore ; anten-
narum articulis primo secundoque viridescentibus, tertio
quartoque nigris, tertio quam quarto vix longiore, hoe apice intus
compresso et dense punctulato (articuli ceteri desunt).
Long. 48, lat. (ad basin elytrorum) 15 mm.
Hab. 8. Madagascar.
Head strongly and unevenly punctured in front, densely
but much more feebly punctured behind; front rather flat
below and with a median groove above between the eyes.
Pronotum with a smooth dark violet and slightly raised area
on each side just in front of the middle and about midway
between the longitudinal groove and the lateral margin; this
On the Specific Rank of Limax cinereo-niger. 225
smooth plaga is surrounded by a much larger depressed area,
which is covered by a yellowish powdery pubescence and is
deepest behind, where it commences a little in front of the
basal margin, but becomes shallower as it widens out around
the smooth plaga, to narrow again in front, where it extends
close up to the antero-lateral angle. The sides of the pro-
notum are subparallel or slightly divergent from the base up
to the beginning of the posterior third, and thence converge
with a rather gentle curve up to the anterior border. Elytra
with numerous punctures, of which the larger ones bear each
a minute tuft of white hairs; the sides are sinuately emar-
ginate just in front of the middle, then slightly curve out, to
narrow again up to the apex, near which each is furnished
with four or five very short teeth; the innermost costa of
each elytron extends back but a short distance from the base
and approaches the suture, the second costa reaches from the
base to the extreme apex, the two outer cost are shorter and
less distinct. Body underneath densely foveolate-punctured,
the punctures bearing small fascicles of whitish hairs. Pro-
sternal process flat, produced and narrowed behind to an obtuse
point, which fits into a corresponding channel extending along
the whole length of the mesosternum ; the pits on its surface
are more or less elongated.
This splendid Buprestid, which is named in honour of its
possessor, to whom I owe the privilege of describing it, seems
at first sight to belong to the group of the Psilopterides ; but
its structural characters show that it is foreign to this group.
Taking all its characters into consideration, I do not see that
it can be better placed than in the genus Chalcophoropsis.
The scutellum, though invisible, has its position marked by a
small opening or depression at the base of the elytra.
XXXVI.—The Specific Rank of Limax cinereo-niger, Wolf.
By Wm. Denison Roesuck, F.L.S., Hon. Secretary to
the Conchological Society.
Tus slug, although very closely allied to LZ. maximus, L.,
differs so markedly from it in several particulars, both as
regards external and internal characters, as to make it
impossible for it to be looked upon as a mere colour-variety,
as is somewhat baldly asserted by Mr. Collinge in a note
in the ‘Annals’ for December last. The two slugs differ
in their dentition, in the anatomy of the genital organs,
226 Mr. W. D. Roebuck on the
and in the sculpture of the body, as well as in the coloration
of the foot-sole and the body and shield; and, although
somie of these differences are individually not great, yet
the sum or aggregate of them affords sufficient ground for
considering LZ. cinereo-niger as entitled to rank as, at the very
least, an incipient species, and for justifying malacologists in
awarding to it the specific or subspecific rank which many
authors give it.
To take the differences which exist, and first those in the
anatomy of the genitalia. The importance of the reproductive
organs of the Mollusca as a criterion for the distinction of
species is universally recognized, and an examination of the
very careful and accurate MS. figures of the genitalia of many
individuals of both L. maximus and L. cinereo-niger which
have been made by Mr. Charles Ashford, of Christchurch,
Hants, shows that there are differences worthy of note.
Mr. Ashford has found that the penis-sheath in L. maximus
is very constant in its form, the upper part being much
enlarged and peculiarly and rigidly flexed or bent upon itself,
while in ZL. cinereo-niger the penis-sheath is longer and of
tolerably equal width throughout, in which respect Mr. Ash-
ford’s figures tally with the one published by Schmidt. ‘The
sperm-duct in L. cinereo-niger is only very slightly attached
to the oviduct in a part of its length, but in L. maximus
moderate force is required to break the attachment. The
relative sizes of other parts, as the albumen-gland, the ovo-
testis, &c., in sexually mature specimens is constantly different
in the two forms. JL. cinereo-niger has a larger and less
deeply coloured ovo-testis, its average length in three speci-
mens from different localities being 29°6 millim., the shortest
one being 15 millim., while the greatest length Mr. Ashford
has noted in L. maximus is 18 millim. and the average in a
number of examples no more than 11 millim, L. cinereo-
niger has a smaller and narrower albumen-gland, its average
length in the same three specimens being 9°6 millim. and the
greatest length noted 13 millim., while in LZ. maximus the
average of sexually mature specimens is 20 millim., that of
all, including both mature and immature, 15} millim. The
difference ‘in the point of origin of the retractor muscle of
the penis is referred to by Dr. Scharff and corroborated by
other writers.
I am well aware that the differences of the two forms
anatomically are not so great nor of such importance as
F. Sordelli, whose paper was my authority for the statement
I made in the ‘ Journal of Conchology ’ in 1883, attached to
them ; yet that there are differences 1s quite certain from the
Specific Rank of Limax cinereo-niger, Wolf. 227
observations, which are both numerous and accurate, made by
Mr. Ashford.
Another point of difference between the two forms under
consideration is afforded by the radula, which is only second
in importance to the reproductive organs as affording a
criterion for the differentiation of molluscan species. Indeed
some authors ascribe to it a much higher value in this
regard. The differences in the radule of Z. maximus and
L. ctnereo-niger, as figured by Heynemann, who was the
acknowledged foremost limacologist of his time, in his paper
“ Ueber Schneckenzungen der Gattung Limax” (Mal. Blitt.
1863, pp. 200-218), are very striking. The teeth of ZL.
maximus are simple in form nearly throughout, only the
extreme outer teeth being shown as bifid, whereas in ZL. cinereo-
niger the extreme outer teeth are simple, and the side-teeth
are throughout bifid and even trifid.
The external characters, as I have pointed out more than
once, are sufficiently distinctive to make ZL. cinereo-niger a
particularly easy species to recognize. I have had ample
opportunity for forming a judgment in this respect, as by far
the greater proportion of the British specimens known have
passed through my hands.
The sculpture of the two species is markedly different in
character. In ZL. maximus the rugosities of the body are
small, fine, and closely set in comparison with ZL. cinereo-
niger, which has them large and coarse, with deep furrows
separating them. In this respect it resembles Arion ater, and
there can be little doubt that it is frequently mistaken for that
species by inexperienced conchologists, more especially as the
colour is often very similar, ZL. cinereo-niger being nearly
always a very dark species, usually black, with but few pale
markings, often none.
The differences in colour are important. One of the most
striking characters of L. cinereo-niger is that the foot-sole is
what may be loosely called “ trifasciated,”’ the two side-areas
(longitudinal) being black, blackish, or dark-coloured, with
the central area white. This is never the case with true
L. maximus, but is a fairly constant character in L. cinereo-
niger. 1 have on one or two occasions seen juvenile examples
with the whole foot-sole white, but never adults.
The coloration of the shield offers another ready mark of
distinction. L. maximus always has this part maculated or
marbled ; but in L. cinereo-niger there are no maculations or
marblings, the shield being invariably of a uniform dark
colour.
A less constant but very convenient character of L. cinereo-
228 Variations of the Lateral Shields in the Stickleback.
niger is that the keel and a line continuing it along the back
to where it joins the shield is the last part of the body to
retain the light ground-colour. Except in the totally black
variety (v. maura) L. cinereo-niger always has this keel and
line more or less evident as a thin pale stripe.
This evidence will suffice to make it clear that, whatever
views we may hold as to the specific or subspecitic rank of
L. cinereo-niger, it is not open for us to acquiesce in the
statement that it “ is simply one of the many colour-variations
of the well-known Z. maaimus, L.”
XXXVII.—WNote on the Variations of the Lateral Shields in
the Three-spined Stickleback (Gastrosteus aculeatus). By
G. A. BOULENGER.
EVER since Cuvier proceeded to divide the Sticklebacks into
species according to the presence or absence and the develop-
ment of the lateral armour, the question of the value of this
character has been much discussed. Most modern European
writers, with the exception of Blanchard and Sauvage, have
refused to accept Cuvier’s species as such, although they
have usually retained them as varieties or subspecies.
These supposed species are, however, maintained provision-
ally by American authors, Jordan not long ago remarking
that he has not yet met with distinctly intermediate forms
either on the Atlantic or Pacific coast. Bonizzi, Day, and
Fatio have published results of investigations into the varia-
tions of the spines and shields in the smooth-tailed form from
one locality; but the differences in the lateral armour in
northern brackish- water specimens do not appear to have yet
been subjected to a thorough statistical examination.
In July last I collected indiscriminately in a tidal pool close
to Ostend Harbour sixty-six specimens of the three-spined
stickleback, with the object of testing their characters, as I
had observed that the three principal forms, viz. G. trachurus,
G. semiarmatus, and G. gymnurus, occurred promiscuously
both in and outside the harbour, ‘The result is inter-
esting, as showing how complete the gradation between the
shielded and the smooth form is and how much the characters
may differ on the two sides of one and the same specimen. I
have therefore recorded the number of lateral shields in all
the specimens, and arranged them in a series from the most
perfectly armoured to the naked specimens. The numbers
given refer to the shields on either side, those of the left side
On a new Porcupine from East Africa. 229
separated from those on the right by a dash, the cross indi-
cating a gap between the thoracic and caudal shields.
The first group represents G. trachurus, with an uninter-
rupted series of shields from the shoulder to the end of the tail ;
the third the G. semdarmatus, with a naked gap between the
thoracic shields and the caudals; the fifth the G. gymnurus,
with the tail completely naked; whilst the second and fourth
groups are of special interest as comprising individuals which
belong to two different forms, according to whether the right
or the left side is examined. It will be observed that the
intermediate specimens (G. semiarmatus) are the most
numerous.
E 85. 144+2—14438.
223]. 6, a7 — 187
> Ea 37. 1447—13+7.
8. 30—29. 38, 1447—11+7.
4-6, 299—29, 39, 124+8—1347.
7,8, 28—29. 40. 13-+-7—12+6.
9-13. 29—28. 41. 12+4+6—12+47.
14-16, 28—28. 42, 124+7—11+46.
43, 19-6117.
: Peneeite
URIs selieaey 48. 10461046.
19. 18-++-7—29. 48. 84-2843.
20. 283—17+10. 49. 746746.
21. 28—144+6. 50, 644-744.
ps , IV.
22. 19+-7—17+8. =
93. 17+7—19-+7. 51. 94+3—9.
24, 17+6—18-+46. -
25. 18+-7—16+7. .
26. 1441+4+6—18+46. 52, 12—12.
ae ile 17-57. 58, 54. 10—10.
98, 16+8—16+5. 55,56. 7—7.
29. 164+8—15+7. 57, 7—6:
30. 15+5—16+6. 58, 59. 6—7.
31. 15+8—15+48. 60. 5—7.
$2. 144+-7—15+47. 61-64. 6—6.
33. 104+2+4—1543. 65. 6—5.
34, 144+6—14+46. 66. 4—4.
ee
XXXVIII.—Description of a New Porcupine from East
Africa. By OLDFIELD THOMAS.
Mr. F. J. Jackson, the discoverer of so many new East-
African mammals, picked up on the beach at Lamu the skull
of a porcupine which, on comparison, appears to me to differ
Ann. & Mag. N. Hist. Ser. 6. Vol. xi. 17
230 On a new Porcupine from East Africa.
equally from the northern Hystrix cristata, L., and the
southern //. africe-australis, Pet. Itis not fully adult, the last
molar being still quite unused, although in position, and the
tooth-change not having yet taken place; this change, how-
ever, occurs so late in life in the genus /ystrix that, judging
by other skulls, we may suppose that this skull would not
have greatly altered in size or form in later life.
The species may be called
Hystrix galeata, sp. n.
Skull approximating to that of H. cristata in the relative
lengths of the nasals and frontals (see measurements below),
but markedly different from that, as also from H. africe-
australis, by the reduced breadths of the same bones, espe-
cially in the interorbital region of the skull ; so that the upper
inflated part of the skull is more or less parallel-sided, instead
of being broadly oval; when viewed from above this appear-
ance is increased by the nasal bones being almost as broad
anteriorly as posteriorly and by the ascending process of the
premaxille being bowed out laterally ; the muzzle therefore
does not decrease evenly in breadth from the orbits forwards,
but is parallel-sided to the level of the anterior point of junc-
tion of the nasals and premaxille. These processes are also
much broader in a vertical direction than in the allied species,
and in this respect approach the Indian HZ. leucura. In con-
sequence also of the greater anterior breadth of the nasals and
of the bowing outwards of the ascending premaxillary processes
the nasal opening is very decidedly larger than in either of the
other African species. Supraorbital edges of frontal straight
or even slightly concave, the broadest point across the frontals
being at the rudimentary postorbital processes, not at the edge
of the lacrymal bones. In the side view the skull is remark-
able for its extreme height at about ™:!, from which point it
slopes rapidly down, both forwards and backwards. Lower
anterior root of zygoma broad and flattened, as in the northern,
not styliform as in the southern species; general size and
form of the ante-orbital foramen also very much as in the
former species,
The molar teeth appear to be rather rounder in section, less
narrow and elongated than in //. cristata; but much more
material is needed before the value of this character can be at
all properly estimated.
Measurements of a Gambian skull of H. cristata, of the
typical skull of H. galeata, and of a Natal skull of H. africa-
australis (the last-named is fully adult, the other two have
each not yet shed their mp.*) :—
ee
Formation of the Skeletal Paris in Echinoderms. 231
H, africe-
H, cristata. H. galeata. australis.
PMU AON OU. uccn cy Orde ose tier < 138 138 151
Zygomatic breadth ............ 86 84 91
Length (round curve) of nasals.. 100 99 91
9 x » oOffrontal 31 33 47*
Length of parietal (to extreme
Baek of Crash yoo. evs sca vs 31 33 33*
Breadth of nasals at anterior end
of naso-premaxillary suture = 87°5 4] 32
Breadth of nasals at posterior end 69°5 55 61
Interorbital breadth at edge of
MROEUIGANN ss oc cite cc box tats Us 73 62°3 76
Interorbital breadth at rudimen-
tary postorbital processes .. 68°5 665 70°6
Height of skull from palate be-
eel iat pe ee 70 76 73°5
Least vertical diameter of lower
anterior zygoma-root ...... 5 55 7
rg 0 pai a a oa (ii § 80 87
Mesial length of premaxille .... 29 25°7 28
These measurements show clearly the conspicuous differ-
ences between /. galeata and its allies in the relative propor-
tions of the two interorbital breadths, anterior and posterior,
in the shape of the nasals, and in the height of the skull, this
last being the character from which is derived the name
selected for the species.
Hab. Lamu, East Africa.
It is much to be hoped that more specimens of this inter-
esting addition to the fauna of our East-African possessions
will soon be brought to Europe, and it may be noted, as a hint
to collectors, that a good series of skulls of different ages
would be of especial value for the proper comparison of the
species with its allies.
XXXIX.—The Formation of the Skeletal Parts in
Echinoderms. By Cari Cuun, of Breslau ft.
THE share of the mesoderm cells of Echinoderms in the
building-up of the elements of the calcareous skeleton is alto-
gether imperfectly understood. After it had already been
shown by the older observers that the calcareous bodies are
formed by the connective-tissue cells of the gelatinous central
* Fronto-parietal suture closed and its position not quite certain.
+ Translated from the ‘Zoologischer Anzeiger, xv. Jahrg., no. 408,
December 26, 1892, pp. 470-474.
fe
232 M. Carl Chun on the Formation of
substance of the larve (it was scarcely likely that Selenka
and Hérouard should still retain the ideas they formerly
expressed as to an ectodermal origin of the calcareous bodies
of the integument), it was recently sought by Selenka and
Semon to determine the finer processes which take place in
the secretion of the calcareous matter. According to Semon’s
account there arises within the skeletogenous mesoderm cell a
tetrahedron, which subsequently develops into the triradiate
body already observed by Selenka and passes out of the cell
enveloped in a delicate membrane. The triradiate body is
then approached by other mesoderm cells, which enlarge it,
and by means of complicated furcations transform it into the
definitive calcareous structure. Semon’s observations have
been confirmed by a careful memoir by H. Théel, which has
just appeared (‘‘ Development of Eehinocyamus pusillus,”
R. 8. Upsala), in so far as Théel also claims the tetrahedron
developing into a triradiate body as the foundation for the
building-up of the skeletal parts of the Pluteus. It is true
that there are material discrepancies in the observations as to
the earliest origin of the tetrahedron. For [according to
Théel] it is formed between at least three cells in a clear
organic basal substance, and therefore from the outset proves
itself to be an intercellular skeletal element, in the enlarge-
ment of which a large number of amceboid cells subsequently
take part.
Now Semon is inclined to regard the tetrahedron, which
develops into a body with three or four rays as the case may
be, as the universal] starting-point of the whole of the skeletal
structures of Echinoderms. This view, then, would also
sanction the converse inference, that the individual calcareous
structures represent the product of a larger number of meso-
derm cells. Nevertheless this assumption does not hold for
all skeletal elements, inasmuch as, e. g., the wheels of the
Auriculari@ and the anchors and anchor-plates of the Synap-
tide, according to the statements of older investigators, which
in essential points were recently confirmed by Semon and
Ludwig, do not exhibit a tetrahedron and body with three or
four rays as a starting-point. Semon therefore believes that
the original condition became obliterated in these instances,
and that the appearance of a star with six rays, which he
gives as the basis of many calcareous wheels, implies a
curtailment of the primitive arrangement. It would lead us
too far, should we wish to discuss this conception here ;
against it the objection may always be advanced that the
calcareous wheels clearly represent quite primitive structures,
which not only furnish the distinctive character of the Auricu-
the Skeletal Parts in Echinoderms. 233
laria-larva and in the case of many Synaptidw persist
throughout life, but also, according to the discovery of Joh.
Miiller, which was confirmed by Ludwig for the Asteriade,
by Semon for Ophiuride, and by Théel for Echinide, form
the basis for the construction of the spines. Nevertheless
it may appear to be open to question whether we are justified
in homologizing with the wheels of the Auricularia the basal
wheels of the spines, which are stated by Ludwig to arise
from a triradiate body, and according to Théel’s latest result
again exhibit a tetrahedron as their starting-point. For,
according to my own observations, the mode of formation of
the larval calcareous wheels is so peculiar that it by no means
allows itself to be forced into the scheme, which was con-
structed on the basis of our previous knowledge of the
building-up of the calcareous skeletal parts from mesoderm
cells.
As material for investigation I made use of the splendid
Auricularie which I caught in different stages of develop-
ment at the Canary Islands in the winter of 1888. As I
shall describe the larvee, which attained a length of 7 millim.,
in another place, I here dispense with an account of the
complicated course of their ciliated bands and of their internal
structure. Let it merely be remarked that the calcareous
wheels appear relatively very late, but then accumulate in
unusual abundance in the aboral tuft-shaped outgrowths of
the lateral regions, further on along the entire dorsal surface,
and much more scantily upon the ventral side. In order to
check my observations the Mediterranean Auricularie were
also examined, which, so far as regards the formation of the
calcareous wheels, exhibit almost identical conditions.
At the time of the appearance of the first calcareous wheels
the cellular elements of the gelatinous substance are sharply
differentiated into skeletogenous and connective-tissue cells.
The latter possess several long processes, which are much
ramified and are interwoven almost after the manner of felt;
the skeletogenous cells, on the contrary, are spherical and
surrounded by a distinct membrane, in consequence of which
they emit no pseudopodia. ‘The sharp histological differen-
tiation of the mesoderm cells, which was certainly preceded
by an indifferent stage, may be essentially due to the fact
that the calcareous bodies originate at a remarkably late
period in comparison with what is found to be the case in
other Echinoderm larve. The skeletogenous cells accumu-
late around the stone-canal and close beneath the ectodermal
pavement epithelium. ‘The latter with its nuclei is always
distinctly discernible, even in Auriculari@ of the largest size,
234 M. Carl Chun on the Formation of
and there is no reason to suppose that it degenerates and is
subsequently replaced (Semon) by means of the cells of the
ciliated band or of the peripheral connective tissue.
A richly vacuolate plasma at once distinguishes the skeleto-
genous oaltd the average size of which is ‘01 millim. They
rapidly grow to twice and thrice this bulk, while simulta-
neously the number of the cell-nuclei increases. In the same
Auricularia we meet with all intermediate stages between
uni- and multinucleate cells, which at first still retain a
rounded contour, but subsequently flatten out on one side and
become cup-shaped. The nuclei measure from ‘003 to ‘004
millim in length, and originally (so long as only from two to
four are present) occupy a peripheral position; they after-
wards increase to from six to eight in the case of the Medi-
terranean Auricularia, and to from twelve to eighteen in that
of those from the Canary Islands, and form a central nuclear
cluster.
When the cells have attained a size of ‘03 millim. there
appears within the old cell-membrane a new one, which has
an undulating outline towards the circular margin and speedily
assumes a star-shaped form. The tubular rays of the star
which grow out are equal in calibre and meet the external
membrane, arching forward somewhat at the points of contact.
The longitudinal extension of the radially arranged out-
growths keeps pace with the increase in the size of the cell,
and finally, when the cell attains a size of from *06 to ‘07
millim., the rays become united by a peripheral membranous
ring. It is now impossible to mistake the mould of the
subsequent calcareous wheel, prepared as it is by the complex
folds of an internal membrane: the central portion with the
cluster of nuclei corresponds to the nave, the tubes running
out like the rays of a star represent the spokes, and the peri-
pheral ring takes the place of the circumference (the felly) of
the future calcareous wheel. Moreover the calx is actually
secreted into this organic matrix formed by the skeletogenous
cell, as into a mould, and in such a way that (as the older
accounts already teach us) calcification takes place first in
the nave, then in the spokes, and finally in the felly of the
wheel. It is likewise in accordance with the theories which
have recently been formulated as to the share of the nuclei in
the vital processes of the cell that, corresponding with this
centrifugal progress of the calcification, the majority of the
cell-nuclei also separate from one another in a centrifugal
direction, and in the case of the Auricular’e from the Canary
Islands come to lie in the acute angles between the spokes.
the Skeletal Parts in Echinoderms. 235
In rare instances they advance as far as the middle of the
spokes or even to the periphery.
No secondary multiplication of the spokes of the wheel
takes place ; their number corresponds exactly with that of
the undulating evaginations of the newly formed internal
membrane, which develop into radiating tubes. As is well
known, the number of the spokes varies ; in the case of the
Auricularia trom the Canaries we find from thirteen to
eighteen,
Since the diameter of the fully formed calcareous wheels is
found to be from ‘09 to ‘1 millim., it follows that a tenfold
enlargement of the diameter of the skeletogenous cells takes
place, since the latter in the stage with a single nucleus only
measure ‘01 millim. Nevertheless after the secretion of the
caleareous wheels they expand still further ; for if we examine
the wheels in alcohol preparations (the delicate points referred
to can scarcely be demonstrated in glycerine and Canada
balsam), we can distinguish a distant periphery formed by a
delicate membrane, from which, alternating with the spokes
and almost equalling them in length, membranous tubes
arranged in the shape of a star run to the periphery of the
wheel, where they usually exhibit flask-shaped expansions.
On careful decalcification of the wheels by means of weak
chromic acid it is easy to show the nuclei and the contour of
the wheel in the shape of a delicate membranous envelope
within the skeletogenous cell.
The above statements as to the formation of the wheels in
the Auricularia reveal a mode of development which at
present appears to be unique. While the skeletal pieces of
Echinoderms were hitherto essentially regarded as intercellular
structures, the formation of which was due to several mobile
amceboid cells (I am well aware that more recent observers
are inclined to attribute the shape of the skeletal elements
without hesitation to directly mechanical influences), we now
find that the form of the calcareous wheel is traced out within
a multinucleate cell by means of an organic membrane which
assumes complex folds, and that in this definitely circumscribed
mould the casting of the hard parts ensues.
236 Mr. EF. J. Allen on the Nephridia and
XL.—Preliminary Account of the Nephridia and Body-
Cavity of the Larva of Palemonetes varians. By EDGAR
J. ALLEN, B.Se., University College, London*.
THE researches of which the present communication contains
a brief summary were carried on during the summer of the
present year at the Laboratory of the Marine Biological Asso-
ciation in Plymouth, where | occupied a table by appointment
of the British Association Committee. The observations were
made chiefly on larvee of Palemonetes varians, but other species
have also been included, and will be mentioned in particular
instances.
I. The Nephridia.
During the greater part of the larval life two pairs of
nephridia are present—the green glands, which open at the
bases of the second antenne, and the shell-glands, which open
at the bases of the second maxille.
The Green Gland.—In a larva of Palemonetes which is a
few days old the green gland has a form similar to that
described by Weldon t and Marchal f for the adult of Virbius,
Pandalus, and Crangon, excepting that the remarkable
enlargements of the bladder, which the former author has
termed ‘‘nephroperitoneal sacs,’ are not as yet developed.
The gland consists of an end sac, which communicates by
means of a U-shaped tube with a very short ureter opening
at the base of the antenna. ‘The distal portion of the tube is
slightly enlarged, and may be called the bladder. At the
time of hatching of the larva the whole gland consists of a
solid mass of cells, in which no cavity has appeared, although
the portions which afterwards form the end sac and the tube
can be distinguished, and the ureter and external opening are
already present. Shortly after the larva has become free the
cells separate and give rise to the lumen of the gland.
The later development of the green gland consists chiefl
in the enlargement of the bladder, which grows at first nwa
* From the ‘Proceedings of the Royal Society,’ vol. lii. no. 318,
pp. 338-842,
+ Weldon, W. F. R., “The Renal Organs of certain Decapod Crus-
tacea,” Quart. Journ. Micr. Sci. vol. xxxii., 1891.
{ Marchal, P., “ Recherches anatomiques et physiologiques sur l’appa-
reil excréteur des Crustacés Décapodes,’ ‘Atoh. Yast expér. vol. x., 1802.
Body- Cavity of the Larva of Palemonetes varians. 237
towards the middle ventral line of the body, then upwards,
within the cesophageal nerve-ring and anterior to the cesopha-
gus, to the middle dorsal line, where it meets its fellow of the
opposite side. The two bladders grow backwards over the
stomach, subsequently fusing in the middle line to form the
unpaired nephroperitoneal sac. ‘This mode of development
confirms the view as to the nature of the latter sac already
arrived at by Weldon and Marchal from a comparative study
of the renal organs of Decapods.
The Shell-Gland.—In a figure of a Callianassa larva in
the Mysis stage Claus * inserts and names the shell-gland,
opening at the base of the second maxilla. ‘This is, I believe,
the only recorded instance of the gland having been recog-
nized in a Decapod, unless, indeed, the “ segmental organ”
described by Lebedinski ¢ as opening at the base of the first
maxillipede of the larva of Hriphya spinifrons and communi-
cating with the body-cavity be the same organ.
In late embryos and at the time of hatching of the larva the
shell-glands are the functional kidneys of Palemonetes and
Palemon, the green gland being still without a lumen. The
shell-gland of Palaemonetes consists of a comparatively short
renal tube with a considerable lumen, which communicates
internally with an end sac, and opens externally at the base
of the second maxilla. ‘he general form of the tube may be
expressed by saying that it is Y-shaped, the two arms of the
Y being in a horizontal plane, with the end sac attached to the
internal one, whilst the leg of the Y¥ is curved in a vertical
plane, the concavity looking downwards and backwards. The
histological structure of both end sac and renal tube is similar
to that described by Grobben { for the green gland of Mysis.
The entrance from the end sac to the tube is guarded, how-
ever, by a valve formed of elongated cells of the end sac,
which does not appear to have been found in other forms.
I have detected no trace of the shell-gland in young adults.
II. The Body- Cavity.
The Anterior Region of the Thorax.—A transverse section
through the region of the second maxille of a Palemonetes
* Claus, C., “ Neue Beitriige zur Morphologie der Crustaceen,” Arb,
Zool. Inst. Wien, vol. vi., 1886.
+ Lebedinski, J., “ Einige Untersuchungen iiber die Entwicklungs-
geschichte der Seekrabben,” Biol. Centralbl. vol. x., 1890,
t Grobben, C., “ Die Antennendriise der Crustaceen,” Arb. Zool. Inst.
Wien, vol. iii., 1881.
238 Mr. E. J. Allen on the Nephridia and
larva which is a few days old shows that the cavity enclosed
by the external chitin and ectoderm may be divided into four
regions:—a dorsal sac, surrounded by a definite layer of
epithelium and within which the cephalic aorta lies, but which
does not itself contain blood; a central cavity, in which the
liver, intestine, and nerve-cord are found; two /ateral cavities,
separated from the central cavities by masses of muscle and
bands of connective tissue, and which, in the region under
consideration, contain the proximal ends of the shell-glands ;
and, fourthly, the cavities of the limbs, which contain the
distal ends of the same organs. ‘The cavities of the limbs
communicate with the lateral cavities, and the latter frequently
communicate with the central cavity by the disappearance of
the connective-tissue bands. ‘The central cavity, the lateral
cavities, and the cavities of the limbs all contain blood.
The Dorsal Sac.—I have found the dorsal sac in Palemon
serratus, Palemonetes varians, and Crangon vulgaris, and in
the adult it attains a considerable size. If a dissection be
made of an adult Palemon the sac is readily seen. Ante-
riorly it appears as an elongated cylindrical tube lying upon
the nephroperitoneal sac, and containing within it the cephalic
aorta. Posteriorly it is very much enlarged, covering the
front part of the ovaries, and running downwards on either
side into the cavity which surrounds the intestine and liver.
A similar condition of things is found in the adult Pale-
monetes.
The dorsal sac does not contain blood. I have been led
to this conclusion for the following reasons :—(1) In a large
number of series of sections, both of larve and adults, I have
never seen a blood-corpuscle within the sac. (2) The sac is
completely closed and has no communication with the blood-
sinuses of the body ; in preserved specimens it contains a clot,
which can generally be distinguished from the surrounding
blood-clot. (3) Lhave observed carefully and for a long time
living larvee, and the space occupied by the sac has always
been perfectly free from blood-corpuscles.
At its anterior end the dorsal sac is surrounded by a mass
of tissue, from the external surface of which blood-corpuscles
appear to be budding off. Before commencing this research
Professor Weldon, to whom the existence of this tissue was
known, and who has indicated it in his figures, suggested to
me this view of its nature, and it is fully supported by my
preparations.
Development of the Dorsal Sac.—In embryos of Palemonetes
in which the cephalic aorta is already formed the cells sur-
one
7”
Body- Cavity of the Larva of Palemonetes varians. 239
rounding the latter are arranged in two layers, an internal
and an external. Before the time of hatching arrives the cells
of the external layer enlarge considerably and give rise to the
appearance of a solid mass of cells upon either side of the
aorta. The dorsal sac is formed by the hollowing out of
these masses of cells. Two lateral cavities are thus formed,
which are separated by the aorta. The protoplasm of the
cells lining these cavities, which is at first gathered into
masses around the nuclei, then spreads out into a thin sheet,
drawing away from the lower portion of the aorta, and causing
the two lateral cavities to unite ventrally and so form a single
sac. In the region of the first and second maxille all the
stages of the process just described may be seen. In the
region anterior to this I have never actually observed the
stage with two lateral cavities, the two having always united
ventrally ; but I do not doubt that the process is here the same
as in the posterior region.
In one series of sections of a larva, preserved very soon
after it had left the egg, the cavity was formed upon either
side of and below the aorta, as far back as the anterior end of
the segment of the first maxilla, where for one section it was
almost completely closed. Behind this the condition with
two lateral cavities was found and persisted through the region
of the first maxille, whilst in the region of the second maxille
no cavity had yet opened, and solid masses of cells still Jay
upon either side of the aorta.
The further development of the dorsal sac consists mainly
of an increase in its size. At its posterior end it grows back-
wards in a pair of lobes, which extend as far as the front end
of the pericardium.
The Posterior Region of the Thoraz.—The central and
lateral cavities are here similar to those of the anterior region,
whilst dorsal to them the pericardial chamber lies. This
chamber is separated from the central body-cavity, as is
already well known, by the pericardial septum, and it contains
the heart. The genital organs are situated at the front end
of the pericardium, immediately below the pericardial septum.
In the just-hatched larva these consist of two masses of cells
with large nuclei, each mass being enclosed in a sheath of
mesoderm. I have not detected any trace of the genital ducts
at this stage.
The Abdomen.—With regard to the abdomen, my _ sec-
tions confirm the accounts given by Milne-Edwards * and
* Milne-Edwards, ‘ Histoire Naturelle des Crustacés,’ Paris, 1834.
240 On the Larva of Paleemonetes varians.
Claus *. There are two main sinuses, which run along its
length: a dorsal sinus, in which the intestine lies, and a ven-
tral one, which contains the nerve-cord. These two sinuses
are generally separated by masses of muscle; but they
communicate at intervals by means of lateral sinuses.
Theoretical Considerations.
The state of the body-cavity in the anterior region of the
thorax of Palemonetes may be compared with that of Peri-
patus, as described by Sedgwick f, at the time when the dorsal
portions of the mesoblastic somites have attained their maxi-
mum development. Bearing in mind that the dorsal sac of
Palemonetes has been formed by the union of two lateral
cavities, which lay on either side of the aorta, the differences
between the two forms are very slight. The dorsal sac repre-
sents the two dorsal portions of the mesoblastic somites, whilst
the central cavity, the lateral cavities, and the nephridia
agree, with the one exception that the two lateral portions of
the nerve-cord of Peripatus have united in the middle line in
Palemonetes, and in the process have passed out of the lateral
cavities. The agreement is so close that it appears to me to
be fully justifiable to homolegize the various parts. If this
be so, it follows that the dorsal sac of Palamonetes is homo-
logous with the dorsal portions of the mesoblastic somites of
Peripatus and that its cavity isa true celom. The central
and lateral cavities, together with the cavities of the legs, will
represent the pseudoceele, and, being filled with blood, may
be termed, with Lankester, hemocele.
Passing now to the posterior part of the thorax, the region
of the heart, we find that the different cavities correspond with
those which persist in the adult Peripatus. Heart, peri-
cardium, and pericardial septum of Palemonetes present
exactly the same relations as in Peripatus, and are clearly
homologous structures in the two animals. The central and
lateral cavities only differ on account of the relative position
of the nervous system, and this difference has already been
accounted for. It must be noted, however, that in this region
no nephridia are found in the cavities of the limbs. Beneath
the anterior end of the pericardial septum are found, as has
* Claus, C., “Zur Kenntniss der Kreislaufsorgane der Schizopoden und
Decapoden,” Arb. Zool. Inst. Wien, vol. v., 1884.
+ Sedgwick, A., “The Development of the Cape Species of Peripatus:
Parts L-IV.,” Quart. Journ. Micr. Sci. vols. xxv.—xxviii., 1885-88.
On Subdivision of the Median Fin in a Dipnoan Fish. 241
been already stated, the genital organs, and here also the
comparison with Peripatus may be instituted. We find a
similar agreement to that which existed in the other regions
compared, and we may, with a considerable degree of proba-
bility, again homologize corresponding parts. The genital
organs of Palemonetes must then be regarded as the represen-
tatives, in this region, of the ccelom.
If the homologies here suggested are valid, the body-cavity
relations of the Crustaceans under consideration may be stated
briefly thus :—both enteroccele (true coelom) and pseudoccele
are present, the enteroccele consisting of the dorsal sac, the
green gland, and shell-gland, or the end sacs of these organs,
together with the genital organs and their ducts, whilst the
pericardial septum may be regarded as equivalent to portions
of the walls of another part of the same structure.
The pseudoccele consists of the heart and arteries, the peri-
cardial cavity, the central cavity of the thorax, with the
lateral cavities and the cavities of the limbs, and the various
sinuses of the abdomen. The pseudoceele is filled with blood,
and hence can be termed a hemoceele.
I hope shortly to publish a more detailed account, with
figures, of the relations described in this communication.
XLI.—WNote on a Case of Subdivision of the Median Fin in a
Dipnoan Fish. By A. Smita Woopwarp, F.L.S.
PALHZONTOLOGY has demonstrated that in the Crossoptery-
gian Teleostomatous fishes the process of subdivision of the
median fin is usually different from that observed in the more
specialized Actinopterygians. Whereas in the latter case
portions of the fin-membrane disappear and the endoskeletal
supports afterwards become atrophied, in the former case it is
the almost universal rule that the supports of the dorsal and
anal region become concentrated in clusters, and thus induce
subdivision of the primitively continuous fin. The proximal
fin-supports (axonosts) in each of these clusters ordinarily
fuse into a single club-shaped element, and the distal supports
(baseosts) are often more or less reduced in number by the
same process”.
* Smith Woodward, ‘Catalogue of Fossil Fishes in the British Mu-
seum,’ part ii. (1891); and “The Evolution of Fins,” Natural Scieuce,
vol. 1. pp. 28-385 (1892).
242 On Subdivision of the Median Fin in a Dipnoan Fish.
Such being the arrangement in the Crossopterygii, it is of
much interest to be able to institute comparisons with the
Dipnoan fishes, which exhibit so many features of resem-
blance to the order just mentioned. Two Devonian genera,
Phaneropleuron and Dipterus, are known to possess a discon-
tinuous median fin ; and quite lately specimens of the first of
these genera have been found displaying all the fin-supports
in undisturbed position. The fossils in question are referable
to Phaneropleuron curtum, were obtained from the Upper
Devonian of Canada, and have been acquired by the British
Museum.
It now appears that, at least in the case of the Dipnoan
anal fin, the process of separation from the caudal is precisely
similar to that observed in the Crossopterygii. As shown by
the accompanying illustration
the axonosts (a) are fused into a
single club-shaped element, and
there are three hour-glass-
shaped baseosts (6) of nearly
equal size. Round the distal
ends of these cartilages the
clusters of dermal fin-rays (/)
overlap in the usual manner.
The same arrangement is also cox
indicated in the figures of Pha- (
neropleuron curtum already pub- i HI"\-y
lished by Whiteaves * ; but,
owing to the considerable dis- ino enithart. foetk 1
placement of the fin-sup ports in , Hamer rene Bay rb.
all the specimens at the dis- (Canada. ” a, axonost; 6, base-
nee of that author, the pecu- _osts; f, bases of fin-rays. [Brit.
iarity now described does not = Mus. no. P. 6785.]
appear to have been observed. !
When specimens of Dipterus showing the fin-supports have
been discovered the same phenomenon will doubtless be met
with also in that genus; but as yet there is unfortunately no
evidence on the subject.
Skeleton of Anal Fin of Phanero-
* J. F. Whiteaves, “ Illustrations of the Fossil Fishes of the Devonian
Rocks of Canada.—Part I.,” Trans, Roy. Soc. Canada, vol. iv. sect. iv.
p- 108, pl. x. fig. 2.
On the Mechanical Genesis of the Scales of Fishes, 243
XLII.—On the Mechanical Genesis of the Scales of
Fishes. By Joun A. Ryver*.
FOURTEEN years ago the present writer suggested that the
slow metamorphosis of the forms of the crowns of the teeth of
Mammalia fT in the course of a vast number of successive
generations might be ascribed to the continuous, slow, and
cumulative action of mechanical strains and pressures in
definite directions, resulting in the production of permanent
stresses and consequent changes in the forms of the crowns,
especially of the molar series. The evidence since accumu-
lated from vertebrate paleontology and anatomy has served
to strengthen the belief that such an hypothesis cannot be
dismissed as useless until a better one has been offered in its
stead. The present paper is an attempt to apply somewhat
analogous reasoning to a somewhat simpler but no less inter-
esting problem in morphogenesis.
The mechanical hypothesis now to be offered respecting the
genesis of the scales of fishes accounts for the origin of such
scales from a continuous subepidermal matrix, which may be
regarded as a basement-membrane. Such a matrix is found
to actually exist in some forms at an early stage just beneath
the epidermis. It is thickest on the dorsal and lateral aspects
of the body, as is seen in sections of the young of the scaleless
Batrachus tau, for example. Such a matrix also exists in
the larval stages of other scale-bearing forms, and may be
continuous with the very attenuated basement-membrane
from which the actinotrichia or primordial fin-rays of embryo
fishes seem to be in part differentiated. Such a matrix is
almost coextensive with the whole epidermal layer of the
young of many types of fishes, just at the time when the
scales commence to be developed.
The hypothesis further accounts for the arrangement of the
scales in longitudinal and in oblique rows in two directions.
The oblique rows are arranged, as is well known, in a direc-
tion from above downward and backward, and also in the
reverse direction from below upward and backward—that is,
the scales may be counted in rows in three directions down-
ward and forward as well as downward and backward, and,
* From the ‘Proceedings of the Academy of Natural Sciences of
Philadelphia,’ 1892, pp. 219-224.
+ “On the Mechanical Genesis of Tooth-forms,” Proc. Acad. Nat. Sei,
Philad, 1878.
244 Mr. J. A. Ryder on the
starting from any scale in any oblique row, they may be
counted either forward or backward longitudinally or in con-
formity with the direction of the axis of the body of the fish.
This is conspicuously the case in Clupeoids and some
Cyprinoids.
In such archaic types as these, approximating the primitive
isospondylous condition, it is also found that the number of
scales in a longitudinal row corresponds on the sides of the
body very exactly with the number of muscle-plates or somites
of the body. It is also found that the myocommata or sheets
of connective tissue intervening between the successive somites
are attached with great firmness to the deeper layers of the
skin or corium. Such a construction, together with the
peculiar arrangement of the muscle-plates at the time the
scales begin to develop, conditions the further growth of the
scale matrix. This is affected in such a manner that the
whole of the integument is thrown into definitely circum-
scribed areole during the ordinary movements of the fish in
swimming. The central portions of each of these areole are
left in a quiescent condition, while their margins are wrinkled
or folded as a result of the current action of the lateral muscles
of the body. In this wise each and every one of the dermal
and epidermal areole are circumscribed by the action of the
fish in the normal act of swimming. In each of the circum-
scribed areola a scale develops ; the continuity of its develop-
ment with its fellows across the margins of the areola is
prevented by the continual bendings or flexures to which the
dermis is there subjected owing to the action of the muscles.
This will be better understood by referring to the accom-
panying diagram, representing the arrangementof the muscular
somites of a Cyprinoid (Carassius) with their intervening
myocommata as seen from the side when the skin with its
scales is removed. Before proceeding further, however, it
may be well to insist upon the fact that the rows of scales are
found to conform to the successive somites. ‘This is of itself
significant. The careful interpretation of the facts from
observation, however, discloses a very remarkable effect due
to the peculiar arrangement of the muscle-plates.
As is well known, the muscular masses of the sides of the
body of a fish are arranged in the form of two longitudinal
trihedral columns separated along the middle line of the side
a to b into a dorsal and ventral half. ‘lhe somites entering
into the composition of these ventral and dorsal masses were
at first absolutely continuous across the longitudinal horizontal
septum a, b. If we suppose the somites of the adult as deve-
loped from a continuous embryonic segment extending the
see. “Fe
Mechanical Genesis of the Scales of Fishes. 245
whole depth of the body, then will somites I, If, II, IV, V,
&c. in the figure form two parallel series of muscular blocks
above and below the line a, 6. Each half-somite is also seen
to present an acute apex directed backward at the points ¢ and
ce’ above and below the line a, 6. Thesomites I, II, ILI, IV,
and V are therefore sigmoid in outline as seen from the
exterior. The myo-
commata or connec- Fic
: ‘ ia
tive-tissue septa 1, Z
2, 3, 4, 5, 6, &e., I DMOIIV V VI vil
which intervene be- Sai VV V\E
tween the somites,
have a correspond- \
ing sigmoid arrange- \
ment. The sigmoid \
or =-shaped myo- \
commata and the
myotomes are recip- ;
rocally coadapted to
each other in con- i
figuration, like a é
nest of S=S=Z=’s
turned upon their
sides. If we fur-
ther supposed that
thin- and thick-legged =’s alternated thuun—g=q aa=-
we might suppose the thin-legged ones to represent the myo-
commata and the thick-legged ones the muscle-plates or myo-
tomes. The muscular fibres of the thick-leggzed S's run
longitudinally from the posterior surface of the myocomma
immediately in front of it, to be inserted into the anterior face
of the myocomma immediately behind it. The muscular
tension is therefore exerted upon the opposite sides of the
myocommata, and is thus propagated along the sides of the
body from the head to the tail, from the first to the last myo-
tome. But the tension upon the inner face of the skin is
along the lines of insertion of the myocommata 1 c’c, 2 cc,
3 ec, &c.; this will serve to wrinkle the skin not only along
the lines 1 e’c, 2 ec, &e., but also to cause wrinkles to appear
along the dotted lines d, d’, e, and e’. In this way it is easy
to see that the whole lateral integument will be thrown into
definitely circumscribed rhomboidal areolz, in which separate
overlapping scales, s, s, s,s, may appear. ‘The wrinkles thus
produced by the tension of myocommata upon the integuments
of the body will cross each other and be reinforced at six
points in the zigzag transverse course of each myocomma, viz.
Ann. & Mag. N. Hist. Ser. 6. Vol. xi. 18
246 Mr. J. A. Ryder on the
twice at each of the points c and c’ and along the line a, d,
where the middle limb of the = rests.
The only point which now remains to be discussed is the
imbrication of the scales. This is also as readily accounted
for as the delimitation of the scale-forming areole, s, s,s, and
their trilinear arrangement in three directions in the convex
surface of the integument by means of the mechanical hypo-
thesis here outlined. If we were to make a longitudinal
transverse section through a fish along the plane of the letters
c, c, ¢, or e’, ce’, c’ of fig. 1, we should get a section like fig. 2,
Fig. 2.
A mmuuuuuMnumM
in which the myotomes M, M, M, &e. (fig. 2) were again
found to be V-shaped on either side of the medial axis or
vertebral column, c, ¢ (fig. 2). This proves that the myotomes
are really cones fitting into one another, and that, if we
suppose the first one to be inserted into the base of the skull
along the line A, 6 (fig. 2), that point becomes the anterior
fulerum or point d’appui of the whole muscular system. The
tensions thus developed upon the skin along the successive
myocommata 1, 2, 3, &e. (fig. 2) are such that the integument
would be flexed inward opposite each myocomma as shown in
the next figure on a larger scale, which represents the foldings
of the integument at the surface of a part of fig. 2. Here in
fig. 3 the myocommata m, 7v’, m, v’, m, &c. are seen to be
inserted upon the
internal face of the
epidermis. The
membrane m, m, m,
m, acted upon by
the muscular fibres
of the myotomes M,
M, M, will have the
effect of pulling the
integument inward
in the direction of
the arrows 7, 1,
from the linear
points of attach-
ment of the myocommata to the integuments at 7, 7, 7, toward
the vertebral bodies V, V, V. In this manner will be deve-
loped the imbrication indicated by the heavy border along the
posterior margins of the scales s, s, s in fig. 1, and in fig. 3 in
longitudinal section through the scale-sacks or pockets at s, s, s.
Mechanical Genesis of the Scales of Fishes. 247
Tt will be clear that in the case considered the arrangement
and imbrication of the scales is determined by the actions of
the segmentally arranged muscles of the body. In other words,
whatever has determined the development of somites has also,
in the most clear and direct manner, determined the segmen-
tally recurrent and peculiar trilinear and imbricated arrange-
ment of the scales of many fishes. It may be urged in
objection that heredity has determined the number, arrange-
ment, and the development of the somites, and therefore the
development of the scales is also a sequence of hereditary
influences working thus indirectly. This view of the case may
be admitted without invalidating the conclusion that, given
the growing mechanism here described, the development of
the scales would under any circumstances have been interfered
with at the points where the integument was being continually
flexed, wrinkled, or folded as it is around the integumentary
areolx wherein the scales are formed, as has been here proved
to correspond with the facts.
Special types of squamation amongst fishes may require an
interpretation different as to details from the foregoing ; but
it is probable that such special cases will rather tend to con-
firm than otherwise the views developed in this sketch of an
hypothesis respecting the mechanical origin of the arrange-
ment and imbrication of the scales of fishes. For example,
one of the most extreme cases, that of the sturgeon, shows
that the smaller integumentary plates between the large
dorsal, lateral, and ventral rows conform to these lines of
tension of the myocommata upon the integument. An even
more instructive example is that of the common eel, in which
the scales are oblong rhombs or parallelograms, arranged with
their diameters in oblique lines, running in two directions
conformably with the tensions, wrinklings, and foldings of
the integuments produced by the oblique insertions of the
muscles when the latter are brought into action. Other cases
where the scales are very fine might be urged in objection,
especially where several oblique rows of scales are found to
correspond to each somite. Such parallel duplication of scale-
rows, however, does not invalidate the principle, since the rows
still conform to the lines of tension of the linear attachment
of the myocommata to the integuments. The hypothesis may
also be extended so as to consistently consider such forms as
the pipe-fishes and other anomalous forms, where sluggish
habits coupled with the almost exclusive use of the dorsal fin
in swimming has rendered the lateral musculature of the
body comparatively subordinate in function, and which may
even lead to secondary fusion of somites and the consolidation
of consecutive pairs or triplets of vertebral centra into single
vertebral bodies,
18*
248 Mr. R. I. Pocock on some Types o7
Two conclusions of prime importance may be drawn from
the hypothesis and the evidence here presented, namely :—
1. The scales of fishes bear a segmental relation to the
remaining hard and soft parts, and are either repeated con-
secutively and in oblique rows corresponding to the number
of segments, or they may be repeated in rows as multiples of
the somites, or segmental reduction may occur which may
affect the arrangement of the scales so as to reduce the number
of rows below the number of somites indicated by the other
soft and hard parts.
2. The peculiar manner of interdigitation of the muscular
somites, as indicated by the sigmoid outline of the myocom-
mata as seen from their outer faces, and the oblique direction
of the membranes separating the muscular cones, has developed
a mode of insertion of the myocommata upon the corium
which has thrown the integument into rhombic areole during
muscular contraction. ‘These areole are in line in three
directions, and the folds separating them, particularly at their
posterior borders, are inflected in such a manner by muscular
tensions, due to the arrangement of muscular cones, as to
induce the condition of imbrication so characteristic of the
squamation of many fishes.
XLIII.— Upon the Identity of some of the Types of Diplopoda
contained in the Collection of the British Museum, together
with Descriptions of some new Species of Exotic lulide. By
R. I. Pocock.
(Plate XVI.]
Part I.—WNotes upon some Types of Diplopoaa.
Lysiopetalum Hardwickii (Newport), Ann. Mag. Nat. Hist. xiii.
p. 267 (1844), is based upon a specimen of L. fetidissimum,
Savi, Upuse. Scient. Bologna, i. p. 334 (1817). There appears
to be no foundation for the supposition that this specimen came
from India.
Lysiopetalum Ltichti (Gray), in Griffith’s Animal Kingdom (Insects,
i.), pl. 135. fig. 4, and further characterized by Newport in
vol. xiii. of the Annals of Nat. Hist., has been recharacterized
by Latzel as LZ. anceps (Myr. Ost.-Ung. Mon. ii. p. 232) and
very possibly as sicanum by Berlese (Acari, Myr. e Scorp.
Ital. pt. vi. no. 7, 1883).
Lysiopetalum rugulosum and lineatum of Newport, Ann. & Mag. Nat.
Hist. xiii. p. 267, are based upon two specimens specifically
identical with each other and with L. lactarium of Say; the
latter name has the priority.
Iulus niger, Leach, Tr. Linn. Soc. xi. p. 3878 (1815) =J. albipes,
C. Koch, &c. ; Leach’s name has the priority.
Iulus punctatus, Leach, loc. cit. p. 379 (1815)= 1. silvarum, Meinert,
Porath, &e.; not punctatus, Meinert, Porath.
Diplopoda in the British Museum. 249
Lulus pilosus, Newport, Ann. Nat. Hist. xiii. p. 267 (1844) =fallax,
Meinert, Nat. Tidskr. (3) v. p. 15 (1868), not fallax, Latzel,
Haase. ‘This species is Leach’s terrestris (Linn.), but not the
terrestris of Linné, Porath, and Stuxberg.
Iulus pilipes, Newport, loc. cit. p. 268 (1844)=J. varius, Fabr.
Spec. Ins. i. p. 528 (1781).
Spirostreptus gracilipes, Newp. loc. cit. p. 269, from the Philippine
Islands, is based upon a specimen of Spirobolus, and has been
redescribed as Spirobolus juloides by Karsch, Zeitschr. Naturwiss.
(3) vi. p. 65.
Spirobolus pulvillatus and simillimus, Newport, loc. cit. pp. 268, 269,
are synonyms ; the species has been recharacterized as giganteus
by Porath, Gifv. Vet.-Ak. Forh, 1872, p. 17, and as ligulatus by
Voges, Zeitschr. wissen. Zool. 1878, pp. 180, 181.
It is extremely common at Lagos.
Spirobolus caudatus, Newport, loc. cit. p. 269, has been redescribed
as P. letus by Karsch, Zeitschr. Naturwissen. (3) vi. p. 70.
Spirobolus roseus (Gervais), Ins. Apt. iv. p. 181 (1847), of which a
co-type is in the British Museum, has been redescribed as Sp.
costulatus by Porath, Bih. Sv. Vet.-Akad. Handl. iv. no. 7, p. 31.
Spirostreptus corculus, Butler, Ann. Mag. Nat. Hist. (5) ix. p. 330
(1882), belongs to the same category of species as the other
known Madagascar forms. It is based upon young specimens,
and may be easily recognized by the two wide longitudinal
flavous bands which run throughout the length of the body ; the
head is flavous, with a conspicuous black band between the eyes.
Loc. Betsileo, Madagascar.
Spirostreptus Cowani, Butler, loc. cit. p. 328, is in reality a Spiro-
bolus (cf. infra). _
Loc. Betsileo (Madagascar),
Spirostreptus avernus, Butler (Ann. Mag. Nat. Hist., June 1876,
p. 445), from Rodriguez, is a small, slender, Iuliform species of
Spirobolus. There are four labral pores on each side; the eyes
are separated by a space about equal to three times a diameter ;
the antennz are very short ; the collum laterally resembles that
of a Spirostreptus, extending as low as the second tergite. The
posterior half of the tergites is elevated and smooth above ; the
median portion is marked irregularly with crescentic impres-
sions ; the pores are situated behind the sulcus on the posterior
part of the tergites. Scobina absent. The anal tergite is pro-
duced into a triangular tail, which overhangs the valves;
valves with margins not compressed.
Sp.globulanus of Karsch, from the Comoro Islands, is very close
to this, but appears to differ in the number of labral pores,
Spirostreptus sorornus, Butler (abid.), from Rodriguez, is a Spiro-
bolus related apparently to Goésii of Porath, but with the median
part of the somites scarcely sculptured above and the copulatory
feet of a different form.
It is closely allied to S. comorensis of Karsch, and to lumbri-
cinus of Gerstiicker, both of which have been recorded by Karsch
from Mayotte.
Spirostreptus simulans, Butler (ibid.), from Rodriguez, is based upon
250 Mr. R. I. Pocock on some
young examples of a species of Spirobolus, which will probably
prove to be the same as avernus.
Spirostreptus Gulliveri, Butler (ibid.), from Rodriguez, belongs to the
same group as all the Madagascar species. ‘The segments are
strongly sulcate, being posteriorly nearly smooth above, but
decorated with an obscure anastomosing pattern of low ridges ;
the anal valves are convex, with their borders uncompressed.
Spirobolus hecate, Butler (zbid.), from Rodriguez, is a Spirostreptus,
allied to all the known Madagascar forms. The posterior part
of its somites is thickly punctulate and striolate, and the
margins of the anal valves are compressed. '
Part I1.—Descriptions of some New Species.
Spirostreptus stenorhynchus, sp.n. (Pl. XVI. figs. 1-1 d.)
Closely allied to S. Lunelit of Humbert, of which kandy-
anus, Humb., is probably the young. Both of these have been
well figured and described, so that a brief diagnosis of this
new form will suffice.
Colour jet-black, shining, and polished, tergites paler in
front ; lower half of head and antenne ferruginous, legs clear
yellow.
First tergite with a fine anterior sulcus in front of the mar-
ginal sulcus; the anterior angle very obtuse in the female,
rounded in male, and a little produced. In the anterior half of
the body the tergites are dorsally smooth or nearly so, but in
the posterior half they are distinctly punctulate and _striolate
dorsally ; the lateral strive do not in any of the segments unite
with the transverse striole of the anterior half. ‘The ventral
grooves are short, and the sterna are striolate. Anal somite
as in Lunelit. Legs with a series of hairs (about 4) on the
lower surface of each segment.
Copulatory feet differing from those of S. Luneliz in that the
anterior lateral lamina bears three teeth, of which the inferior
is the longest and the upper the shortest; in S. Lunelii the
middle one is much longer than the inferior, and the upper one
of S. stenorhynchus is absent ; the protrusible lamina is distally
bifid, the external ramus is short, curved, and sublaminate,
the internal very long, curled upon itself, and flagelliform.
Loc. Ceylon (Cuming); Punduloya (Ceylon), collected
and presented by Mr. EK. KE. Green. Also another example
without locality.
Spirobolus erythrocephalus, sp. un. (Pl. XVI. figs. 2-2 d.)
Colour. Head, legs, and antenne blood-red; first tergite
almost entirely of the same colour, but with a large black
patch on each side above the lateral angle, the two patches
connected across the middle line by an ill-defined shadowy
fuscous band, which is darker on the middle; the second,
third, and fourth tergites black at the sides, blood-red above,
new Species of Exotic Lulide. 251
with a median black spot; the following tergites in the
anterior half of the body similarly coloured, but the dorsal
red marks gradually lose their distinctness, becoming first
brown, then black ; anal somite deep black-brown.
Head smooth and shining, the sulcus distinct below ; labral
pores obsolete. yes separated by a space that is about equal
to a diameter and a half. Antenne long, much longer than
the head.
First tergite projecting inferiorly almost as low as the
second, acutely angled, with marginal sulcus, but without
other sulci. The rest of the tergites with the posterior portion
higher than the anterior, the transverse sulcus complete, the
upper surface in front of the sulcus and behind it smooth and
polished, the lateral surface thickly and longitudinally striate
below, the striz gradually disappearing above and taking a
vertical direction on the anterior part of the tergite; sterna
and inferior portion of anterior surface transversely striolate ;
pores conspicuous, above the middle of the side, in front of the
sulcus. Anal somite nearly smooth, tergite acutely angled
behind, covering the summit of the valves ; valves prominent,
the borders strongly convex, almost the posterior half com-
pressed; the sternite large, distinct, and triangular, its angle
nearly a right angle.
Legs long, with a single seta beneath each segment, ter-
minal segment padded.
Copulatory feet very wide (see fig. 2 5).
Length about 146 millim., width 10.
Number of somites 52.
Loc. N. Madagascar (Rev. R. Baron).
Closely allied to Sp. Cowant (Butler), from Betsileo. In
Cowant, however, the anal somite is blood-red, and there is
a wide flesh-coloured band along the dorsal surface of the
somites. ‘The somites, too, are not so elevated behind and
are finely striolate.
Spirobolus urophorus, sp.n. (Pl. XVI. figs. 3, 3 a.)
Colour. Head and legs flavous ; posterior half and inferior
portion of somites also flavous; anterior half fuscous; a
median dorsal flavous band more conspicuous posteriorly.
Antenne infuscate.
Body moderately robust, much narrowed posteriorly.
Head smooth, sulcate and striolate above, with two pores on
each side of the labrum. Antenna little longer than the head,
the segments, except the first and last,subequal in length. The
greatest diameter of the eyes is from above downwards ; the
distance between them much less than twice this diameter.
The collum not projecting so low laterally as the second
somite, narrowed laterally, with marginal sulcus, the angle
252 Mr. R. I. Pocock on some
rounded. The rest of the somites laterally striate, the upper
surface lightly striolate. The transverse sulcus shallow, the
pores immediately in front of it. Scobina absent. Anal
somite small; tergite produced into a conspicuous, high, com-
pressed, apically upeurled caudal process ; valves very lightly
convex, their margins largely but not deeply compressed ;
sternite rounded.
Legs moderately slender, the distal segment with many
hairs beneath, the rest with only one.
Number of somites 52.
Length about 42 millim., width 4:5.
Loc. Mahé Island, Seychelles (47.1/.S. ‘Alert’).
Spirobolus Naresti, sp.n. (Pl. XVI. figs. 4-46.)
Closely allied to Sp. Goést?, but differing in colour and in
the form of the copulatory feet.
Colour. Legs and antenne flavous and contrasting strongly
with the slate-grey tint of the rest of the body; the posterior
borders of the tergites ochraceous.
Head, eyes, antenna, collum, anal somite, pores, &c. as in
Goésii; the dorsal surface of the posterior portion of the
somites much Jess closely sculptured, with a transverse row of
ring-shaped or crescentic impressions just in front of the
sulcus.
Number of somites 49-52.
Length up to 40 millim., width 3.
Loc. Mahé Island, Seychelles (H.IL.S. ‘Alert’).
Spirobolus hoplurus, sp.n. (Pl. XVI. figs. 5-5 b.)
¢. Colour (in alcohol). Head lightly fuscous above, flavous
beneath, first tergite and anal somite blood-red, rest of the
tergites black in front, with posterior third blood-red; an-
tenne and legs flavous.
Body slender, parallel-sided.
Head striolate and convex above, flat and smooth below,
with two Jabral pores on each side. yes separated by a
space a little larger than a diameter, with inner border nearly
straight, composed of about six transverse rows of ocelli.
Antenne about as long as the head.
First tergite smooth, produced laterally considerably below
the level of the second, narrowed, the anterior border and
angle evenly rounded, with strong marginal sulcus ; the pos-
terior angle also rounded, the border above it emarginate.
The rest of the somites smooth, polished, the posterior half
slightly elevated and separated from the anterior by a shallow
groove, very faintly striate below. Pores about the middle
of the side, just in front of the transverse groove, Sterna
lightly striate transversely. Without scobina.
Anal tergite produced above intoavery long, straight, caudal
new Species of Exotic lulide. 253
process, which is wide at the base and narrowed towards the
apex ; valves moderately convex, with their margins deeply
compressed above, less deeply below; sternite distinct, tri-
angular.
Legs very long, each segment furnished with a single seta
beneath, the distal segment padded in front below.
Copulatory feet as in fig. 5b.
Number of somites 48.
Length about 80 millim., width almost 5.
Loc. N.W. Borneo (&. Everett). A second example
(Borneo, Lords of the Admiralty) has 54 somites and measures
about 115 millim.
In its slender body, long legs, and produced collum this
species superficially resembles a Spirostreptus. It is, how-
ever, unquestionably a Spdrobolus, and appears to be a well-
marked species, belonging to the same category as Sp. phranus
of Karsch and rufo-marginatus of Témisvary.
Spirobolus caledonicus, sp.n. (Pl. XVI. figs. 6-6 d.)
3 ?. Colour fusco-olivaceous, with the hinder borders of
the tergites ferruginous or flavous; legs rarely narrowly
testaceo-annulate, usually black.
Head convex, smooth and shining, the median sulcus widely
interrupted in the middle; on each side about five labral
pores, of which the three internal are set close together. Hyes
composed of about twenty ocelli set in six transverse series,
with an acute inner angle, separated by a space equal to about
twice a diameter. Antenne a little shorter than the head,
scarcely reaching to the hinder border of the first tergite.
First tergite smooth above, narrowed laterally, and extend-
ing inferiorly as low as the inferior portion of the second, the
angle acute but rounded, the anterior edge lightly sinuate but
scarcely emarginate, with scarcely a vestige of sulcus. The
rest of the tergites without scobina, smooth and polished
behind, the anterior (covered) portion being minutely striolate,
and the lateral portion just above the base of the legs finely
striate ; the transverse sulcus almost obsolete, the pores very
minute, situated above the middle of the side and just upon
the feeble transverse sulcus. Sterna striate.
Anal tergite smooth, its posterior border produced into an
angle which just overlaps the top of the valves; the valves
with their borders lightly compressed or convex and without a
trace of compression ; sternite with subcircular hinder border.
Legs with a seta on the lower surface of each segment,
male without tarsal pads, the cox of the third to fifth pairs
a little enlarged.
Copulatory feet as in fig. 6d.
Number of somites 42 (40-44).
254. Mr. C. J. Gahan on new
Length from 52-73 millim,
A number of examples from New Caledonia.
Very closely allied to Sp. albidicollis of Porath, from the
same locality, but undoubtedly differing in being smooth and
polished instead of coriaceous. The colour, too, is distinct,
the collum being by no means largely white.
EXPLANATION OF PLATE XVI.
Fig. 1. Spirostreptus stenorhynchus, sp. n.; head and first tergite of male.
la. Ditto of female. 14, Anal somite. 1c, Anterior aspect
of copulatory apparatus. 1d. Protrusible portion of copu-
latory apparatus.
Fig. 2. Spirobolus erythrocephalus, sp. n.; head and first tergite of male.
2a. Anal somite. 20, Anterior aspect of copulatory appa-
ratus.
Fig. 3. Spirobolus urophorus, sp. n.; head and first tergite. 3a, Anal
somite. -
Fig. 4. Spirobolus Naresii, sp. n.; head and first tergite. 4a, Anal
somite. 44. Anterior aspect of copulatory apparatus.
5. Spirobolus hoplurus, sp. n.; head and first tergite. 5a, Anal
somite. 5b. Anterior aspect of copulatory apparatus.
Fig. 6. Spirobolus caledonicus, sp. n.; head and first tergite. 6a. Anal
somite. 66. Anterior aspect of copulatory apparatus.
N.B.—AII the figures are enlarged.
y*
iW.
XLIV.—Descriptions of some new Genera and Species of
Longicorn Coleoptera. By C. J. GAHAN, M.A., of the
British Museum (Natural History).
‘HE genera and species described in this paper are all from
Asia.
Philus costatus, sp. n.
Flavo-testaceus; prothorace quam longiore sesqui-latiore, dorso crebre
punctulato, lateribus subrotundatis, postice sat distincte carinatis
elytris crebre punctulatis, utrisque costis quatuor paullo elevatis,
quarum duabus internis distinctioribus, obtectis ; articulis anten-
narum a tertio ad decimum subequalibus, apice intus angulatis. »
g. Antennis quam corpore longioribus; femoribus subtus longi-
tudinaliter canaliculatis, canaliculis pilosis.
9. Antennis medium elytrorum haud superantibus; femoribus
subtus versus apicem subplanatis, vyix canaliculatis.
Long., ¢ 20, 9 22-26 millim.
Hab. Siam (J. C. Bowring).
Head thickly punctured, with the punctures somewhat
coarser towards the occiput; the front divided by a median
longitudinal groove, and separated from the clypeus by a
rather deep transversely arcuate groove. Prothorax densely
but feebly punctured, with a very narrow space along the
Genera and Species of Longicorn Coleoptera. 255
middle, which is smooth, nitid, and impressed with a faint
line ; sides distinctly carinate in an oblique direction on their
posterior half, the carina becoming obsolete in front of the
middle. Scutellum and sides of mesonotum rather thickly
punctured. Elytra closely punctulate, each with three or four
raised lines or costa, of which the two innermost are usually
very distinct and unite posteriorly, to be continued a little
further back as a single line; the third costa is less distinct,
and a little before the apex bends in to join the posterior end
of the common stem of the two inner cost; the fourth line
is still less distinct and is sometimes almost quite obsolete.
A very faint pubescence, consisting of very short suberect
fulvous hairs, is present over all parts of the body.
PARAPHILUS, gen. nov.
Allied to Vesperus, but differing from that genus by the
following characters:—Head shorter, gradually narrowed
from the eyes backwards, and without a distinct constriction
orneck. Prothorax slightly rounded at the sides, and scarcely
narrower anteriorly than at the base, its lower front margin not
deeply emarginate ; prosternal process somewhat wider, so
that the anterior coxe are less approximated. First abdo-
minal sternite of the female more sharply pointed in front.
Third joint of the tarsi less deeply cleft.
The great resemblance which the unique species of this
genus presents to some of the species of PAclus is further
evidence in favour of the view that Vesperus is nearly related
to, and should be placed near, the Monodesmid group of
Prionide.
Paraphilus duplex, sp. n.
Testaceus, fulvo tenuissime pubescens; elytris pallidioribus ; pro-
thorace paullo transyerso, supra subnitido, sat dense minuteque
punctulato ; elytris crebre punctulatis, pallidis, versus basin sub-
nitidis testaceis, (¢) apicem abdominis paullo superantibus,
(2) segmentum primum abdominis ventrale vix excedentibus.
Long. ¢ 17-21 millim.
Hab. China: Pekin (Swinhoe) ; Foochow (J. de la Touche).
Head dull reddish testaceous, closely and minutely punc-
tured. Prothorax somewhat nitid above, minutely and rather
densely punctured, the sides slightly rounded in the middle,
and without any trace of a lateral carina. Elytra in the male
completely covering the abdomen, closely punctulate, pale
yellowish testaceous in colour, with the base more reddish
and slightly nitid, each narrowed and subacuminate at the
apex ; in the female scarcely reaching beyond the second
ventral segment of the abdomen, dehiscent at the suture, each
rounded at the apex.
256 Mr. C. J. Gahan on new
Antenne of the male a little longer than the body, first
joint short, third to tenth subequal to one another, each nearly
three times as long as the first, the eleventh a little longer
than the tenth; the joints from the third are all slightly flat-
tened and area little thickened, but scarcely angulate at their
distal ends. The antenne of the female extend beyond the
apex of the elytra, but do not reach to the tip of the abdomen,
and, besides being shorter, are slenderer than in the male.
PSEUDOPHILUS, gen. nov.
Head strongly exserted, with the upper side oblique, and
slightly depressed between the eyes ; front rather short, sub-
vertical, with its upper portion longitudinally impressed in
the middle, its lower part with a somewhat elliptical impres-
sion circumscribing a narrow transverse area. Mandibles and
palpi of moderate length. Inner border of each of the anten-
nary condyles forming a slightly projecting process above.
Antenne of the male about equal in length to the body; first
joint slightly curved, thickened towards its apex, third joint
scarcely longer than the first, the fourth much shorter, the
fifth distinctly longer than the fourth, the sixth to eleventh
gradually increasing in length. Antenne of the female
reaching to very little beyond the middle of the elytra. Eyes
coarsely faceted, emarginate, with the lower lobes rather large.
Prothorax slightly constricted and transversely impressed
near the base and apex, with its sides slightly rounded in the
middle ; its greatest breadth about equal to its median length.
Scutellum somewhat concave from side to side ; the anterior
part of the mesonotum, which is separated from the scutellum
by a transverse groove, has no stridulating surface, but bears
instead an impressed line along the middle, on each side of
which it is rather coarsely and thickly punctured. ‘The
elytra, which are a good deal broader than, and about three
and a half times as long as, the prothorax, are rounded at the
apex. ‘The anterior cotyloid cavities are angulate externally
and are somewhat open behind. ‘The legs, which slightly
increase in Jength from the anterior to the posterior, have the
femora somewhat compressed and broader towards the middle
of their length.
‘This genus seems to me to be best placed in the group of
the Hesperophanides, of which it possesses all the most essen-
tial characters. It has some resemblance in general appear-
ance to the Prionid genus Philus, and, like the Prionide in
general, it is without a stridulating surface to the mesonotum.
Pseudophilus testaceus, sp. n.
Brunneo-testaceus, fulvo sparse erecteque setosus; capite dense
punctato; prothorace subrugoso-punctato, disco nonnihil leviore,
Genera and Species of Longicorn Coleoptera. 257
linea media elevata plus minusve distincta; elytris fulvo tenuis-
sime pubescentibus et sparse setosis.
Long. 27-31 millim.
Hab. Valley of the Tigris and Euphrates.
Prothorax rather strongly and somewhat rugosely punc-
tured at the sides; the disk sometimes rather smooth, espe-
cially posteriorly, and provided with a median, slightly elevated
line or carina, which in some examples is almost obsolete.
The elytra each bear one or two feeble or almost obsolete
longitudinal coste, and are furnished with small punctures
of two kinds :—1st, the very minute pits from which the
short hairs of the pubescence arise, and, 2nd, the somewhat
larger and sparser pits from which the longer sete spring; they
are of a testaceous colour, somewhat darker towards the base.
The female of this species, which was taken at Fao, on the
Persian Gulf, has been presented to the Museum by Mr. W. D.
Cumming ; for the male, taken at Bussorah, I am indebted to
the liberality of Mr. R. Wyle Lloyd. A much damaged
specimen, which has long been in the Museum collection, was
brought from near the ruins of Nineveh.
Niphona tibialis, sp. n.
N. cylindracee similis, sed prothorace lateraliter minus valde den-
tato, tibiis intermediis ( g ) versus apicem intus crassatis et valde
spinosis. ;
Long. 22 millim.
Hab, Canara, South India (Bell).
Pubescence greyish, with a number of small scattered rust-
coloured spots. Prothorax rather darker than the rest of the
upperside, with two short feeble ridges running close together
along the middle of the disk, and, external to these, two or
three small tubercles on each side of the disk. There is a
rather small but distinct tooth low down on each side a little
behind the anterior margin of the prothorax, and a larger
blunter tooth or tubercle higher up and nearer to the middle.
The elytra are thickly and strongly punctured, though the
punctures are to some extent obscured by the close pubescence ;
each has at the base a few small tufts of dark brown hairs,
and, just below the shoulder, two very short dark brown
vittee ; a pubescence of a paler grey forms two somewhat
irregular and ill-defined bands crossing the elytra, one before,
the other behind, the middle. In the male the fifth abdominal
sternite has a peculiar form ; the posterior half is marked by
two grooves, which divide off two lateral portions from a
median portion; the median part is flattened, and is trans-
versely truncate behind; the lateral portions are slightly
eurved downwards, and are produced behind into two pro-
cesses, Which arch over and meet in the middle line. ‘The
second abdominal segment has a narrow transverse depression
258 On new Genera and Species of Longicorn Coleoptera.
on each side anteriorly, which is covered over by the fringe
of fulvous hairs attached to the hind margin of the first
segment. The anterior tibiz of the male are strongly bent
inwards from about the middle of their length, and each bears
a spine on the inner side between the middle and apex; the
tibie of the middle pair are strongly bent inwards towards
their apex, and each has a strong sharp spine placed on the
inner side of this somewhat thickened apical portion. The
trochanters of the middle pair are also produced into a small
spine at their extremities.
This species has a great resemblance to Niphona cylin-
dracea, White; but the latter is more strongly toothed on
the side of the prothorax, its middle tibize in the male are not
bent in strongly at the apex, and are not spined, nor are the
trochanters of the same pair spinose at their extremities ; the
lateral processes of the last ventral segment of the male are
also shorter, and in those examples that I have seen are not
curved downwards towards the middle line.
Auipus, gen. nov. (Niphonide).
Prothorax angulate or feebly tuberculate at the middle of
each side, Mesonotum without stridulating surface, but with
a smooth polished line along the middle. Prosternal and
mesosternal processes with their opposed faces snbvertical,
both processes transversely rugulose and longitudinally cana-
liculate ; the horizontal face of the prosternal process some-
what dilated posteriorly to fit into corresponding depressions
on the sides of the cox. Eyes rather small, subdivided.
Antenne of the male nearly one half longer than the body ;
those of the female about equal in length to the body ; first
and third joints subequal in length, the fifth longer and
slightly curved, each ot the succeeding joints, the eleventh
excepted, subequal to or slightly shorter than the third.
This genus may be recognized by the rather exceptional
character, for this group, which it possesses in having the
sides of the prothorax tuberculate in the middle.
Alidus biplagiatus, sp. 0.
Dilute fuscus, maculis punctisque flavo-albidis sparsim aspersis,
prothoracis dorso plaga ovali fusco-velutina utrinque ornato,
Long. 23—25, lat. 9-11 millim.
Hab. China (J. C. Bowring) and Assam (Atkinson).
Dark brown, with darker brown points and numerous
minute scattered yellowish-white points and spots, the most
conspicuous of which are a spot on the base of the fourth
antennal joint, a small patch on the middle of each tibia, and
a few spots forming a narrow zigzag band between the middle
and posterior thirds of each elytron. Prothorax with a large
Geological Society. 259
oval dark brown velvety area on each side of the disk, which
is bounded by a narrow pale yellowish margin. The elytra
are furnished with some sparsely scattered granules towards
the base, and with some sparse punctures, each of which is
marked in position by a small dark spot.
This species may be easily recognized by the two large
oval and dark brown velvety areas on the pronotum.
PROCEEDINGS OF LEARNED SOCIETIES.
GEOLOGICAL SOCIETY.
December 21, 1892.—Prof. J. W. Judd, F.R.S.,
Vice-President, in the Chair.
The following communications were read :—
1. “On a Sauropodous Dinosaurian Vertebra from the Wealden
of Hastings.” By R. Lydekker, Esq., B.A., F.G.S.
In addition to Hoplosaurus armatus and Pelerosaurus Conybearet,
there is evidence of another large Sauropodous Dinosaur in the
Wealden, now known as Morosaurus brevis. Up to the present time
it has been impossible adequately to compare Hoplosaurus armatus
with Morosaurus brevis; but recently Mr. Rufford has sent to the
British Museum an imperfect dorsal vertebra of a large Sauropodous
Dinosaur from the Wealden of Hastings, which enabled the required
comparison to be made. The Author describes the vertebra, con-
trasts it with that of Hoplosaurus armatus, and gives presumptive
evidence that it should be referred to the so-called Morosaurus
Becklesi (Marsh), which apparently cannot be separated from
M. (Cetiosaurus) brevis. He has not been able to compare Mr.
Rufford’s specimen with the dorsals of the American Morosaurus,
in order to discover whether the English Dinosaur is correctly
referred to that genus.
2. “ On some additional Remains of Cestraciont and other Fishes in
the Green Gritty Marls, immediately overlying the Red Marls of the
Upper Keuper in Warwickshire.” By the Rev. P. B. Brodie, M.A.
The vertebrate remains occur in a very thin band of marly friable
sandstone lying between two beds of green marl, though in some
places the same bed has itself no admixture of sandy material.
Bones and teeth are so numerous that it might almost be called a
bone-bed. It does not exceed three inches in thickness. It
contains ichthyodorulites of Cestraciont fishes, abundant palatal
teeth of Acrodus keuperinus, ganoid fish-scales, and abundant
broken bones, some of which may belong to fishes, others to laby-
rinthodonts, and amongst the latter a fragment of a cranial bone.
3. “ Calamostachys Binneyana, Schimp.” By Thomas Hick, Esq.,
B.A., B.Sc. Communicated by J. W. Davis, Esq., F.G.S., F.L.S.
A number of preparations lately placed in the hands of the
Author by Mr. W. Cash, of Halifax, give much additional in-
formation on many important points of detail connected with the
260 Miscellaneous.
structure and characters of the sporiferous spike known as Calamo-
stachys Binneyana, and also throw considerable light on the question
of its affinities. The Author fully describes the structure of the fossil,
and diseusses its systematic position. His description shows the
close resemblance between the structure of this spike and that of
Equisetum ; and it is noted that Carruthers, in 1867, did not hesitate
to describe Calamostachys Binneyana as the fruit of Calamites,
and that Binney, when making it the fruit of Calamodendron
commune, Was in practical agreement with Carruthers. William-
son, however, realizing the difficulty of reconciling the supposed
vascular central part of the axis of the spike with a Calamitean
affinity, suggested that the relationship lies rather with the Lyco-
podiacew. Now the establishment in the present paper of the
existence of parenchymatous pith in the axis of the spike removes
the chief ground for Williamson’s suggestion ; and the demonstra-
tion of the existence of primary vascular bundles with carinal canals
affords positive proof of the closeness of its relationship with the
Calamite. The Author gives reasons for concluding that there is
much probability that the fossil described in the paper is a spike of
a Calamites of the type of Arthropitys.
4. “Notes on some Pennsylvanian Calamites.” By W. 8.
Gresley, Esq., F.G.S.
The Author believes that a group of specimens from the Pottsville
conglomerate series demonstrates that the so-called Calamite-casts
in sandstone are not always to be regarded as casts of pith-cavity
only, but are often casts of the vascular cylinder as well; also that
these specimens show that some Calamites, at all events, possessed
furrowed exteriors or barks. A specimen from the Pittston bed of
anthracite in the Wyoming basin also seems to suggest that the true
bark had not always a smooth exterior.
A specimen from the same geological horizon as the last specimen,
and from a neighbouring locality, indicates a Calamite of gigantic
proportions. Another specimen, from the Pittsburgh bed, is re-
markable as showing two branch-scars not on, but below the nodes
of the Calamite.
MISCELLANEOUS.
The Dates of Moore's ‘ Lepidoptera Indica,’
To the Editors of the ‘ Annals and Magazine of Natural History,’
GrntTLEMEN,—The extreme importance of the exact dating of publi-
cations which contain new generic or specific names is now so
generally recognized that no apology is needed for bringing before
the notice of your readers a particularly gross case, whereby the
author eventually obtains six months or more priority over other
writers. The foolish practice of throwing away the wrappers of
separately published parts of a serial work, instead of binding them
up in place exactly as issued, is so common that those who pre-date
eee
Miscellaneous. 261
their publications are invariably assisted in their deception. Fortu-
nately in the Natural-History Museum a sharp look-out is kept for
cases of this kind, and covers are always considered when a book is
sent to the binders.
The book that I wish to call attention to is F. Moorr’s ‘ Lepi-
doptera Indica,’ of which thirteen parts have already appeared, and
of which the proper dates are the following :—
Pt. 1. pp. i-viii, 1-24, April 1890.
2. ,, 25-56, June 1890.
3. 3/-80, Sept. 1890.
4. ,, 81-96, Nov. 1890.
5. ,, 97-112, Dec. 1890.
6. ,, 113-128, Feb. 1891. Not Aug. 1890, as stated in the
signatures !
i 129-152, Aug. 1891. Not Aug. and Nov. 1890 and
July 1891, as stated in the
signatures !
8. ,, 153-176, Oct. 1891. Not July 1891, as stated in the
signatures !
9. 177-208, Jan. 1892. Not Aug. and Sept. 1891, as
stated in the signatures !
10. ,, 209-240, April 1892. Not Oct. 1891 and Jan. 1892,
as stated in the signatures !
11. ,, 241-272, Sept. 1892. Not Jan., Mar., and Apr. 1892,
as stated in the signatures !
12. ,, 273-310, Dec. 1892. Not May and June 1892, us
stated in the signatures !
1... 1-32, Jan. 1893. Not Aug. 1892, as stated in the
signatures !
{ do not know whether the author or the publisher is responsible
for this deceptive dating ; but it is to be hoped that neither Mr. Lovell
Reeve nor Mr. Moore will any further indulge in such objectionable
vagaries.
C. Davies SHerborn
Natural-History Museum, (Index gen. et spec. anim.).
Cromwell Road, S.W.
Feb. 18, 1893.
Diffuse Pigmentation of the Epidermis of the Oyster due to prolonged
exposure to the light: Regeneration of Shell and Loss of Adductor
Muscle.
Prof. Ryder reported on behalf of Prof. R. C. Schiedt that oysters
which had the right valve removed and exposed to the light in this
condition, in a living state, for a fortnight or so, developed pigment
over the whole of the epidermis of the exposed right mantle and on
the upper exposed sides of the gills, so that the whole animal from
this cause assumed a dark-brown colour. Animals so exposed not
only attempted to reproduce the lost valve and hinge, but also partly
succeeded in so doing, even re-establishing the insertion of the
Ann. & Mag. N. Hist. Ser. 6. Vol. xi. 19
262 Miscellaneous.
diminutive pedal muscle upon the inner face of the imperfectly
reproduced right valve, which was deformed owing to the lack of
support of the right mantle, because of the removal of the original
right valve. As a consequence the right mantle was rolled up at the
edge, and this deformation of the mantle was reflected in the
attempted regeneration of the lost right valve. The pigment
developed during exposure to light in the mantle and gills in oysters
with the right valve removed which were kept alive in the aquaria at
Sea Isle City by Prof. Schiedt was wholly confined to the epidermis
as it normally is at the mantle-border in the unmutilated animal in
nature. The inference to be drawn from these facts is that the
development of pigment in the mantle and gills was wholly and
directly due to the abnormal and general stimulus of light over the
exposed surface of the mantle and gills, due to removal of the right
valve, and that the mantle-border, the only pigmented portion of the
animal, is pigmented because it is the only portion of the animal
which is normally and constantly subjected to the stimulus of light.
Oysters which had the right valve removed were found to live
perfectly well in the marine aquaria at Sea Isle, and would no
doubt have survived till now had Prof. Schiedt been able to continue
his experiments there. The most remarkable results obtained as a
consequence of these experiments were that the adductor muscle
was soon attacked by bacteria and destroyed by putrefaction, while
the great ganglion underlying it remained uninjured. The peri-
cardiac cavity was also torn open, exposing the heart completely, in
some instances. In these cases the heart continued to beat and
propel the blood through the other organs of the body as if nothing
untoward had happened. The maximum rate of pulsation of the
heart noted was 52 per minute, which is much greater than the
rate hitherto reported.
The anus was also retracted into a new and more anterior
position, owing to the loss of support which it had suffered in conse-
quence of the sloughing away of the adductor muscle. Whether
the adductor muscle thus sloughed away would ultimately be repro-
duced was not determined, since the experiments were interrupted
before the animals had time to present evidence of such regeneration
of the lost muscles.
These experiments open up a most suggestive line of investigation
upon other univalve and bivalve mollusca, viz.: experimental
researches as to the effect of removing the valves and exposing
them to the light. Many other species, both marine and fresh-
water, might obviously be experimented upon with very instruc-
tive results as respects the questions raised by the present
communication.—Proc. Acad. Nat. Sci. Philad. Nov. 15, 1892.
The Hermaphroditism and Viviparity of the Oysters of the
North-west coast of the United States.
Prof. J. A. Ryder also reported on behalf of Prof. R. C. Schiedt,
of Franklin and Marshall College, Lancaster, Pa., the latter's
discovery of the fact that the oysters native to the north-west coast
Miscellaneous. 263
of the United States are hermaphrodite and viviparous. Specimens
from the coast of Oregon and Washington show that the same
condition exists in the reproductive follicles as in those of Ostrea
edulis of Europe. The presence of eggs and of spermatoblasts and
spermatozoa in the same follicles is the invariable rule. The ova, like
those of O. edulis, are much larger than those of O. virginica, though
perhaps not quite so large as the former, The embryos are
fertilized in the gill and mantle cavities, where they undergo
development.
These north-west-coast oysters also resemble the oysters of Europe
in that they are small and have little or no indication of purple
pigment on the impression or point of insertion of the adductor
muscle, which is so conspicuous a feature in Ostrea virginica of our
eastern coast.—Proc. Acad. Nat. Sci. Philad. Nov. 15, 1892.
Large Variations in the Metamorphosis of the same Species.
_ Anelaborate memoir entitled, ‘‘The Embryology and Metamorpho-
sis of the Macroura,” by W. K. Brooks and F. H. Herrick, makes 140
pages quarto of the fifth volume of the Memoirs of the U.S. National
Academy of Sciences, and is illustrated by 57 plates. The species
microscopically investigated and here reported upon are of the genera
Gonodactylus, Alpheus, and Stenopus. ‘The authors mention, in the
introductory pages, as one remarkable result of their study of the
genus Alpheus, the discovery that while the larval stages of different
species are similar, the individuals of a single species sometimes
differ more from each other as regards their metamorphoses than
the individuals of two very distinct species, and make on this point
the following remarks :—
This phenomenon has been observed by us and carefully studied
in two species—Alpheus heterochelis and Alpheus Sauleyi—and it is
described in detail, with ample illustrations, in the chapter on the
metamorphosis of Alpheus. In the case of the first species the
difference seems to be geogrdphical, for while all the individuals
which live in the same locality pass through the same series of
larval stages, the life-history of those which are found at Key
West is very different from that of those which live on the coast of
North Carolina, while those which we studied in the Bahama Islands
present still another life-history. In the case of the second species
—Alpheus Sauleyi—the difference stands in direct relation to the
conditions of life. ‘The individuals of this species inhabit the tubes
and chambers of two species of sponges which are often found
growing on the same reef, and the metamorphosis of those which
live in one of these sponges is sometimes different from that of those
which inhabit the other. In this species the adults also are
different from each other, but as we found a perfect series of
transitional forms there is no good reason for regarding them as
specifically distinct ; and in the case of the other species— Alpheus
heterochelis—we were unable, after the most thorough and minute
comparison, to find any difference whatever between adults from
North Carolina and those from the Bahama Islands, although their
264 Miscellaneous.
life-histories exhibit a most surprising lack of agreement. In fact,
the early stages in the life of Alpheus heterochelis in the Bahama
Islands differ much less from those of Alpheus minor or Alpheus
Normant than they do from those of the North Carolina Alpheus
heterochelis ; and, according to Packard, the Key West heterochelis
presents still another life-history.
In the summer of 1881 I received the ‘ American Naturalist’ with
Packard's very bricf abstract of his observations at Key West upon
the development of Alpheus heterochelis, and read with great surprise
his statement that this species has no metamorphosis, since, while
still inside the egg, it has all the essential characteristics of the
adult. As I had under my microscope at Beaufort on the very
day when I read his account a newly hatched larva of the same
species and was engaged in making drawings to illustrate the meta-
morphosis of which he denies the existence, and as my experience
in the study of other Crustacea had taught me that all the larve of
a species at the same age are apparently facsimiles of each other
down to the smallest hair, Packard’s account seemed absolutely
incredible, and I hastily decided that, inasmuch as it was without
illustrations and was written from notes made many years before,
it involved some serious error and was unworthy of acceptance.
This hasty verdict 1 now believe to have been unjust, since my
wider acquaintance with the genus has brought to my notice other
instances of equally great diversity between the larve of different
specimens of a single species.
The phenomenon is, however, a highly remarkable one and
worthy the most thorough examination, for it is a most surprising
departure from one of the established laws of embryology—the law
that the embryonic and larval stages of animals best exhibit their
fundamental affinities and general resemblances, while their specific
characteristics and individual peculiarities make their appearance
later.—Am. Journ. Sci., Feb. 1893, pp. 166, 167.
Absorption in the Actinise and the Origin of the Mesenterial Filaments.
By Vicror Wittem, Assistant in Zoology at the University of
Ghent.
If albumen stained with carmine is administered to specimens of
Actinia or Sagartia it is found that at the end of a few hours
particles of carmine are present in the cells of the lining of the
enteric cavity. Properly speaking this absorption does not take
place in the enteroids, as is stated by Krukenberg * and Metschni-
koff +, but at first and chiefly in the region of the radial partitions
which adjoins these filaments, a region where the epithelium forms
a thickening parallel with the edge of the septum ¢; the grains of
* Krukenberg, “ Ueber den Verdauungsmodus der Actinien,” Vergl.-
phys. Studien an der Kiiste der Adria, 1 Abth., 1880.
t Metschnikotf, “Ueber die intracellulare Verdauung bei Ccelente-
raten,” Zool. Anz. 1880, p. 261.
} Vide Hertwig, “ Die Actinien,” Jenaische Zeitschrift, 1879, Taf. xxi,
fig. 13,
Miscellaneous. 265
carmine are again met with in the cells of the grooves which sepa-
rate either the lateral ciliated bands from the median band which is
provided with nematocysts, or the lateral bands from one another *.
If nutriment tinctured with carmine is supplied in great abund-
ance, not only does the red coloration become more intense in the
regions which have just been indicated, but we also find grains of
carmine in large numbers in the cells of the whole endodermic
lining. In Sagartia again, contrary to the assertion of Metschni-
koff, these may be easily observed in the acontia, throughout the
entire zone which is devoid of nematocysts and gland-cells f.
Moreover, identical results are obtained by using mussel’s liver ;
the yellowish-brown coloration of the fatty globules contained in
the cells of this organ enables us to recognize the same progression
in the absorption of these droplets. The examination of sections of
Actinie which have been fed abundantly upon the fat-body of cater-
pillars shows, in the cells of the entire endodermic lining and in
those of the regions of the mesenterial filaments or of the acontia
which I have indicated above, the presence of numerous fatty
droplets, which are often of enormous size relatively to the dimen-
sions of the elements which contain them.
In the numerous fresh or prepared specimens which I have
examined I have never seen a particle of carmine or a fatty globule
in a cell of one of the three swellings which enter into the compo-
sition of a typical mesenterial filament.
Absorption therefore belongs in itself to the whole of the
endodermic lining, and if (as, for instance, when the animals were
supplied with nutriment stained with carmine) it appears to be more
evident in certain parts of the enteroids and in their neighbourhood,
this is due to two causes: in the first place to the fact that the zones
in question, being closer to the spot § at which the disaggregation of
the ingested matters takes place, capture the grains of carmine as
soon as they are set at liberty by the dissolution of their sub-
stratum ; those only which penetrate into the radial divisions are
absorbed by the general lining. In the second place it is due to
the existence along the enteroids of a thickening in which the more
elevated cells accumulate more of the carmine particles.
The localization of the absorbent cells brings out an important
point in the disposition of the different tissues in the Actiniw, which
is in relation to their nutrition and the absence of an actual circu-
latory system. All the regions of the body, of however little extent,
include cells in which there takes place an intra-cellular digestion
and the elaboration of the assimilable substances which are destined
for the elements in the immediate neighbourhood: the outer wall,
including the tentacles and the stomodeeum itself, is clothed with an
endodermic lining; the swellings of the enteroids are separated by
bands of absorbent cells, while the acontia exhibit on one of their
* Ibid. Taf. xxi. figs. 10 and 14. + Ibid, Taf. xvi. fig. 11.
{ These are shown in the Hertwigs’ figures as constituted by a granular
epithelium.
§ The central] region of the general cavity.
266 Miscellaneous.
faces a zone which may be termed the nutritive zone of the acon-
tium; in the immediate neighbourhood of the mesenterial filaments
—at the spot where the median swelling alone persists—the epithe-
lium of the septa exhibits a special development, which seems to me
to be in direct relation to the nutrition of these organs of secretion.
These physiological facts appear to me to contribute towards
clearing up the origin of the convoluted filaments in ontogeny.
Heider *, relying on the identity of their histological structure,
admitted the ectodermic origin of the enteroids in Certanthus, and
E. B. Wilson considered it to be very probable that the lateral
lobes are the homologues of the ectodermic bands of the Alcyonaria,
while in his opinion the central lobe is of an endodermic nature.
A. Andres + thought that he had succeeded in deriving certain fila-
ments of the Actiniaria from ectodermic proliferations of the
stomodeum. But it seems to follow, from certain observations of
H. Y. Wilson §, that this ectodermic origin ought to be attributed
not only to the lateral lobes, but also to the median one, at the very
least in the case of the primary mesenteries.
However it may be with regard to supposing, as appears to me to
be very probable, that the three lobes composing a typical filament
are of an ectodermie nature, it is not less probable that, relying on
the identity in histological structure and physiological function, we
ought to consider as endodermic the regions of the filaments which
separate the lobes from one another and the nutritive zone of
the acontia.— Zoologischer Anzeiger, xvi. Jahrg., no. 409 (January 9,
1893), pp. 10-12.
On Phagocytosis observed, in the living Animal, in the Gills of
Lamellibranch Mollusea\|. By M. px Bruyne.
In a communication of Noy. 3, 1891 (*‘ Annales de la Faculté de
Médecine de Gand’), I mentioned incidentally some phenomena of
phagocytosis which had been observed in the normal ciliated
epithelium of the gills and mantle of Lamellibranch Mollusks ;
these observations related exclusively to fixed preparations. Since
then I have studied the phenomenon in the living animal, by
operating upon four very common forms—Mytilus, Unio, and
Anodonta, which are very suitable for this kind of observation, and
Ostrea, which is much less so, probably on account of the thickness
of its gills.
If a fragment of gill be removed from a freshly opened specimen
of one of these animals (Mytilus by preference), and examined
under the microscope, the observer is struck by the clearness with
* V. Heider, “Certanthus membranaceus,” Sitzgsber. Akad. Wiss. Wien,
Bd. lxxix. (1879).
+ E. B. Wilson, “The Mesenterial Filaments of the Alcyonaria,”
Mittheil. zool. Stat. Neapel, Bd. v. (1884).
t A. Andres, quoted in the foregoing paper.
§ H. V. Wilson, *‘ The Development of Manicina areolata,” Journal of
Morphology, 1889.
|| The investigations were conducted at the Laboratory of Normal
Histology of the University of Ghent.
Miscellaneous. 267
which all the structural details are seen. I have obtained the best
results by using Zeiss’ No. 4 eyepiece and F objective (magnifying-
power 1010). Under these conditions the globules of the blood
appear as little amceboid corpuscules with very distinct nuclei. Their
protoplasm is sometimes perfectly hyaline, with a few scattered
granulations which exhibit an active Brownian movement; some-
times finely granular; or again packed with hyaline globes of
variable dimensions, which may or may not contain in their turn
some very refringent spheres. The pseudopodia, which may be
seen to appear and disappear, are either numerous and delicate,
always perfectly hyaline and extended in all directions, or else on
the contrary isolated, broad, and true lobopodia. It is possible to
follow the blood-corpuscles in their peregrinations ; in this way, on
the confines of the epithelium, one of these migratory cells is often
seen to leave the connective tissue and penetrate between the
ciliated cells, separating and depressing them, and thus producing
behind it a solution of continuity which goes on increasing. But
we soon find that it is something more than a simple separation or
depression: the actual substance of the cells is broken into, and
exhibits unequivocal signs of alteration. The protoplasm diminishes
in places and appears to be eaten away at spots where it has been
in contact with a leucocyte. In no case was I able to discover
that the amceboid globule here contained fragments of epithelial
cells. The question arises whether we have to deal with digestion
at the surface of the leucocyte, or at least with dissolution pre-
paratory to digestion. However this may be, a cavity is thus
produced which is hollowed out by the leucocytes at the expense of
the epithelium. This cavity continues to increase in size, and soon
extends to several neighbouring cells; it has very distinct limits,
which are sometimes exceedingly irregular, and successively assumes
the most varied shapes.
Since it most often happens that the same phenomenon takes
place at adjoining spots, the result frequently is that the last
remaining cells which still separate the cavities disappear in their
turn by the same process; extensive lacune are thus formed by
means of fusion, in which there move a larger or smaller number of
leucocytes, each of which acts as a phagocyte on its own account *.
The free portion of the ciliated cells is most often intact, and there
always persists a border constituted by the fragments of protoplasm
which have escaped from the devastation. The body of the phago-
cytes grows considerably, in most cases becomes packed with hyaline
spheres, and continues to move freely in the lacuna. Here and
there they finally cross the border itself at some spot or other, and
leave the lacuna to reach the surface of the branchial mucous
membrane; where they are speedily carried away by the current
which is produced in the water by the motion of the vibratile cilia.
The very interesting phenomenon which I have just described,
and which, so far as I am aware, has not hitherto been recorded,
* In these lacune it is also not unusual to meet with one or more
leucocytes which have come from the depths, and are charged with
matter of a yellow or dirty brown colour.
268 Miscellaneous.
is in the living animal usually observed only in the epithelium of
the lower (free) border of the gills (it also occurs pretty frequently
in the buccal tentacles); but in most cases, over the entire extent
of the epithelium, it is possible to witness the penetration of
blood-corpuscles, which come from the lacunar connective tissue and
are charged with granulations or hyaline spheres of variable, and
sometimes very considerable, dimensions. ‘They advance slowly,
thrusting out pseudopodia between the epithelial cells, and thus
opening a path for themselves to the free surface, where they undergo
the same fate as those which leave the lacune: 7%. e. they are
carried away by the current. The presence of phagocytes among
the epithelial cells has recently been mentioned by Dr. Janssens
(* La Cellule,’ t. ix.).
It may be asked what is the physiological object of this pheno-
menon. Without pretending to solve the question at present, and
reserving to myself the right of reverting to it at greater length,
I wish to point out one way of looking at it. In preparations
fixed by means of Flemming’s or Hermann’s fluid, I have often
discovered, either in the phagocytes, or in the tissues, the presence
of degenerate leucocytes. They appeared in the most varied shapes ;
but they were always composed of a sphere or of an irregular body,
but little or not at all stainable, and serving as a substratum for
one or more corpuscles which took a decided saffron stain: the
substratum was of protoplasmic origin, the chromatic element
was derived from the nucleus. In the same way in the phagocytes,
the spheres, which were hyaline in the fresh state, seemed, from
their low degree of colourability, to be composed of more or less
altered protoplasm. Given these details, the question arises
whether it would not be possible to find in the present case an
application of the views of Metschnikoff and others, on the subject
of a continual strife between the cells of the same organism,
which results in the sweeping of the tissues, and in the removal
of anatomical elements which are enfeebled, sickly, or decaying,
by means of ameeboid cells still in the full vigour of active life. The
phagocytes which have come from the depths and contain spheres
or coloured matters would have loaded themselves with feces and
dead cells, met with in their diapedeses, and would transport
them to the exterior through the epithelium; on the other hand,
the migratory elements which hollow out the epithelium would
have been summoned thither by chemiotaxy: the vibratile cells of
the lower border of the gill-lamelle are, by their very position,
more than all the others exposed to all kinds of destructive
influences; as a Consequence they would rapidly become worn
out and enfeebled, and their debilitated bodies would attract the
leucocytes.
N.B.—The way in which the tissues thus injured are renewed
still remains to be examined; this will form the subject of investi-
gations to which I intend to devote myself later on. In conclusion,
I do not think it necessary to dwell upon the fact that the lacune,
which have been mentioned so often in the present communication,
cannot be confounded with the lattice-like clefts, which permit the
passage of the water into the interbrancbial spaces.— Comptes Rendus,
t. exvi. no, 2 (Jan. 9, 1893), pp. 65-68,
a
Ann.d Mag. Nat. Hist. §
er ee
6 Vol..W. Pl. VI.
) Ann.& Mag. Nat. Hist 8.6 Vol VE. PL. VIL
THE ANNALS
AND
MAGAZINE OF NATURAL HISTORY.
[SIXTH SERIES. ]
No. 64. APRIL 1893.
XLV.—Preliminary Account of the Freshwater Medusa of
Lake Tanganyika. By Rh. 'T. Ginruer, B.A.
[Plates XIII. & XIV.]
In August 1891 Mr. F. L. M. Moir, Manager of the African
Lakes Company, informed Dr. A. Giinther that he had
observed Medusz in large numbers in one of the bays of Lake
Tanganyika. ‘‘ They were about ? of an inch in diameter,
and their rounded top was so clear as to give the idea they
were merely circular bodies and not the shape of an ordinary
Medusa ; tlie more opaque ovaries gave this appearance.” At
the same time Mr. Moir kindly undertook to obtain specimens
properly prepared with osmic acid; and, thanks to the assist-
ance of Mr. A. J. Swann, these specimens arrived in good
condition at the end of last year. This material was placed
by my father in my hands for examination and description.
Although Mr. Moir has thus the merit of having been the
first to succeed in having specimens brought to Europe, he is
not the original discoverer of the species. In fact, from April
and during our summer months the Meduse seem to be
exceedingly abundant about the bays of the lake, so much so,
that they attract the notice of all visitors.
In December 1883 Professor E. von Martens * communi-
* Sitzgsb. naturf. Fr. Berl. 1883, p. 197.
Ann. & Mag. N. Hist. Ser. 6. Vol. xi. 20
270 Mr. R. T. Giinther on the
cated to the ‘Gesellschaft naturforschender Freunde zu
Berlin’ the substance of a letter from Dr. R. Bohm, then
travelling in Africa, relating to the discovery of a Craspe-
dote Medusa in Lake Tanganyika. ‘The information to be
gathered trom Béhm’s account was restricted to the following
points :—that the Medusa was provided with a velum; that
it had a small short broad stomach, at the base of which the
genital organs are situated; and, finally, that the tentacles
were very numerous and of unequal length. On account of
the complete physical isolation of Lake Tanganyika from all
other localities from which Meduse are known, Bohm very
justly considered the newly discovered species to be unknown
to science, and accordingly proposed the specific name of
tanganjice, leaving the generic name, and at the same time
deploring that the absence of all necessary literature prevented
him from further determining the zoological position of the
species.
Evidently the same animal has since been observed by
H. von Wissmann, who, when describing his passage
across the lake on the 13th of April, 1887*, says, ‘ I was
ereatly astonished to observe a number of sea-nettles surround
our boat for about half an hour. They were transparent,
of the shape of a disk, and like a mark-piece in size; round
the edge was a milky circle, hanging down in fibres, by
means of which they swam.”
Such, then, are the sole facts which, so far as I know, have
been hitherto published concerning what is probably the most
aberrant of the very few freshwater Meduse at present known
to us.
The specimens were preserved in strong alcohol and were
much blackened by the osmic acid which had been employed
as a fixing agent, and which had acted especially upon the
margin of the umbrella and the manubrium. In size the
individuals varied from 1 to 1°8 centimetre across the bell,
while the largest specimen was as much as 2°2 centim. in
diameter.
The umbrella is characterized by its flattened shape, the
diameter being about four times as large as the greatest
thickness. ‘The central portion of the umbrella, measurin
about two thirds of the whole diameter, is much thickened waa
has the shape of a nearly hemispherical lens (PI. XLV. fig. 4),
while the peripheral portions are very much thinner. The
margin of the umbrella is bent over and incurved towards the
mouth. ‘The exumbrella is smooth and covered with an epithe-
* H. von Wissmann, ‘ Through Equatorial Africa’ (English transla-
tion), p. 253.
Freshwater Medusa of Lake Tanganyika. 271
lium of flattened polygonal cells. The velum is well developed
in some specimens, but not so conspicuous in others. ‘he
circular muscles of the velum (fig. 5, m.v.) are arranged
on the inner side in separate concentric bands, varying in
number according to the breadth of the velum. The muscle-
bands near the free margin are always more highly developed
than those situated near the attachment of the velum.
The gastrovascular system differs from that of all other
Medusz hitherto described in the relative size of its parts.
The mouth and stomach are both circular, widely open to the
exterior, and of so great a diameter—two thirds that of the
umbrella—that the lips of the mouth probably never com-
pletely close the stomach in the adult animal. In all the
specimens examined the central portion of the shallow stomach-
cavity is almost completely filled with the large central lens-
shaped boss of the umbrella (fig. 4), so that the stomach is
reduced to a circular trench all round the central boss inside
the manubrium, and can hardly be said to possess any central
cavity at all. It might be suggested that this extraordinary
condition is merely due to post-mortem changes; but the
fact that the relative dimensions of the parts do not vary
appreciably in any of the specimens examined, seems to me to
be sufficient reason for regarding the condition described as
the normal one, at any rate when the animal is not feeding.
A possible explanation of this curious dilatation of the mouth
and stomach was suggested to me by Mr. G. C. Bourne. It
is obvious that any increase in the diameter of the mouth and
stomach would involve a corresponding increase in the circum-
ference of the manubrium, which, as will appear in the sequel,
is the bearer of the reproductive organs. ‘The large size of
the mouth would appear, then, to be correlated with an enlarge-
ment of the area upon which the reproductive organs, sexual
or asexual, are developed.
The slightly thickened margin of the mouth is quite
simple, without processes of any kind. The radial canals are
four in number in the majority of individuals; but instances
of the occurrence of five or six seem to be by no means rare,
specimens with six being rather more numerous than those
with five. The radial canals radiate from the stomach in a
more or less horizontal plane, but assume a more vertical
direction just before they enter the ring-canal, in consequence
of the inflexion of the margin of the umbrella. The ring-
canal varies in size in different individuals ; it is lined by an
epithelium of small squarish cells, similar to those lining the
radial canals. On the outer side of the canal, however, these
cells are in very great numbers, forming a lobulated mass
20*
272 Mr. R. T. Giinther on the
projecting into and partly blocking up the lumen of the canal
(fig. 5). Hollow cavities often occur in this lobulated mass
of endoderm-cells.
As in Limnocodium, the tentacles are very numerous and
may amount to over two hundred in number. In some of the
smaller specimens the four primary perradial tentacles are
conspicuous on account of their greater length and thickness,
but in most of the larger examples they are almost equalled
in point of size by the interradial and even the adradial
tentacles, so that they are no longer readily recognizable.
Alternating with these primary, secondary, and tertiary series
tentacles of the fourth and even of the fifth and sixth orders
arise with great regularity, and it is only when the seventh
order of tentacles are intercalated that the typical regularity
of arrangement cannot be further traced. In preserved speci-
mens the relative length of the tentacles varies to a great
extent, indicating a considerable capability of contraction and
extension. The tentacles are hollow, the lumen of the
larger tentacles at any rate being clearly continuous with the
lumen of the ring-canal. The lumen of the tentacles is lined
with large, thin-walled, columnar endoderm-cells, which are
continuous with the endodermic lining of the ring-canal
(fig. 5). The transition from the tentacular endoderm-cells
to those of the ring-canal is very abrupt. The thread-cells are
of small size and generally arranged in little wart-like groups
or batteries (fig. 5), which are more or less disposed in whorls
and which are especially closely set at the tips of the tentacles.
The proximal ends of the tentacles are all adnate to the
margin of the umbrella for some considerable distance
(figs. 4 and 5, te.), the older tentacles being adherent along a
greater length than the younger ones. At the points of
fusion the tentacles become partly imbedded in the substance
of the margin of the umbrella, while the tentacular ectoderm
becomes continuous with that of the exumbrella, so that in a
section across the line of fusion an ectoderm lamella is seen
between the mesogloea of the tentacle and that of the
umbrella (fig. 5, ect.).
On the side of the ring-canal opposite to that at which the
tentacles arise and just external to the origin of the velum is
situated a circle of sense-organs. ‘These vary consider-
ably in number and are arranged at irregular intervals, so
that neither their number nor grouping can be brought into
relation with that of the tentacles. ‘Lhe structure of these
marginal bodies is peculiar.
They are refringent egg-shaped bodies attached to one side
of a round or ovate capsule, the walls of which are lined with
a it te
ap a er tA
Freshwater Medusa of Lake Tanganytka. 273
a flattened epithelium (fig. 7; s in figs. 4 and 5). These
egg-shaped bodies are composed of numerous cells and appa-
rently attached by a thin stalk or thread to that side of the
capsule which is nearest the ring-canal. The basal cells of
the marginal bodies have a granular and opaque appearance,
being coloured dark brown by the action of the osmic acid.
The apical cells, however, are quite clear and refrinzent
(fig. 8). ‘These extraordinary bodies at once recall the corre-
sponding structures in Limnocodium described by Professor
Ray Lankester*, to which they bear a remarkably close
resemblance. At the same time they differ in structure from
all other sense-organs hitherto described in Medusz, the chief
difference between the marginal bodies of Limnoenida (as
I shall call the Tanganyika Medusa) and those of Limno-
codium consisting in the absence of any tubular extension of
the capsules into the adumbral ectoderm layer of the velum
as is the case in Limnocodium. Another less important
point of difference lies in the marked contrast between
the granular basal cells and the refringent apical cells
in the marginal bodies of Limnocnida. However, in spite
of these differences there is a very great similarity of
structure in the organs of both, and probably the marginal
bodies of Limnocnida will be shown to be endodermal in
origin, just as the “refringent bulbs” of Limnocodium have
been proved to be.
Among the individuals of Limnocnida examined some
were found to have the outer wall of the manubrium quite
smooth, while others have the proximal half of the manu-
brium covered with small swellings visible to the naked eye.
When subjected to microscopic examination the individuals
with the smooth manubriums turned out to be males and
females with the external wall of the manubrium covered with
spermatozoa or ova in all stages of development.
The other set of individuals with the small swellings proved
to be examples showing stages of bud-formation. Buds may
originate anywhere on the basal half of the manubrium, so
that in some specimens several rows of buds occur, completely
encircling that organ.
As far as the method of bud-formation has been observed
at present, it appears that the young buds first arise as out-
growths on the external wall of the manubrium. Besides
these, a certain number of older buds with rudimentary
tentacles occur, projecting into the stomach and _present-
ing the appearance of having actually grown through the
* “On Limnocodium Sowerbii, E. Ray Lankester,” Quart. Journ, Mier.
Sci. vol. xx., 1880.
274 On the Freshwater Medusa of Lake Tanganyika.
wall of the manubrium. I hope, however, to be able to
supply further details of this interesting process of bud-
formation in a subsequent paper.
As the Tanganyika Medusa is still without a generic name,
I propose that of Limnocnida for it, with the following cha-
racters :—
LIMNOCNIDA, gen. nov.
Umbrella flat, almost disk-like, about four times as broad
as high, with a lens-shaped thickening in the middle. Ten-
tacles very numerous, hollow, with bases partly imbedded in
and adnate to the umbrella margin. Marginal sense-organs
situate along line of attachment of velum, composed of a
multicellular mass of refringent cells enclosed in a roundish
capsule. Mouth round, two thirds the diameter of animal.
Manubrium very short. Stomach shallow, of same diameter
as mouth. Radial canals four. Reproduction by budding.
Sexual organs situated on the manubrium.
Limnocnida tanganjice (Bohm).
Size: diameter of umbrella j inch.
Loc. Lake Tanganyika.
As my knowledge of the significance of several points
observed in the general anatomy of this freshwater Medusa is
as yet very imperfect, I must reserve a more detailed account
of these, as well as all discussion of the probable affinities and
systematic position of Limnocnida, for another paper. If a
place has to be found for Limnocnida in Heckel’s System of
Meduse we are beset with almost the same difficulties as those
which presented themselves in the case of Limnocodium. The
same battle will have to be fought over again, only the field
is changed. Those who considered Limnocodium to be one
of the Leptomeduse will refer Limnocnida to the Antho-
medusx, while those who regarded Limnocodium as one of
the Trachomeduse will have to place Limnocnida among the
Narcomeduse.
EXPLANATION OF THE PLATES.
Piate XIII.
Fig. 1. Side view of Limnoenida tanganjice, X 4.
Fig. 2. Ventral view of ditto, x 3.
Fig. 3. Ventral view of a portion of the periphery of the disk, x 9.
On a Specimen of Sowerby’s Whale. 275
PLATE XIV.
Fig. 4. Diagram of a transverse section passing along a radial canal on
the right hand side but not on the left.
Fig. 5. A radial section cut a little to one side of the opening of a radial
] canal into the ring-canal, x 60.
Fig. 6. A portion of a tentacle, showing the arrangement of thread-cells.
Fig. 7. A tangential section through two adjacent marginal sense-organs.
Fig. 8. A marginal body seen in longitudinal optical section.
c.c. Circular canal. ms, Mesogloea.
ect. Ectoderm. m.v. Muscles of velum.
e.l, Endoderm lamella. R. Radial tentacle.
end. Endoderm. r.c. Radial canal.
em. Medusa buds. s. Marginal sense-organ.
R. Interradial tentacle. te. Tentacle.
mn. Manubrium. v. Velum.
XLVI.—Notes on a Specimen of Sowerby’s Whale (Mesoplodon
bidens*), stranded on the Norfolk Coast. By T. SOUTHWELL,
F.Z.S., and Sripney F. Harmer, M.A., F.Z.S.
[Plate XV.]
On the 19th December, 1892, we received intimation that a
“large fish ” accompanied by a young one had come ashore
on the previous day at Overstrand, near Cromer, and on the
following day we visited Overstrand together with the expec-
tation of seeing one of the commoner species of Cetacea, but
were agreeably surprised to find that the animal which had
been stranded was a fine adult female specimen, 16 feet
2 inches in length, of Sowerby’s Whale. It had been disco-
vered on the morning of the 18th December left in shallow
water by the receding tide, still alive but feeble, and, after
being anchored to the shore for security, had been placed on
a trolly and carried to the top of the cliff; it died, however,
before it was taken from the water. Previously to our arrival
on the scene it had been eviscerated, and was lying in a shied
on the top of the cliff, a foetus, 5 feet 2 inches long, having
been extracted from it.
* Professor Sir W. Turner, in a recent paper on its occurrence in the
Firth of Forth, has adopted the generic name Micropteron for this animal,
as proposed by A. Wagner in 1846, and used by Eschricht and G. Cuvier;
but for the reasons assigned by Professor Sir W. Flower, in a footnote to
his paper on the “ Recent Ziphioid Whales” (Tr. Z. S. viii. p. 208), we
prefer to retain the more familiar term Mesopludon, as applied by M. P.
Gervais in 1850.
276 Messrs. T. Southwell and S. F. Harmer on a
We are informed that both mother and young are being
stuffed and the skeleton of the former mounted for the Hon.
Walter Rothschild, for whom they were purehased, being
intended for his museum.
Eighteen examples of this species have been previously
recorded, all from the North Atlantic, but in only one previous
instance has it been met with in England, namely at Spurn
Point in 1885*, and with five exceptions all have been males ;
nor has any perfect specimen in a fresh condition ever been
fully described, unless, indeed, the young female taken at
Atlantic City in March 1889 should prove an exception; but
of this we have at present received no particulars. We were
therefore glad of an opportunity of examining so good a
specimen under comparatively favourable, although not alto-
gether satisfactory, circumstances, for the animal was unfor-
tunately lying in such a position that it was impossible to
obtain a photograph of it, and one side only could be seen.
We regretted this the more as, although several figures of
Sowerby’s Whale have already been published, most of them
are not completely satisfactory in all respects, and we should
have been glad to have secured a good photograph; but the
accompanying figure (Pl. XV.) has been carefully recon-
structed from sketches and measurements made on the spot.
Of the original published figures the best with which we
are acquainted is that of Reinhardt +, which is unfortunately
contained in a journal not easy of access to English readers ;
other figures more or less satisfactory have been given by
Sowerby ¢, Dumortier§, Andrews||, &c., and a figure of a head
has been recently published by Sir William Turner § from a
photograph which differs in several noteworthy particulars
irom the specimen examined by us, especially in the propor-
tions of the rostrum and lower jaw. ‘The coloration of the
body also differs considerably in the various descriptions; but
to these we shall have to refer later on, and will proceed to
give the results of a careful examination of the specimen
under consideration,
The two remarkable mandibular tecth so characteristic of
the adult male of this species were not visible in our specimen,
* Southwell and Eagle Clarke, Ann. & Mag. Nat. Hist., Jan. 1886,
». 53.
: + Oversigt over d. Kong. Danske Vid. Selskabs, Copenhagen, 1880,
t J. Sowerby, ‘The British Miscellany,’ 1804-1806, vol. i, pl. i.
§ B.C. Dumortier, “ Mém. sur le De/phinorhynque microptére,’ Nouv.
Mém. de Acad. Roy. de Bruxelles, xii., 1839.
| W. Andrews, “ On Ziphius Sowerbyi,” Trans. Roy. Irish Acad. xxiv.
1871, pl. xxv.
§| Proc, Roy. Phys. Soc. Edinburgh, x., 1888-89.
Specimen of Sowerby’s Whale. 277
which was a female. The jaws were apparently completely
edentulous, and although it was possible to feel through the
gums a slight prominence on either side in the position of the
teeth of the male, we could not by this means definitely satisfy
ourselves with respect to this point, nor were we able to
ascertain the presence of any other rudimentary teeth in either
jaw. ‘The evidence which exists on this subject is favourable
to the view that the female of this species is not provided with
any teeth which are large enough to pierce the gums. Both
the rostrum and the lower jaw were distinctly longer and more
slender than in Sir William Turner’s specimen, in our com-
parisons with which the difference in sex and age must be
borne in mind. The rostrum and lower jaw together were
nearly cylindrical when the mouth was closely shut and both
were evenly rounded at the tip, the lower jaw projecting about
half an inch beyond the rostrum, The contour of the mouth
also differed considerably from that of the specimen figured by
Sir W. ‘Turner—the mandible passed backwards in a nearly
straight line for a considerable distance, the lower edge of the
mouth then curving upwards and again downwards. ‘The
gape was small, the angle of the mouth being nearly 4 inches
in front of the point of junction of the upper and lower lips,
which, behind the angle of the mouth, bounded a groove
continuing the direction of the mouth. Beneath the lower
jaw were the two singular gular sulci characteristic of the
members of the subfamily Ziphiine ; these were expansible
straight grooves in the skin 11? inches long, converging at
their anterior ends, which were only half an inch apart, and
diverging to their posterior ends, which were separated by an
interval of 94 inches.
The eye was contained in a horizontal slit 14 inch long,
the iris being brown; the pupil, a long oval with its major
axis horizontal, appeared to be black. It was situated at a
distance of 74 inches from the termination of the furrow
formed by the mounth-slit and 114 inches behind the actual
angle of the mouth.
The aperture of the ear was very minute and not easy to
discover in the adult specimen; a comparison with the fcetus,
however, enabled us to convince ourselves that a minute
aperture situated about 3 inches behind the eye in the adult
specimen, and into which we were able to pass a fine bristle,
was really the external auditory meatus. This aperture was
distinctly below the level of the eye, as shown in Reinhardt’s
figure, and not in a line with it as in Sir W. Turner’s *
figure.
* Loc. cit.
278 Messrs. T. Southwell and 8S. F. Harmer on a
The blow-hole was slightly in advance of the eye and was
ecrescentic in shape, the horns of the crescent pointing
forward. The major portion of the blow-hole was situated
to the left of the medial line of the head, but we could not
detect any other asymmetry in the arrangement of this aper-
ture. In Sir W. Turner’s specimen the right limb of the
crescent was slightly in front of the left.
The head was very prominent dorsally for a short distance
in front of the blow-hole.
The pectoral limb was relatively small and was situated at
a low level on the animal, convex on both its borders and
somewhat sharply pointed. A cutaneous groove passed
forwards from its anterior end, continuing the line of its lower
margin *, the length of the lower border, measured along the
curve from the anterior end of this groove, being 1 foot
94 inches, whereas the length of the upper border (also
measured along the curve) was 1 foot 54 inches.
The dorsal fin was situated considerably behind the middle
of the animal (not quite two thirds) and was strongly faleate
behind ; its base measured 1 foot 1? inches and its height was
74 inches. The commencement of the fin was 10 feet
2 inches from the tip of the rostrum, and the posterior end of
its base was 5 feet 11 inches from the middle of the border of
the dorsal fin, both measurements following the curve of the
back.
The greatest dorso-ventral diameter (3 feet 5 inches) was
about midway between the pectoral and dorsal fins. The
large size of the middle of the body, which tapered to much
smaller dimensions at either end, was one of the most obvious
of the external features of this specimen, and in this it agreed
closely with the published accounts of other individuals; it
must not, however, be forgotten that the Overstrand specimen
was in a pregnant condition when captured.
The body became laterally compressed near the tail, and
both the dorsal and ventral edge formed a conspicuous ridge
or medial keel passing a short distance along either surface of
the tail, which was entire on its posterior border, the place
of the medial notch, as in Hyperoodon, being taken by a
slight convexity, which in this specimen had, however, been
somewhat abraded by the rope used in its ‘capture. The
flukes of the tail measured 3 feet 8 inches from tip to tip.
* This could not be shown in fig. 1, in which what is apparently the
insertion of the limb is really the termination of the cutaneous groove
mentioned above.
=: age
Specimen of Sowerhy’s Whale. 279
Coloration.
Previous observers have described this animal as being
lighter beneath than above. This was distinctly not the case
in the specimen under consideration, which was of a uniform
black colour (with the slight exceptions shortly to be men-
tioned), the skin being very smooth and polished, as has been
described in other instances; and the fishermen in charge
who had assisted in its capture informed us that at first there
was a perceptible bluish tint on the skin in a good light.
But the most remarkable feature was the presence of a number
of curiously shaped marks sparsely distributed over the body,
but most numerous on the side and ventral surface. These
spots were most irregular in size and figure, some being
annular, others mere blotches, others again having the appear-
ance of splashes or smears varying in size up to that of a
man’s hand. One peculiar mark in the dorsal region consisted
of several horizontal lines crossed by two others in a nearly
vertical direction. These curious markings, which seem
somewhat to resemble the markings on Grampus griseus,
have been frequently referred to, and seem to differ consider-
ably in different individuals; but certainly in this instance
there was very little resemblance to the “ vermicular streaks ”
described and figured by Sowerby *, and with which he stated
the sides of his specimen were ‘ completely covered.” We
have not attempted to reproduce these markings in the
accompanying figure.
The colour was not appreciably lighter on the belly than on
the back ; the anterior edges of the tail-flukes were grey, that
colour shading off gradually into the general black hue of the
rest of the tail. The lower jaw was grey in front, but became
distinctly white near the posterior end of the slender part of
the jaw, the white colour extending some distance backwards
along the edge of the mouth; the upper jaw was greyish, its
edges being white, the inside of the mouth black, and the
small tongue, which was only free at the point, flesh-coloured.
The foetus (Pl. XV. fig. 2) had been removed from its
membranes and the umbilical cord severed close to the
body ; it resembled the adult in its general characters,
but of course was much less robust in appearance; the
grooves of the throat were well developed and the fins
were identical in position and form with those of the
parent, the most noteworthy point of difference being the
* Sowerby, ‘ British Miscellany,’ pl. i.
280 Messrs. T. Southwell and S. F. Harmer on a
relative shortness of the jaws as compared with the adult. |
The Ziphioid elongation of the beak had not yet been acquired,
and no trace of teeth could be detected in either jaw.
The colour of the foetus was black above and red below,
the latter colour being due to the effusion of blood into the
skin, which was doubtless white when the foetus was removed
from the uterus. Parts of the head, including a horizontal
streak passing through the eye, were bluish grey, and the
first two inches of both the upper and lower jaws were black.
Table of Measurements.
ADULT. Farvs.
re—n ne—_—™
feet. in. feet. in.
Extreme length in straight line from tip
of rostrum to middle of tail........ 2 i) sat
Girth of head round eminence in front of
DSW Oe an ceases hs tae eon ee ee 2 8
Girth in plane of blow-hole............ 3.4
Girth in plane 43 inches behindeye .... 4 5
Transverse diameter of blow-hole ..... ‘ 1
From tip of lower jaw to angle of mouth. 1 653 5}
eet sy i posterior end of
the groove running back from the
angle of the mouth .............. 1 9} 7
From tip of upper jaw to angle of mouth. 1 4 5
From eye-slit to angle of mouth........ 11}
Antero-posterior diameter of eye-slit .... 1}
From eye-slit to vertical line passing
through external auditory meatus .. 3
From horizontal line produced back from
eye-slit to external auditory meatus 2
Greatest dorso-ventral diameter of the
art of the head in front of the blow-
CO er ete 1 3S
Length by which mandible projects be-
VON TOALMAM . ic sins ciple se Rule Sineiete 4
Height of dorsal fin .........0.+.ss000- 7% 3t
Length of base of dorsal fin............ 7 oe 4
From tip of upper jaw to front end of
base of dorsal fin (measured along :
CGTye Or DAGK)) ops's fing s Suman 10 2 3.0,
From posterior end of base of dorsal fin
to middle line of tail (measured along
PVE AMON) ore oS wiw's t Sik bp.e Ce ete Sr is 1 8}
From tip to tip of flukes (measured in a
BURA EID does o's bye oa oe ba cole > a
Length of slits fur mamme ............ 23
Length of lower border of pectoral fin
(measured along curve from front end
of groove extending forwards from
hase of fil) Swiss aeeies ohv3ee Wine 1 9} 9
Specimen of Sowerby’s Whale. 251
ADULT. Fatus.
m——_—~™ NN
feet. in. feet. in.
Length of upper border of pectoral fin
(measured along curve)............ 1 65} 8
Length of grooves beneath lower jaw .. 11%
Distance between front ends of the above
GPTOOVES: “aictewietttir. cniels ches wine e's «1c! 3
Distance between hind ends of the above
OV OO VOR es trariosnig nle's sree sofa tober tia oye 93
Greatest dorso-ventral diameter (2 feet
8 inches in front of dorsal fin)...... 3.6
Of the general history of the Ziphioid whales Sir W. H.
Flower has given an admirable summary in his paper “ Ona
the Recent Ziphioid Whales,” which will be found published
in the ‘Transactions of the Zoological Society of London,’
vol. viii. p. 203 (1871), and “ A Further Contribution to the
Knowledge of the existing Ziphioid Whales: Genus Meso-
plodon,” “which appeared in the same publication, vol. x.
p- 415, in 1877: and of the anatomy and, to some extent, of
the external appearance Sir William Turner has contributed
most valuable papers, which have appeared from time to time
in the ‘Journal of Anatomy and Physiology ;’ but unfortu-
nately the specimens which came under ‘the notice of this
excellent anatomist have never been in such a condition as to
enable him to give the exhaustive description of the exterior
appearance of the animal which we should certainly have
received had the material been at his disposal.
The same may be said to a greater or less degree of the
continental naturalists, and Sowerby’ 8 type specimen was
described by him at second hand; the two Bandon specimens
also were so mutilated as to be of little service to science. It
thus happens that the skeleton and soft parts of this species
are better known than its external appearance. This defi-
ciency we have endeavoured, so far as the Overstrand
example is concerned, to supply ; but, as so many discrep-
ancies are noticeable between’ our observations and those
which have previously appeared, we venture to append some
observations on the published descriptions and original figures
of this singular animal. How far these discrepancies may be
due to differences of age and sex we have not suflicient
evidence to venture an opinion.
Sowerrsy, James. ‘The British Miscellany,’ 1804-1806, vol. i. pl. i.
Physeter bidens 3, Elginshire.—The figure in outline closely
resembles the Overstrand specimen, the general shape of the
body being almost identical, but the head is relatively shorter
282
Messrs. T’. Southwell and S. F. Harmer on a
and the lower jaw more massive, with of course two teeth,
indicative of the male sex; the tail is shown with a mesial
notch, and the conspicuous keels, both on the upper and lower
surfaces of the termination of the body, are absent ; the dorsal
fin is much less falcate and the pectoral limbs longer, narrower,
and less pointed than in the Overstrand specimen. The colo-
ration, “black above, nearly white below,” and the sides
“completely covered with white vermicular streaks in every
direction,” differs materially from the specimen we had the
opportunity of examining.
Rernarpt, J. “ Mesoplodon bidens, en Tilvext til den danske
Havfauna,” Oversigt over d. K. D. Vid. Selskabs, 1880 (Kj6ben-
havn), p. 63, tab. 11. (adult female, side view, and dorsal view
of head and part of the body).—This appears to be a very good
figure; the external auditory meatus is shown in much the
same position as in the Overstrand specimen, but the contour
of the head differs in having a deep depression at the base of
the skull and in the body being less fusiform. The animal had
been dead for over a month when Reinhardt examined it *,
and, except in a few places, nearly all the cuticle was removed,
which may account for the auditory meatus being so con-
spicuous, probably also for the occipital depression already
mentioned. The only reference to colour is the statement that
the remains of the epidermis and the interior of the mouth were
blackish. On the shrinking of the integuments two small
functionless teeth about the size of a pin’s head were seen on
each side of the upper jaw, so loosely lodged that they were
freely movable ; a third could be felt on cutting into the skin,
and, more posteriorly, apparently a fourth, but no trace of
teeth could be found in the mandible, though the author suggests
that they also were probably present in this specimen.
Anprews, W. “On Ziphius Sowerbyi (Mesoplodon Sowerbiensis,
Van Beneden),” Trans. Roy. Irish Acad. xxiv., 1871 [read
8th April, 1867], p. 429, pl. xxv. figs. 1, 2, 3: 1, side view of
frontal portion of head; 2, upper portion of head; 3, under
portion, showing the throat-furrows ; all from photographs.
g.—Teeth very conspicuous and throat furrows figured as
uniting in front; irides said to have been blue.
Dumorrier, B. C. “ Mémoire sur le Delphinorhynque microptere
échoué & Ostende,” Nouv. Mém. de l’Acad. Roy. de Bruxelles,”
tome xii., 1839, pl. i., Mesoplodon bidens 2, stranded at
Ostend, Aug. 21, 1835.—The figure is on the whole good, but
the following criticisms may be made. The shape is much the
same as that of the Overstrand specimen, but the attachment
of the caudal fin is not accurately drawn, and both the dorsal
* We are indebted to Sir William Turner’s paper in the ‘Journal of
Anatomy and Physiology, 1882, p. 459, fora translation of the substance
of Reinhardt’s remarks.
Specimen of Sowerby’s Whale. 283
and pectoral fins appear to be too large. The mouth is widely
open, but no groove running backwards from its angle is shown.
The length is stated to have been 3 metres 45 centim. (=11 feet
4 inches), and the animal was kept alive out of the water for
two days; it made a noise like a cow. ‘The colour was
*brunitre plombée,” except the ventral surface, which was
‘‘blanchitre et cendré.” Plate ii. shows skeleton, plate iii.
hyoid and other details.
The first known example of this species was met with in
the year 1800; after twenty-five years two others were pro-
cured in France; an interval of ten years elapsed (1835)
before the next specimen was obtained on the Belgian coast
at Ostend ; nothing more was heard of the species till 1864,
in which year, also in each of the years 1866, 1867, 1869,
1870, and 1872, a single individual was procured; the next
example was obtained in 1880; 1881 produced two, 1885
three, 1888 one, 1889 one, and 1892 one—nineteen examples
in all. Of seventeen instances in which the sex was known
or believed to be known eleven were males and six females ;
we mention this preponderance of males as being the reverse
of what occurs with regard to the nearly allied Hyperoodon,
the adult male of which has never been met with on our coast,
although large numbers of both sexes congregate in summer
in the seas between Iceland and Jan Mayen and the females are
regular spring and autumn visitors in the North Sea on their
way toand from their summer habitat; but the line of migra-
tion of the males appears to be different to that followed by
the females.
Again, with regard to the seasons at which it has been met
with, Sowerby’s Whale displays none of the regularity which
marks the movements of the Hyperoodon ; an analysis of the
dates shows that it has been met with in February once, March
twice, April once, May twice, June once, ‘‘summer”’ once,
August twice, September twice, October twice, and December
once, the bulk having been stranded on the shores of the
North Sea and of the English Channel, and on the south-
west coast of Ireland; 7. e. from the Sketland Isles to the
north of France. It may be that the February and De-
cember examples were belated wanderers which had lost
their way ; but the occurrence of so many examples between
the months of March and October would seem to indicate that
the waters of the North Sea represent the extreme northern
boundary of the wanderings of this species; their southern
habitat is quite unknown, except that two have been met with
on the western side of the North Atlantic, at Nantucket Island
284 Mr. E. A. Smith on the
and Atlantic City, the latter in the monthof March. Judginz
from the hopeless way in which it becomes entangled in the
sandbanks and shoals of a shallow coast, it would seem to be
unaccustomed to such impediments, and is probably a deep-
water species. It is also worthy of note that the Overstrand
Mesoplodon was within a short time of giving birth toa young
one, whereas the young of the Hyperoodon are produced in
the months of May and June.
Notwithstanding the very great advance which has been
made in our knowledge of the Cetacea of late years, we are
still very ignorant with regard to the habits and distribution
of many species, and it is most desirable to place on record
every circumstance which may tend, even indirectly, to throw
light on a subject of so much interest.
EXPLANATION OF PLATE XV.
Fig. 1. Mesoplodon bidens, 2, seen from the left side. Seale 1:40. The
arrow @ indicates the position of the blow-hole ; 4, gular sulcus;
c, eye; d, external auditory meatus,
Fig. 2. Foetus which belonged to the above specimen, seen from the left
and slightly from the ventral side. From a photograph taken
by Mr. W. D. Harmer. Scale 1:12. band eas above; e, right
pectoral limb.
XLVII.—Note on the Genera Geothauma and Gyrostropha.
By Epaar A. SMITH. ;
THE genus Geothauma has recently been proposed by
M. Crosse * for the reception of that most wonderful little
shell described by Lieut.-Col. Godwin-Austen under the name
of Opisthostoma grandispinosum. M. Crosse considers the
trochoid form, the spinous ornamentation, and the manner in
which the last whorl is produced upward to the top of the
spire of sufficient importance to distinguish it generically from
the typical species of Opisthostoma from India. Many of the
characters given in the diagnosis of the genus are specific
rather than generic. ‘This may have arisen from the fact
that M. Crosse appears to have had but a single species in
view at the time. At all events, it is unsatisfactory that he
has not expressed his views regarding the other known species
of Opisthostoma from Borneo, namely O. de Crespignyt,
H. Adams, O. Wallace’, Ancey, and O. pulchellum and
* Journ, de Conch. 1892, p. 282.
ee
-
Genera Geothauma and Gyrostropha. 285
O. Hosei of Godwin-Austen. In addition to these I have
just described five other species from the same island.
A careful study of all these species seems to prove that the
genus Geothauma is not required, at all events at present.
The form is sometimes as “ pupiform”’ as in the Indian
species, e. g. O. baritense, Smith ; the last whorl in O. busa-
nense, Smith, ascends and presses against the penultimate,
and the sculpture varies from the finest lamell, as in O. bari-
tense and O. Wallace’, to much stronger lamellations, as in
O. Jucundum, Smith. In O. pulchellum the lamelle are pro-
duced at the middle of the whorls, occasionally forming hollow
spine-like projections. In O. Averett’, Smith, these produc-
tions are still more developed, and in O. mirabile, Smith,
they are as remarkable as in O. grandispinosum.
The height to which the last whorl may be produced
upward is also a matter of degree. For instance, in O. Hose
and Everett? it rises almost as high as the top of the spire,
whereas in O. baritense it does not ascend beyond the penul-
timate whorl. Most conchologists would at a glance confess
that these ten species from Borneo are evidently modifications
of one type; and, even supposing that they differed materially
from the Indian forms, which is not the case, a new generic
name was not required for them.
In 1865 Mr. H. Adams* described the genus Plectostoma
for the O. de Crespigny?, and this, as it does not appear to
have been preoccupied in any other branch of zoology, is still
available. However, until some distinguishing differences
are discovered in the anatomy of the animals, it seems to me
altogether impossible with reason to separate the Indian and
Bornean species on conchological grounds fF.
In my opinion the genus or subgenus Gyrostropha is
equally useless. It was suggested by M. Ancey f to include
O. pauluccie, Crosse and Nevill, and O. perakensis, Godwin-
Austen and Nevill, from Perak. He considered that they
should be separated from the typical Indian forms, because
the summit of the spire was supposed to be roundly obtuse or
subglobose and the aperture situated almost in a vertical
plane parallel to the axis. Can anything be more trivial than
this? We might as well separate generically shells having
but five instead of five and a half volutions!
* Ann. & Mag. Nat. Hist. 1865, vol. xv. p. 177.
+ Mr. W. T. Blanford has already expressed a similar opinion (Ann. &
Mag. Nat. Hist. 1867, vol. xix. pp. 305, 306).
} Bull. Soc. Mal. France, 1887, p. 275.
Ann. & Mag. N. Hist. Ser. 6. Vol. xi. 21
286 Mr. W. E. Collinge on the Variety
XLVIII.—On the Variety cinereo-niger, Wolf, of Limax
maximus, Z. By Watrer E. CoLiince, Demonstrator
of Biology in Mason College, Birmingham.
In the ‘Annals’ for March Mr. Roebuck objects to my
classing the Limaz cinereo-niger, Wolf, as a colour-variation
of the well-known ZL. maximus, L., and puts forward an
argument for the retention of this slug as a distinct species or
subspecies.
He contends that it differs from LZ. maximus in its “ den-
tition,” the anatomy of the reproductive organs, and in its
external form and coloration. When I stated * that there
were no differences of importance in the general anatomy, I
did not think it necessary to enter into details as to the slight
variations and modifications to which all the Mollusca are
subject, according to age, season of the year, habitat, &c.,
for both Simroth + and Scharff} had previously described the
anatomy, and I stated that my investigations agreed with
the accounts given by them.
It seems, however, since Mr. Roebuck has become con-
vinced that he was wrong in stating that there were important
differences in the reproductive organs, solely upon the obser-
vations of Sordelli§, that he now intends to uphold its
specific distinction upon some drawings &c. made by Mr.
Charles Ashford which exhibit some slight variations from
one another, and upon some observations on the lingual
ribbon.
The lingual ribbon, which Mr. Roebuck regards as of such
importance, is, for generic or specific distinction, of little or no
value; indeed, such is its unreliability that not a few mala-
cologists totally ignore it.
Simroth, who has had more experience with the anatomy
of the slugs than any other living investigator, years ago
showed the great variation it was subject to, and that it “ was
perhaps the most unsuitable portion of the slug’s body ” that
could have been chosen for purposes of classification. Indeed
a large number of species constituted upon differences in the
lingual ribbon he conclusively proved by a series of careful
anatomical investigations to be but varieties of L. maximus,
whose lingual ribbon is subject to endless variations. ‘The
* Ann. & Mag. Nat. Hist. 1892, vol. x. p. 425.
+ Zeitschr. f. wiss. Zool. 1885, vol. xlii.
t Trans. Royal Dublin Soc. 1891, ser. 2, vol. iv. p. 518.
§ I think Mr. Roebuck’s statement “important differe mees” is hardly
borne out by Sordelli’s original account.
>
cinereo-niger, Wolf, of Limax maximus, L. 287
only malacologists I can call to mind who use it to-day are
Binney and Pollonera, and both give it quite a secondary
place. I therefore am of opinion that, from a consideration
of its development and variation, it is absolutely useless for
specific or generic distinction.
Respecting the reproductive organs, I regret Mr. Roebuck
has not a knowledge from personal examination. Accurate
as Mr. Ashford’s drawings may be—and I see no cause to
doubt the particulars given—personal observation of the
reproductive organs of a few slugs would, I think, at least
have shown him that to separate species of slugs upon any
single system is unsafe, if not unscientific. My contention is
that the general anatomy—the reproductive, nervous, diges-
tive, and muscular systems—of cénereo-niger is the same as
that of LZ. maximus. All who have had any experience in
the anatomical examination of the Mollusca know the endless
minor variations that occur in the form, size, and colour of
the reproductive organs, according to age, season, habitat, &c.
The writings of Simroth, Lessona, Pollonera, Scharff, Jour-
dain, Binney, and others will bear out my statement. It is
therefore needless for me to dwell upon these slight variations
Mr. Roebuck has seen in Mr. Ashtford’s drawings.
I have frequently of late dwelt upon the external variations
of the slugs in form and colour, and I was careful to point
out that I agreed with Jourdain that species could not be
founded or separated from each other upon these minor
differences. Mr. Roebuck says he has examined “ by far the
greater proportion of British specimens known”; and yet it
seems to me that he is still ignorant of very many of the
variations to which L. mazimus is subject. Seeing that the
actual published records of cinereo-niger are but few, I can
only regard his statements re variation as founded upon an
insufficient knowledge of the subject. Personally I have
only seen about one hundred examples of cinereo-niger and
some seven or eight hundred of L. maximus, most of which I
have subjected to a careful examination, and many anatomi-
cally. Amongst these I have met with both LD. maximus and
its variety cénereo-niger with the marblings on the mantle
and with a white foot-sole also.
In conclusion, I fail to see any points which are of suffi-
cient importance to establish the specific identity of ctnereo-
niger, and until such are shown to exist I shall, with the
above-mentioned malacologists, continue to regard it as a
variety of L. maximus.
21%
288 Mr. G. Lewis on
XLIX.— On some Japanese Scaphidiide.
By G. Lewis, F.L.S.
THE following notices record the larger species only of the
Scaphidiide 1 found in Japan. To complete the list nnme-
rous species of Scaphisoma have yet to be examined, and at
present only three species have been described by Reitter;
the names of these are given in the list below. There are
doubtless more species to be found in Japan by resident
collectors; but the species are easily found, for they can
scarcely be said to habitually secrete themselves, and I think
it unlikely the number will be greatly increased. Numeri-
cally in all countries the Scaphidiide are always far below
the other families which infest fungi, such as the Staphylinide,
Erotylidz, and Endomychide. Some of the maculate species
of Scaphidium resemble in colour and general markings certain
Carabide (Lioptera) and Erotylide (piscapha), which also
congregate in arboreal fungi; these species in Japan are
limited to five. I have placed all the species possessing a
hairy sternum in the male in the genus Scaphidium, but I
have excluded all the others, as I find their structure, espe-
cially that of their sternal plates, necessitates separation.
List of described Species.
Ascaphium sulcipenne. Scaphidium femorale.
tibiale. Reitteri, Lewis.
apicale. rufopygum.
Scaphium optabile. incisum.
Episcaphium semirufum. Cyparium sibiricum, Solsky.
ruticolle, var. ? Toxidium japonicum, Feitter.
Scaphidium emarginatum. Scaphisoma hemorrhoidale, Rett.
longipes. rubrum, Reitier.
japonicum, Rertter. —— castaneipenne, Reitter.
ASCAPHIUM, gen. nov.
Body narrowly oval or elliptical, rather convex; the head,
neck, and anterior part of the thorax somewhat narrow; an-
tennz and palpi slender, each joint of the first cylindrical and
setose, the last five joints of nearly equal length. The thorax
has the usual transverse line of punctures; the scutellum is
semicircular posteriorly ; the elytra punctate-striate. The
mesosternum is anteriorly rounded off in a vertical direction
on each side, leaving a prominent median keel on the same
plane as the metasternum; the metasternum is margined
some Japanese Scaphidiide. 289
anteriorly with an arched stria, and the edge of this plate is
feebly bowed in outline ; from the base to beyond the middle
is a median cuneiform depression (e. g. A. sulerpenne), or the
limit of the depression is indicated by two striw (e. gi
tibiale) ; the legs are slender, and in two species the tarsi
alone give a sexual character.
Ascaphium sulcipenne, sp. n.
Angustato-ovatum, nigrum, nitidum ; ore, antennis pedibusque rufis ;
elytris striis profundis et punctato-striatis.
L, 63 mill.
Narrowly ovate, black, shining ; epistoma, mouth-organs,
and antenne reddish ; the head irregularly punctulate between
the eyes, neck smooth; the thorax finely and sparsely punc-
tulate on the disk, nearly smooth behind the transverse
punctures ; the elytra—strie seven, very deep, with a single
row of punctures along each, the sutural stria continues in
the form of punctures along the base, two strie next to the
sutural are a little abbreviated apically, the sutural margin is
a little elevated ; the legs are without any conspicuous differ-
ences in the male, but the tarsi are slightly wider and serve
to distinguish the sexes.
Hab. Nikko and Miyanoshita. Scarce, six examples only.
Ascaphium tibiale, sp. n.
Angustato-ovatum, nigrum, nitidum ; elytris punctato-striatis ; tarsis
rufis, ¢ tibiis anticis longioribus.
L, 5-52 mill.
Elliptical, black, shining, mouth-organs and tarsi reddish ;
the head distinctly but not densely punctured between the
eyes, neck nearly smooth; the thorax punctured like the
head, but less densely ; the elytra punctate-striate, but much
less deeply than in A. sulctpenne, the outer stria punctiform
and evanescent; the sutural continues along the base, the
two striz next the sutural are of equal length and shortened
before the base; the pygidium rufous at apex; the legs in
the male are elongate and swollen rather nearer the tarsi
than the middle. ‘The five apical joints of the antenne are
black, the others pitchy red.
Distinguished from A. suleipenne by the smaller size,
black legs, elytral striae much less deep, and by the mascu-
line character in the tibie.
Hab. Miyanoshita, Subashiri, Nikko, and Oyayama, Not
uncommon.
290 Mr. G. Lewis on
Ascaphium apicale, sp. n.
Angustato-ovatum, nigrum, nitidum; antennis articulis apicalibus
rufis; tibiis nigris vel rufo-piceis.
I. 33 mill,
Elliptical, black, sometimes faintly metallic ; mouth-organs,
six basal and the apical joints of the antenne red; the head
clearly, not densely punctulate; the thorax similarly punc-
tured ; the elytral seulpture resembles that of A. tibiale; the
legs are black or pitchy red.
This species is smaller than A. tibéale; the apical joint of
the antenna is pale or red and the anterior tibie are alike in
both sexes.
Hab. Found in the same localities as the last species.
Rare.
Scaphium optabile, sp. n.
Oblongo-ovatum, nigrum, nitidum; supra punctulatum; antennis
pedibusque concoloribus ; metasterno inciso.
L. 4 mill.
Oblong-oval, black, shining; head robust, somewhat closely
punctulate; the thorax more distinctly punctured and the pune-
tures less close, the stria behind the neck very fine, leaving a
narrow smooth margin ; the elytra with the sutural stria punc-
tate, entire, and continuing along the base, surface punctured
more finely than the thorax, but without linear arrangement ;
the propygidium has an elevated margin at the sides, and
the base of the pygidium is laterally somewhat similar ; both
segments are finely punctulate; the claws are pale. The meso-
sternum is formed like that of a species of Scaphidium, trans-
verse, margined anteriorly, and acutely produced in front; and
the metasternum is truncate anteriorly, with a median longi-
tudinal canaliculation, and behind, the edge is conspicuously
incised in the middle and rounded off on either side of the
incision.
This curious species, being unique in my collection, is
assigned to Scaphium because of the incision in the meta-
sternum, but provisionally only, as in Scaphium the meso-
sternum is robust and triangular and in this insect it is formed
more like that of Scaphidium.
Hab. Ichiuchi, in Higo, May 1st, 1881.
EPISCAPHIUM, gen. nov.
Body oval, convex; the head, neck, and anterior part of
some Japanese Scaphidiide. 291
thorax narrowed ; the antennz not very slender, the last five
joints are thickened but not compressed, ninth and tenth equal
in size, but not transverse like those of Scaphidium; eyes
large and a little prominent; scutellum semicircular poste-
riorly ; the thorax with transverse line of punctures; the
mesosternum is much less wide than in Scaphidium and the
anterior process is much more produced, has parallel sides,
and is carried forward in the form of a keel; the meta-
sternum is margined with an arched stria. The elytral strie
are all punctiform except the sutural.
Episcaphium semirufum, sp. n.
Oblongo-ovatum, nigrum, nitidum ; elytris, pygidio tarsisque rufis.
L. 43-5 mill.
Oblong-oval, convex, shining, black, with elytra, abdo-
minal segments, tarsi, and basal joints of antenne red; the
head clearly, evenly, and somewhat closely punctured; the
thorax with the punctures faint and scattered ; the elytra with
the sutural stria punctate, complete, and continuing along the
base, surface with punctures arranged in lines more or less
broken, the rows nearest the humeral angles being the best
defined ; the legs black, without sexual differences.
Hab. Nikko, Kiga, and Miyanoshita. Not very common.
Note-—I took a series (seventeen specimens in all) in
Kiushiu, at Yuyama, Ichiuchi, Konose, and other places with
the thorax, except a narrow rim behind the neck, red, to
which I wish to apply the name of A. ruficolle. It may bea
variety, but, if so, it is a constant one in South Japan.
The five species following have maculate elytra :—
Scaphidium emarginatum, sp. n.
Ovatum, convexum, nigrum, nitidum ; elytris duabus maculis den-
tato-sinuatis, rubris; ¢ tibiis anticis intus minute emarginatis.
L. 63-7 mill.
Oval, convex, black, shining, each elytron with two red
denticulate fascie ; the head feebly and sparsely punctured
between the eyes; the thorax more distinctly punctate before
the basal punctiform line, smoother behind it; the elytra with
a somewhat stronger punctuation, especially in the dorsal
region, sutural stria rather fine and punctulate, deeper with
large punctures along the base; anterior fascia touches the
292 Mr. G. Lewis on
base below the humeral angle, leaving on the humeral angle
a black nearly circular space; the red colour does not touch
the base on the sutural side; posteriorly the fascia has three
distinct tooth-like projections ; the posterior fascia is smaller,
with only two denticulations, which point towards the head ;
the antenne with the last five joints broad and densely black,
ninth and tenth nearly equal in size, basal joints pitchy red.
Male anterior tibiw swollen apically on the inner edge, and
in the middle of the bulge there is a small but very distinct
emargination.
This species differs from S. japonicum, Reit., in having
broader apical joints to the antenne, in the outline of the
denticulate elytral fascia, and in the tibix of the male being
emarginate,
Hab. Occurs on the mountains in Kiushiu, and has been
found at Chiuzenji and on Ontaki-san.
Scaphidium longipes, sp. u.
S. japonico simillimum, sed pedibus multo longioribus ; femoribus
anticis minute denticulatis; elytris duabus maculis sinuatis
rubris.
L. 7 mill.
Oval, black, shining, each elytron with two red fascie ; the
fascize are much less wide than in S. emarginatum and agree
more closely with those of S. yaponicum ; the anterior band
has two ill-defined denticulations on the hinder edge, the
posterior band usually has two denticulations similar to that
of S. emarginatum, but not so clearly marked; the legs are
much longer and the anterior thighs have a small tooth on
the underside nearer the tibie than the middle of the limb.
‘Lhe mesosternum also is wider. I see no other characters to
separate it from S. yaponicum, except that it is a little broader
in its general build.
Hab. Higo, and on the main island at Kiga and Miyano-
shita.
Scaphidium japonicum, Reitter.
Hab. Main Island and Kiushiu.
‘The commonest species in Japan.
Scaphidium femorale, sp. u.
Ovatum, convexum, nigrum; elytris duabus maculis flavo-rubris ;
d femoribus robuste dentatis.
L. 44-53 mill.
some Jupanese Scaphidiide. 293
Oval, convex, black, shining, each elytron with two (often
somewhat wide) yellowish-red fasciw ; the head, punctuation
between the eyes sparse and irregular; the thorax more
evenly but not thickly punctured, transverse punctures largest
before the scutellum; the elytra with similar punctuation,
basal fasciz rather broad, somewhat irregular in outline, but
usually divided into two lobes, in some examples (apparently
varieties) the spots are small and confined to the outer edge
of the wing-case ; the antenne are black, basal joints a little
pitchy.
&. The anterior tibie are swollen on the inner edge and
the hindermost thighs have a conspicuous blunt tooth on the
inner side not far from the middle of the limb, but nearest to
the tibia.
flab, Main island and Kiushiu.
Scaphidium Reittert, Lew.
Scaphidium Reitteri, Lew. Ann. & Mag. Nat. Hist. 1879, iv. p. 460.
This is the only species recorded from Japan that has part
of the thorax red; in this respect it resembles an American
form. Beyond the hairy sternum it has no conspicuous
masculine character.
Hab. Vries Island, Kashiwagi, Chiuzenji, and several
places in Higo. I also met with it on Maiyasan, near Kobé,
in almost the exact spot where I found it ten years previously.
The following species have the elytra wholly black :—
Scaphidium rufopygum, sp. 0.
Oblongo-ovatum, convexum, pygidio rufo ; pedibus nigris vel rufis.
L. 44-43 mill.
Oblong-ovate, convex, black, shining, pygidium bright
red ; the head feebly and sparsely punctured; the thorax
more clearly punctured, especially on the disk before the
transverse punctures ; the elytra, sutural stria well marked
and continued along the base nearly to the humeral angle,
dorsal area irregularly punctured, some of the punctures
assume a linear arrangement. Beyond the hairy metasternum
and a slightly increased breadth of the anterior tarsi there are
no visible masculine differences; the legs are usually black,
but frequently reddish.
Hab. Yuyama, Kiga, and Nikko.
294 Mr. R. H. F. Rippon on a new
: Saat bor
Scaphidium incisum, sp. n.
Angustato-ovatum, nigrum, nitidum, supra punctulatum; ¢ tibiis
antice inconspicue emarginatis.
L. 43-43 mill.
Somewhat elliptical, black, shining ; the head and thorax
clearly, not thickly punctured, transverse punctures of the
latter well-marked and similar throughout; the elytra with the
sutural stria punctate, continuing as in the last species along
the greater part of the base, dorsal surface clearly and evenly
punctured, with little or no indication of linear arrangement ;
legs and antenne black, male with enlarged anterior tibia,
the enlargement being feebly emarginate on the inner edge ;
tarsi brownish; antenne pitchy red or black at the base.
Hab. Miyanoshita, Nikko, and Mayebashi. Apparently
scarce.
Cyparium sibiricum, Solsky.
Hab. S. Yezo, abundantly, and on the main island as far
south as Chiuzenji, more rarely. One example in Higo,
L.—Description of a new Transitional Form of Ornithoptera
belonging to the Subgenus Priamoptera. By Roperr
H. F. Rippon.
In the August number of the ‘Annals’* I described a
beautiful and most interesting new species of the genus
Ornithoptera closely allied to O. aruana, Feld., which
evidently linked the genus most closely with its section
Priamoptera. I have now the privilege, by the courtesy of
Mr. Elwes, of describing a still more important and inter-
esting novelty discovered by Mr. Doherty in the Island of
Talaut, which may ultimately throw much light on the
phylogenetic history of the genus, inasmuch as it may be
regarded as one of the transitional forms, some of which have
probably long since perished, which would enable us to link
together in a natural series all the many nominal species
which we find it necessary to admit into the subgenus Pom-
peoptera, and even to suggest, on the side of the female at
* ‘Annals,’ Aug. 1892, p. 193; also in part v. of my ‘Icones
Ornithopterorum,’ where a plate of coloured figures is given.
Transitional Form of Ornithoptera. 295
least, a closer relationship with the true Ornithoptera than at
one time might have seemed probable.
We owe a debt of gratitude to Mr. Doherty for the faithful
and admirable work he has for a long time been doing among
the islands and islets of the Sunda Straits and adjacent
localities. In this respect he has been rapidly entitling
himself to be regarded as a second Wallace. ‘The novelty
which is now before us appears to me to be one of the most
useful contributions to the Rhopalocerous fauna which he has
been enabled to send to us.
I have therefore much pleasure in dedicating this new
species to him, partly at the suggestion of Mr. Elwes, and
partly because I think it is right to give what honour we can
to those who faithfully spend their health and energies in
difficult regions of the earth, for the purpose of discovering
those lovely things which furnish us with fresh materials for
one of the most delightful studies, and with an increased
knowledge of God’s works.
Ornithoptera (Priamoptera) Doherty?, sp. n.
g. Primaries an intense silky black, almost like a bur-
nished black, with a faint green gloss in certain lights,
which gives it an exceedingly rich appearance. The grey
discal rays which enclose the nervules are so obscurely
represented as to be scarcely visible except when the light is
transmitted from the underside. The veins are all well
expressed, being somewhat lighter on the dark and much
darker on the light parts of the wing; the costa is also of the
same tint, especially near the apex when viewed opposite the
light; the costal outline is nearly straight to two thirds of
its length, or at the termination of the costal nervure, when
it curves considerably to the apex, which is so much rounded
as to pass insensibly into the posterior margin ; the outline of
the posterior margin curves somewhat irregularly, with an ont-
ward trend much greater in proportion at the termination of
the second and third median nervules than at any other part,
not excepting the apex; it is very delicately and equally
sinuate along its entire outline, with very fine reddish-
ochreous fringe-lunules between the veins as usual.
Secondaries entirely glossy black, slightly greenish or
bluish according to the position in which they are viewed ;
the veins sufficiently defined in a lighter tint; the abdominal
marginal sexual pouch or fold is a faintly redder black; the
posterior margin regularly and sufficiently scalloped, and
296 Mr. R. H. F. Rippon on @a new
without the white fringe-lunules. Hence the entire upper
surface of the wings is absolutely immaculate.
Under surface of primaries velvety or silky black, less
dark and more glossy towards the posterior margin; within
the discoidal cell, at the termination of the third pseudoneurus,
are a few scarcely visible grey atoms; the discal grey rays
enclosing the nervules are strongly defined in a slightly
greenish grey, rendered darker by a graduation with black
atoms towards the cell; the rays between the third and fourth
subcostal nervules are only faint streaks of grey, and only
one occurs on the upperside of the third median nervule; the
posterior margin almost entirely outlined by the white fringe-
lunules, which at the apex slightly encroach upon the costa ;
veins strongly defined. Secondaries a deep silky black; the
abdominal fold or pouch browner, with a fringe of delicate
hairs as long as the width of the pouch; the discoidal cell
immaculate ; between the first and second subcostal nervules
is a subquadrate patch of yellow atoms; between the second
subcostal and third median nervules a broad discal band of
lemon-yellow, forming four subrectangular patches of unequal
length, strongly divided by the nervules, the first broadly
indented from the direction of the cell, sinuate without, the
second indented outwardly and enclosing an elongate black
spot, the third indented without and enclosing a. slightly
interrupted elongate black spot, which nearly divides the
yellow area, the fourth Junate without and sharply indented
above ; viewed obliquely with a side light this yellow band
becomes a rich emerald-green, not unlike the green of the
typical Ornithoptera; the hind margin is without the ochreous
fringe-lunules.
Head: antenne black; eyes chestnut-brown ; haustellum
silky black. Prothorax with a crimson collar ; thorax above
velvety black, beneath deep black, with lateral crimson spots ;
legs black. Abdomen pale brown, bordered with piceous
brown ; the first segment entirely piceous brown, the subanal
segment also of the same colour, margined with creamy
white; the anal valves creamy white, with the usual dorsal
black termination; laterally the abdomen is of a creamy
ochreous white; subdorsal slightly yellower at its junction
with the metathorax, more ochreous near the anal valve, the
first and second segments well divided by black; the lateral
black dots fairly large.
The space of wing enclosed by the two branches of the
precostal nervure is more equal in width than is usual till it
reaches its junction with the subcostal nervure, when, of
course, it becomes narrower. This is really a very striking
Transitional Form of Ornithoptera. 297
character, the nearest resemblance to which is found in the
male of P. hephestus so tar as [ can discover; it is also
nearly as rectangular in the female.
Expanse of costa 69 millim.; width of upper wing 57 ;
length of hind wing 44, width 31; length of abdomen or
antenne 31; of legs, first pair, femur 12, tibia 9, tarsi 12;
second pair, femur 12, tibia 12, tarsi 15; third pair, femur 11,
tibia 13, tarsi 15 millim.
@. Primaries: costa equally arched but more rounded at
the apex, the outline insensibly passing into the posterior
margin, which is not so irregularly curved as in the male;
inner margin nearly a straight line. Wings light fuscous
brown; within the discoidal cell are faint atomic indications
of grey markings somewhat like those of the female of P. hali-
phron; the grey discal rays very indistinct except those
enclosing the first and second median nervules (which are
fairly though faintly defined), broadly separated by the brown
margins of the nervules; the pseuwdoneura are only just
visible; posterior margin with whitish fringe-lunules.
Secondaries slightly darker fuscous brown; within the cell
a faint ochreous rufous-yellow spot close to the discocellular
nervules; four very short narrow rufous-yellow discal marks,
the first 6 millim. long, close to the second discocellular
nervule, the second the same length from the cell, the space
between the second discocellular and first median nervules
broadly incurved and (on the right wing only) enclosing a
small brown dot; the third 7 millim. long from the cell, in-
dented two thirds its length, enclosing a brown spot (on the
right wing) ; the fourth 5 millim. long from the cell, indented
(on the right wing only about half the size) ; extremely
faint indications of a submarginal band of small spots; the
submedian area of wing is clouded pale reddish ochreous,
meeting an anal, angular, ochreous, sublunate spot, from which
two rays run partly up the abdominal margin; margin suffi-
ciently scalloped, with ochreous fringe-lunules.
Under surface: primaries the same colour as above; the
grey rays and discocellular markings similar but much
broader, a brighter ochreous red at the outer margin, greenish
grey towards the base and confluent between the second dis-
coidal and second median nervules ; the marks within the cell
are formed of microscopic atoms, and are divided into three
submarks—the first broad, one third of the cell in length, the
second very narrow and divided by the second pseudoneurus,
the third or lower one nearly as broad and long as the first
and divided by the third pseudoneurus; the marginal fringe
with a nearly continuous row of ochreous lunules. Second-
298 Ona new Transitional Form of Ornithoptera.
aries same colour as above, but slightly darker; a large
discocellular pale spot clouded with atoms close to the disco-
cellular nervules; the discal marks corresponding with the
four on the upper surface are much paler and extend much
further down between the nervules—the first is partly pure
and partly formed by atoms, the second is indented with
brown, modified by atoms, the third much larger and with a
long brown indentation, and the fourth very long, but so
interrupted with brown modified by atoms as to form an
ochreous ray close to the nervules extending nearly to the
marginal border; outside the third median nervule is also
an ochreous atomic ray close to the nervule, almost hastate,
followed by a second longitudinal stripe extending lower
down nearly to the outer margin ; the submedian nervule and
the abdominal margin have similar ochreous stripes as above,
but more prominently developed; there is a broad submar-
ginal band indicated by narrow ochreous stripes and cunei-
form spots, so meeting and enclosing the brown areas between
the nervules, especially from the second subcostal and third
median, as to give them almost the appearance of arches, the
bases of which are the fringe-lunules of the posterior margin.
Head: antenne deep black; eyes chestnut-brown; hau-
stellum black. Thorax and prothorax dark brown, with
crimson collar ; lateral crimson spots, and subdorsal with less
black. Abdomen pale fuscous brown, with lateral borders of
reddish ochreous ; subdorsal the same, with strongly outlined
or entirely fuscous brown segments ; anal tuft dark brown.
Expanse of costa 82 millim.; width of upper wing 61;
length of lower wing 51, width 45; length of antenne or
abdomen 31, of thorax with head 21. ‘The legs, all but one,
are absent from the specimen.
Hab. Isle of Talaut (Doherty). Feb. to March 1892.
Some examples of the female are entirely free from discal
marks above, as in the male, and in one example more than
half the under surface of the primaries is entirely fuscous
white, the veins only being dark.
Setting aside the remarkable precostal character of the
male, which allies it to P. hephestus, the nearest affinity of
the male is with P. Platent on the upperside, appearing like
an immaculate var. of that Philippine species. On the under
surface of the lower wing the yellow marks suggest an approach
to those of haliphron; the temale also on the upper wing,
and somewhat on the under surface, comes nearer to the hali-
phron tacies*, ‘The outline of the male primary wing is unlike
* Since writing the above I have been able to compare the females
with a female of Priamoptera iris, Robers, and am struck with the
resemblance of one or two examples of P. Dekerty: with the latter.
Hon. W. Rothschild on Apteryx Haastii. 299
that of any other; nevertheless we have in this species a close
ally of Platent and possibly an old form from which perhaps
several of the varieties of the haliphron and helena groups
have been derived by differentiation—a transitional species I
have long expected would be discovered. After awhile other
transitional forms will, I hope, be found, and we may then be
able to understand some of the geographical development
history of the whole genus.
I am much indebted to Mr. Elwes for the cession of a large
and interesting series of this novelty, the types of which, with
several varieties, will occupy two plates in the sixth part of
my ‘ Icones Ornithopterorum,’ with extended information.
24 Jasper Road,
Upper Norwood, 8.E.
LI.—Note on Apteryx Haastii.
To the Editors of the ‘Annals and Magazine of
Natural History,’
GENTLEMEN,—In the February number of your Magazine
Mr. H. O. Forbes answers my notes on Apteryx Haastii. I
deeply regret that he considers I have been guilty of an indis-
cretion in writing about opinions which, although openly
expressed, he never published in print, and I hope he will
accept my full apology.
I must, however, in justice to myself answer his objections
in full. In the first place he contends that, contrary to my
statement, Apteryx Haastti has only been recorded from the
South Island, and that in single specimens in localities only
where A. australis and A. Oweni are found together. In
reply to this I have to state that the first four specimens of
A. Haastiit I ever received were young birds of the size of
A, australis, and were so distinct that I had at first thought
they were a new species; but I now, since my last article,
find they are not. These four birds were collected by a
German botanist on Stewart’s Island and were sent me by
Sir Walter Buller. Then Sir Walter Buller sent me alive a
young Apierya Haastii without a locality ; and, lastly, he
sent me fourteen or fifteen specimens, all collected on the
west coast of the North Island, among which were two live
birds, male and female, and a chick and rotten egg, all dug
out of one hole.
300 Hon. W. Rothschild on Apteryx Haastii.
In addition to these I have notice of some A. //aastit sent
to me lately by a man named Danneford, which also came
from the North Island. Moreover, Professor Reichenow
exhibited before the German Ornithological Society a speci-
men of A. Haast?i from the North Island.
Mr. Forbes has stated further, in support of his theory,
that the cecum of A. Haastiz, being different in the two
sexes, was rather a sign of hybridity than otherwise. Unfor-
tunately he has read my notes quite wrongly. I said:—
*‘ Another distinctive character will be found in the cecum,
which seems to differ not only in the two sexes, but also in
all the various species of the genus.” By this I meant to
say that a further difference lay in the distinction of the
cecum in each species, not in the sexes, for, as I have
proved, the male and female ceca are distinct in each
species. Professor Owen was the first to describe this strange
difference in his article on the anatomy of Apteryx australis,
in the ‘ Transactions of the Zoological Society.’ As regards
my argument upon the size of the beak of A. Haastii, I
never mentioned the subject of length; I can only say that
as to length I have several A. Mantellii with longer beaks ;
but what I said was that, “though not so long as that of
A. maximus, it is very much stouter even than that of the
recently discovered extinct species.” This is a fact, as my
largest female A. Haastii has a beak nearly if not quite
twice as large in circumference at the base as any known
Apteryx.
Finally, I must reiterate the fact that not only have I found
the ceca of Apteryges differ in the two sexes, but also most
of the Struthionide present this difference; and I have to
point out that the cecum of the Emu (Dromeus) is about
3 inches long, while that of the much smaller Rhea americana
is nearly 7 feet long, and Apteryx Mantellii has a cecum
9 inches long, while in A. maximus it is barely 4 inches
long *.
WALTER ROTHSCHILD.
* If, however, it should be proved, which I for one very much doubt,
that the two specimens of A. Haastii described by Potts are hybrids,
both Scuth-Island, North-Island, and Stewart’s-Island specimens of large
grey Apteryx will have to be described as new species; for I know they
come from districts where no A. Owenii have ever been found, and are
totally unlike anything which could be produced by crossing.
Development of the Tooth-Rudiments in Rodents. 301
LII.— Contributions to the Development of the Tooth-
Rudiments in Rodents. By PAaut Freunp*.
In the autumn of 1890, at the suggestion of Prof. Born, I
commenced to investigate the development of the dentition of
various Rodents, with a view to discovering whether in the
diastema between the fully developed teeth it was still possible
to find remains of rudiments of such as had disappeared. It
was only after making certain discoveries bearing upon the
question that I learnt that a paper by MM. Pouchet and
Chabry ¢ contains a series of statements with reference to
this subject. Nevertheless, when I have furnished a more
precise critical comparison of the results of these authors with
my own, it will be seen that what are really very material
points have escaped their notice.
The dentition of the Rodents attracts attention in the first
place owing to the great reduction in the number of the
teeth. The largest number is found in the Leporide, which
possess twenty-eight teeth, namely @. 7, c. 2, m. 3, the smallest
in the Muride, in which in the majority of cases the number
is sixteen, namely 7.1, ¢- 2, m.2; while in Hydromys we
actually find only twelve teeth (7. {,c.°, m. 3). The incisors
are always separated by a wide diastema from the molars or
premolars, as the case may be. As further peculiarities we
must regard the absence of the milk-dentition in many species
and also the circumstance that the incisors invariably, and
the molars usually, belong to the category of continually
growing rootless teeth.
It is assumed by all authors who have written concerning
the origin of the Rodents that the latter have been derived
from forms with a fuller and, above all, more completely
equipped dentition, and that consequently the reduction in
the number and the disappearance of certain kinds of teeth,
such as the canines, are phenomena of a secondary character,
connected with the special adaptation of the dentition to a
peculiar sort of food, which could hardly be turned to account
without this modification of the teeth. In other words, the
* Translated from the ‘ Archiv fiir mikroskopische Anatomie,’ Bd. 39,
p- 525-554 (Bonn, 1892): from a separate impression communicated by
Br. Forsyth-Major.
+ G. Pouchet and L. Chabry, “Contributions a l’Odontologie des
Mammiferes,” Journal de ]’Anatomie et de la Physiologie: Paris, 1884.
Ann. & Mag. N. Hist. Ser. 6. Vol. xi. 22
302 Herr Paul Freund on the Development
reduction in the number of the incisors (in the mandible there
is always only one on each side, in the upper jaw there are
two in the Leporide only, in all other cases there is likewise
one) and the disappearance of the canines, and, at any rate,
of a portion also of the premolars, are related to the modifica-
tion of the persisting incisors to form chisel-like gnawing-
teeth, by means of which the animals are able to consume
even very solid vegetable food-substances, such as the bark of
trees and grains, the latter of which possess an especially
high nutritive value. It has already been demonstrated by
numerous authors that the peculiarity of the continuous
growth of the gnawing-teeth also belongs to the complete
efficiency of the entire arrangement. ‘The effectiveness of
this form of dentition in the struggle for existence is at once
evident from the great wealth of species possessed by the
order Rodentia, the universal geographical distribution of its
representatives, as well as the enormous number of indi-
viduals composing the various species. The latter pheno-
menon naturally results from the proverbial fertility of the
Rodents; but this, as we know, is directly proportional to
the abundance of food and the ease with which it is obtained.
It is characteristic that the advantages in the struggle for
existence, which are ultimately occasioned by the form of the
dentition, are so great that Nature has almost entirely dis-
pensed with the additional gift of other means of defence.
The Rodents are among the most defenceless of Mammals.
Our views as to the reduction of the dentition in particular
depend upon the conceptions which we have formed as to the
phylogenetic derivation of the Rodents. A whole series of
such conceptions, however, have already been formulated by
various authors. ‘The fullest and most recent treatment of
this subject, which is based upon an exhaustive comparison
of the dentition, of various parts of the skeleton, the uro-
genital system, and the mammary organs, is furnished by
A. Fleischmann *.
Fleischmann is inclined to derive the Rodents from mar-
supial-like ancestors, with a complete insectivorous dentition.
In his seventh chapter, which he terms a ‘ Phylogenetic
Sketch,” he sums up his results as follows :—‘‘ In the organ-
ization of the Rodents two different stages are accordingly
recognizable, which are full of meaning for the history of
Mammals. On the one side many peculiarities are empha-
sized which are of functional importance in a lower stage,
* «Embryologische Untersuchungen von A. Fleischmann, Privat-
docent in Erlangen,’ ii. Heft.: A. ‘‘ Die Stammesgeschichte der Nage-
thiere.”
of the Tooth-Iudiments in Rodents. 303
represented by the Marsupialia; on the other many conditions
of form are approximated to the type of the higher Placen-
talia. The Rodents stand as it were as ancient monuments
of a time which has long since disappeared, and unfold before
us the representation of the modification which, perhaps as
early as the Cretaceous period, elevated the Protheria into
the condition of placental Mammals. Nevertheless the facts
with which we are at present acquainted do not suffice to
prove a direct relationship to the Marsupials. It is only
possible to maintain that animals very similar in their organi-
zation to the structure of the Marsupialia were the ancestors of
the Rodents. The forms of phylogenetic importance are not
concentrated in one group, but are distributed over all four
divisions of the Rodentia; for, in accordance with the special
conditions of existence, with which we are still very imper-
fectly acquainted, certain organs have retained their primitive
characters in one division, and others in another.” In direct
reference to our results in the case of Lepus I would also like
to quote here the concluding sentence of the same chapter :—
“As regards the Lagomorpha, it seems to me that the theory
founded by Schlosser is established, namely that they did not
branch off from the primitive stock simultaneously with the
true Rodents, but that they have existed as placental Rodents
only for a relatively short period.”
In other respects Schlosser’s admirable paleontological
and general investigations* upon the organization and
historical development of the Rodents have already been
exhaustively discussed in Fleischmann’s treatise.
Especial stress must also be laid upon the fact that,
although certain of the herbivorous Marsupials possess a
dentition which is in the highest degree similar to that of the
Rodents (Phascolomys even has continuously growing incisors,
and withal the dental formula 7. ;, c. ?, p. }, m. 4), Fleisch-
mann nevertheless expresses himself very decidedly against
a direct derivation of the Rodents from these animals. He
writes :— The herbivorous Marsupials do not stand in a
closer phylogenetic relation to the Rodents, but are a pecu-
harly differentiated branch of the Metatheria. My object in
minutely describing the dentition of various Marsupialia was
merely to demonstrate that the reduction of an originally
complete dentition in the herbivorous Diprotodontia occasions
arrangements which are very similar to the dentition of the
* M. Schlosser, “ Die Nager des europaischen Tertiirs nebst Betracht-
ungen iiber die Organisation und die geschichtliche Entwicklung der
Nager iiberhaupt”: Cassel, 1885. ‘ Paleeontographica,’ 31 Bd. Dritte
Folge, 7 Bd.
22*
304 Herr Paul Freund on the Development
Rodents. Consequently, if in a branch of the Mammalian
stem it is possible to recognize in actual examples the deve-
lopment of an upper and lower pair of incisors, accompanied
by the loss of the rest of the incisors and the canines, it is also
a probable assumption that in the ancestors of the Rodents a
similar process occasioned an analogous result. In the group
which was undergoing modification the adaptation of the
cutting-teeth to their heightened function would have
advanced further, since the rooted incisors attained a pris-
matic shape and the faculty of permanent regeneration.”
Another view is supported by Cope *, who derives the
Rodents from the Bunotheria, as represented by the suborder
Tillodontia, which, while possessing a more complete den-
tition, exhibit a decided rodent-like formation of individual
incisors.
With reference to the results of our embryological investi-
gations, it seems worth remarking that, according to this
derivation, the large gnawing-teeth of the existing Rodentia
would correspond to the second incisors of what Cope supposes
to have been their ancestors, since in the latter the first
incisor is already rudimentary and small, while the second
is developed into the gnawing-tooth.
An altogether separate position is taken up by Baume f in
his speculations.
This position is in the first place based upon the fact that
this author is desirous of regarding continuously growing
teeth as the prototypes of Mammalian teeth in general. In
forming this conception he relies upon the view that the
continuously growing rootless tooth is of relatively simpler
construction. At the same time he considers that the organ-
ization of such teeth is adapted for the production of a mass
of tooth-substance. This property is likewise considered by
Baume to be of a primitive character, since, as he urges, in
the whole animal series we may recognize the law of a pro-
gressive reduction in the formation of tooth-substance.
Moreover, according to Baume, with the exception of the
Rodentia, animals with continuously growing teeth belong to
old groups, the still existing representatives of which are
perhaps on the way to extinction. It is evident that by
* Cope, “ The Mechanical Causes of the Development of the Hard
Parts of the Mammalia,” Journ. of Morphology, vol. iii. 1889 (Boston),
I was unfortunately unable to consult the actual original communications
of this author upon the present subject in the ‘ American Naturalist’ of
1883-84 and the ‘ Report of the United States Geological Survey.—Ter-
tiary Vertebrata,’ 1885.
+ R. Baume, ‘Odontologische Forschungen, Theil i—Versuch einer
Entwicklungsgeschichte des Gebisses’: Leipzig, 1882.
of the Tooth-Rudiments in Rodents. 305
piping Baume’s view we avoid a difficulty of the following
kind: if we derive continuously growing rootless teeth from
rooted teeth with limited growth, we are compelled to assume
a repetition of the same process (convergence) in represen-
tatives of the most widely different families, which have no
direct connexion whatever one with another; for, according
to Baume, we find continuously growing teeth “in a motley
series among the Carnivora, Cetacea, Prosimiz, Multiungu-
lata, Sirenia, Ruminantia, Rodentia, and Marsupialia. They
are, as appears from the above comparison, widely distributed,
but also very scattered, when relationships are considered.
The various representatives occupy a number of isolated
positions.”
Mahn* and Fleischmann have disputed the justice of
Baume’s assumptions, adducing weighty arguments in oppo-
sition to them.
Moreover this conception of Baume’s is in accordance with
the other peculiar views advanced by this author, the most
prominent of which is his theory that the milk-teeth should
not be regarded as constituting a special dentition. Accord-
ing to this idea we should only get a pseudo-diphyodontism
in Mammals. The milk-teeth would belong to the same
series as the permanent ones ; they would merely be feebler
structures, developing more quickly. The stronger (perma-
nent) teeth develop more slowly according to Baume, but
afterwards displace the quickly developed feeble (milk-) teeth.
This is not the place for the elucidation of the question as to
what are the facts of comparative anatomy and embryology
upon which Baume bases his conception. I still have to
refer incidentally in the special portion of this paper to the
results of Baume’s investigations into ontogeny. At any
rate, even Baume coincides with all other authors in assuming
that the dentition of the Rodentia has arisen by means of
reduction from a richer and completer series of teeth.
How extensive the reduction must be supposed to have
been is evident from a comparison of the existing dental
equipment of the Rodents with the fundamental type of the
dentition of the Eutheria, as the latter is formulated by
Schlosser ¢ after Oldfield Thomas. According to the theory
referred to, the ancestors of the Placentalia would have
possessed five incisors, one canine, four premolars, and four
molars, though it must certainly be remarked that such an
* R. Mabn, “ Bau und Entwicklung der Molaren bei Mus und Arvi-
cola,” Morphol. Jahrb. Bd. xvi. Heft 4.
+ M. Schlosser, “ Ueber die Deutung des Milchgebisses der Siiuge-
thiere,”’ Biol, Centralbl. Bd. x.
306 Herr Paul Freund on the Development
abundance of teeth is not found to be realized in any recent
or known fossil form.
The question now arises as to how far traces of the process
of reduction, and remnants of the teeth which have disap-
peared, still admit of embryological demonstration in the case
of the Rodents.
This query naturally further divides itself into three in the
following manner, each of which questions must be separately
investigated for the upper and lower jaw :—
(1) Since, with the exception of the premaxilla of the
Leporide (in which two teeth are present), the recent
Rodentia possess only one incisor, we are confronted with the
question whether vestiges of the other incisors which have
disappeared are traceable in the embryo. Herein, especially
in connexion with the views of Cope which have been men-
tioned above, regard must be paid to the position of possible
embryonic rudiments with reference to the largeincisor. Are
the rudiments situated on the inside or outside of the latter?
And is this consequently to be considered as 7. 1 or as ¢. 2?
(2) Do vestiges of tooth-rudiments exist in the diastema ?
This question divides itself, according to the three stages
which are observable in the embryonic development of teeth,
into three subquestions :-—
(a) Do we find only a dental fold (Zahnleiste) in the
diastema ?
(5) If such a structure is present, are enamel-organs formed
upon it ?
(c) Is enamel produced within the enamel-organs, and
dentine by the papille ?
In the event of (>) and (ec) being answered in the affirma-
tive, this supplementary question arises:
Are the tooth-rudiments which are observed connected with
the incisors or the premolars, or do both conditions occur ?
(3) Since in the Rodent series the number of the molars
descends from 6 (in the upper jaw of Lepus) to 3 in the case
of the Muride (in Hydromys even to 2), in forms with a
reduced number a search would also have to be made for
remnants of the molars which have been lost. It is clear that
this task may partly coincide with one subdivison of the fore-
pong question (rudiment at the posterior end of the diastema).
wish to mention at once that in the present paper I have
not devoted any special attention to this third question.
Since the development of the molars in the Muride has been
closely studied by Mahn under Fleischmann’s direction, and
since that author says nothing about rudiments of the vanished
molars, which certainly would not have escaped his notice,
of the Tooth-Rudiments in Rodents. 307
such an investigation seems at the very outset to offer but few
prospects of success.
The material investigated by me is derived from embryos
of Lepus cuniculus, Sciurus vulgaris, Cavia cobaya, Cricetus
Srumentarius, Mus decumanus, and Mus musculus. My
methods consisted in the preparation of continuous series of
sections through the heads of embryos according to well-known
precepts. In particular I made use of the technique intro-
duced by Prof. Born in the Breslau Anatomical Institute, as
described by himself in his paper entitled “ Noch einmal die
Plattenmodellirmethode ”’ *.
Plate-models were prepared of the most important stages.
These naturally contained the epithelial tooth-rudiments, as
well as the epithelium of the oral cavity from which they had
proceeded ; it is evident that Mahn (loc. cit.) worked in a
precisely similar fashion.
I will take first the two species in the investigation of
which positive results have been obtained, i. e. Lepus and
Sciurus. The remaining forms, in which, in the sense of the
queries which we have propounded, nothing was discovered,
can subsequently be dismissed in a few words.
LEPUS.
With reference to this form a series of statements are con-
tained in Pouchet and Chabry’s paper (loc. cit.), which it is
necessary for me to cite, since [ have partly to coafirm, partly
to amplify, and partly to refute them. ‘The French authors
confirm the older observations, that in the premaxilla of the
Rabbit three larger incisors are formed on each side, which,
however, do not stand side by side, but one behind the other.
The second of the series are deciduous, and drop out shortly
before or shortly after birth. The ¢. 3f are formed much
* Zeitschrift fiir wissenschaftliche Mikroskopie, Bd. v.: Braunschweig,
1888. In this paper Prof. Born writes as follows:—“ As is well known,
the essence of the plate-model method is that from each of a series of
sections of equal thickness the parts which are to be modelled are drawn
upon plates, which are exactly so much thicker than the section as the
sketch is larger than the original. The parts drawn are cut out and stuck
one upon another; in this manner a plastic reconstruction is obtained of
the structure under investigation” (p. 445). .... “ Formerly I scratched
the outlines of the section upon prepared (cast) plates. I now use plates
of waxed paper, in accordance with Strasser’s suggestion, and after
drawing upon the paper, I cause thisto be thickened with wax until plates
of the desired thickness are obtained” (p, 446).
+ For convenience’ sake here and subsequently di. 2 (according to the
customary nomenclature) is designated shortly 7. 2, while the permanent
7, 2 is termed 2. 3.
308 Herr Paul Freund on the Development
later than 7. 1 and 7. 2. From my own experience I have
nothing to add to these statements, since the rudiment of ¢, 3
(the permanent ¢. 2) in the largest embryo examined by me
(10 centim. in total length) was but little further developed
than in the embryo from which fig. 20 of the French authors
is derived. The rudiment of this tooth was represented by a
clavate enamel-organ, indented only by quite a flat papilla,
and situated ventrally and posteriorly to the greatly developed
deciduous 7, 2. As stated by Pouchet and Chabry, 7.2 and
7. 3 lie in one and the same alveolus.
As Fleischmann convinced himself from the examination
of transverse sections, the dz. 2 are enamel-crowned teeth
with two roots of dentine.
In the mandible the rudiment of only one large incisor is
formed. In addition to this, however, Pouchet and Chabry
discovered in Lepus two new deciduous teeth, which are
situated in both jaws in front of the large 7. 1. They found
these teeth in embryos measuring from 28 to 45 millim. in
length, and in a rudimentary condition, to which nothing
analogous is known in other Vertebrates. The abridged
account of their discovery runs somewhat as follows :—In
embryos measuring from 28 to 45 millim. and something over
in total length, there is found in front of the well-developed
enamel-organ of the large incisor, and in direct connexion
with its anterior face, an epithelial mass, which, in a cavity
which is completely closed towards the exterior, contains a
thimble-shaped cusp of dentine. The innermost columnar cells
of the epithelial sac stand directly upon the surface of this
cusp, and its pulp-cavity is filled with a certain number of
blood-corpuscles and embryonic cells. ‘The enamel-organ (for
the epithelial sac can be considered as nothing else) of this
dentine cusp is distinguished, according to the French authors,
on the one hand by its compact texture (the enamel-pulp is
absent), and on the other by the fact that it completely sur-
rounds the cusp. The enamel-organ is connected by a cord
of epithelium with the posterior side and the lower border of
the sunken epithelial wall (“mur plongeant”’). (Such is the
term bestowed by these authors upon the sunken epithelial
masses, which are found in many embryos in the region of
the future groove between the lip and the alveolar margin.
Projecting masses of epithelium, to which the authors apply
the term “mur saillant,” correspond approximately to the
thickenings of the epithelium of the oral cavity at the edge of
the jaw, which Germans call the jaw-wall [| Kieferwall |.)—
Our cusp of dentine disappears long before birth, since it is
either absorbed or drops out. According to Pouchet and
of the Tooth-Rudiments in Rodents. 309
Chabry the enamel-organ of the large incisor is also connected
by a cord of epithelium with the mur plongeant, but with its
posterior and upper border. ‘The authors referred to found a
similar little rudimentary tooth in the upper jaw only in
embryos which measured not less than 32 to 40 millim, in
length. “It lies immediately in front of the large incisor,
and its tip appears in the midst of the epithelium of the mur
plongeant”’ (fig. 21).
The earliest stages which I was able to examine thoroughly
consisted of embryos of Lepus cuniculus measuring 15 millim.
in total length, of which the head occupied 9 millim. For
earlier ones I had at my disposal only the series of sections
from an embryo with a head 6 millim. in length. Since this
shows only the dental fold, which is apparently continuous in
the upper jaw, but in tle lower one on the coutrary is inter-
rupted behind the region of the incisors, while the enamel-
organs are as yet entirely absent, 1 shall defer the investiga-
tion of these youngest stages until I have more abundant
material at my disposal.
First of all I deal with the region of the incisors in embryos
with heads measuring somewhat more or somewhat less than
lcentim. ‘The length of these embryos from the crown of
the head to the root of the tail, which, for reasons which will
be readily understood, is much more variable, fluctuates be-
tween 1°5 and 3 centim. Fig. 13* gives a representation of
the model which I have prepared of the epithelium of the
oral cavity, with the tooth-rudiments of the upper jaw of the
right side, of an embryo measuring ‘9 centim. in cephalic
length and 2-1 centim. in all; the model is seen from above
and somewhat from outside; I have not figured a second
model showing the rudiments of the incisors in the upper and
lower jaws of an animal of about the same size.
In the lower jaw the rudimentary tooth discovered by Chabry
and Pouchet may be seen. Fig. 1 gives a distinct view of this,
from which it appears that the enamel-organ of the tooth is
not closed on all sides, as asserted by Chabry and Pouchet,
but is open on the underside. Through the aperture a richly
nucleated pulp enters the interior of the tooth. The centre of
the pulp is occupied by a wide blood-vessel, while its peripheral
cells form a well-developed layer of odontoblasts, which has
already secreted a solid hook-shaped cap of dentine. The
latter, in the preparation upon which the drawing was based,
has withdrawn a little from the well-developed enamel-epi-
* The figures referred to in this paper will be found in vol. 39 of the
‘ Archiy fiir mikroskopische Anatomie.
310 Herr Paul Freund on the Development
thelium. In short, apart from the absence of enamel, it is a
thoroughly typical representation of a small tooth-rudiment,
such as we are otherwise wont to find in lower Vertebrates.
The connective cord, which unites the enamel-organ of the
rudimentary tooth with the epithelium of the oral cavity,
actually extends with its anterior end, as stated by the French
authors, as far as the posterior circumference of the mur plon-
geant, which is here but little developed. Further back-
wards, however, this connective cord proceeds, as is shown in
fig. 1, from the unthickened epithelium of the oral cavity.
Moreover, we see from the figure that a strong epithelial pro-
jection, which is directed inwards, starts from the spot where
the connective cord passes into the actual enamel-bell
(Schmelzglocke). ‘This arrangement is repeated more or less
decidedly in each section. According to the customary con-
ception the tract a—6 in fig. 1 would have to be regarded as
the “ dental fold ” (‘‘ Zahnleiste ’’), on the outer side of which,
close above the end without a neck, the enamel-organ of the
rudimentary tooth would be situated. Whether this mode of
interpretation is here correct, can only be determined by the
study of younger stages, which I am reserving to myself.
Now the enamel-organ of the large incisor of the lower jaw is
directly connected with the posterior side of the enamel-organ
of the rudimentary tooth ; in the same manner the cord which
connects the enamel-organ of the rudimentary tooth with the
epithelium of the oral cavity is posteriorly prolonged directly
into the similar connective cord of the enamel-organ of the
large incisor, If, as has been indicated above, we regard
this cord as a dental fold, this continuity of the dental fold
from one enamel-organ to the other would be nothing unusual,
—on the contrary, it is in accordance with the general rule.
The direct connexion, however, of two enamel-organs, as,
moreover, also shown with great distinctness in fig. 19 (longi-
tudinal section) of Pouchet and Chabry, is unusual. ‘The
enamel-organ of the permanent incisor is still at a very low
stage of development. It is a lumpy epithelial mass, which
on the posterior side appears somewhat flattened and indented.
This shallow hollow subsequently deepens into the bell of the
enamel-organ, which is open towards the rear. In the rudi-
ment of the large incisor no trace of tooth-substance is yet
developed.
Behind the large incisor the dental fold in the lower jaw
entirely ceases. In order to dispose of this question at once,
it may be mentioned that in the case of Lepus, in this, as in
all subsequent stages, an absolute diastema is found in the
ror
of the Tooth-Rudiments in Rodents. 311
lower jaw between the incisor and the molars, in which
distinct vestiges of a dental fold can never be detected,
Now as regards the incisors of the premaxilla in rabbit-
embryos of about 1 centim. in cephalic length (cf. fig. 13): con-
trary tothe statements of Chabry and Pouchet, the rudimentary
tooth in the premaxilla is also already completely developed
in embryos measuring 1 centim. in cephalic length, and about
2 centim. in all. As is shown in fig. 13, it is situated upon
the epithelium of the oral cavity close behind the mur plon-
geant, which is still but little developed, in such a manner that
a special connective cord between its enamel-organ and the
oral epithelium cannot be distinguished. The enamel-organ
is longer than in the lower jaw; the longitudinal axis appears
to be directed obliquely upwards and outwards. At the
upper end, displaced a little towards the front, we find the
very narrow aperture, surrounded by aswollen margin, which
leads into the interior of the pulp-cavity. The solid dentine
tooth is developed just as beautifully as in the lower jaw.
Owing to the absence of a special connective cord, the rounded
tip of the very long tooth is separated from the surface of the
epithelium of the oral cavity only by a few layers of cells.
In the upper jaw also the enamel-organ of the rudimentary
tooth is in direct continuity with that of the large incisor.
In the model it is easy to recognize the peculiar conditions
which are presented by the attachment of the first large
incisor to the epithelium of the oral cavity,—conditions which
can hardly be understood from the examination of sections,
and to which it isonly with difficulty that justice can be done
in description also (in the figure, too, they can be seen but
imperfectly). For behind the spot at which the enamel-organ
of the large incisor is attached, the roof of the oral cavity
bulges out downwards in the form of a step. (A similar step
follows further back at the spot at which the enamel-organ of
7. 2 is attached.) Moreover, the connective cord of the
enamel-organ of the large incisor has a peculiar shape ; for it
consists of two plates, one placed transversely, the front of
which, like the greater portion of the enamel-organ itself of
7.1, is fused with the enamel-organ of the rudimentary tooth,
and another in the shape of a sagittal plate, attached at right
angles to the inner edge of the former, and prolonged back-
wards as far as the enamel-organ of 7. 2. In the case of the
latter tooth the same conditions are repeated. We can also
express the facts by saying that the connective cords, which
unite the enamel-organs of ¢. 1 and ¢. 2 with the epithelium
of the oral cavity, are hollowed out from behind and some-
what from inside. ‘lhe connective cord of the first incisor is
312 Herr Paul Freund on the Development
fused in front with the enamel-organ of the rudimentary tooth,
and behind passes into the connective cord of ¢, 2, while, as
we shall see, the latter is prolonged backwards into the dental
fold of the diastema. This, together with the steps, produces,
as has already been stated, highly complicated appearances.
What we have to remember is, that the enamel-organs of 7, 1
and the rudimentary tooth are in direct continuity, while ¢. 1
and 7. 2 are united by a structure which may well be regarded
as a dental fold. The epitlielial mass of the enamel-organ of
¢. 1 itself is lumpy, and when seen from the upper side behind
somewhat flattened and indented. ‘The margins of this hollow
subsequently become more decidedly elevated, and so assume
the characteristic bell-shape. The enamel-organ of ¢. 2 is, as
may be seen from fig. 13, in not nearly so developed a con-
dition.
We will here at once mention the changes which take
place in the region of the incisors in subsequent stages. With
reference to the lower jaw we have to observe that the sunken
epithelial wall (mur plongeant), which marks out the subse-
quent groove between the lip and the alveolar border, becomes
much more strongly developed and extends further back-
wards. In the foremost sections it appears as a bifurcate
and deeply sunken epithelial mass, which to a certain extent
ensheaths the anterior end of the Meckelian cartilage and the
covering bones lying thereon. On the inside of the epithelial
mass is attached the common connective cord of the enamel-
organs of the rudimentary tooth and of the large incisor. ‘The
latter in embryos measuring 1°5 centim. in cephalic length and
about 4 centim. from crown to rump, already assumes the
well-known bell-shape, in the manner which has been indi-
cated above. Since the size of this bell, which is enormously
expanded behind, and within which the tooth develops in the
usual way, by far preponderates, it now appears as though
the tip of the rudimentary tooth were implanted in the cord
connecting this great bell with the epithelium of the oral
cavity, while only the lower end of the tooth, surrounded by
its enamel-organ, projects freely downwards in front of the
great bell. ‘lhe conditions which are now present are conse-
quently similar to those which are shown in fig. 14 in the
case of the squirrel. ‘The rudimentary tooth is indeed far
behind the large incisor in growth, but nevertheless in older
animals, e.g. in rabbits measuring about 2°5 centim. in
cephalic length and etrca 7 centim. in all, it has decidedly
increased in size in comparison with the younger stages. Its
pulp-cavity is never closed towards the exterior, as is stated
by Chabry and Pouchet, but always remains in connexion
of the Tooth-Rudiments in Rodents. 313
with the surrounding mesoderm, though it is trué not in the
simple manner which is seen in fig. 1. In larger rabbits the
size of the rudimentary tooth appears decidedly diminished,
as though it were in process of absorption. In still later
stages I could no longer find any trace of it, although I am
unable to make any precise statement as to the manner of its
disappearance, any more than Chabry and Pouchet were able
to do.
In the premaxilla the changes are precisely similar to those
in the lower jaw. A strong sunken epithelial wall is formed,
which indicates the region of the subsequent cleft between the
two halves of the lip and the anterior end of the jaw. It
consequently appears simple in the foremost sections, and
bifureate further back. One half of it is to be seen in fig. 2,
and a longitudinal section in the combination-picture shown
in fig. 8. In this enormous epithelial mass the anterior step
on the roof of the oral cavity, as described above in the
younger stage, completely disappears. On the inside of the
lateral tip of this epithelial mass is situated, as is shown in
fig. 2, the enamel-organ of the rudimentary tooth (fig. 2
belonged to a series through the head of a rabbit embryo
measuring 5'l centim. in all and 2-3 centim. in cephalic
length). The enamel-organ of the first incisor develops into
a huge bell, which is open behind, and, in accordance with the
conditions previously described (ef. figs. 2 and 13), the rudi-
mentary tooth now appears to lie in the cord connecting this
bell with the mur plongeant. Owing to the development of
the mur plongeant the tip of the tooth has naturally pene-
trated very deep down. The combination-picture fig. 3,
which is constructed from longitudinal sections through the
head of a rabbit embryo of the same size as that of fig. 2,
shows these conditions perhaps most distinctly.
The histological conditions of the rudimentary tooth may
be recognized in fig. 2; its dentine casing appears broken
through on the outside in this figure. The connexion between
the pulp and the mesoderm takes place through this breach,
not directly however, but in a very complicated fashion, since
the enamel-organ appears as if cleft by strands of connective
tissue, producing conditions of which the detailed description
belongs to another place. In the same way I have no inten-
tion of entering into a discussion of the histological conditions
in the enamel-organ of the large incisor. I would merely
refer the reader once more to fig. 2, which in some degree
exhibits the peculiar penetration of vascular outgrowths into
the outer wall of the enamel-organ, which has already been
dealt with by so many investigators. ‘lhe rudimentary tooth
314 Herr Paul Freund on the Development
of the premaxilla persists for about as long as that of the
lower jaw. Iam at present unable to say anything as to the
manner of its disappearance. In embryos with a cephalic
length of 1:7 centim., and measuring 4°5 centim. in all, the
dental fold between the first and second incisors was still
present, although it is true that its connexion with the oral
epithelium had almost everywhere disappeared. The enamel-
organ of 7.2 had become distinctly campanulate. In an older
series the dental fold between the two large incisors had
already entirely disappeared, although in the model of the
stage in question the step upon the palate was still to be
seen, to the anterior margin of which the cord connecting the
enamel-organ of 7. 2 with the oral cavity was previously
attached, while, in all probability as a remnant of this con-
nexion, we still find an epithelial projection jutting out
slightly into the connective tissue.
While up to this point, with the exception of immaterial
modifications, I was able to confirm the statements of Chabry
and Pouchet on the subject of the development of the teeth in
the rabbit, the results at which I have arrived with reference
to the diastema in the upper jaw are quite different from those
obtained by the French authors. The latter assert that in
the rabbit the dental fold is entirely interrupted in the
diastema in both jaws. As I have already mentioned, I can
confirm this statement as regards the lower jaw. In the
upper jaw, on the contrary, I find the dental fold already
typically developed, in the youngest stages investigated by
me (measuring about 1 centim. in cephalic length and from
2 to 3 centim. in all), throughout the entire length of the
diastema between the rudiments of the incisors and those of
the molars. A good representation of this is given in fig, 18.
The dental fold is situated in precisely the same spot as that
in which it is found in mammals with a complete dentition,
7. e. internally to the furrow made by the approximately hori-
zontal roof of the mouth in bending round into the perpen-
dicular mucous membrane of the cheeks. The spot at which
the dental fold is attached is in the region of the diastema, as
in that of the rudiments of the molars, somewhat depressed,
towards a protuberance of the surface of the palate situated
on the inner side of it (jaw-wall?). It is perhaps expedient,
in addition to the measurements, to give yet other character-
istics of the stage in which the fold is found to be fully
developed. ‘The stage is that in which the closure of the
palate has just been completed, in which we still find distinct
remains of epithelium in the perpendicular palatal suture, and
in which, in the horizontal palatal suture (between the nasal
of the Tooth-Rudiments in Rodents. 315
septum and the hard palate), a characteristic arrangement of
the connective-tissue elements is still evident; by this, how-
ever, it is in no way intended to imply that the epithelial
fold is not already present before this stage when the palatal
cleft is open, and subsequently when the palatal suture has
disappeared. My series furnish me with suflicient proofs of
both cases. I merely wished to characterize the stage at
which the fold appeared to me to be most developed. The
epithelial fold at its anterior commencement is very shallow
and inconspicuous, as it is also reproduced in model 1, and is
connected with the enamel-organ of ¢. 2. In the region of
Stenson’s canal it is sometimes so inconsiderable that it is
with difficulty detected. Behind this it speedily increases in
height, and attains its greatest elevation at the transition into
the rudiments of the molars. In the sections it appears as an
epithelial cord, which proceeds from the epithelium of the
oral cavity, inclined somewhat obliquely inwards, and is
buried in the mesoderm. In the neighbourhood of the oral
epithelium it appears narrow, having a breadth of only two
very low rows of cells; the sunken end is thickened into a
button-shape, owing partly to the increased height of the
peripheral cells and partly to the intercalation of several cells
in its interior. ‘The form and size naturally vary very con-
siderably according to the direction of the section and other
accidental circumstances. Fig. 4 gives a representation of
the epithelial fold in a series im which it appears particularly
broad. In the other series it looks decidedly more slender
and longer. In the section figured (fig. 4) the length of the
epithelial depression amounts to about 100 w, the breadth of
the thick end to about 60 yw, and that of the narrowed neck to
30 pw. In the series from which the model shown in fig. 13
is constructed, the height of the epithelial fold at about the
middle of its length amounts to nearly 120 pw, the breadth at
the thickened end only to something over 20 yw, and at the
narrowed neck only to about 10 p.
Even in an embryo measuring 11 millim. in cephalic length
and 34 millim. in all, but much more distinctly and much
more extensively in an older one of 17 millim. and 44 millim.
respectively, the dental fold in the diastema exhibits a remark-
able modification. At first only for a limited space, but in
the older embryo almost throughout its entire length, the neck
of the epithelial fold appears broken through by the mesoderm.
The appearances in this connexion are precisely similar to
those seen in other places, where a strand of epithelium is
displaced by the adjacent connective tissue. The breach
sometimes affects only the portion beneath the button-shaped
316 Herr Paul Freund on the Development
end, so that a small cone is attached to the oral epithelium,
followed by connective tissue, in the arrangement of the cells
of which we can still frequently trace the direction of the
epithelial cord which has disappeared. Deep down we then
see the severed thickened end as a circle filled with epithelial
cells, which is distinguished by the fact that, like other
epithelial remnants undergoing degeneration in the same way,
it takes up carmine very freely, and consequently appears of
a dark pak colour. It is clear that this process is the same as
that also undergone by the dental fold in mammals with a
complete dentition, after the enamel-organs have been formed
and constricted off from it. The dental fold in the diastema —
of the rabbit, however, perishes without ever having produced
enamel-organs. In an embryo rabbit 23 millim. in cephalic
and 51 millim. in total length every trace of the fold has
disappeared in the anterior portion of the diastema; but in
the posterior portion, on the contrary, in the neighbourhood
of the molars, there appears in each section at the same spot
in the connective tissue of the palate a small red circle, which
I am inclined to regard as the last vestige of the dental fold.
My reasons for doing so are the following :—In the first place,
it is found in the posterior portion of the diastema in every
section at the same spot, and this spot, so far as I can see,
corresponds to that at which the epithelial fold is found in
younger stages. Secondly, in the connective tissue there
frequently proceeds from the red spot to the oral epithelium
a kind of cord of cicatricial tissue, ¢. e. a clear streak, in which
we find but few nuclei, or none at all, while at the edges of it
the nuclei are closely packed. ‘Thirdly, the epithelial circle
can be traced as far as the dental fold of the first premolar,
though I am unable to assert with absolute certainty that a
direct connexion exists. If my interpretation is correct, it
would result therefrom, that the dental fold in the diastema
in the rabbit disappears from in front backwards.
With regard to the rudiments of the molars in Lepus cunt-
culus, I have no special statement to make. In fig. 13 we see
only three, and these still in a very undeveloped condition :
the enamel-organs have the shape of flat shells. Another
model, which is not figured here (prepared from a series of
longitudinal sections through the head of a rabbit embryo
2°3 centim. in cephalic and 5°1 centim. in total length), shows
the rudiments of five molars. In the second and third we
find the characteristic bell-shape, with the projections directed
towards the pulp. Nos. 1 and 4 are less developed; the fifth
rudiment appears in the form of a button-shaped epithelial
thickening, with only quite a shallow depression, at the
posterior end of the dental fold.
of the Tooth-Rudiments in Rodents, 317
ScCIURUS.
Of this very interesting form, as it proved to be, I suc-
ceeded, in spite of a considerable expenditure of trouble and
expense, only in obtaining one pregnant female with seven
embryos. However, I reserve to myself the working out of
older and younger stages, since I have at any rate learnt the
way in which these may be obtained with certainty next year.
Fortunately the stage which I have before me corresponds
fairly accurately with that to which the most important state-
ments of Chabry and Pouchet refer. Since I differ from them
very considerably, it will be well to give a summary of the
statements of the French authors, which are somewhat diffi-
cult to understand. They runas follows :—“ The dental fold
commences in the upper jaw at the level of the incisors, and
extends to beyond the molars.... In front of the canals of
Stenson the dental fold is entirely wanting.... In the
mandible the dental fold is continuous throughout the entire
extent of the diastema.”’
Now follows a sentence which does not seem quite intelli-
gible after what bas gone before. It runs :—“ Consequently
the dental fold in all Rodents which we have studied is com-
posed of two halves, which are not united in the middle. In
the squirrel in particular the incisors arise from the same
epithelial invagination (involution) as the molars.”
According to what was previously stated, this would be
correct only for the lower jaw. For in the upper jaw the
dental fold is said to be completely wanting even betore the
canals of Stenson.
The explanation of the figures shows that the statements
of Chabry and Pouchet refer to embryos of the length of
4centim. ‘The embryos studied by me were 2 centim. in
cephalic length and 4°7 centim. in all, and were consequently
only slightly larger than those examined by the French
authors.
The foilowing are the results which I obtained :—
(1) Like the rabbit, the squirrel possesses in front of the
large incisor a rudimentary tooth, which seems to have been
overlooked by Chabry and Pouchet.
(2) There is found in the lower jaw, in the diastema be-
tween the incisors and molars, and extending only for a very
short space, about as far as may be seen in fig. 15, in con-
tinuation of the dental fold of the incisors, a small epithelial
cord, which is detached from the epithelium of the oral cavity,
and is to be regarded as a remnant of the dental fold. Inthe
whole of the large space behind this, as far as the molars, no
Ann. & Mag. N. Hist. Ser. 6. Vol. xi. 23
318 Herr Paul Freund on the Development
trace of it can be detected. Whether we may assume that in
the only slightly younger stages which were investigated by
Chabry and Pouchet, a complete dental fold is, as stated by
these authors, present throughout the entire diastema of the
lower jaw, is a question which I must postpone until I have
more ample material at my disposal. Ido not think it pro-
bable that itis so.
(3) Inthe upper jaw very extensive remnants of the dental
fold are found in the diastema. The foldis not once wanting
in any section from the incisor to Stenson’s canal: compare
figs. 8-10, and the model (fig. 14). In most places the epi-
thelial cord is detached from the epithelium of the oral cavity,
in others it is connected with it. It frequently appears to lie
parallel to the surface of the palate; the nuclei of the con-
nective tissue are disposed in a regular concentric arrange-
ment around the epithelial remnant. This is therefore directly
contrary to what is stated by Chabry and Pouchet.
Behind the orifice of Stenson’s canals we find for a space
no vestige of the epithelial fold: then it is present in certain
sections once more, or in its place there appears a kind of
cicatrix in the connective tissue. Even before the middle of
the space between the orifices of the canals of Stenson and
the first molar, we observe a remnant of the epithelial fold as
an element which is again regularly visible in every section.
At the posterior end it is easy to distinguish the way in which
this remnant of the dental fold of the diastema passes into
the dental fold of the first molar. It may further be remarked
that in this tract also there is much variation; for the epi-
thelial cord may appear larger or smaller, perpendicular or
bent round, connected with the epithelium of the oral cavity
or isolated. After this discovery it appears scarcely open to
doubt, that in younger stages of Sczwrus the diastema in the
upper jaw is traversed by a continuous dental fold, precisely
as in the case of Lepus.
(4) On the outer side of the dental fold in the diastema
there are situated two little enamel-organs with distinct
papillee,—one close in front of, and the other in the plane of a
transverse section through, the orifices of the canals of Stenson
(cf. model, fig. 14). ‘These two enamel-organs naturally lie
one close behind the other. In both the aperture of the
enamel-bell is directed forwards and upwards. The anterior
of these two enamel-organs is the better characterized as such,
in that it contains enamel-pulp (cf. fig. 7). But that the
posterior rudiment is also of the nature of an enamel-germ
(cf. fig. 10) is scarcely open to doubt. It is curious that in
their fig. 26, MM. Chabry and Pouchet have given a perfectly
of the Tooth-Rudiments in Rodents. 319
characteristic representation of the posterior of the two rudi-
ments (the section also passes through the region of the orifice
of the Stenson’s canal) : compare for this purpose my fig, 10.
But according to the explanation of the figures they interpret
the rudiment quite wrongly as that of the large incisor,
which, as a glance at the model, fig. 14, will show, is much
further developed, and besides this also has quite a different
position.
It now remains for us to make more special observations
with reference to certain of the above four paragraphs.
The dental fold wpon which in the lower jaw the enamel-
organs of the rudimentary tooth and of the large incisor are
situated (cf. for this figs. 5 & 6, and model, fig. 15) proceeds
from the commencement of a peculiar mur plongeant, which
envelops as a deep case the anterior end of the Meckelian carti-
lage together with the superincumbent covering-bone. It is
precisely the same formation as that described and figured by
Chabry and Pouchet for the rat. Together with the epithe-
lium of the oral cavity, it appears in a median transverse
section as a closed circular epithelial arch, in the interior of
which the dental rudiments, the bone, and the cartilage are
contained. Upon the surface of the mucous membrane of the
oral cavity, the spot from which the epithelial fold and be-
side it the mur plongeant proceed is distinguished by a groove.
For posteriorly the epithelial fold ceases to be in continuity
with the epithelium of the oral cavity (cf model, fig. 15).
That from the fold an epithelial cord runs backwards for a
short distance has already been mentioned, and may be seen
from the model. Now on the outer side of this epithelial fold
the enamel-organs of the rudimentary tooth and of the large
incisor are attached close together, each by means of a long
neck, in such a way that only the somewhat clavate end ot
the epithelial fold projects a little further downwards and
inwards than this point of attachment. For, as is shown both
by the figures and the model, the neck of the rudimentary
tooth is a particularly elongated structure, which reaches so
far downwards and backwards, that the enamel-bell of the
rudimentary tooth comes to lie beside the Meckelian cartilage
(ff. fig. 6). Neither the neck of this enamel-organ, the
stouter and more massive one of the incisor lying behind it,
nor lastly also the dental fold itself have smooth outlines ; on
the contrary, they are beset with projections and depressions
of the most varied kind (cf for this figs. 5 & 6), so that in
many places the appearance presented reminds us of that of
the proliferating rudiment of a gland, or still better of that of
a@ carcinoma.
23*
320 Herr Paul Freund on the Development
The opening of the enamel-bell of the rudimentary tooth
is, like that of the large incisor, directed backwards. ithin
it we do not find, as in the case of Lepus, a well-formed tooth
of dentine, but a small, inconsiderable, almost solid dentine
cusp, into which from the inner side only a narrow cord of
connective tissue enters. This cusp, too, appears to be no
longer enveloped on all sides by epithelium. With reference
to the neck of the enamel-organ of the incisor, which follows
after it, there is nothing more to be said than that at its
starting-point from the dental fold it is broadly fused with
the neck of the enamel-germ of the rudimentary tooth.
The position of the germ of the rudiment in the upper jaw
is explained by model fig. 14. We observe that it is situated
in front of the enamel-germ of the large incisor. The enamel-
germ of the rudimentary tooth of the upper jaw is likewise
attached to a long neck, which passes obliquely upwards and
unites with the massive neck of the large incisor of the pre-
maxilla. Here also, though certainly less decidedly than in
the lower jaw, the irregularity of contour is repeated. The
opening of the enamel-bell of the rudimentary tooth is directed
backwards and upwards. Inside it the pulp surrounds a
delicate little cap of dentine. Figs. 11 and 12 will explain
these statements. Fig. 11 gives a representation of the ar-
rangement magnified nearly fifty times. We see from the
figure that the enamel-germ of the rudimentary tooth and that
of the large incisor are surrounded by one and the same thick-
ened envelope of connective tissue. Fig. 12 shows from the
same section the enamel-germ of the rudimentary tooth mag-
nified nearly two hundred times.
With reference to paragraph no. 4 the following remarks
must be made :—Fig. 7 shows, magnified one hundred times,
a section through the anterior rudimentary enamel-organ,
which is situated close in front of the orifice of Stenson’s
canal. We also get a sectional view of the portion of the
dental fold belonging to it; but in this section the fold is in
connexion neither with the epithelium of the oral cavity nor
with the transversely-directed neck of the enamel-organ.
Both, however, lie together in an envelope of connective
tissue, which is packed with nuclei and consists of concentric
layers. In the interior of the enamel-bell distinet enamel-
pulp is found: a dentine germ could not be distinguished.
The most superficial nuclei of the pulp appeared clearer: this
is somewhat exaggerated in the figure. ‘They were separated
from the deeper-lying dark ones by a transverse cleft, which
perhaps represents a blood-vessel.
Fig. 8 shows the arrangement of the structures, magnified
of the Tooth-Rudiments in Rodents. 321
twenty-six times, about nine sections (15 w) further back.
The section passes through the anterior circumference of the
orifices of Stenson’s canals. Of the enamel-bell of the
anterior rudiment only the posterior margin is still cut, while
inwards from it is seen the dental fold detached from the
epithelium.
Fig. 9 lies three sections further backwards (same scale).
The section passes through the narrow interval between the
two rudimentary enamel-organs. The dental fold is to be
observed, and to the side of it a thickening of the epithelium
of the oral cavity, with which the fold at other spots is con-
nected. Lastly, fig. 10 lies only two sections behind the fore-
going, and shows the second rudimentary enamel-bell. This
is by far less distinctly characterized as such than is the
former one. For, in the first place, it is directly attached to
the outside of the epithelial fold, and is united to the epithe-
lium of the oral cavity by means of a short thick connective
cord. Secondly, the enamel-pulp is absent from its interior,
and, moreover, the connective-tissue-pulp, or tooth-papilla, is
not so thickly nucleated as in the case of the anterior rudi-
ment. Yet the whole structure is enveloped in a concentric
membrane of connective tissue (not visible in the figure), and,
especially after examination of the model, fig. 14, it is hardly
possible to doubt that the rudiment is of the nature of an
enamel-organ.
CAVIA.
Of Cavia I was able to examine a small specimen of the
cephalic length of 4°5 millim. This presented no special
interest for our subject, since the tooth-rudiments were still at
the very commencement of their development*.
Then come two series, one through the head of an embryo
measuring 1°4 centim. in cephalic length and 2-9 centim. in
all, while the other was derived from a somewhat younger
specimen, which Prof. Froriep, of Tiibingen, was kind enough
to make overtome. ‘The stages were so far very favourable in
that they corresponded to those of the rabbit in which the rudi-
mentary tooth-germs are most distinct. The result of the
investigation was, however, an almost negative one: no trace
of a rudimentary tooth and no trace of a germ for the second
incisor. In the diastema there were found here and there
thick cones of epithelium, which projected downwards for a
* Iam indebted for this specimen to the kindness of Dr. Keibel, of
Freiburg i. B., who placed at my disposal a whole series of guinea-pig
embryos. Most of these, however, were too young and consequently
not available for the purposes of my investigation. :
322 Herr Paul Freund on the Development
certain distance from the epithelium of the upper jaw, and
were also frequently continued backwards through a few
sections as detached round epithelial circles, after which, how-
ever, they always came to an end. It is very possible that
these represent remnants of the dental fold, though it is true
that a more definite proof cannot be advanced. We might do
this if we could trace such an epithelial cone into continuity
either with the dental fold of the incisors in front, or with
that of the molars behind. _ But we can do neither. Yet one
argument which supports the idea that these cones are of the
nature of a dental fold is to be found in the place from which
the ingrowths start, which agrees fairly well with the typical
position of the dental fold in Lepus. It is true that in Cavia
the relations of the epithelium of the oral cavity and of the
mucous membrane of the palate are somewhat different from
those in Lepus; but this is a question which I cannot here
proceed to discuss. Moreover we might establish a proof by
the process of exclusion ; besides rudiments of a dental fold,
the epithelial cones could only be germs of glands. It is,
however, improbable that the germs of small oral pituitary
glands would be already so distinct in the stage which we are
considering. But this attempt at a proof is not sufficient.
CRICETUS FRUMENTARIUS, Mus MuscULUS, and Mus
DECUMANUS.
Few words are needed to dispose of these animals.
Although we had at our disposal continuous series of at least
the two last-named species, the investigation of them never-
theless produced absolutely negative results. Not a trace of
rudimentary tooth-germs was found either in the neighbour-
hood of the incisors or in the diastema.
The discussion and valuation of my results may be divided
into heads according to the questions with which we
started :—
(1) Are embryonic tooth-rudiments found in the neighbour-
hood of the incisors in Rodents? To this we reply :—In
Lepus we find a rudimentary tooth in the upper and lower
jaw in front of the large incisor, as already discovered by
Chabry and Pouchet. This remarkable observation appears
to have escaped the notice of most of the subsequent authors,
but I was able to confirm it and to add that evidently the
same tooth-rudiment is also present in the squirrel.
Now Dr. Fleischmann has led me to inquire whether this
rudimentary tooth represents the remnant of an ¢. 1—in which
case the large incisor would have to be designated 7, 2, which
of the Tooth-Rudiments in Rodents, 323
agrees in a remarkable manner with Cope’s derivation of the
Rodents, as quoted above,—or whether it must be considered
as the remnant of the germ of a precursor of the large incisor.
Unfortunately the material furnished by the actual results
does not entirely suffice for a decision between these two
alternatives.
The development of the rudimentary tooth, as I was able
to follow it continuously in Lepus, by no means agrees in its
main features with that of a milk-tooth, whether, in respect of
the relation between the development of a milk-tooth and that
of the permanent one, we accept the older view, which is
reproduced in the text-books, or that which is held by Baume.
So far as I am aware, no case is known in which the enamel-
organ of a milk-tooth appears directly united with that of its
successor, as MM. Chabry and Pouchet have already shown
to be the case in Lepus, at least with regard to the rudimen-
tary tooth of the lower jaw. In the premaxilla the French
authors failed to observe the stage in question; I was able to
show that here, at any rate at first, the same thing takes place
in Lepus.
In a superficial examination of the question, the conditions
pertaining to the second deciduous incisor in the premaxilla
of Lepus tell very much against the assumption that the
rudimentary tooth likewise belongs to the milk series. The
deciduous incisor referred to is clearly a true precursor of the
second and smaller permanent incisor. It attains a high
degree of development at a time when the germ of the per-
manent 7. 2, which at all events proceeds from the adjoining
portion of the dental fold, is still quite small and insignificant.
It is not until the last stage of foetal life that the permanent
7.2 becomes more strongly developed and displaces the
deciduous 7. 2, which drops out shortly before or after birth.
Now are we to suppose that in Lepus the second incisor has
retained a true milk-tooth as its precursor, while in the case of
the first incisor the milk-tooth remains in quite a rudimentary
condition, and is outstripped unusually quickly by the germ
of the permanent tooth, which appears at the same time as, or
even earlier than, that of di. 2? It is true that in explanation
of this divergence we can adduce the necessity for a more
speedy development of the permanent 7.1 (which, indeed,
represents the actual large incisor), whereby to a certain
extent its precursor is deprived of the necessary material for
development, and is prevented from passing beyond a rudi-
mentary condition.
Finally, we must also confess that it is more easily
imaginable that the enamel-organs of a milk-tooth and its
324 Herr Paul Freund on the Development
successor, which in some other way at any rate also stand in
closer relation one to another, should under the special con-
ditions in question (especially great and therefore also more
rapid development of the permanent tooth) fuse together, than
that two tooth-germs situated one behind the other, which
have otherwise nothing whatever to do with one another,
should enter into intimate mutual relations of this kind.
Another telling point is that the rudimentary tooth and the
incisor lie together not only in an alveolus, but also, as may
be seen with especial distinctness in the case of the squirrel,
in one tooth-sac of connective tissue. I do not wish to attach
too great weight to the relative position of the tooth-germs.
It is true that the rudimentary tooth lies in front of the
incisor, but the deciduous 7.2, which we must yet certainly
regard asa milk-tooth, also lies, at least according ,to the
representation of MM. Chabry and Pouchet, in front of the
germ of the second permanent tooth. This is connected with
the peculiar conditions of space in the jaws of Rodents. It
appears to me to be of more importance that in the squirrel,
in which the conditions in question seem to be to a certain
extent less abnormal than in the rabbit, the enamel-germs of
the rudimentary and permanent tooth are connected together
by their “necks” (and this both in the upper as well as in
the lower jaw) precisely as this is usually described for the
milk-tooth and its successor. Altogether I am personally
more inclined to the view that the rudimentary tooth is to be
considered as the rudiment of a precursor of the large incisor
in process of active degeneration, though I am ready to admit
that this can by no means be regarded as finally established.
It may further be specially pointed out that in Sedwrus, which
in many respects has proved even more conservative than
Lepus, no distinet trace could be discovered of 7.2, which
Lepus has preserved in the shape of the milk-tooth and its
successor. It will be shown later on that the anterior of the
two enamel-organs, which I have proved to exist near the
orifices of Stenson’s canals, can only with difficulty be regarded
as 7.2, Further investigations will have to determine
whether the rudimentary teeth, as they appear in the stage of
Sciurus examined by me, have exactly arrived at the height
of their development, whether they develop still further, or,
lastly, whether they are not perhaps already in process of
degeneration.
(2) In the Rodents examined are germs of teeth found in
the diastema? ‘To this our discoveries enable us to reply :—
In Lepus a well-developed dental fold is found throughout the
of the Tooth-Rudiments in Rodents. 325
entire length of the diastema, but only in the upper jaw, which
was expressly denied by Pouchet and Chabry.
Upon this dental fold, however, enamel-organs are never
developed in Lepus. After having lasted for a somewhat
lengthy period, it disappears without leaving a trace behind.
In the upper jaw of Se/urus in the stage we examined a
dental fold was present, which was interrupted for a certain
distance behind the canals of Stenson, but was otherwise
greatly extended and continuous. According to Chabry and
Pouchet it is just in front of Stenson’s canals that the fold is
wanting. Moreover it is asserted by the French authors,
though their statements upon this point are certainly some-
what confused, that in the lower jaw there is a complete
dental fold throughout the entire diastema. In our stage,
which was but a little older, it was possible to distinguish
mere traces of a prolongation of the dental fold close behind
the rudiment of the incisor in the lower jaw, so that the state-
ment of MM. Chabry and Pouchet does not appear to me to
be a very credible one. On the other hand, we may with
confidence assume that in younger stages of the squirrel the
dental fold of the upper jaw will be found perfectly con-
tinuous in the diastema.
In addition to this we find in Sciwrus near the canals of
Stenson two enamel-organs lying one close behind the other,
of which the anterior is quite undeniably characterized as
such, while the posterior bears somewhat less distinct, but
nevertheless sufficiently definite, indications of its character.
It is remarkable that Chabry and Pouchet have figured one
of these rudiments, but, as we have seen, have interpreted it
quite incorrectly.
How are these structures to be regarded? ‘The anterior
enamel-organ can scarcely be regarded as a remnant of the
7.2 which is present in Lepus; for in this animal the cord
connecting ¢. 2 with the epithelium of the oral cavity is
attached tar in front of the orifices of Stenson’s canals; the
interval which is visible in fig. 14 (of the squirrel) between
this rudiment and that of 7.1 appears much too great for it
to be possible to assume that we are here dealing with two
tooth-rudiments following directly one after the other. It is
true that this question too will not be finally decided until we
have examined younger stages, since the possibility of a
secondary divergence of the two rudiments is also not excluded,
although it is not probable that this has taken place. More-
over it will be determined only by further investigations
whether a deposition of dentine still takes place in these
326 On the Development of the Tooth-Rudiments in Rodents.
rudiments or not, and how they disappear. Taking the
conditions as we find them in the stage under consideration,
we might regard these two enamel-organs as remnants of
rudiments of a posterior incisor and a canine tooth which
have been strangely preserved; their interpretation as remnants
of premolars is opposed by their position beside the orifices
of Stenson’s canals.
The reader may here once more be reminded that in Cavia
only very doubtful remnants of the dental fold were distin-
guishable in the diastema, while in Cricetus and Mus no trace
of a rudimentary embryonic dental germ was visible.
In conclusion, it may be also pointed out that our results
correspond to Fleischmann’s view, according to which the
degeneration of the tooth-rudiments has advanced further
in the lower than in the upper jaw. Apart from the doubtful
statements of MM. Chabry and Pouchet with regard to the
squirrel, no dental fold was found in the diastema in the
lower jaw; in the upper jaw, on the other hand, a structure
of this kind was distinguishable in a condition of greater or
less completeness in Lepus, Cavia, and Sciwrus, while in the
case of the last-mentioned form it is even provided with
enamel-organs. It also agrees very well with the general
views of Schlosser and Fleischmann, that the Lagomorpha
represent an especially old and conservative Rodent type, and
that after these the Sciuromorpha come next in order; that in
representatives of these groups it was possible to demonstrate
the most distinct and most widely extended embryonic remains
of tooth-rudiments which have disappeared; while in the
more highly modified Myomorpha, in so far as these were
investigated, the embryonic reduction was also more com-
plete. Lastly, stress may likewise be laid upon the fact that
the discovery of a dental fold in the diastema in the case of
Lepus and Sciurus is in conformity with the similar disco-
veries of other authors in the case of different Mammals with
incomplete dentitions. Thus the presence of a dental fold in
the diastema in the premaxilla of Ruminantia was not long
since finally determined by Mayo*. For the Edentata the
same was proved by Chabry and Pouchet, and it is well
known that in the case of the whalebone whales Geoffroy
St.-Hilaire has demonstrated the existence not only of the
dental fold, but even of entire embryonic teeth, which merely
fail to cut the gum and are absorbed within the jaw.
* Mayo, “The Superior Incisors and Canine Teeth of Sheep” (two
plates), ‘ Bulletin of the Museum of Comparative Zoology at Harvard
College,’ vol. vili. (Cambridge, 1886-188).
Mr. J. H. Hart on the Habits of a Species of Trigona. 327
At the end of my paper I will add a few words of thanks.
In the first place, I am most deeply indebted to Prof. G.
Born, Prosector to the Royal Anatomical School of Breslau
and Director of the Embryological Section. It was he who
first induced me to undertake these exceedingly interesting
and instructive studies, he initiated me into the always
difficult technique which work of this kind demands, and he
assisted me by word and deed wherever he could, shunning no
trouble and no expense. In return for his extraordinary
amiability and self-sacrificing care I trust I may be permitted
once more to express here my heartiest thanks to Prof. Born.
I am likewise under very special obligations to Prof. Hasse,
Director of the Royal Anatomical School of Breslau, for the
permission which he most readily accorded to me to work in
the Embryological Laboratory of the School.
Lastly, my best thanks are due to Dr. A. Fleischmann, of
Erlangen, who drew my attention to a series of highly
remarkable facts, introduced me to the paleontological litera-
ture of the subject, and furnished me with very valuable
statements and observations for my memoir.
LIII.— On the Habits of a Species of 'Trigona.
By J. H. Hart, F.L.S., Royal Botanic Gardens, Trinidad.
AMONG the several species of Trigona, or ‘‘ wild bees,” which
are common in Trinidad none is more interesting in its habits
than a peculiar small dark species which is often found in the
proximity of dwelling-houses, under slates, or in crevices of
the woodwork of out-buildings.
The specific name of the insect has not been ascertained,
but it has been found that it belongs to the genus Zrigona.
It was first observed in 1887 in the walls of a dwelling-house,
owing to its building a peculiar entrance-tube. This tube
was about ? of an inch in diameter, about 4 inches long, with
the entrance at the bottom, through a small hole in the
centre of the wax disk which closes it.
When the office of the Royal Botanic Gardens was under
repair quite recently the same species was discovered making
its home between two walls,
An attempt was made to house them, and a small box
prepared, with a small cut for entrance allowed, in a similar
manner to that usually seen in the common bee-hive, viz. at
the base. The bees took to the box and commenced work ;
328 Mr. J. H. Hart on the Habits of a Species of Trigona.
but after the first day they closed the lower entrance ver
completely with a sticky kind of wax, and adopted a ant |
crack in the upper portion of their box as the entrance. To
the inside of this crack they attached a tube similar to the one
first observed, but completely adapted to the new position, by
first building it along the crack and afterwards in a pendulous
manner downwards.
Wishing to have the insects more completely under obser-
vation, I built a small glass-sided box with sliding covers,
made an entrance for them in the top gable, and transferred
them thereto. They again took very kindly to their new
quarters and commenced work by rapidly sealing up every
crevice, making their home practically air-tight. To the
entrance they again attached the entrance-tube, which in this
case was brought from the entrance inwards, but built in the
same manner as the previous ones. It was, however, sup-
ported by wax stays, by which it was held at about ? inch
distant from the inside wall.
During the removal from their former home opportunity was
taken to examine the construction of their peculiar entrance-
tube, and it was found on making a section that it was con-
stricted in several places by disks, leaving only sufficient
space in the centre for the passing of one bee at a time; and,
if beaten back from the first, they have still the chance of
holding the inner ones in succession. These constrictions and
the sealing-up are evidently adopted by the insects as a means
of defence against their enemies. A further defensive
measure may be seen if the nest is examined after nightfall,
when it will be found that the orifice which admits of ingress
and egress during the day is sealed completely over, all but
imperceptible orifices being left in the closing sheet of wax,
we suppose for the admission of air. This safeguard is regu-
larly removed in the early morning near daybreak, and again
closed each night after nightfall. The honey-cells of this
bee are distinct from the breeding-cells, are ovate in shape,
over 2 inch in length, and somewhat pointed at the closed
apex; and the food- or pollen-cells are of the same form and
size, but situated at a different part of the hive. The honey-
cells are separated from the rest of the nest; they are affixed
closely together, and are somewhat irregular in size, but
firmly fixed to the side-wall of the hive in one layer only,
cones pointing upwards. ‘The pollen-cells are spread over
the floor-space, reminding one of sacks of grain in a granary.
The breeding-cells are not more than % inch in length, regu-
larly oval and in single tiers, held in position separate from
each other by small wax stays, which leaves each cell sepa-
On Two new “ Pocket-Mice” of the Genus Heteromys. 329
rate and distinct from its neighbour, though generally on the
same or nearly the same horizontal plane.
In our nest there are several layers one above another, the
main support being thin walls of wax built up the sides and
throughout the mass at intervals in a perpendicular manner,
and reaching to the top of the hive.
The queen has a very large body, very much larger in
comparison with the workers than that of the queen of the
common hive-bee, and very similar to some of the termites.
A small dipterous insect was observed in the nest when it was
first taken, but this disappeared after it was fully sealed.
This may prove to be the natural enemy’ of this insect.
Our bee has no sting, which renders it particularly easy to
handle and observe, and the want of which doubtlessly
occasions it to use such careful means of defence to secure its
home from the attacks of its enemies.
The honey is perfectly sweet and wholesome, very clear,
and of a nice flavour, but when squeezed out soon becomes
sour.
Another species of 7rigona is found in hollow or decayed
trees in our gardens, in large nests 2 to 3 feet in diameter,
formed of black, gummy, waxy, or resinous matter. This
species is very pugnacious, and attacks persons coming near
it, with a buzz and hum similar to that of the common honey-
bee ; but it is powerless to harm, as it has no sting. It,
however, fixes itself in the hair of the head or beard, and
produces a peculiar tickling feeling, which quickly induces a
sensation of fear in those who know the result of the attack
of an angry common hive-bee; and even when its cha-
racter is known the attack (almost unconsciously) causes the
intruder to retreat.
LIV.—Deseription of Two new “Pocket-Mice”’ of the Genus
Heteromys. By Ouprietp Tuomas.
WHILE attempting to determine a specimen belonging to the
genus Heteromys obtained by Dr. Audley Buller in Jalisco,
Mexico, | have found it necessary to make an examination of
all the species of the genus, and I find that they are readily
divisible into groups by the characters of the soles, whether
naked or hairy, with five pads or six. These characters do
not seem to have been known to Mr, Alston when, in his
330 Mr, O. Thomas on Two new
great work on the Mammals of Central America, he lumped
under one heading four species described by Gray and one by
Peters ; for two of the five, namely H. troratus and albo-
limbatus, are hairy-soled, with five pads, and the other three,
H. melanoleucus, longicaudatus, and adspersus, are naked-
soled and have six pads. To the first group there also belong
H. Alleni, the South-Texan species, and HH, Bulleri, the
Mexican one now described; while to the second, besides
those mentioned, there should be added the Trinidad species,
H. anomalus, Thomps., and Gray’s H. Desmarestianus.
Finally, the second new species now described, H. Salvint,
forms a third, annectant, group, with the hairy sole of the first
and the six pads of the second.
Heteromys Bulleri, sp. n.
Intermediate in size between H. Allent and H. trroratus.
General colour above the usual smoky grey, grizzled and
lightened with yellowish; sides with an indistinct yellowish
band along the junction of the back and belly colours ; lower
surface from mouth to anus pure white; ears dark, appa-
rently not white-edged ; hands and feet white ; arms to elbow
also white, but on the hind limb the dark colour of the rump
passes down to the heel and all round the ankle, but the inner
side of the upper leg is white.
Claws, especially the anterior ones, very long. Soles of
hind feet hairy for their posterior half, the hairs pale brownish ;
sole-pads five in number; the large posterior pad circular in
outline. Tail rather longer than the head and body, well-
haired throughout, clearly bicolor, brown all along its upper
surface, white on the sides and below.
Mamme 1—2=6.
Skull strong and stoutly built, differing mainly from that
of H. Allent in its much greater size, and from that of
H. trroratus by its differently shaped interparietal, which,
more like that of HH. Allenz, is comparatively rounded, with
its longitudinal 62°5 per cent. of its transverse diameter, and
has its anterior edge very convex forwards.
Teeth much worn in the only specimen, but in their shape
and proportions they do not seem to differ from those of
H. Allent, Equally worn teeth of H. irroratus are not avail-
able for comparison.
Dimensions of the type (an adult female in alcohol) :—
Head and body 114 millim.; tail 120; hind foot 28°5*
* Without claw.
“Pocket-Mice” of the Genus Heteromys. 331
ear, above head 10°5, from notch 15; heel to front of last
foot-pad 12°5.
Skull: basal length 29°5; greatest length 34:5; greatest
breadth 16°8; nasals, length 13°5; interorbital breadth 8°5 ;
interparietal, length 4°0, breadth 6°4; diastema 9°5; palate,
length 21; length of upper tooth-series 5°3.
Hab. La Laguna, Sierra de Juanacatlan, Jalisco, Mexico,
7000 feet. Coll. Dr. A. C. Buller, December 1892.
A male specimen of H, Allent from Brownsville, Texas,
the typical locality, has a caudal length of 110 and a basal
length of 26 millim., while a full-grown skull has a greatest
length of 30°3 millim., from which it will be seen that that
species is considerably smaller than /Z. Bulleri.
Heteromys Salvini, sp. n.
Heteromys longicaudatus, Alst. Biol. Cent.-Am., Mamm. p. 167, pl. xvii.
fig. 2 (1880) (mec Gray).
Size about equal to that of H. Buller’, but the feet are
decidedly shorter, being as short as in H. Allent. Fur spiny
as usual. General colour blackish, rather darker than in
most of the other species, grizzled with yellowish on the back.
Lower surface pure white, as usual. Outer sides of forearm
with a narrow slaty-grey edging, more conspicuous than in
H. Allent and Bulleri, less than in H. longicaudatus. Hind
foot very short, as short as in H. Allend; posterior half of
sole covered with short brownish hairs; pads six, as in the
larger naked-soled species, the two posterior pads very close
together in the median line of the foot. Tail thinly haired,
bicolor, but not so sharply and decidedly as usual, brown
above, whitish below.
Skull in size and shape very like that of H. Bulleri, but
the interparietal is much more extended transversely, while
only of about the same antero-posterior diameter; its longi-
tudinal percentage is therefore only 44 of its transverse.
Molars proportionally small and slender.
Measurements of the type (an adult male in skin) :—
Head and body (c.) 115 millim.; tail (imperfect at tip)
95+?; hind foot 26°5; heel to front of last foot-pad 11.
Skull: basal length 28; greatest length 33°6; greatest
breadth 15-2; nasals, length 13-1; interorbital breadth 7°6 ;
interparietal, length 4°5, breadth 10-2; diastema 8-9; palate,
length 19°2; length of upper tooth-series 4:4.
Hab. Dueiias, Guatemala. Coll. O. Salvin, July 31,
1873.
332 Mr. W. T. Calman on a new Pedalion.
The type specimen of H. Salvini is the very one referred
to and figured by Mr. Alston in the work above quoted.
It is with great pleasure that I connect with this interesting
annectant species the name of Mr. Osbert Salvin, its disco-
verer in 1873, and one of the editors of the great work in
which it is so beautifully figured.
A Heteromys from Costa Rica, in the Museum collection,
also appears to belong to H. Salvini; its hind feet agree
precisely in their structure with those of the type, but are
even shorter, measuring only 25 millim.
LV.—A new Pedalion. By W. T. CALMAN,
University College, Dundee.
THe remarkable rotifer Pedalion mirum, discovered by
Dr. Hudson in 1871, has been the source of much interest
and speculation to zoologists on account of its aberrant form
and supposed arthropod affinities. ‘Though it has since been
found in several localities, both in this country and on the
continent *, it has always been a great rarity, and no allied
Sar ul AAW,
PICS Oe me
se
Chip!
Yi %300
Pedalion fennicum, Levander.—d, dorsal appendage ; v, ventral
appendage; d./., dorso-lateral appendage ; v./., ventro-lateral appendage.
forms were known until Dr. K. M. Levander, of Helsingfors,
announced in the ‘ Zoologischer Anzeiger’ for October lastT
7 Be . . 4
* See Imhof, “Notiz ii. d. Vorkommen yon Pedalion mirum, Huds.,’
Zool. Anz. 13 Jahrg. pp. 609-611 (1890).
+ K. M. Levander, “ Eine neue Pedalion-Art,” Zool. Anz. xv. pp. 402-—
404 (1892) ; “ Zusatz z. meiner Mitth. iiber P. fennicum,” ibid. xvi. pp. 26,
27 (1893).
Mr. W. T. Calman on a new Pedalion. 333
the discovery of a second species of the genus in Finland.
By the kindness of Dr. Levander the Dundee Museum has
lately received some excellently preserved specimens of the
new Pedalion fennicum, from which the annexed figure is
taken. Dr. Levander will no doubt in due course figure the
species and publish his completed researches on its structure ;
but meanwhile this figure and the following notes may be
temporarily of interest.
Although bearing a great resemblance to P. mirum, the
new species differs from it in many details, of which the
following are the most conspicuous :—
(1) The two stylate processes at the posterior end of the body,
which form a conspicuous feature of P. mirum, are
here wanting.
(2) The dorso- and ventro-lateral appendages are nearly equal
in size. In P. mirum the dorsal is the shorter.
(3) The unpaired ventral appendage hardly reaches beyond
the posterior end of the body. In P. mirum it is
very much longer,
(4) The bristles of the ventral, and less markedly those of the
other appendages, for the most part arise in pairs
from a common base ina manner similar to that
shown in Dr. Schmarda’s figure of Hexarthra polyptera.
The length is about 23 millim. Only females were found,
many bearing one or two reddish eggs attached to the hinder
end of the body. In a few cases a larger number (about ten)
of small eggs was seen, which Dr. Levander conjectures with
great probability to have been male eggs.
With regard to the habitat, Dr. Levander says :—“ The
animal occurred in numbers in a pool of water about two yards
square by a foot deep, on a bare granite cliff on the little
islet of Léf6, about 12 km. south-west of Helsingfors, in
company with Daphnia pulex, Chydorus sphericus, and
Cyclops, sp. The little pool was about 10 yards from the sea-
shore and not a yard above the sea-level ; no vegetation; grey
detritus on the bottom ; the water moderately clear, sweet.”
ann. & Mag. N. Hist. Ser. 6. Vol. xi. 24
334 Bibliographical Notice.
BIBLIOGRAPHICAL NOTICE.
British New Guinea. By J. P. Toomson.
G. Philip and Son: London, 1892.
Tue illustrations and general style of this handsome volume are
highly creditable to the publishers ; while as regards the matter, we
can at least say that we are presented with a useful and readable
summary of the state of our knowledge of Papua, fairly up to date.
It is of course to be regretted that reports of Sir William Mac-
gregor’s explorations of the Bamo—an important affluent of the Fly
River—as well as of the country to the eastward, should not have
reached Mr. Thomson in time for insertion; but that is not the
fault of the compiler: for, to prevent misunderstanding, we may
say at once that the book is a mere compilation by one who, we
believe, has never visited New Guinea, but, from his abode in Bris-
bane has sung the achievements of his ‘ fellow-officer,” the Adminis-
trator of British New Guinea. Far be it from us to undervalue
the work done by Sir William Macgregor; but when we find that
the surveys made by his predecessors—distinguished naval officers
like Capt. Moresby, Commanders Pullen and Field, and others—
are treated without due sense of proportion, while every thing is
attributed to “the Administrator,” a certain feeling of antagonism is
aroused, and this is increased by the inflated tone adopted by the
author. We will quote the heading to the first chapter, for it seems
to furnish a keynote to the book :—* It appears to me to be a noble
employment to rescue from oblivion those who deserve to be eter-
nally remembered, and, by extending the reputation of others, to
advance at the same time our own.—Puinius minor.” If Sir
William is pleased with the manner in which his reputation has
been extended he must have a strong stomach.
An “ Historical Sketch ’—far too brief—brings us to the year
1888,when British sovereignty was proclaimed at Port Moresby; after
which the Administrator visited the Louisiade Archipelago, the
D’Entrecasteaux group, the southern coast of Papua as far as the
Dutch limit, and the northern shore up to the German _ protecto-
rate. These tours of inspection are not badly described, and the
account of the attainment of the highest peak in the Owen Stanley
Range (12,452 feet) affords some important details respecting the
configuration of the mountain mass; but we must add our protest
to those of others against the substitution of the name ‘ Mount
Victoria ” for that bestowed upon the peak many years ago. From
a scientific point of view the most valuable portion of the work is
the Appendix, which contains some succinct general notes on the
Flora of British New Guinea, by Baron von Miiller; a full report
(occupying 50 pp.) on the Insects, by Mr. Henry Tryon; an inter-
esting chapter on the Reptiles, by Mr. C. W. De Vis ; ; several
important vocabularies, &c. ‘To the above-named gentlemen and
some other “ collaborators” Mr. Thomson expresses his gratitude,
and without doubt it is well deserved.
a
Miscellaneous. 335
MISCELLANEOUS.
Classification of the Pelecypoda.—Emendatory Note.
By B. B. Woopwarp, F.G.S. &e.
By an oversight, when rearranging Fischer’s families of the
Pelecypoda to suit Pelseneer’s classification, the Tancrediide, which
by Fischer are placed between the Unicardiide and the Donacide,
were grouped with the former instead of with the latter. In the
table given, therefore, in the February number of the ‘ Annals,’
pp. 158, 159, the Tancrediide should stand as Fam. 42 instead of
49, and the Fams. there numbered 42 to 48 should be 43 to 49.
A Contribution to the Developmental Cycle of the Compound
Ascidians. By Jowan Hysort, of Christiania.
I.
(1) Like Metschnikoff * and Della Vallet+ I have always found
that the earliest rudiment of the bud of Botryllus is in the shape of
a two-layered vesicle, of which the inner layer proceeds from the
parietal layer of the peribranchial cavity, while the outer one is
derived from the ectoderm of the larva. I have never been able to
confirm the repeated assertions of Giardt and Herdman§ that in
the Botryllide a ‘“stolonial” gemmation occurs, in that the buds
are produced from the stolons of the mantle. Owing to the very
numerous transitional stages it is even possible to explain the
usually widely separated buds of colonies of Botryllus as having
arisen in consequence of a “pallial” gemmation. While, however,
Della Valle regards the inner vesicle of the bud-rudiments which
is formed in the course of this pallial gemmation as endodermal, I
myself feel bound, in accordance with the investigations of Kowa-
levsky |} and Van Beneden and Julin] and studies recently made
by Willey **, to consider this vesicle as ectodermal, since, according
* “ Entwicklungsgeschichtliche Beitrage,’ Mélanges Biologiques tirés
du Bull, de l’Acad. de St. Pétersbourg, t. vi.
+ “Sur le bourgeonnement des Ascidies composées,”’ Archives Itali-
ennes de Biologie, 1882.
¢ “Recherches sur les Ascidies composées,” Archives de Zoologie
expérimentale, 1872; also Br ea Rendus,’ 1891.
§ Report of the ‘ Challenger’ Expedition, vol. xiv.
|| ““ Weitere Studien iiber die Entwicklung der einfachen Ascidien,”
Archiv f. mikr. Anat. 7 Bd., 1871.
™ “Recherches sur la morphologie des Tuniciers,’ Archives de
Biologie, t. vi., 1886.
** “On the Development of the Hypophysis in the Ascidians,”
Zoologischer Anzeiger, xv. Jahrg. no. 400, pp. 352-334.
336 Miscellaneous.
to the investigations referred to, the peribranchial vesicle of the
larva proceeds from its ectoderm. The entire bud-rudiment of the
Botryllide consequently consists, as in the case of the Bryozoa, of
two epithelial lamellw of ectodermal origin, with mesodermal cells
scattered in between. It follows from this that the gemmation of
the Botryllide differs in this respect from that of the other Asci-
dians, such as Perophora, Clavelina, Amaroucium, Didemnium, and
Distaplia, since in all these cases, either directly or indirectly
(through the epicardium and the stolonial septum), the inner vesicle
is formed from the endoderm.
(2) From the inner vesicle are developed the intestinal tract, the
peribranchial vesicle, and the nervous system.
The process by which the peribranchial vesicle is formed is ushered
in by the upheaval of the ventral wall of the inner vesicle, com-
mencing from in front, into two folds which project into the
interior. By this means from the single inner vesicle there is
formed a median one, which communicates posteriorly by an opening
on each side with two lateral ones, the peribranchial vesicles. Now
Della Valle asserts that three separate vesicles arise and that the
two lateral ones form by fusion the single peribranchial cavity. I
have been unable to confirm the occurrence of this secondary fusion ;
on the contrary, the peribranchial cavity is formed from the
beginning as a saddle-shaped double vesicle, which becomes con-
stricted off dorsally from the median vesicle.
The earliest rudiment of the neryous system takes the shape of
a nearly hemispherical evagination, situated at about the centre of
the dorsal wall of the median vesicle. This evagination soon grows
out into a tube, which is directed anteriorly and ends blindly in
front. The closed anterior end of the cecum fuses afresh with the
median vesicle ; an aperture is formed here, and we now find upon
the dorsal side a tube with both ends opening into the median
vesicle. The anterior opening lies not far from the oral aperture,
which now arises as a fusion between the ectoderm and the enteron ;
the posterior opening of the tube communicates with the dorsal
portion, which connects the two lateral halves of the saddle-shaped
peribranchial vesicle. While the latter, as has been described,
becomes constricted off from the intestine, this posterior opening
also closes up, and we simply find a dorsal tube communicating in
front with the intestine.
(3) At this stage we already observe a thickening of the ventral
side of the tube, which increases considerably during the subsequent
development. Moreover it becomes more and more constricted off
from the dorsal section of the tube, while delicate fibres are differen-
tiated in its interior.
This thickening develops into the permanent ganglion, while the
tube, which gradually becomes longer and thinner, persists as the
hypophysis.
I would here point out that Kowalevsky *, in the case of Didem-
* “Ueber die Knospung der Ascidien,” Archiy f. mikr. Anatomie.
Miscellaneous. 337
nium styliferum, Perophora*, and Amaroucium, likewise derives the
nervous system from the inner vesicle, although in a somewhat
different fashion. Consequently both in the forms alluded to and
in Botryllus the intestinal tract, the peribranchial cavity, and the
nervous system proceed from the inner vesicle of the two-layered
bud-rudiment.
While, however, in the first-mentioned forms this inner vesicle is
endodermal, in Botryllus it is, as has been shown, of an ectodermal
nature.
As is evident from what has been stated above, these results of
mine are diametrically opposed to the conceptions of Seeliger tT and
Salensky. In the case of the buds of Clavelina and Pyrosomat the
nervous system is derived by Seeliger from immigrant mesoderm
cells, while in the latter form it is stated by Salensky § to arise as a
thickening of the ‘‘ outer vesicle.” With the last-mentioned author,
however, I am so far in agreement in that I also have found a
common origin for the hypophysis and the ganglion.
rr.
The mode of formation of the ganglion in the buds of Botryllus
led me also to study the development of the same organ in the
lJarve of compound Ascidians. It will be seen from what follows
that owing to these investigations I have discovered several parallels
between the development of buds and larve.
In the cerebral vesicle of the larva of Distaplia magnilarva there
appears at an early stage a difference between the left and right
sides. Nearly in the middle of the right side of the cerebral vesicle
there arises an evagination, which soon exhibits the most manifold
differentiations, and from which in later development proceeds the
larval brain, which has been so exhaustively described by Van
Beneden and Julin |}.
The left wall, however, in the meantime maintains its indifferent
cellular character: at first consisting of a single layer, it gradually
increases in thickness.
In front of the described evagination of the right wall of the
cerebral vesicle the latter becomes tubular and joined to the intes-
tine. This anterior portion possesses cells of the same constitution
as that of those of the left wall.
* Kowalevsky, “Sur le bourgeonnement du Perophora Listeri”’ (transl.
Giard), Rey. d. Sc. nat., Sept. 1874.
+ “Ejibildung und Knospung yon Clavelina lepadiformis,” Sitzgsber.
d. kais.-kgl. Akad. d. wiss. Wien, 1882.
t “Zur Entwicklungsgeschichte der Pyrosomen,” Jenaische Zeitschrift,
23 Bd.
§ “ Beitrige zur Embryonalentwicklung der Pyrosomen,” Zoologische
Jahrbiicher, 1891.
| “Le systéme nerveux central des Ascidies adultes et ses rapports
ayec celui des larves urodéles,” Archives de Biologie, t. v., 1884.
338 Miscellaneous.
During the development which succeeds this stage the following
important changes set in :—
(1) The anterior portion of the cerebral vesicle, which was joined
to the intestine, acquires an opening into the latter, and thus forms
a communication between intestine and cerebral vesicle which
persists throughout the whole of the larval period.
(2) The multilaminate left wall of the cerebral vesicle produces
nearly at its middle a thickening, which gradually becomes con-
stricted off, and goes to form the permanent ganglion.
The left wall itself, which hitherto formed an undivided mass in
conjunction with the permanent ganglion, after the latter has
become constricted off assumes an epithelial character. In front
it is directly continuous with the anterior portion of the cerebral
vesicle. This portion also becomes epithelial, elongates, and repre-
sents the well-known ciliated pit.
(3) Now while the larval brain (formed by the evagination of
the right wall of the cerebral vesicle) becomes constricted off and
disappears, the epithelium-like left wall of the original cerebral
vesicle bends round into a tube, which represents an immediate
prolongation of the ciliated pit just described, and in which we
recognize the hypophysis.
(1) In the development of the larva, as in that of the bud, we
consequently find that the hypophysis and the persisting ganglion
have a common origin.
(2) The common rudiment is tubular in both modes of develop-
ment, and in both cases the ganglion is formed as a thickening of
the tube.
(3) The larval cerebral cavity opens, as described by Kowalevsky *
and disputed by Van Beneden and Julin +, through the hypophysis
into the intestine.
(4) In the adult animals, produced from the larva, there persists
from the lumen of the larval cerebral vesicle only the lumen of the
hypophysis.
The above investigations were carried out in the Zoological
Institute in Munich and in the Zoological Station at Naples. It is
a pleasing duty to express to my revered teacher, Prof. Hertwig,
and to Dr. Dohrn my best thanks for all the kindness shown to me.
—ZLoologischer Anzeiger, xv. Jahrg. no, 400 (Sept. 12, 1892),
pp. 328-332.
Munich, June 29, 1892.
* “Weitere Studien iiber die Entwicklung der einfachen Ascidien,”
Archiy f. mikr, Anat. 7 Bd., 1871,
+ Op. cit.
Miscellaneous, 339
The Development of the Intestinal Gregarines of Marine Worms.
By M. Lovts Lferr.
The Gregarines with a single segment, which are very frequently
met with living freely in the digestive tract of a large number of
Annelids, have hitherto been considered to be Monocystidea.
The study of the development of Doliocystis nereidis, parasitic in
the intestine of Nereis cultrifera, and of Doliocystis polydore from
the intestine of Polydora Agassiz, shows that these Gregarines are
in reality Dicystidea, exhibiting in their earliest youth the intra-
cellular stage, followed by a stage of budding which gives rise to
the Gregarine proper. During the budding stage the Gregarine
always exhibits two segments—the intra-cellular segment, or epi-
merite, and the extra-cellular segment, in which the nucleus is
contained, It is therefore at this moment only that the Gregarine
appears as atrue Dicystid; but this condition does not last long.
At a very early period the young individuals drop their epimerite
and become free in the intestine, when they exhibit all the cha-
racters of true Monocystis, with which they have hitherto been
confounded.
In order to study the development of Doltocystis nereidis it is
necessary to examine with much care the elements of the epithelial
lining of the digestive tract of the Merets, especially in its anterior
third. Individuals will then be met with which are extremely
young and still in the Coccidiid stage, th::t is to say, in the state of
a simple spherical nucleated mass, situated between the nucleus of
the cell and the surface. In the subsequent stages the primitive
Coccidiid has budded forth a segment, which makes its way into
the lumen of the digestive tract, and which is destined to form the
Gregarine proper. Finally, in a still more advanced stage the
Gregarine is definitively constituted; the extra-cellular bud has
considerably increased in size, and a layer of transverse muscular
fibrils has already become differentiated, while the intra-cellular
portion, on the contrary, is reduced to the condition of a simple
little knob (epimerite). Conditions such as this are met with pretty
frequently in the preparation, and we may even observe free indi-
viduals of which the epimerites are still capped with the shrivelled
epithelial cells ; this is the Cephalin stage, which is soon concluded
by the falling off of the epimerite, to give place to the stage of the
Sporadin. The Gregarine is henceforth free in the intestine, in the
form of an ovoid or elongated Monocystis, more or less drawn out
into a point at one of the poles.
The development of Doliocystis polydore, a new species which I
have met with in the Bay of Marseilles, takes place in precisely the
same manner ; only the epimerite is developed to a greater extent
than in the preceding species ; moreover it persists for a long time,
the result of which is that Cephalins of this species are commonly
encountered. This epimerite is in the form of an inverted frustum
of a cone, and it is directly continuous with the anterior extremity
340 Miscellaneous.
of the second segment, which is elongated in the shape of a neck.
At the moment of the falling-off of the epimerite, which is easily
observed under the microscope, a broad wound is produced by which
granulations of the endocyte make their escape ; but this promptly
cicatrizes, and the Gregarine soon no longer exhibits any traces of
its first segment—it has passed into the Monocystis stage.
The development of these two species is thus identical with that
of the genus Schneideria, which we may justly consider as the most
perfect type of the Dicystid Gregarines. The only difference con-
sists in the fact that the epimerite always remains very simple and
rudimentary in Doliocystis, while in Schneideria it attains a certain
degree of complication. Moreover it is interesting to note that, in
a general way, the epimerites of the marine Gregarines never
exhibit so high a degree of differentiation as do those of the majority
of the terrestrial forms; they all belong, at least in the species
which I have studied hitherto, to the group of the regular simple
eprmerites.
The free stage is succeeded in Doliocystis by encystment and
sporulation, which take place normally as in the other Polycystidea.
The cysts of Doliocystis nereidis, which I have succeeded in culti-
vating in spite of their extremely small size, give rise when mature,
by means of simple rupture, to oval corpusculated spores, measuring
7p in their long axis by 5» in their short axis. They exhibit a
reneanle thickening of the wall at one of the poles, which is a
very important character in my opinion, since it is also common to
the spores of the genus Schneideria.
Thus we see that the Gregarines with a single segment from the
digestive tract of marine worms exhibit the same mode of develop-
ment and the same form of spores as do the typical Dicystidea; it
is therefore in this group that they must in future be placed, and it
is necessary to distinguish them henceforth from Monocystis proper,
the deyelopment of which is entirely different, and which, more-
over, inhabits almost exclusively the general body-cavity.
While retaining the generic name of Monocystis for these latter
forms, I propose to unite the former, that is to say the pseudo-
Monocystis of the digestive tract of worms, under the common
generic name of Doliocystis; as regards their specific name, this will
be very well indicated by the name of the host which harbours them.
We shall have Doliocystis nereidis for the Gregarine of Nereis, Dolio-
cystis polydore for that of Polydora, &c.
Thus the confusion resulting from the union under the same
name of genera essentially distinct will cease, and the group
Dicystidea will therefore comprise two important genera :—
(1) The genus Schneideria, peculiar to the digestive tract of
terrestrial Arthropods ;
(2) The genus Doliocystis, peculiar to the digestive tract of
marine worms.— Comptes Rendus, t. cxvi. no. 5 (January 30, 1893),
pp. 204-206.
yl
May. Nat. Hist. 8.6. Vol. 1M. PLATT
t
Anmn..c.
Mintern Bros. hth.
Ann. Mag. Nat List. S06 Vol 171 PLXIV.
Werntern Bros . hth.
FT TAAARTOVC RIITNA TARTAR. ITSO
Anid May. Nat. Mist, (7am 5) a, 0 a ia baa, 41
Ann .& Mayg.. Nat. Hist. S.6.Vol.XI. PU. XVI.
R.1LP. del Nintern Bros -hth.
THE ANNALS
AND
MAGAZINE OF NATURAL HISTORY.
{SIXTH SERIES. ]
No. 65. MAY 1893.
LVI.—On some new Bornean Mammalia.
By OLpFieLp THOMAS.
Tue British Museum has recently received from Mr. Alfred
Everett a further considerable consignment of small mammals
from Sarawak and North Borneo, and in working them out
in conjunction with others received of late years from the
same region the following species prove to need description :—
Cynopterus maculatus, sp. n.
Size very small, the smallest of the genus. Muzzle with
its central groove almost obsolete. Ears small, oval, laid
forward they barely reach to the posterior corner of the eye;
their tip rounded ; their outer base with a distinct rounded
lobe, as in C. brachysoma; their colour black, but with a
distinct white or yellowish spot on the centre of their inner
margin. Thumb involved in membrane for half its length.
Calcar and interfemoral membrane fairly well developed.
Tail wholly absent. Palate with about twelve simple ridges,
the anterior ones much curved, the posterior two or three
directly transverse.
Fur above thinly covering the membranes near the body
and the base of the forearm, but not extending on the tibia,
although the backs of the feet are fairly well clothed.
Colour of fur above sooty grey, paler, with a tinge of
Ann. & Mag. N. Hist. Ser. 6. Vol. xi. 25
342 Mr. O. Thomas on some
rufous, on the back, darker, almost black, on the head. Below
dull uniform grey, paler than the back.
Ground-colour of ears and membranes black, but on all the
finger-joints and on the membranes between them there are
numerous prominent yellow spots, forming a most obvious
and peculiar characteristic. As already noted, there is also
one of these spots on the anterior margin of each ear.
Skull with the usual short muzzle of Cynopterus and
showing no tendency towards that of Xantharpyia.
Teeth different in number to those of any other known
species, viz.:—I. 3, C. 7, P. 3, M. x 2=380.
Inner upper incisors nearly twice the length of the outer
ones ; canines short, not much exceeding in length the middle
premolar. Anterior premolar and last molar both very
minute, about equal in section to one of the inner upper
incisors. Lower anterior premolar and posterior molar also
very small, equal to each other, and about two or three times
the size of the corresponding upper teeth.
Dimensions of the type (an adult male in spirit) :—
Forearm 41 millim.
Head and body 65; ear from notch 10; thumb (including
claw) 16°6; index finger (including claw) 31°5; metacarpal
of third finger 80; lower leg 14°53; calcar 4:4; depth of
interfemoral in centre 4.
Skull: basal length 20°6; greatest length 23; greatest
breadth 15:7; interorbital breadth 5:2; palate length 11:9.
Hab, Sarawak. Coll, A. Everett, June 1892.
This most remarkable little species, of which six specimens
were obtained, is distinguished from all others by its extra
posterior molars, two instead of four lower incisors, and by
the characteristic spotting of the wing-membranes and joints.
In other respects, in colour, size, and the absence of a tail,
it appears to agree with C. melanocephalus, Temm., which
Dobson erroneously placed with the tailed species of the
genus, and to which it is probably most nearly allied. That
species, a native of Java, is as yet only represented by the
typical examples in the Leyden Museum, and these are so
faded that little stress can be laid on their coloration. At
the same time Dr. Jentink has kindly reexamined them at
my request, and informs me that all the three skulls in that
museum have four lower incisors as usual, that none of them
have any trace of the minute extra molars in the upper jaw,
and that, so far as he can tell, the wing-membranes are quite
without the white spots present in OC. maculatus.
new Bornean Mammalia. 343
Tupaia ferruginea longipes, subsp. n.
Hind feet conspicuously longer than in the typical
Sumatran form. General colour less ferruginous above, but
more so below; the shoulder-streak also, instead of being
yellowish or whitish, is rich rufous. Upperside of tail con-
colorous with the back, instead of being markedly greyer.
Skull and teeth apparently as in the typical subspecies.
Dimensions of the type (a skin) :—
Head and body 192 millim.; tail 190; hind foot 48-5.
Hab. N.W. Borneo. Coll. H. Low, 1876.
This long-footed form of J. ferruginea has been long
known to me, and, as all the further specimens from the same
region fully agree with Mr. Low’s example, I take the
present opportunity of describing it, and am only doubtful
whether it should be described as a species or subspecies.
The specimen which, on account of its having been the first
collected, I have selected as the type, has no exact locality,
but others since sent home by Messrs. Hose and Everett show
the same elongate feet and define more precisely the range of
the subspecies. Thus, there is a skin from Spitang, opposite
Labuan (Zverett), with a hind foot 48 millim. in length, one
from Baram (Hose) of exactly the same length, and, finally,
Mr. Everett has sent a full-grown male in spirit from Sarawak
which presents the tollowing dimensions :—Head and body
215 millim., tail 210, hind foot 52°5. The type specimen of
T. ferruginea typica has a hind foot of only 42 millim., and
this length does not appear to be exceeded by any other of
our Sumatran or Malaccan examples.
It seems probable that this form represents 7. ferruginea
throughout Sarawak and Northern Borneo, and perhaps over
the whole island, as I have seen no Bornean specimens of
the usual Sumatran type.
Crocidura (Pachyura) Hoset, sp. n.
Size very small, scarcely exceeding that of the minute
Indian species C. Perottett, Hodgsoni, &c., and belonging
evidently to the same group, which has not hitherto been
recorded from the Malay region. Fur close, crisp, and
velvety. Colour deep smoky grey, finely grizzled with
whitish ; belly rather paler ; ears, feet, and tail dark brown.
Tail very short and slender, closely haired, with a few longish
hairs as usual scattered among the shorter ones.
Anterior part of first incisor short and thick ; posterior cusp
about equal in size to one of the two posterior unicuspids ;
25*
344 Mr. O. Thomas on some
first unicuspid large, its tip reaching the same level as the
first incisor and the tip of the large premolar; second and
third unicuspids about equal in size, the second slightly longer
but slenderer than the third; fourth well developed, its tip
just visible externally, and about equal in height to the basal
process of the large premolar and about half the height of the
third. Anterior lower incisor long and slender, without
denticulations; second lower unicuspid just exceeding in
height the anterior cusp of the large premolar.
Dimensions of the type (an adult skin) :—
Head and body (probably stretched) 59 millim.; tail 20;
hind foot 8°6.
Front of &! to back of ™% 6:4; breadth of palate outside
m.1 4+], inside ™1 1°6; distance from tip of &! to tip of the
large premolar 3:1; length of lower tooth-row 5°8.
Hab. Bakong River, Baram, Kast Sarawak. Coll. Charles
Hose, Dec. 1891.
This species is larger than any of the Indian pigmy shrews,
while it is far smaller than any other Oriental Pachyura as yet
described. Its short tail seems also to distinguish it from all
its allies.
Mr. Everett has also collected a minute shrew in Sarawak,
but it proves to be a true Croctdura (s. s.), and, judging
merely from the description, I do not at present feel justified
in distinguishing it from the Javan C. monticola, Pet.
Chiropodomys major, sp. 0.
Colour and proportions very much as in Oh. glirotdes, but
size conspicuously greater, especially so far as the skull is
concerned (see dimensions below). Upper surface fawn, the
bases of the hairs slate-coloured ; whole of lower surface pure
white. Ears large, naked. Tail long, hairy, and pencilled
as usual; uniformly brown above and below.
Skull with a flatter profile than in Ch, glirotdes ; anterior
palatine foramina very short, ending half their own length in
front of the molars. Molars broad and rounded.
Dimensions of the type (an adult female, stuffed) :—
Head and body 100 millim. ; tail 109; hind foot 21:5;
ear from notch 14.
Skull: upper length 30; breadth of brain-ease 14; length
of nasals 10; interorbital breadth 5:3; interparietal, length
5°2, breadth 10°4; anterior zygoma-root 3°1; palate, length
15:2, breadth outside ™-! 5°8, inside 1 3:4; diastema 8°5;
anterior palatine foramina 3°8; length of upper molar
series 4°4,
new Bornean Mammalia. 345
Hab, Sadong, Sarawak. Coll. A. R. Wallace, Esq.
A second specimen from the same place agrees with the
type in every respect.
Chiropodomys pusillus, sp. n.
Size smaller than in CA, gliroides. Ears and feet deci-
dedly smaller and tail shorter than in that species. Fur
crisp, close and velvety. General colour tawny fawn, head
and centre of back darker, sides paler, outer sides of arms
and legs like back, but the wrists and ankles greyish, a
colour which also extends upon the metatarsus; fingers and
toes white; under surface from chin to anus pure white; no
darker markings on face ; ears small, evenly oval, practically
naked. ‘Tail but little longer than the head and body com-
bined, uniformly brown above and below, its terminal tuft of
hairs of about the same thickness, but less extended and
commencing more abruptly than in the allied species.
Skull smaller and rather more delicately built than in the
other species and showing even more markedly the roundness,
simulating immaturity, characteristic of the genus; supra-
orbital bead but slightly developed; anterior palatine fora-
mina very short. Molars small, their structure as usual.
Measurements of the type (skin) :—
Head and body 76 millim.; tail 81; hind foot 15°8; heel
to front of last foot-pad 7-2; ear from notch 11°5.
Skull: upper length 22°2; breadth of brain-case 11°6;
nasals, length 7:2; interorbital breadth 4:2; interparietal,
length 4°2, breadth 9°2; anterior zygoma-root 2°1; diastema
6°2; anterior palatine foramina 2°7; combined lengths of ™1
and ™-? (™-3 is unfortunately lost) 2°5; length of lower molar
series 31.
Hab, Mount Kina Balu, 1000 feet.
This species is founded on the specimen referred by me in
1889* to Ch. gliroides, a reference mainly induced by the
peculiar rounded and immature appearance of the skull; but
this appearance has since proved to bea characteristic of the
whole genus, and an examination of the teeth shows that
the specimen is after all fairly adult. This being the case,
the marked differences in the dimensions of the ears, feet, and
tail will readily distinguish it from the older known species.
* P. Z.S. 1889, p. 235.
346 On some new Bornean Mammalia.
Mus Margarete, sp. n.
Size, form, and general appearance very much as in Van-
deleuria oleracea. Whiskers numerous and _ prominent,
black. Ears small, oval, practically naked. Colour, so far
as can be made out in a specimen in spirit, deep rufous
chestnut, mixed on the back with the grey of the bases of the
hairs, but clearing on the sides, where it seems to form a
rufous lateral band. Chin, chest, and belly white. Hands
and feet also white, but the metapodials with darker
median patches. Thumbs prominent, opposable, with a large
nail; claws of fingers short and curved. Hallux also oppo-
sable, its claw reduced to a minute conical point, not sur-
passing in length the pad below it; other toes all with their
claws very short and curved, and surpassed in length by the
prominent terminal pads. Soles naked, with six large rounded
pads. Tail very long, slender, finely haired, almost naked ;
scales very small, averaging about seventeen to the centi-
metre, their colour a sort of pale greenish grey, the same
above as below. Mamme 1—2=6. Palate-ridges 3—5.
Skull with a very peculiar and noteworthy resemblance to
that of Chiropodomys, agreeing with that of Ch. gliroides so
closely that it is not until a close examination is made that
the differences become apparent. General proportions short
and broad, the brain-case especially broad and rounded.
Interorbital space flat, its edges sharply beaded, converging
strongly forwards. Interparietal very large, its anterior edge
nearly directly transverse, its posterior one boldly bowed out
backwards. Anteorbital plate of zygoma not projected
forwards at all. Anterior palatine foramina very small and
narrow. Molars of very much the rounded shape and
general proportions of those of Chiropodomys, but their strue-
ture as in Mus.
Dimensions of the type (an adult female in alcohol) :—
Head and body 76 millim. ; tail 144; hind foot 19°7; ear,
above head 11, from notch 13; length of head 28.
Skull: basal length 21°5; upper length 25°5; zygomatic
breadth 13; breadth of brain-case 12°2; nasals, length 7°6;
interorbital breadth 4°1; interparietal, length 4:2, breadth 9;
anterior zygoma-root 23; palate, length 13; diastema 6°8 ;
anterior palatine foramina 3°6; length of upper molar
series 3°7.
Hab, Penrisen Hills, Sarawak. Coll. A. Everett, June
1892.
This very remarkable species will no doubt need in the
future the erection of a special genus or subgenus for its
On Land-Shells from the Philippine Islands. 347
reception, and will perhaps prove to be congeneric with Mus
chiropus, lately described by me *, which also has the molar
teeth of Mus combined with an opposable hallux and a general
Chiropodomys- or Vandeleuria-like form. Pending further
investigation, however, I prefer to do as I did then, and leave
the new form in the genus Mus.
I have taken the liberty of naming this beautiful little
species, which looks as if it would make a most enchanting
pet, in honour of Her Highness the Ranee of Sarawak, a lady
whose interest in the zoology of that country is scarcely
inferior to that of her husband the Rajah.
LVII.—On a Small Collection of Land-Shells from Palawan
and Balabac, Philippine Islands, By EpGar A. SMITH.
[Plate XVIIL}
Tue British Museum has recently obtained a small collection
of land-shells, collected by Mr. A. Everett in Palawan and
Balabac. Of the thirteen species it contains five appear to
be new. ‘This large proportion of undescribed species is not
altogether surprising, as neither of these islands has been
thoroughly searched for Mollusca. A list of the known
species was given by the Rev. A. H. Cooke in the Proc. Zool.
Soc. 1892, pp. 461-463. It comprises the species quoted in
the various papers by Hidalgo in the ‘ Journal de Conchylio-
logie,’ 1887, pp. 36, 37-58, 93-192, 1888, pp. 30-97, and
in Dohrn’s “ Beitrag zur Conchylienfauna des philippinischen
Insel Palawan ” f.
Including the new Lagochilus similis here described four-
teen species of land-mollusks have now been recorded from
Balabac. The Amphidromus quadrasi, although appearing
in Mr. Cooke’s list, was not previously known from Balabac,
but only from a small adjacent island called ‘ Isla de Can-
daramanes ”’ (Hidalgo). Another species trom this island
quoted by Hidalgo¢ and not in Mr. Cooke’s list of the
Balabac shells is Corasia zamboange, Uombron and
Jacquinot.
* Ann. Mus. Genoy. (2) x. p. 884 (1891), and p. 935 (1892), pl. xi.
figs. 4-7.
“+ Nachrichtsblatt deutsch. mal. Gesell. 1889, pp. 53-63.
t Mem. Acad. Madrid, 1890, vol. xiv. p. 151.
348 Mr. E. A. Smith on Land- Shells
Thirty species are enumerated from Palawan or Paragua,
and if Tauptecka boholensis, Pfr., be distinct from L. cebuensis,
Millendorff, it will add one more to the total. It is quoted
from Palawan by Hidalgo*, but not in Cooke’s paper.
The most interesting addition to the fauna occurring in the
present collection is the new species of Lagochilus, a genus
hitherto unknown from Balabac or Palawan. It occurs,
however, sparingly in other ad of the Philippine group and
also to the south in Borneo &e.
1. Macrochlamys pseustes. (Pl. XVIII. figs. 1-3.)
Testa angustissime perforata, depressa, orbicularis, superne fusca,
infra pallidior, nitidissima, subpellucida; anfractus 6, regulariter et
lente crescentes, convexi, sutura profunda discreti, lineis incrementi
supra distinctioribus quam infra sculpti, ultimus ad peripheriam
acute rotundatus; spira brevissima, ad apicem obtusa ; apertura
oblique lunata ; perist. tenue, margine columellari leviter incras-
sato, supra umbilicum angustissimum breviter reflexo.
Diam. maj. 21 millim., min. 19; alt. 10.
Hab. Palawan.
This species is remarkable for the convex whorls and the .
deep suture. It is of a rather rich brown colour on the upper
surface, but considerably paler beneath. The lines of growth
are rather strongly marked above, so that they present an
almost subplicate appearance. ‘The deep suture is bordered
with a very narrow dark brown line, which, however, is not
noticeable without a lens.
The genus Macrochlamys has not hitherto been recorded
from Palawan, but there are several allied species found in
North Borneo which have been placed in the section Hverettia
by Godwin-Austen (Proc. Zool. Soc. 1891, pp. 33-36).
2. Lamprocystis chlororhaphe. (Pl. XVIII. figs. 4-6.)
Testa parva, angustissime perforata, depresse conoidea, tenuis,
pellucida, nitida, cornea; anfractus 5-6, lente accrescentes, con-
vexiusculi, infra suturam anguste albido-marginati, vix striati,
ultimus ad peripheriam acute rotundatus ; spira breviter conoidea,
ad apicem obtusa ; apertura oblique lunata; peristoma simplex,
margine columellari leviter incrassato, sed vix reflexo.
Diam. maj. 6 millim., min. 5}; alt. 4.
Hab, Palawan.
The minute perforation and the palish line at the suture
© 50) pS,
Srom the Philippine Islands. 349
are the most noticeable features of this little species. It is
about the same size as L. lucidella, Pfr., but differs from it
in having the body-whorl narrower, the columella less reflexed
over the perforation, and the suture narrowly margined with
white.
3. Trochonanina paraguensis. (Pl. XVIII. figs. 7-9.)
Testa imperforata, depresse conoidea, acute carinata, unicolor,
cornea, vel interdum utrinque carinam anguste fusco zonata ;
anfractus 7—8, superne vix convexiusculi, lente et regulariter
crescentes, incrementi lineis oblique arcuatis striisque spiralibus
paucis sculpti, ultimus haud descendens, acute carinatus, infra
convexus, haud spiraliter striatus ; carina supra et infra vix com-
pressa ; apertura angusta; peristoma album, magnum, margine
supero leviter inecrassato et subexpanso, basali fortius incrassato.
Diam. maj. 183 millim., min. 17; alt. 8.
Hab, Palawan.
This species is closely allied to 7. labuanensis, Pfeiffer. It
may be distinguished by the spire being rather less elevated,
the keel not so acute and not compressed above or below; the
umbilical region is not so impressed, and the peristome, espe-
cially the basal margin, is distinctly thickened. This species
is much more distinct from 7. labuanensis than the latter is
from YZ. conicoides, Metcalfe, which indeed are considered
synonymous by Godwin-Austen. 7. conicoides, however,
has a more elevated and conical spire than labuanensis, and
the coil of the whorls is different, so that in the latter the last
whorl seen from above is conspicuously narrower than in
conicotdes.
T. sylvana, Dohrn and Semper, has a higher spire than
paraguensis, a sharper and more compressed keel, and has not
the basal lip of the aperture thickened. It is all but identical
with labuanensis.
4, Trochomorpha Metcalfei, Pfr.
Hab. Palawan (Hidalgo, Everett).
This species is very variable and is met with on several of
the islands of the Philippine group. With Hidalgo (2. c.
p- 112) I agree in considering 7. boholensis of Semper as a
variety.
5. Hemiplecta densa, var.
The variety of this species from Palawan, namely that
described by Pfeiffer as Helix Schumacheriana, has already
350 Mr. E. A. Smith on Land- Shells
been quoted by Dohrn (Nachrichtsbl. deutsch. mal. Gesell.
1889, p. 57).
Two forms of it were obtained by Mr. Everett, also in
Palawan, one in which the last whorl is much more acutely
keeled and more finely sculptured beneath than in the other.
Considerable variation in these respects occurs also in the
Bornean examples in the British Museum. ‘The very acute
specimens from Palawan are very like 1. ceymatium of Benson
from Penang. Two specimens of the latter, marked in
Pfeiffer’s handwriting from Pulo Zancavi, Malacca, appear
to me quite inseparable from the present species, but two
others from Penang, presented by Mr. De Burgh, although
very like the var. Schumacheriana, appear to have an addi-
tional whorl and to be less rapidly coiled, so that the last is
narrower than in the Bornean shell.
6. Helix (Hadra) Traillit, Pfeiffer.
The typical form of this species is more conical than the
specimens obtained at Palawan by Mr. Everett, and much
more coarsely striated. The transverse bands also are not
nearly so dark-coloured as in the specimens just received,
Most of the latter are covered with a thin bright yellowish
epidermis, with which the dark zones contrast strongly.
They vary considerably in the elevation of the spire, but
none are so depressed as HH. monochroa or its variety pala-
wanica. Occasionally the ground colour above is of a dark
reddish tint. The bands on the body-whorl are usually four
in number, namely a narrow one at the suture, a broader one
at the periphery, a still broader one below it, and a faint one
above it. ‘The last is sometimes absent.
7. Cochlostyla satyrus, Broderip.
The series sent home by Mr. Everett from Palawan con-
firms the opinion of Dohrn and Cooke that C. palawanensis,
Ptr., C. cinerosa, Ptr. C. librosa, Pfr., and C. Graellst of
Hidalgo are mere varieties.
Mr. Everett also obtained this species at Banguey Island,
situated between Borneo and Palawan.
8. Amphidromus quadrasi, Hidalgo.
(Pl. XVIII. figs. 10-13.)
Amphidromus quadrasi, Hidalgo, Journ. de Conch. 1887, p. 36, pl. ii.
fig. 2. :
Hab. Balabac (£verett) ; island of Candaramanes or Cara-
mandanes, near Balabac (//idalyo).
From the Philippine Islands, 351
Numerous specimens of the typical form of this species
were collected by Mr. Everett, besides a number of other forms
which probably merely indicate colour-variation. They may
be thus described :—
a. Green, with a deep rose-coloured band at the suture and
around the base of the lilac columella (typical form).
(Fig. 10.)
b. Yellow, with similar band at suture and base; colu-
mella lilac.
e. Yellow, without sutural band, with a broad green band
on the back of the body-whorl behind the lip; colu-
mella pale pink. (Fig. 11.)
d. Uniformly yellow, faintly streaked with green; colu-
mella white.
e. Yellow, with the spire ornamented with oblique, wavy,
rich brown lines, forming blotches above the suture,
and with a similar green band as in the preceding
form; columella pale pink. (Fig. 13.)
f. Very dark chestnut, almost black, but pale at the suture,
with oblique wavy white streaks, which are inter-
rupted at the middle of the body-whorl. A yellowish
zone occurs below the periphery and a pinkish one
around the base of the columella. The lip is blackish
and the columella bluish white. (Fig. 12.)
The above are a few variations in colour illustrated by six
specimens selected from a hundred. Other slight variations
were observed in many other examples. The ground-colour
varies from bright green to bright or pale yellow, with all
shades of intermediate tints. ‘The form f, of which there was
but one specimen, is most strikingly abnormal. It has a
black lip, whereas in all the rest it is white. However, in
two specimens in the Museum marked Borneo, which I believe
belong to this species, it is almost black.
Similar great differences in colour occur in A, Adamsit,
Reeve.
9. Opisthoporus quadrasi, Hidalgo.
Hab. Palawan (Hidalgo, Dohrn, and Everett).
10. Leptopoma vitreum, Lesson.
Several varieties of this species occur in Palawan. Some
are white with fine pellucid spiral lines; in others the latter
$52 Mr. E. A. Smith on Land- Shells
are pale brownish and more or less interrupted or dotted, and
the variety named ZL. bicolor, Pfr., is also among the speci-
mens sent by Mr. Everett, who also obtained a similar series
of varieties at Balabac.
11. Leptopoma palawanensis. (Pl. XVIII. figs. 20, 21.)
Testa turbinata, umbilicata, mediocriter tenuis, albida, epidermide
tenui flavescente induta, flammulis obliquis irregularibus nigro-
fuscis subconfertis supra picta, infra peripheriam lineis angustio-
ribus zigzag-formibus et zona saturatiore prope medium ornata ;
spira conica, mediocriter acuta; anfractus 53, primi 13 flaves-
centes, sequentes duo saturate fusco-purpurei, omnes convexi
celeriter crescentes, spiraliter distanter lirati vel carinati, ultimus
in medio subangulatus, postice carinatus (carina versus aperturam
sensim obsoleta), supra peripheriam carinis subsquidistantibus
tribus cinctus, infra medium fere levis; apertura haud circularis,
triangulariter rotundata; peristoma album, leviter expansum,
margine columellari arcuato, recedente, superne callo tenui labro
juncto.
Diam. maj. 18 millim., min. 14; alt. 153. Apertura cum perist.
10 lata, 94 longa.
Hab. Palawan.
Perhaps L. regulare, Pfr.*, is nearer this species in some
respects than any other yet described. It is, however, con-
siderably smaller and has more numerous keels. It is also
taller in proportion to the width and has a rounder mouth.
Besides the spiral keels in the present species, faint traces
of intermediate spiral strie are observable, and the lower
part of the body-whorl not unfrequently exhibits a slender
raised line at a little distance from the more pronounced
central carina.
12. Lagochilus similis. (Pl. XVIII. figs. 14-16.)
Testa LZ. ciliato similis, sed major, levior, haud spiraliter striata,
inferne haud picta; anfractibus superioribus fortius carinatis,
umbilico carina vel angulo haud cireumdato.
Diam. maj. 16 millim., min. 12}; alt. 13. Apertura cum perist.
8 alta et lata.
Hab. Balabae and Palawan.
This species, although so very like L. ciliatum of Sowerbyt,
* Reeve’s Conch. Icon., Leptopoma, figs. 14-14 b.
+ Thesaurus Conch, vol. i. p. 127, pl. xxx. figs. 237,238; Pfeiffer, Conch.-
Cab. ed. 2, Cyclostoma, p. 150, pl. xx. figs. 26-27 ; Reeve, Conch. Icon.,
Leptopoma, pl. vii. fig. 30,
from the Philippine Islands. 353
may be distinguished thus:—It is larger and has a more
glossy appearance, caused by the absence of the fine spiral
striae which mark the surface of that species. The flame-like
painting which ornaments the upper surface never extends
beyond the periphery, the base being of a uniform light or
dark horny tint.
In the three specimens of L. ciliatum in the Museum
collection the base and especially the umbilicus exhibit vivid
flammulations. Also two out of these three specimens
have a strong carina bordering the umbilicus, a feature
entirely wanting in all the examples of the present species.
In the latter the peripherial keel revolves up the spire and is
visible just above the suture, and a second thread-like carina
occurs upon the middle of the penultimate and two preceding
whorls, but is almost obsolete upon the last. In ZL. ctliatum
these keels or liree are much less conspicuous.
The ground-colour varies from reddish to whitish horn-
colour, and the amount of flame-like painting upon the upper
surface is more or less extensive, in some cases forming
merely a series of blotches at the suture.
The typical examples of L. ciliatum were obtained in the
island of Luzon by Mr. Cuming; but the occurrence of the
ae in the Philippine Islands has not since been con-
firmed.
13. Cyclotus euzonus, Dohrn. (PI. XVIII. figs. 17-19.)
Hab. Palawan (Dohrn and Everett).
A single specimen of this species was obtained by Mr. Eve-
rett, differing from the type in being entirely of a chestnut-
colour with the exception of a yellow narrowish zone at the
periphery. In form this species exactly resembles C. Boxalli
of Godwin-Austen, from North Borneo, but may be distin-
guished by difference of colour and its smooth thin epidermis,
which, in the Bornean shell, is thick, opaque, and striated.
EXPLANATION OF PLATE XVIII.
Figs. 1-3. Macrochlamys pseustes.
Figs. 4-6. Lamprocystis chlororhaphe.
Figs. 7-9. Trochonanina paraguensis.
Figs. 10-13. Amphidromus quadrasi,
Figs. 14-16. Lagochilus similis.
Figs. 17-19. Cyclotus euzonus.
Figs. 20,21. Leptopoma palawanensis.
354 Mr. G. Lewis on new Japanese Coleoptera.
LVIII.—On new Japanese Coleoptera (Silphide)*.
By G. Lewis, F.L.S.
In 1881 I had about thirty beetles given to me by Mr. Snow,
who had, in an interval of seal-shooting, gathered them a
short time before on the sea-beach of Ketoi, one of the
smaller islands of the Kurile group. There were four or
five specimens of Nebria Snow?, Bates, a few examples of
Geotrupes levistriatus, Motsch., some Curculionide, and the
rest of the specimens belong to the genus Lyrosoma. The
last are now described here, together with two species in the
allied genera Pteroloma and Pelates taken by myself on the
main island of Japan. The fauna of the Kurile Islands is
doubtless a very limited one; but the discovery of four
species of Lyrosoma amongst so few specimens shows it to
be one of a very peculiar character.
List of Species.
Lyrosoma tripartitum. Lyrosoma ovipenne.
Snowi. Pteroloma discicolle.
suturale. Pelates striatipennis.
Lyrosoma tripartitum, sp. n.
Rufo-brunneum, subopacum; antennis pedibusque concoloribus ;
capite inter oculos carinato.
L. 62-7 mill.
Reddish brown, elytra somewhat darker; the head, area
before the eyes and the epistoma are flat and on the same
plane, posterior limit of this area semicircular in outline ;
between the epistoma and the head is a distinct suture, which
is joined to a longitudinal carina between the eyes, thus
dividing the upper surface of the head into three parts,
surface of head somewhat opaque, owing to a very fine
sculpture; the thorax widest before the middle, and from the
midille narrowed to the hind angles, angles a little obtuse,
behind the neck is a rather wide median impression, surface
microscopically sculptured, uneven, with a few irregular and
ill-defined punctures ; the scutellum triangular, very minutely
and transversely rugose; the elytra margined laterally,
margin and first stria leave a rather wide interstice from
* For the first part of the “Silphide of Japan” see Ann, & Mag.
Nat. Hist. 1887, xx. p. 338,
Mr. G. Lewis on new Japanese Coleoptera. 355
behind the humeral angle to the apex ; striae feebly punctu-
late, interstices flat, with a fine sculpture, which renders them
opaque.
Hab. Ketoi. Under stones on the shore.
Lyrosoma Snowt?, sp. n.
Obscuro-brunneum vel piceum; capite inter oculos subarcuatim
elevato, irregulariter punctato.
L. 6 mill.
Dull reddish brown or piceous, head distinctly darker, legs,
antenne, and thorax concolorous; the head is irregularly and
rather roughly punctured, with the median carina and the
suture, as noted in L. tripartitum, scarcely visible, but between
the eyes there is a crescent-shaped elevation; the thorax
more distinctly punctured than in the last species and less
narrowed behind, with the hind angles less obtuse, impressed
behind the neck ; the scutellum slightly arched at the sides
and densely and distinctly sculptured; the elytra margined
laterally, lateral interstice narrow and continuing to the base,
strie very feebly punctulate, interstices somewhat less flat
than in L. tripartitum.
Hab. Ketoi.
Lyrosoma suturale, sp. n.
Rufo-brunneum, vix nitidum, sparse sed distincte punctatum ;
elytris piceis, interstitiis internis rufo-brunneis.
L, 43 mill.
Reddish brown, somewhat shining; the head uneven, with
two rather large shallow impressions on each side near the
insertion of the antenne, distinctly punctured in certain
places, most free from points on the disk ; the thorax punctured
and uneven like the head, hind angles slightly acute, less
distinctly narrowed behind than in Z. Snow7, impressed behind
the neck ; the scutellum triangular, wholly sculptured, with a
few rather large punctures scattered over it; the elytra are
piceous, except on the two sutural interstices, the striz
distinctly punctate, especially in the scutellar area; the lateral
interstice 1s narrow and ceases before the apex.
Hab. Ketoi,
Lyrosoma ovipenne, sp. n.
Rufo-brunneum, vix nitidum ; elytris oviformibus, striis obsoletis
punctatis.
L. 5 mill.
356 Mr. G. Lewis on new Japanese Coleoptera.
Reddish brown, somewhat shining, elytra a little darker ;
the head uneven above and irregularly punctured ; the thorax
widest about the middle, uneven and punctured like the head,
impression behind the neck shallow, in one example almost
effaced; the hind angles are obtusely rectangular; the
scutellum triangular, feebly arched at the sides, wholly sculp-
tured on surface, with a few punctures dispersed over it ; the
elytra are ovate, marginal interstice narrow and of even width
throughout ; striz obscurely punctured, interstices densely
sculptured and feebly rugose or uneven.
Hab. Ketoi.
The sculpture noticed in these species is peculiar to the
genus; when it is feeble it resembles the surface of tanned
leather, when dense and well-marked, as it is on the scutellum
of L. Snow, it resembles the sculpture so familiar to ento-
mologists on the wing-cases of Agrile.
Pteroloma discicolle, sp. n.
Rufo-brunneum, vix nitidum; thorace disco nigro, angulis anticis
prominentibus.
L. 4 mill.
Reddish brown, somewhat shining; epistoma and disk of
the thorax black or piceous ; the head impunctate but uneven ;
thorax, thehind angles rectangular, sides feebly bowed, anterior
angles prominent, the thorax being cut out in front to receive
the head; the disk is black and convex, the sides longitu-
dinally impressed and spread outwards; the scutellum trian-
gular and without sculpture ; the elytra, margin elevated,
marginal interstice narrow, the second and third strie from
the outer margin join at the base, the strie are somewhat
fine and puuctulate ; the wing-cases are wider than the thorax
at the base, with a somewhat elliptical outline to the base.
I have one example, evidently a variety, in which the
antenne, head, and legs are infuscate.
Hab. Main island. I took six or seven specimens from
under small stones on the summit of Nantaisan, 20th August,
1881.
Pelates striatipennis, sp. n.
Oblongo-ovatus, rufo-brunneus, nitidus ; thorace conspicue punctato ;
elytris punctato-striatis.
L, 52-6 mill.
Oblong-oval, reddish brown or piceous, but with the mar-
gin of the thorax and elytra paler; the head sparsely punctu-
On the Skull of Pisodus Oweni. 357
late, with the median area a little raised; the thorax convex
behind the neck, transverse, arched at the sides, anterior
angles obtuse, lateral margin narrow, very clearly punce-
tured, punctures least close on the disk ; the elytra punctate-
striate, striae deep and well-marked; the legs, palpi, and
antenn are concolorous with the paler parts of the body.
Hab. Main island. At Nikko an old tree, rotten as touch-
wood and ivy-grown, yielded eight specimens. Single
a ae were taken fortuitously in other places by beating
oliage.
LIX.—Description of the Skull of Pisodus Oweni, an Albula-
like Fish of the Eocene Period. By A. SmirH Woopwarp,
F.L.S.
[Plate XVII. }
Ir is now a well-established fact that many types of Tele-
ostean fishes have undergone very little change since the
Eocene, or even since the latter part of the Cretaceous period.
Several well-defined genera seem to date back thus far, and
others are represented by forms that differ in but small
particulars. Moreover, a few of the most remarkable speciali-
zations in piscine skeletal anatomy characterizing the existing
fauna are already recognizable in certain closely related
Eocene types, and the progress of discovery is continually
adding to the number of known examples. A most striking
new case has lately been met with by the present writer
among the fishes trom the London Clay (Lower Eocene), and
this forms the subject of the following notes.
So long ago as 1845 Sir Richard Owen described and
figured the tritoral dentition of an unknown fish from the
London Clay of the Isle of Sheppey under the name of Pisodus
Oweni (ex Agassiz, MS.)*. The original specimen is pre-
served in the Museum of the Royal College of Surgeons, and
exhibits an ovate pavement of small rounded or polygonal
teeth firmly fixed in shallow sockets upon a plate of true
bone. Appearances suggested to Sir Richard Owen that the
fossil had been attached ‘‘ to another bone of the skull, most
probably, as in the Glossodus and Sudis, to a median bone of
the hyoid system.” Agassiz, who first examined the specimen,
* R. Owen, ‘ Odontography,’ p. 138, pl. xlvii. fig. 3 (1845).
Ann. & Mag. N. Hist. Ser. 6. Vol. xi. 26
358 Mr. A. S. Woodward on the
supposed it might pertain to a so-called Pycnodont Ganoid * ;
and in Owen’s ‘ Paleontology’ (edit. 2, 1861, p. 174)
Pisodus is also doubtfully quoted as a “ Ganoid ” of uncertain
position,
It now appears from a nearly complete skull in the British
Museum that the problematical fossil in question is the para-
sphenoid dentition of a fish remarkably similar in cranial
characters to the recent Clupeoid Albula. The fact has
already been incidentally mentioned in a record of the disco-
very of Pisodus in the Middle Eocene of Belgium f; and it
only remains to justify, by a detailed description and figures,
the recognition of an A/bula-like fish at so remote a period as
that of the Lower Eocene. Dr. Shufeldt’s admirable
description of the skull of the recent Albula vulpes fortunately
suffices for requisite comparison tf.
The fossil skull in question is imperfectly preserved in
several respects, though exhibiting most of the principal
characters. It is shown of one half the natural size, from
four aspects, in the accompanying Plate XVII. The rostral
region is wanting, the projecting lateral portions are broken
away, and the basioccipital and basisphencid bones are in
great part removed by fracture. The general form of the
cranium, however, is recognizable, and the great expansion of
the parasphenoid, with its tritoral dentition, is especially well
displayed. ‘The cranial roof (fig. 1) is flattened and termi-
nates abruptly behind in a straight occipital border. The
occipital face (fig. 2) is nearly vertical; and the basioccipital
axis is approximately parallel with the cranial roof, meeting
the dentigerous portion of the parasphenoid in a sharp angu-
lation (fig. 4).
On the occipital face (fig. 2) the two exoccipital elements
(ex.occ.) are observed immediately above the foramen
magnum (f. m.), and seem to have been originally apposed
in the median line except at their upper extremity. ‘These
bones are deeper than broad, so far as preserved, quadrilateral,
and with a slightly concave outer surface. Their superior
internal angles are truncated for the reception of the lower end
of the large triangular supraoccipital (s.occ.), which is flat-
tened, but shows remains of a T-shaped, backwardly directed,
mesial crest (c) in its upper two thirds.
* L. Agassiz, Poiss. Foss. vol. ii. pt. ii. p. 247 (1844).
+ Smith Woodward, ‘‘ Notes on some Fale eerniies from the Lower
Tertiary and Upper Cretaceous of Belgium,’ Geol. Mag. [8] vol. viii.
p: 108 (1891).
t R. W. Shufeldt, “The Osteology of Amia calva &e.,” Ann, Rep.
U.S. Fish Commission, 1885, pp. 61-67, pls. xii., xiii.
Skull of Pisodus Oweni. 359
The supraoccipital bone also appears as a narrow band at
the hinder margin of the cranial roof (fig. 1, s.occ.) bordering
the pair of very short and broad parietal elements’ (pa.).
These bones are not symmetrical, the mesial suture being
bent and distorted to the right. They are much smaller than
the squamosals (sg.), of which that of the left side is pre-
served toa large extent. The frontal bones (fr.) are rela-
tively enormous, and all the smaller roof-bones by which they
would be bounded laterally and anteriorly are unfortunately
removed. While for the most part flat, this pair of bones is
mesially depressed in the hinder two thirds of its extent, and
thus originates a great hollow gradually deepening forwards
to a sudden termination.
Of the bones in the otic region only portions of the pro-otic
and opisthotic remain, and these are not sufficiently well
preserved for description.
The three teeth displayed in their sockets in the para-
sphenoid bone (fig. 3, pas.) have been much worn during the
hfe of the fish, all the gano-dentine being removed from the
top of the flattened crown.
Nothing further need be added to indicate how closely the
cranium thus described resembles that of Albula treated by
Dr. Shufeldt in the memoir already referred to. So far as
preserved the fossil agrees with the recent skull even to minute
particulars, and, apart from size, there is little to indicate
specific, much less generic, differences. The rostral region
may, however, have presented a different configuration from
that of the existing Albula; and in the absence of all know-
ledge of the trunk it is impossible to arrive at any definite
generic determination. The Eocene fish from Sheppey must
therefore retain its provisional name of Pisodus Owent, and
the description of the skull now published merely establishes
its family position.
EXPLANATION OF PLATE XVIL
Pisodus Oweni, Owen (ex Agassiz, MS.). Skull shown from four aspects,
one half the natural size.—London Clay, Isle of Sheppey. | Brit.
Mus. no. 39439. ]
Fig. 1. Cranial roof. c, base of crest on supraoccipital ; fr., frontal; pa.,
parietal ; s.occ., supraoccipital ; sg., squamosal.
Fig. 2. Outline of occipital face. ex.occ., exoccipital; f.m., foramen
magnum,
Fig. 3. Base. pas., parasphenoid, showing three teeth and sockets of
others.
Fig. 4. Side view. orb., position of eye.
26*
360 Herr B. Lwoff on the Formation of the
LX.—On the Formation of the Germinal Layers in Verte-
brates. By Basttius Lworr, of Moscow University *.
EXTENSIVE comparative investigations upon the formation of
the germinal layers in Vertebrates have led me to conclude
that all the theories at present prevailing on the subject of
gastrulation and germinal-layer-formation in this division of
the animal kingdom are not in accordance with the facts.
All these theories regard the invagination, which is charac-
teristic for all Chordata, as nothing else than gastrulation,
although im the majority of cases this invagination bears no
relation to the formation of the intestine. The consequence
of this is that in the higher Vertebrata we have nothing to
help us to distinguish the primary germinal layers. Starting
from the conviction that the true endoderm is always formed
by invagination, authors have proceeded so far in this direc-
tion as to regard the lower layer in the case of the Amniota,
from which the intestine is formed and which is homologous
with the endoderm of the lower Vertebrates, not as the endo-
derm, but as something that admits of no homology with
what is found in the lower Vertebrates (paraderm ec.) ; as
endoderm, however, are designated the ectoderm cells which
are invaginated or grow inwards, but take no part in the
formation of the intestine. Or, again, it is assumed that both
the cells of the lower layer, from which the intestine is
formed, and the invaginated ectoderm cells are to be con-
sidered as endoderm; but the lower Jayer, from which the
intestine arises, is designated secondary or cenogenetic endo-
derm, while to the invaginated cells, which represent the
rudiment of the notochord and mesoderm, the term primary
or palingenetic endoderm is applied. It therefore follows
according to this interpretation that the intestine is formed
from the cenogenetic and the notochord from the palingenetic
endoderm. In order to realize how untenable this concep-
tion is we need only ask which is phylogenetically the older,
the intestine or the notochord ?
Since we would regard the process of invagination which
is characteristic of all Vertebrates as gastrulation in its
original simplicity, it is natural that we should desire to
discover a gastrula mouth in these supposed gastrule also,
and to determine in all Vertebrates the homology of the
dorsal and ventral lips of the blastopore. But the views of
* Translated from the ‘ Biologisches Centralblatt, xiii. Bd., no. 2,
Feb, 1, 1898, pp. 40-60; and no. 3, Feb, 15, 1893, pp. 76-81.
a
Germinal Layers in Vertebrates. 361
different investigators with reference to these questions are
so very divergent as to furnish the best proof of the vague-
ness of the conceptions as to what is to be designated as
gastrulation. Just as many differences of opinion prevail on
the subject of the gastrula mouth. According to the view of
certain embryologists the blastopore in meroblastic ova corre-
sponds to the margin of the epibolic growth which surrounds
the yolk. Other embryologists, on the contrary, insist that
the margin of the epibolic growth in no way corresponds to
the blastopore ; they maintain that it isa peculiarity of mero-
blastic ova, and so forth. It is proposed by this school to
designate as blastopore that spot in the germ at which an
invagination of cells takes place (in the Selachians the poste-
rior portion of the margin of the germinal disk, in the
Amniota the primitive streak and the primitive groove).
But this is not all. There are also embryologists who assume
that the blastopore is always turned towards the neural side
of the animal, where it closes up along a median line, which
is termed the gastrula suture (‘Gastrularaphe”’). In this
manner, according to this conception, the nervous system
develops in the place of the gastrula suture, since the margins
of the blastopore become transformed into the medullary
folds!
In this brief communication it is indeed impossible for me
to enter into a discussion of all the theories which trace the
corresponding developmental processes of Vertebrates to
gastrulation, or which, in other words, would discover in
gastrulation a universal explanation of the formation of
germinal layers, as though no other processes could exist in
these stages. Many attempts have been made to apply the
gastrulation theory in the case of all Vertebrates, but all
these theories are In my opinion forced and unnatural. We
have yet to inquire whether it is not possible to interpret
these processes somewhat differently without unduly extending
the range of the theory of gastrulation.
My investigations have led me to the conclusion that a
conception such as regards the process of invagination not as
gastrulation, but as a phenomenon which is characteristic of
all Chordata, is not merely possible, but is absolutely neces-
sary, if we would compare the earliest developmental pro-
cesses of different Vertebrates one with another, and at the
same time retain the strict homology of the primary germinal
layers. I have studied the formation of the germinal layers in
the following animals :—Amphioxus and Petromyzon; Azolotl,
among the Amphibians ; Pristtwrus and Torpedo among the
Selachians ; Labrax, Julis, and Gobius among Teleostean
362 Herr B. Lwoff on the Formation of the
fishes; and Lacerta among the Reptiles. My investigations
upon the development of Amphiowus have already been
published*. In the following pages it is my intention to
describe the results of my studies upon these developmental
processes in the above-mentioned Vertebrates, so far as is
necessary in order to establish my theory. Although I owe
this theory just as much to an attentive study of the literature
of the subject as to my own investigations, I shall never-
theless in the present paper leave the literature almost entirely
out of the question, in order not to overstep the limits of a
provisional communication. The discussion of the literature
dealing with the subject, as well as the full description of my
investigations, will follow in a detailed paper which is shortly
to appear.
Before proceeding to the account of my results I would
draw attention to certain points which are of importance in
this question. In the first place let us consider whether the
school which is at present predominant is correct in regarding
the invagination in Vertebrates as gastrulation. In order to
answer this question it is necessary to ascertain what is
actually to be understood by the term gastrulation, and to
discover what portion of the process is typical and what is
adventitious and unessential.
Gastrulation is the term which was applied to the process
of invagination which leads to the formation of the enteric
cavity, whereby a distinct gastrula is constituted. But the
process of the formation of the intestine is not always accom-
plished by means of invagination ; it often happens that this
process takes place in such a way that the cells which subse-
quently form the intestine (endoderm cells) are overgrown
by the external (ectoderm) cells, and the enteric cavity after-
wards arises through divergence of the endoderm cells. This
process of enveloping growth is regarded by many investi-
gators as homologous with typical gastrulation, and, as I
think, justly, since in both cases the process essentially con-
sists in the fact that the endoderm cells which constitute the
intestine come to lie in the interior, and are surrounded by
the cells of the ectoderm. Whether this process is accom-
plished in the form of invagination or of enveloping growth
is of subordinate importance. It follows from this that the
only process which can be designated as gastrulation is that
by which the endodermal elements are invaginated or over-
grown, by which consequently above all the formation of
the intestine is ushered in. We must hold fast to this con-
* Biol. Centralblatt, Bd. xii, nos, 23 and 24,
Germinal Layers in Vertebrates. 363
ception if we would believe that gastrulation has any definite
meaning at all. This is the first point to be considered in
this important question.
The second point concerns the discrimination of the primary
germinal layers. If we would derive the bilaterally symme-
trical Chordata from a Gastrula-like form with radial symmetry
we must adhere to the homology of the ectoderm and endo-
derm of such a Gastrula-like form with the external and
internal germinal layers of the Chordata. Since, however,
the endoderm of the Gastrula-like form in the first place
gives rise to the intestine, our first step in the determination
of the germinal layers is to elucidate the question by what
elements or what layer the intestine is formed. To this layer
the term endoderm must be applied, it matters not whether
something besides the intestine is formed from these cells or
not.
I will not here enter into the question whether typical
gastrulation, ¢. e. invagination, represents a primary or a
secondary mode of formation of the endoderm. Nevertheless,
in the interpretation of the conditions which are found in
Vertebrates I assume that the bilaterally symmetrical Chor-
data are derived from a Gastrula-like form with radial
symmetry, since in the development of the lower Chordata it
is impossible not to recognize a gastrula, although a some-
what modified one. At the same time, in homologizing the
germinal layers of the Chordata we must not lose sight of
our thesis, that the inner layer (the endoderm) of the gastrula
forms the intestine, while the ectoderm constitutes the outer
covering ; otherwise the homologization loses all its meaning,
Therefore we shall designate as endoderm cells those from
which the intestine arises, no matter whether something else
is also formed from these cells or not.
If from this point of view we compare the conditions which
exist in Amphioxus and the Vertebrates, we arrive at the
following results :—The segmentation of the ovum proceeds
in such a way that in holoblastic ova, as a result of the
segmentation, we get a blastula, one half of which is com-
posed of smaller blastomeres (micromeres) and the other of
larger blastomeres (macromeres). ‘The difference between
the micro- and macromeres has arisen in consequence of the
fact that the former multiply more rapidly than the latter.
Since the more rapid multiplication of the micromeres also
continues after the formation of the blastula, the micromeres
commence to spread out over the macromeres and to grow
round them. In cases where we find a single-layered
blastula (in Amphiorus) this process takes place in such a
364 Herr B. Lwoff on the Formation of the
way that the macromeres become invaginated; where the
blastula consists of several layers (as in Petromyzon and
Amphibia) the micromeres simply grow round the macro-
meres. Since these macromeres form the intestine, they may
with perfect justice be designated as endoderm cells; the
micromeres, on the contrary, which differ noticeably from the
macromeres and give rise to the outer covering, are to be
termed ectoderm cells. I see no reason to distinguish an
ectodermal and an endodermal half after the first divisions of
the ovum have taken place, or it may be after the first equa-
torial furrow (as is the practice of certain investigators).
Such a distinction seems to me to be lacking in foundation at
this stage, since each supposed endoderm cell of this kind may
divide into two or more cells, of which one will subsequently
become an ectoderm cell by position, while the other will
belong to the endoderm. ‘The distinction of the ectoderm
from the endoderm is only possible when the blastula is
already formed and the macromeres are being overgrown by
the micromeres. In this connexion it makes no difference
whether the macromeres are invaginated or are overgrown by
the smaller cells. I term them endoderm cells not on account
of invagination, but merely because they form the intestine.
But the process in consequence of which the endoderm cells
come to lie within and become surrounded by the cells of the
ectoderm can be homologized with that of gastrulation.
This process is to be regarded as preparatory to the formation
of the intestine. But besides this process of gastrulation, by
which the formation of the intestine is ushered in, there
appears on that side which subsequently becomes the dorsal
side of the animal another process, which represents the
invagination of the ectoderm cells, and which I will term the
dorsal invagination. This dorsal invagination is quite inde-
pendent of the gastrulation, and has nothing to do with the
formation of the intestine; it gives rise to the common ecto-
blastogenous rudiment of the notochord and the mesoderm.
This conception, which was mainly arrived at in the
investigation of the development of Amphiowus, Petromyzon,
and Awolotl, I was also able to follow out in the interpretation
of the developmental processes of the ‘Teleosteans and
Selachians, and it is moreover confirmed in the clearest
manner by the development of the Amniota.
After these preliminary remarks I will proceed to the
account of my results.
Germinal Layers in Vertebrates. 365
My investigations upon the formation of the germinal layers
in Petromyzon commence with the stage of the blastula. The
blastula includes an excentric cavity, lying nearer the upper
pole, the roof of which is constituted by smaller blastoderm
cells and the floor by several layers of larger cells, richer in
yolk, which develop into endoderm. I was also able to
observe the multiplication and extension of the blastoderm
(ectoderm) cells over those richer in yolk (endoderm cells).
Numerous mitoses may be observed in the ectoderm cells. I
counted the mitoses in a series of sections through a stage of
this kind, and found that the endoderm cells contained five
and the superficial ectoderm cells twenty-three mitoses. In
this manner smaller ectodermal cells surround the larger
endodermal elements, which thus come to lie within and
become enveloped by the former. Simultaneously with this
on one side, which becomes the dorsal surface of the embryo,
a particularly active multiplication of the ectoderm cells may
be noticed, and here, at a spot which marks the posterior end
of the embryo, the invagination of the ectoderm cells com-
mences. By this means a cavity is formed, which is usually
termed the cavity of the gastrula, or archenteron, and the
dorsal wall of which is constituted by the invaginated ecto-
derm cells. But this invagination forms only the dorsal wall
of the cavity, and I therefore term it the dorsal invagination.
The ventral wall of the cavity is formed by the endoderm
cells, which are not invaginated, but were previously situated
here. ‘These conditions can be seen with great distinctness
in median sections through stages of this kind. In such
sections we observe that the ectoderm cells grow inwards from
the dorsal margin of the enveloping layer, and form the dorsal
wall of the cavity ; the ventral wall, on the other hand, is
formed by the cells of the endoderm. We also observe the
different character of the cells of the dorsal and ventral walls.
The former are arranged in epithelial fashion, and form the
continuation of the ectoderm cells, which grow inwards from
the margin of the enveloping layer. The cells of the ventral
wall of the cavity have a roundish or polyhedral form, and
are not arranged in epithelial fashion. ‘They acquire the
regular epithelium-like character later on, when the intestine
arises from them. In the enumeration of the mitoses in a
series of sagittal sections it was found that the endoderm cells
contained four (chiefly in the anterior portion of the cavity),
the ectoderm cells twenty-four (chiefly upon the dorsal side ;
among these were seven at the margin of the enveloping layer) ,
and the invaginated ectoderm cells four mitoses.
I must lay quite special emphasis upon the fact that the
366 Herr B. Lwoff on the Formation of the
endoderm cells which subsequently form the intestine are not
invaginated ; they only experience certain displacements in
consequence of the dorsal invagination, whereby the segmen-
tation cavity becomes obliterated. Elements alone are
invaginated from which the notochord and the mesoderm
arise, and moreover this invagination takes place in such a
way that the ectoderm cells grow inwards from the margin of
the enveloping layer and form the continuous dorsal plate,
the ectoblastogenous rudiment of the notochord and mesoderm,
I see no reason for designating the cells of the dorsal wall of
the cavity as endodermal, merely because they grow inwards
(become invaginated). Similarly I see no reason for terming
the cavity at this stage the gastrula or archenteric cavity,
At this stage an enteric cavity is not yet present: it is con-
stituted subsequently, when the endoderm cells diverge one
from another and form the epithelial wall.
From the central portion of the dorsal plate is differentiated
the rudiment of the notochord; the two lateral portions,
together with the adjoining cells of the endoderm, form the
rudiments of the mesoderm. Asa rule no sharp dividing line
can be drawn between the ectoblastogenous and endoblasto-
genous cells of the mesoderm, for the cells which are invagi-
nated grow inwards, while they closely adjoin the cells of the
endoderm. After the rudiment of the notochord has separated
off from the lateral mesodermal rudiments, the margins of the
endoderm grow towards one another, to close the enteric
wall; but before this takes place the rudiment of the noto-
chord is temporarily included in the dorsal wall of the intes-
tine. Since the margins of the endoderm continue to grow
beneath the notochord, the latter is excluded, the edges of the
endoderm unite, and the enteric cavity closed on all sides is
now constituted.
I was able to observe the same developmental processes in
the case of Agolotl also. In this case certain points tell even
more in favour of my conception than in Petromyzon. While
the larger endoderm cells are being overgrown by the smaller
ones of the ectoderm the dorsal invagination begins, This
invagination is nothing else than the ingrowth of the ecto-
derm cells, which grow inwards from the margin of the enve-
lope and form the continuous dorsal plate of cells which, as
in the case of Petromyzon, represents the ectoblastogenous
rudiment of the notochord and of the mesoderm. This
invagination in no way assists in the formation of the enteric
—
Germinal Layers in Vertebrates. 367
cavity. The intestine is formed by the divergence and
displacement of the endoderm cells, which are not invaginated,
but were previously situated here, and may be distinguished
all the time from the smaller ectoderm cells by their relative
size, by the amount of the yolk-granules which they contain,
and to a certain extent by their want of pigment. While the
dorsal invagination is taking place, the cavity which subse-
al becomes that of the intestine is gradually formed by
ivergence of the endoderm cells. The formation of the wall
of this cavity proceeds in precisely the same manner as in
Amphioxus and Petromyzon, i. e. the endoderm cells separate
in such a way that they first give rise to the ventral and
lateral boundary of the intestine, while the dorsal portion is
still open. Thus it comes to pass that the rudiment of the
notochord, which has separated from the lateral mesodermal
rudiments, temporarily takes part in the limitation of the
enteric cavity. Subsequently the endoderm cells unite
beneath the notochord, which in this manner is again
excluded. As regards tlie lateral portions of the dorsal plate-
like ingrowth, which represent the ectoblastogenous rudiment
of the mesoderm, they are from the first excluded from the
delimitation of the enteric cavity owing to the intervening
endoderm cells, part of which attach themselves to the ecto-
blastogenous mesoderm cells in order to furnish their contri-
bution to the formation of the mesoderm.
I must not conceal the fact that these conditions are not
exhibited by all Amphibians. According to the statements
of certain investigators the Anura are an example of an
instance in which the notochord is excluded from the outset
from the delimitation of the enteric cavity. Unfortunately I
have no experience myself in this direction, since I have not
studied the development of the Anura. But if this statement
is correct, the Anura in this respect approach the Selachians
and Teleosteans, as will be shown further on.
It is sufficiently clear from what has been stated that in
the case of the Amphibia also a distinction must be drawn
between the dorsal invagination and the formation of the
intestine. They are two different processes, which have a
certain relation one to another only because the formation in
one organism of two neighbouring organs always shows
certain points of contact or connexion. As regards the forma-
tion of the mesoderm, this arises from ectoderm as well as
from endoderm, and moreover not only in the lateral meso-
dermal rudiments to which reference has already been made,
but also in the so-called ventral mesoderm, in which also no
sharp dividing line is to be drawn between the ectoblasto-
Herr B. Lwoff on the Formation of the
genous and endoblastogenous mesoderm cells. In this brief
communication, however, devoid as it is of figures, I cannot
enter more closely into these details, and must postpone an
account of them until the publication of the longer paper
which is to follow.
In passing on to the description of the corresponding deve-
lopmental processes in the meroblastic ova of Selachians and
Teleostean fishes, I must at the outset join issue with those
embryologists who, following the example of Heckel, would
regard the yolk merely as a store-room from which the germ
derives nutritive matter, and who would deny to the vitelline
elements all participation in the formation of the embryo.
Although this view must now be regarded as an anachronism,
there are still some investigators even at the present time
who assume that the yolk in Teleosteans and Selachians
takes no share in segmentation. In contradiction to these
embryologists I have to state that, according to the results at
which I have arrived, the entire endoderm (the definitive
intestine and the endoblastogenous mesoderm) in Teleostean
fishes as well as in Selachians owes its origin to the vitelline
elements.
I will commence the description of my investigations with
the Teleostean fishes, which, owing to the fact that their ova
contain less yolk, approach the’ Amphibians more closely in
these processes than do the Selachians, in which the same
developmental processes are more modified, probably in con-
sequence of the larger quantity of yolk. In the case of all
the Teleosteans studied by me (Labrax, Julis, and Gobius),
no sharp boundary can be distinguished between the blasto-
derm cells and the yolk. The lower blastoderm cells are so
intimately connected with the subjacent yolk that no dividing
line is to be seen. ‘These cells divide in an equatorial direc-
tion, and, moreover, in such a way that the uppermost cell
which is thus produced becomes constricted off and joins the
blastoderm cells, while the lower one, on the contrary, remains
in connexion with the yolk. I have observed no trace of a
segmentation cavity in any of the Teleosteans which I have
investigated. After the blastoderm has been formed and the
blastoderm cells commence to grow round the yolk we may
observe upon the surface of the latter a continuous layer of
protoplasm with nuclei, around which the outlines of cells
are sometimes to be seen. ‘This is the intermediate layer of
anthors, the nuclei of which, which are direct descendants of
TS
Germinal Layers in Vertebrates. 369
nuclei of the lower blastoderm cells, were termed merocytes
or periblast nuclei. The presence of so many nuclei without
the outlines of cells is probably to be explained as being due
to rapid nuclear division without corresponding cell-division.
IT must lay stress on the fact that these nuclei do not perish,
as some investigators would have us believe, but form new
cells, which take part in the building-up of the embryo. I
shall designate these nuclei simply as yolk-nuclei.
After the formation of the blastoderm a more or less distinct
boundary may be noticed between the blastoderm and the
yolk, or, more strictly speaking, between the blastoderm and
the intermediate layer which belongs to the yolk. Yet we
may observe al] the time the equatorial division of the yolk-
nuclei and the formation of new cells, which become con-
stricted off from the yolk and join the blastoderm cells. Now,
after the overgrowth of the yolk by the blastoderm cells has
commenced it is time to distinguish the primary germinal
layers. I consider the entire blastoderm, the cells of which
grow over the yolk, as the ectoderm, but the yolk with the
yolk-nuclei must be termed the endoderm, since the intestine
arises from these vitelline elements.
In passing on to the description of the further develop-
mental processes which usher in the formation of the noto-
chord and the mesoderm it is necessary for me to draw
attention to the fact that in none of the Teleosteans which I
have examined have I observed an invagination. It is well
known that certain investigators maintain that the mesoderm
in Teleostean fishes is formed by invagination, while others,
on the contrary, assert that it arises by cleavage of the blasto-
derm cells. Although I myself have not observed any
invagination, I think J may assume that no essential differ-
ence exists between the two processes as they are described
by different investigators; for the invagination in this case
also has no connexion with the formation of the intestine,
and is nothing else than the forward growth of the reflected
edge of the blastoderm. In both cases therefore the same
blastodermal or ectodermal elements are engaged, and it is
a matter of secondary importance whether the cells are from
the outset arranged in such a way that the continuous rudi-
ment of the notochord and mesoderm arises by splitting off
from the upper portion of the blastoderm (the rudiment of the
nervous system), or whether this rudiment is formed by the
forward displacement of the cells starting from the margin of
the enveloping layer. Whether by means of the process of
splitting-off or by that of so-called invagination, the con-
tinuous rudiment of the notochord and mesoderm is equally
370 Herr B. Lwoff on the Formation of the
formed from the same elements (ectoderm cells) as those from
which the nervous system is also developed. In this rudi-
ment the central portion (the rudiment of the notochord)
becomes differentiated from the lateral regions, from which
the mesoderm arises. But here also it may be seen that the
endoderm cells take part in the formation of the mesoderm,
since the cells which arise from the yolk-nuclei become con-
stricted off and join the rest of the mesoderm cells. Thus in
the present case, as in that of other Vertebrates, we can
distinguish a twofold source whence the mesoderm arises °
(ectoblastogenous and endoblastogenous), While the forma-
tion of the notochord and mesoderm is taking place there is
formed upon the surface of the yolk from the yolk-nuclei the
continuous layer of cells from which the intestine is deve-
loped. In this manner in Teleosteans also the intestine does
not owe its origin to an invagination, but arises from the
derivatives of yolk-nuclei.
While the cells of the blastoderm are growing round the
yolk, the yolk-nuclei which are in process of multiplication
commence to spread out over the latter, so that soon the
entire periphery of the yolk is provided with these nuclei. I
must emphatically controvert the view, which is held by
many investigators, that these nuclei perish without taking
part in the formation of the embryo. It has already been
mentioned that the yolk-nuclei give rise to the intestine and
participate in the formation of the lateral mesodermal rudi-
ments; but I must add that the nuclei also which are situated
in the lower (ventral) portion of the yolk take a share in the
formation of the embryo, for in my preparations I notice
mitoses in this region, and it may be observed that the cells
which are here formed from the yolk-nuclei become con-
stricted off and join the blastoderm cells which have grown
round the yolk. I am inclined to assume that these cells are
homologous with the ventral mesoderm of the Amphibians.
In subsequent stages it may be seen that the yolk-nuclei take
part in the formation of the liver, since a portion of the cells
which arise from these nuclei is directly transformed into
hepatic cells. ‘lhe importance of the réle of the yolk-nuclei
cannot therefore be open to doubt.
I will now pass on to the Selachians.
Although at the present time Selachians are the favourite
subject of embryologists, the latter are nevertheless not agreed
as to many important developmental processes in these
animals, and, above all, as to the origin of the intestine.
While certain investigators regard the intestine as originating
through invagination, the rest assert that it is formed from
Germinal Layers in Vertebrates. 371
the yolk-nuclei. On the other hand, while in this manner an
important rd/e is ascribed to the yolk-nuclei by some investi-
gators, the others deny that they take any share in the
formation of the embryo. From my own results I am led to
suppose that not only the intestine, but also the entire endo-
derm, arises from the yolk-nuclei, while the blastoderm cells
represent the ectoderm. ‘The formation of the intestine from
the yolk-nuclei is so distinct in all stages that I cannot under-
stand how this fact can be disputed. But, on the other hand,
I must corroborate the assertion that a kind of invagination
(blastodermal fold) is to be observed at the posterior margin
of the germinal disk. My results consequently reconcile to a
certain extent the statements of different investigators, for I
confirm both the origin of the intestine from the yolk-nuclei,
as also the presence of the invagination. But in the present
instance also this invagination or doubling-down of the blasto-
derm cells has nothing to do with the formation of the
intestine. I therefore see in it no process of gastrulation, but,
as in the case of other Vertebrates, the ingrowth of the ecto-
blastogenous rudiment of the notochord and mesoderm, which
proceeds in a forward direction from the posterior margin of
the enveloping layer.. The Selachians agree with the Tele-
ostean fishes also in the fact that not only is the inversion of
the blastoderm cells to be observed at the posterior margin of
the enveloping layer, but also the splitting-off of cells here
and there. ‘These two different processes—the formation of
the intestine from the yolk-nuclei and the ectoblastogenous
invagination—are accomplished simultaneously in such a way
that, while the ectoderm cells are invaginated at the posterior
margin of the enveloping layer, the continuous sheet of endo-
derm is formed from the yolk-cells, which give rise to the
intestine. The invaginating blastoderm cells in no way con-
stitute the intestinal wall; they merely lie, while they grow
inwards, so closely upon the true endoderm cells which have
arisen from the yolk-nuclei, that the two different rudiments
—the invaginated ectodermal one and the endodermal which
has developed on the spot from the yolk-nuclei—are intimately
connected one with another. It is the more difficult to draw
a dividing line between the two rudiments, since in the lateral
rudiments of the mesoderm both kinds of cells are in close
connexion. :
The rudiment of the notochord is excluded from the limita-
tion of the intestinal cavity by the actual endoderm cells, and
forms for a time an appendage of the dorsal wall of the intes-
tine. As regards the mesoderm, we may follow Riickert in
distinguishing the axial and peripheral portions of this layer.
372 Herr B. Lwoff on the Formation of the
In the formation of these parts of the mesoderm, which differ
only in position, the cells of both ectoderm and endoderm
participate. These two sources of the mesoderm cells are very
distinctly visible; but the detailed demonstration must be
reserved for my subsequent paper. I will here but briefly
allude to one more point. It is evident from what has been
stated that I by no means regard the Selachians as primitive
forms, from which other animals are to be derived. I see no
reason to derive the Amphibians from the Selachians, as is
done by certain investigators. This amounts to deriving
simple and primitive conditions from such as are complicated
and modified. Although the Selachians in many respects
exhibit primitive characteristics, their conditions are never-
theless so much modified in consequence of the superabun-
dance of the food-yolk, that they can be regarded as the
aneestral type of other animals just as little as the Teleostean
fishes.
In the case of Lacerta no difficulty is experienced in distin-
guishing the two primary germinal layers, since we here get
a bilaminate germinal disk as the result of segmentation. At
the posterior margin of the germinal disk we may observe an
active multiplication, and in consequence of this a prolifera-
tion of the ectoderm cells. This proliferating spot in the
ectoderm, which in the case of Amniota is termed the primitive
tubercle (“ Primitivknopf’”’), primitive plate, primitive streak,
and I know not what besides, gives rise to the ectoblasto-
genous rudiment of the notochord and mesoderm, which,
starting from this point in a forward direction, grows in
between the two primary germinal layers as a continuous
plate of cells. In the middle of the primitive streak, but
nearer its anterior end, there may be noticed a depression by
which the invagination is ushered in. In this manner there
arises the margin of the enveloping layer at which the cells of
the rudiment of the nervous system and those of the rudiment
of the notochord bend round into each other, and where nume-
rous mitoses are to be seen. The direction of these mitoses
shows in the clearest manner that the cells of the rudiment of
the notochord grow forward from the margin of the over- -
growth. ‘This invagination in no way leads to the formation
of the intestine, since all the invaginated cells are absorbed in
the production of the notochord and mesoderm; but the
intestine arises from the cells of the lower primary germinal
layer. I have no reason in this case to designate the process
Germinal Layers in Vertebrates. 373
of invagination as gastrulation, for this process here bears no
relation to the formation of the intestine. It is true that
certain investigators describe a sac-like invagination in the
case of Reptiles, and consider its cavity to be that of the
archenteron. But, in my opinion, this discovery does not
prove what these authors wish to prove, but rather exactly
the opposite, since it is evident from their description that,
although the cavity produced by invagination exists, never-
theless its walls do not participate in the formation of the
intestine, and that the cavity of invagination in no way
becomes that of the intestine.
Consequently here also we see in the invagination a process
characteristic of the Vertebrata, namely the formation of the
ectoblastogenous rudiment of the notochord and mesoderm.
In this case also the notochordal rudiment, after it has sepa-
rated from the lateral rudiments of the mesoderm, enters into
a temporary connexion with the endoderm cells, since it
becomes intercalated among them; but subsequently the
notochord is once more excluded. A correct description of
this process has already been furnished by many investigators.
As regards the mesoderm, we can here distinguish the axial
and peripheral regions of this layer. The axial mesoderm
develops on both sides of the notochord from the ectoblasto-
genous rudiment already described; but the cells which split
off from the endoderm also take part in the formation of these
mesodermal rudiments. The peripheral mesoderm is formed
from the endoderm cells. To the peripheral mesoderm like-
wise belongs that at the posterior margin of the primitive
streak, which also arises from the cells of the endoderm.
The following important conclusions result from my inves-
tigations :—
(1) In the invagination in the case of Amphiorus two
different processes are to be distinguished :—(a) ‘The invagi-
nation of the endoderm cells which form the intestine (the
palingenetic process or gastrulation); (b} the dorsal invagi-
nation of the ectoderm cells, which constitutes the ectoblasto-
genous rudiment of the notochord and mesoderm (the ceno-
enetic process).
(2) In all Vertebrates these two processes can be distin-
guished—the origin of the intestine from the endoderm cells,
and the formation of the rudiment of the notochord and
mesoderm from a proliferation of the cells of the ectoderm.
Ann. & Mag. N. Hist. Ser. 6. Vol. xi. 27
374 Herr B. Lwoff on the Formation of the
In no Vertebrate is the intestine formed by invagination.
The endoderm cells are overgrown by those of the ectoderm,
and the intestine arises through divergence of the former. In
the process of overgrowth a modified gastrulation may be seen.
But the further this process extends the more it is suppressed,
In meroblastic ova, where the yolk is very large (Selachians,
Sauropsida), the process of overgrowth is so greatly modified
in consequence of adaptation to the new conditions that
gastrulation is practically entirely suppressed. The dorsal
ectodermal invagination, on the contrary, exists in all Verte-
brates as a characteristic process, which has nothing to do
with the formation of the intestine, and is to be regarded as
a new formation *, that is, as the ingrowth of the ectoblas-
togenous rudiment of the notochord and mesoderm. This
rudiment develops from the cells of the ectoderm and has a
common origin with the rudiment of the nervous system.
(3) In no Vertebrate is a true gastrula to be observed ; it
is therefore impracticable to search for the gastrula mouth in
its proper sense, and to endeavour to discover its dorsal and
ventral lips, since the primitive conditions are practically
entirely obliterated by the new formations. Accordingly I
leave this question out of consideration. I consider all
attempts to homologize the dorsal and ventral lips of the
gastrula mouth in different Vertebrates as forced and un-
natural,
(4) There is only one spot in the germ the homology of
which can be traced in all Vertebrates, namely that trom
which the invagination + of the ectoderm celis proceeds—
in other words, where the ingrowth of the ectoblastogenous
rudiment of the notochord and mesoderm commences.
(5) The so-called neurenteric canal appears to me in a very
different light. It is not a tube of communication between
the nervous system and the intestinal tract. Just as the
margin ot the enveloping layer represents a transitional region
between the cells of the rudiment of the nervous system and
the rudiment of the notochord, so this canal is to be regarded
as the morphological expression of this transition, ‘There-
fore it may be regarded as a canal of communication between
the tube of the nervous system and the notechordal groove or
notochordal cavity, and may be termed the newrochordal
* It will be evident from this that I virtually dissent from the cus-
tomary view, according to which the posterior end of the germ, where the
invagination is to be observed, exhibits more primitive conditions.
+ In the majority of cases it is really not a true invagination, but
merely the ingrowth of the cells or the turning-in of the margin of the
enveloping layer.
Germinal Layers in Vertebrates, 375
canal, This canal owes its origin to the genetic connexion
between the rudiment of the nervous system and that of the
notochord,
(6) The fact that the rudiment of the notochord and of the
mesoderm arises as a continuous mass of cells from a rudi-
ment which also gives origin to the nervous system may be
turned to account for the purposes of phylogeny. It follows
from this that, in attempting to derive the bilaterally symme-
trical Chordata from Cceelenterate-like forms with radial
symmetry, we must remember that the notochord and the
mesoderm situated on either side of it in no way arise from
the dorsal portion of the intestine, but from the ingrowth of
the proliferating ectoderm cells. Since this ectoblastogenous
mesoderm is chiefly utilized for the formation of the muscu-
lature, we may apply the term chordo-muscular to the entire
ectoblastogenous rudiment. Since this rudiment has a
common origin with that of the nervous system, the Chordata
exhibit a direct connexion with the Annelids, in which,
according to the investigations of Kleinenberg, the muscu-
lature also arises from the common neuro-muscular rudiment.
In my opinion the resemblance is still further increased by
the presence of the cephalic shield (or ventral shield, as the
case may be) in Annelids, which, in its position and its
relation to the nervous system and the muscles and in its
structure, is very similar to the notochord. It might then be
assumed that this continuous neurochordo-muscular rudiment
had already made its appearance in the animals which are to
be regarded as the common ancestors of the Chordata and
the Annelids. The notochord may have arisen in these forms
from an ectodermal rudiment as an axial structure which
served for the attachment of the musculature on the one side
and of the nervous system on the other. ‘The further deve-
lopment and differentiation of different portions of this con-
tinuous ectodermal rudiment might have taken different roads.
In the Annelids the notochordal rudiment experienced no
further development. Perhaps, nevertheless, in the case of
certain Annelids a homologue of the notochord may be proved
to exist in the epithelial vesicular tissue *, which, in addition
to a certain similarity of structure, also exhibits some resem-
blance to the notochord in the fact that it serves for the
attachment both of the nervous system and of the lateral
musculature.
The Ascidians have followed another path. In these
* I shall later on make a special communication on the subject of this
notochord-like structure in Annelids,
27*
376 Formation of the Germinal Layers in Vertebrates.
animals the notochord is only found in the tail.. We might
imagine such an animal as an Ascidian larva to have been
developed in the following manner :—The proliferation arising
at the posterior end by multiplication of the cells, which
represents the chordo-muscular rudiment, instead of growing
forwards between the intestine and external covering, as in
Vertebrates, formed an appendage in which the notochord
and the lateral musculature were differentiated from the
common rudiment. It must be pointed out that the lateral
muscles are present only on either side of the notochord, @. e,
only in the tail of the larva; in the body of the Ascidian,
where there is no notochord, they are entirely absent. This
shows with sufficient clearness that the lateral musculature
and the notochord here by no means arise from the elements
of the endoderm (dorsal portion of the intestine) as is
commonly supposed, but, as I think, from a special continuous
ectodermal rudiment. In Amphioxus and the Vertebrates the
proliferation of the ectoderm cells proceeds from the hinder
margin in such a way that the chordo-muscular rudiment
grows forward between the two primary germinal layers, and
thus forms a plate of cells which is situated between the
nervous system and the intestine. The ingrowth of this
rudiment from the posterior margin of the enveloping layer
may be regarded as an ancestral feature, since in the case of
the Ascidians also the proliferation takes place at the posterior
margin. But this process was subsequently capable of being
so tar modified that in certain Vertebrates, in addition to
growth from the margin of the enveloping layer, splitting-off
of the ectoderm cells is also observable. From this point of
view the dorsal wall of the gastrula of Amphioaxus, the dorsal
wall of the corresponding stages in Petromyzon and the
Amphibia, the blastoderm of the Selachians and Teleostean
fishes, and the ectodermal portion of the germinal disk with
the primitive streak and cephalic process of the Amniota,
each represent a continuous rudiment, from which the nervous
system, the notochord, and the lateral musculature are deve-
loped.
a) The fact that the notochord, although it arises from an
ectoblastogenous rudiment, nevertheless enters into a tran-
sient connexion with the cells of the endoderm, may also be
turned to account for the purposes of phylogeny. It points
to the conclusion that the notochord soon after its origin
entered into intimate relations with the intestine, and perhaps
also commenced to serve for its attachment. ‘This con-
nexion, however, is to be regarded as being of a secondary
nature,
ee ——— ———
a a ai ata i
od
Mr. C. J. Gahan on new Longicorn Coleoptera. 377
I desire to set up this conception in opposition to the theory
of gastrulation which at present prevails. I would mention
once more that I reserve for my subsequent paper the full
description of my investigations and the detailed statement of
proof, as well as the consideration of the literature of the
subject. In the present communication I wished to substan-
tiate my view as shortly as possible.
Moscow, Noy. 1892.
LXI.—Descriptions of some new Longicorn Coleoptera from
the Indian Region. By C.J. GAHAN, M.A., of the British
Museum (Nat. Hist.).
[Plate XIX. figs. 4~-7.]
THE new species described in this paper form part of some
recent accessions to the British-Museum collection. Those
from the collections made by Mr. G. Q. Corbett in Burma
and by Mr. A. Bell in South India were presented through
the intermediation of Dr. Andrewes; while most of the re-
maining species described are from the collections made by the
late Mr. E. 'T. Atkinson.
MHolesthes basicornis, sp. n.
Aureo-sericeo pubescens ; prothorace lateraliter in medio subangn-
lato et leviter tuberculato ; dorso transversim undulateque rugoso,
versus medium sulcis duabus antice conjunctis longitudinaliter im-
presso ; elytrorum apicibus anguste truncatis, utrisque bidentatis ;
antennis quam corpore triplo longioribus, articulis 1° 3°que trans-
versim rugulosis, hoc apice subabrupte crassato.
Long. 38, lat. 10°5 mm.
Hab. Andaman Islands (Atkinson Collection).
The underside of the head is crossed by a deep intergenal
groove, which is followed by a second shallower and some-
what ill-defined groove; both grooves are slightly bowed
forwards in the middle. The prothorax is somewhat con-
stricted anteriorly ; it is obtusely angulate and provided with
a very small tubercle at the middle of each side; it is wrinkled
above over almost the whole surface, the rug having a wavy
course in a general transverse direction; two longitudinal
impressions, bending towards the middle line to unite ante-
riorly, cut off a central space which is wrinkled like the rest
378 Mr. C.J. Gahan on new Longicorn Coleoptera
of the surface. The silky pubescence of the elytra is so
arranged as to present the appearance of numerous brighter
and less bright areas varying in lustre with the direction in
which the light falls upon them. The antenne of the male
are about three times as long as the body ; the first joint and
the third are transversely rugulose, the third joint has at the
apex an abrupt ring-like thickening which projects rather
more on the outer side; the fourth and fifth joints are also
slightly thickened at the apex; each of the joints from the
fifth to the eighth is furnished with a small acute spine at the
outer apex, and each from the sixth to the tenth is denticu-
lately produced at the inner apex ; the third joint is slightly
longer, the fourth a little shorter, than the first; the fifth 1s
distinctly longer than the third.
Margites sulcicollis, sp. n.
Niger, griseo sparse setosus ; elytris piceo-brunneis, griseo tenuiter
pubescentibus et fulvo sat dense setosis ; prothorace basi apiceque
constricto, lateraliter rotundato, dorso utrinque quadrisulcato,
medio sparsim punctato; corpore subtus pedibus antennisque
piceis vel nigris, griseo tenuiter pubescentibus; antennis (2 )
quam corpore brevioribus, articulis 1°, 3°, 5°que subsqualibus,
utrisque quam quarto paullo longioribus, articulis 6° et sequentibus
gradatim crescentibus,
Long. 13 mm.
Hab. Paungdé, in Burma (G. Q. Corbett).
Prothorax strongly enough rounded at the sides in the
middle, narrowed at the base and apex; the disk with two
very distinct longitudinal grooves which cut off a median,
somewhat oblong, and sparsely punctured area from two
lateral plage, each of which is traversed longitudinally by
three or four narrower and less distinct grooves, the intervals
between which are narrow and cariniform, the sides below
these areas are intricately and not strongly rugose. The
elytra bear punctures of two kinds—larger and less closely
crowded punctures from which the suberect fulvous sete
spring, and minute closely crowded punctures occupying
the intervals and giving rise to the finer hairs of the laid
pubescence.
This species is to be recognized by the peculiar sculpturing
of the prothorax.
Nyphasia apicalis, sp. n.
Fulvescens ; elytris apice chalybeato-violaceis, utrisque ad suturam
breyiter spinosis ; antennis ( ¢ ) quam corpore sesqui-longioribus,
a
1 nee ype
Dn ail
from the Indian Region. 379
articulis 3° ad 6" nigris, apice extus spinosis, articulo 7° brevius
spinoso, hoe et sequentibus testaceo-fuscis ; corpore subtus pedi-
busque rufo-fulvescentibus.
Long. 15 mm.
Hab, Canara, in South India (A. Bell).
This species agrees closely in structural characters with
N. Pascoet, Lac., and N. fuscipennis, Gah., but has the pro-
thorax somewhat more strongly tuberculate at the middle of
each side, the scutellum more prolonged behind, and the elytra
each furnished with a short sutural spine at the apex. It may
be further distinguished by its coloration, being entirely of a
somewhat reddish fulvous, with the exception of a small
portion at the apex of the elytra which is violet-blue, and
of the antenne, of which the two proximal joints only are
fulvous.
Although this species has the elytra spined at the apices, it
accords in other respects much better with Lacordaire’s second
section of the genus, the remaining species of which have the
elytra unarmed at the apex.
Cleonice bivittata, sp. n.
Viridis, pedibus antennisque atro-ceeruleis ; elytris creberrime punc-
tatis, viridi-opacis, vitta dorsali utrinque wneo-viridi, apice con-
junctim subacuminate rotundatis; antennis (¢) quam corpore
multo longioribus ; articulo tertio quam quarto sequi-longiore,
articulis primo tertioque sulcatis.
Long. 19, lat. 4°75 mm.
Hab, Baram, in North-west Borneo (Everett).
Head rather sparsely punctured in front, more densely
above and on the sides ; impressed along the middle in front
with a linear groove which extends backwards above, between
the antennz and eyes, as far as the middle of a small sparsely-
punctured dorsal area. Prothorax thickly punctured above,
except along the anterior margin, on a small centro-dorsal
area, and over each of two feebly raised and obtuse tubercles
which are placed towards each side. LKlytra very closely
punctured, of a rather dark dull green colour, but each with a
dorsal vitta of a somewhat brassy and brighter colour. The
body underneath isalso green, but is clothed with a faintsilvery-
grey pubescence. ‘The legs and antenne are dark blue, the
peduncles of the femora only being greenish. ‘The antennae
of the male are more than half as long again as the body ;
the first and third joints only are sulcate along their anterior
or inner surface; the third is about half as long again as the
fourth.
380 Mr. C. J. Gahan on new Longicorn Coleoptera
The colour and punctuation of the elytra in this species are
very suggestive of the South-African genus Ochimus. Iam
not quite certain that the species is congeneric with Cleonice
vestita, Thoms. (= Seuthes sericatus, Pasc.), as I have no
example of the latter by me with which to compare it.
Epepeotes andamanicus, sp. n.
Niger, griseo tenuissime pubescens ; prothorace transverso, lateraliter
utrinque spinoso, dorso medio leviter transversim rugosulo ;
elytris punctatis, punctis versus basin asperatis, apicibus sinuato-
truncatis.
Long. 25 mm.
Hab. Andaman Islands (Prof. Meldola and Atkinson Coll.).
Black, with a faint slate-grey pubescence. Head and pro-
thorax almost impunctate; the latter feebly wrinkled over
the middle of the disk, armed with a sharp tubercle or spine
on the middle of each side. Elytra densely punctured, the
punctures being more closely crowded and asperate on the
basal portion, and becoming gradually smaller towards the
apex.
P This species has some resemblance to Paragnoma nigra,
Thoms., with which it was confounded in the collection of the
late Mr. E. Atkinson, but may be distinguished by the shorter
and more transverse prothorax and the lateral spines with
which it is armed.
Epepeotes uncinatus, Gahan.
A variety of this species, represented by a single female
specimen taken by Mr. Alexander Bell at Canara in South
India, differs from the described form in having the black
spots of the elytra much more elongated; those along each
side of the suture especially, forming for some distance an
almost continuous vitta. ‘This variety has a remarkable re-
semblance to Macrochenus tigrinus, Oliv., which occurs also
in South India as well as in Ceylon. An analogous case of
resemblance is furnished by Macrochenus birmanus, Thoms.,
and Lpepeotes guttatus, Guér.
Macrochenus Atkinsont, sp. n.
Niger ; capitis fronte, lateribus vittaque dorsali, vittis tribus protho-
racis (una dorsali, una utrinque laterali), scutello et maculis
irregularibus elytrorum fulvo-pubescentibus ; corpore subtus
nigro, vitta utrinque abdominis et maculis thoracis fulvo-pubes-
centibus ; apicibus elytrorum sinuato-truncatis. (¢ ) Prothorace
TE
From the Indian Region. 381
quam latiori sesqui-longiori. (9) Prothorace quam latiori vix
longiori.
Long. (3) 23, (2) 29 mm.
Hab. Andaman Islands (Atkinson Collection),
Black, covered with a faint dark grey or brownish indu-
mentum. Sides and front of the head and a dorsal median
vitta fulvous. Prothorax with three fulvous vitte, one medio-
dorsal, and one, low down, on each side; transversely wrinkled
above and rather sparsely punctured. Elytra rather strongly
punctured at the base aud for a short distance behind each
shoulder, more feebly and sparsely punctured over the rest of
their surface, especially towards the apex; adorned with a
somewhat variable number of spots of fulvous pubescence
which are of different sizes and very irregular in shape. ‘The
apices of the elytra are sinuately truncate, with the outer
angles very feebly denticulate.
This species agrees in its general structure with JL Guerini,
White, but may be easily distinguished by its different style
of marking.
Thestus armatus, sp. n.
Pube fulvo-ferruginea obtectus; prothorace lateraliter in medio
valde spinoso ; elytris basi minute sat denseque granulatis, fasciis
duabus cinereis transversim arcuatis ornatis ; antennis (¢ ) quam
corpore duplo longioribus,
Long. 35, lat. 11:5 mm.
Hab. Sandakan, in North-west Borneo (Atkinson Col-
lection).
This species seems to have the closest resemblance in
colour and pattern of marking to Thestus oncideroides, Pasc.
The granules on the basal portion of the elytra are closer
together than they are represented to be in the figure of the
latter species. ‘The present species differs chiefly, however,
in having a strong spine on each side of the prothorax. In
T’. oncideroides the prothorax has but a feeble tubercle or tooth
on the middle of each side.
Eutenia Corbett, sp. n.
Nigra; capitis fronte lateribusque, fasciis duabus prothoracis medio
interruptis, scutello et maculis vittisque elytrorum, fulvescente-
pubescentibus ; corpore subtus pedibusque plus minusve fulvyes-
centibus ; antennis (¢) quam corpore multo longioribus, nigris,
articulo primo crasso, apice late cicatricoso, articulis 3° ad 5%™
subzequalibus, utrisque quam primo sesqui-longioribus, articulis
6° 7°que paullo decrescentibus (ceteri desunt),
Long. 18 mm,
382. Mr. C. J. Gahan on new Longicorn Coleoptera
Hab. Tharawaddy, in Burma (G. Q. Corbett).
Deep black ; with the front and sides of the head, two
transverse bands of the prothorax, one anteriorly, the other at
the base, both broadly interrupted in the middle, the scutellum,
six spots and two posterior vittea on each elytron, covered
with a dense yellowish-tawny pubescence. Of the six spots
on each elytron three form a transverse row a little behind
the base, the innermost spot being much smaller than either
of the two exterior; two spots are placed obliquely near the
middle, with the inner spot farther back than the outer; the
remaining spot is small and triangular and is placed close to
the suture in a line with, and a little farther back than, the
innermost spot of the anterior row. The two posterior vitta,
of which that nearest the suture is shorter and broader,
narrow behind and unite at the apical border. The black
portions of the prothorax and elytra, which are made opaque
by a black powdery or scaly pubescence, are seen to be rather
thickly and strongly punctured. ‘The body underneath and
parts of the legs are clothed with a less dense pale fulvous
pubescence.
The spots and vitte of this species will probably be found
to vary. ‘The entirely black antenne, together with the style
of marking, distinguish it at once from other species of the
genus.
POLYTRETUS, gen. nov.
Head with the front rectangular, slightly higher than its
breadth, with the antennal tubercles rather prominent, slightly
divergent, and at their base approximate, so that the concavity
between them is deep and rather narrow. Eyes moderately
large, their upper lobes narrow, their lower lobes extending
downwards more than halfway to the base of the mandibles.
Antenne about half as long again as the body, with the scape
moderately elongate and gradually and not strongly thickened
to the apex, where it is provided with a narrow and completely
margined cicatrice ; third joint about half as long again as
the scape; the fourth and following joits gradually dimi-
nishing in length ; the distal halves of the third and fourth
joints thickly fringed with hairs on the outer side. Prothorax
slightly transverse, furnished at the middle of each side with
a rather short conical tubercle which points somewhat obliquely
upwards. Elytra rather elongate, with the sides subparallel
and the apices rounded. At the base, between the scutellum
and the shoulder, each elytron presents a small angular lobe
which projects forwards and slightly overlaps the base of the
Srom the Indian Region. 383
prothorax. Legs subequal to one another in length. Inter-
mediate tibie with their outer margin entire. Claws of the
tarsi broadly divergent, but not strictly divaricate. Pro-
sternal process almost horizontal, being but feebly raised
towards the middle of its length ; it is very slightly widened
posteriorly and just at its hind extremity bears on each side
a very small process which does not reach to the free end of
the epimeron ; so that the anterior acetabula remain slightly
open behind. ‘The mesosternum is gradually narrowed pos-
teriorly, and bears a small but distinct tubercle.
This genus must, in accordance with the system of
Lacordaire, be placed in the group of the Monohammides,
from all the other members of which it may be very readily
distinguished.
Polytretus cribripennis, sp.n. (Pl. XIX. fig. 4.)
Piceo-niger ; antennis articulis tertio quartoque nigro-fimbriatis ;
prothorace nitido fere impunctato, disco ineequali, postice tuberculo
medio paullo elevato et supra planato munito; elytris fulvo-tes-
taceis, crebre foyeato-punctatis, utrisque lineis tribus—duabus
dorsalibus, tertia laterali—paullo elevatis.
Long. 11°5, lat. 3°5 mm.
Hab, Mungphu, in Sikkim (Atkinson Collection).
Head, prothorax, body underneath, legs, and antenne
black, with a slightly reddish tint at the margins of the pro-
thorax and of the abdominal segments. The antenne are
dull in colour, being covered with a faint blackish pubescence ;
the third and fourth joints are thickly fringed with blackish
hairs on their outer and lower border throughout the greater
part of their length. ‘The disk of the prothorax has on each
side, in front of the middle, a small convex polished area,
which is separated from its fellow by a median slightly
depressed area ; between the middle and the base there is a
median slightly raised area which is flattened and not very
smooth above, and which has a rounded and sharply-limited
posterior border. The elytra are very strongly and closely
punctured throughout, and are entirely of a fulvous-testaceous
colour ; they are rounded at the apex, and each bears three
slightly, and not very distinctly, raised lines—two dorsally
and one towards the side.
Anamera fulvescens, sp. n.
Fusca, supra pube fulvescente sat dense obtecta; elytris dense
punctatis, viridi-fuscis, fulyo-pubescentibus ; corpore subtus fulyo-
cinereo-pubescente.
Long. 23, lat. 8°5 mm.
384 Mr. C. J. Gahan on new Longicorn Coleoptera
Hab. South India (Atkinson Collection).
Head and prothorax very sparsely punctured, covered with
a rather close fulvous pubescence ; their derm, where exposed
by the rubbing away of the pubescence, is seen to have a
dark brown colour. The elytra are closely punctured, and
are clothed with a pale fulvous pubescence. ‘The underside
of the body is paler than the upperside, being covered with
an ashy fulvous pubescence. ‘The first joint of the antenne
is dark brown with a grey pubescence, the third and following
joints are testaceous with a pale grey pubescence.
Two species only have hitherto been placed in the genus
Anamera, viz. A. alboguttata, Thoms., and A. concolor,
Lacord. (Gén. des Coléop. ix. p. 883. 1), in both of which
the labrum is arcuately emarginate and the upperside of the
thorax and elytra furnished with rather long, erect, black
hairs—characters not found in the present species. But when
this species is compared in its general structure with A. con-
color there seems little reason to doubt that the two are con-
generic. The genus Anamera was placed by Lacordaire in
the group of the Mesosides; but why in this group, rather
than in the Monohammides, I fail to understand: the cica-
trice of the antennal scape is completely margined, the lateral
spines of the prothorax are median in position, the interme-
diate tibie grooved, and the claws of the tarsi divaricate.
PARAGNIA, gen. nov.
Head with the front widening towards the base, with the
antennal tubercles prominent, slightly diverging and separated
from one another by a rather narrow wedge-shaped interval
above. Eyes emarginate, with their lower lobes rather small.
Antenne a little longer than the body; with the scape
thickened, slightly curved, and furnished at the apex with a
cicatrice the limiting carina of which is incomplete; third
joint about half as long again as the scape, distinctly thick-
ened throughout nearly the whole of its length; fourth a
little shorter than the third, the fifth and following gradually
decreasing in length. Prothorax slightly transverse, some-
what narrowed towards the base; its sides unarmed. LElytra
rounded at the apex. Legs subequal to one another in length,
Intermediate tibiz with a distinct oblique groove just below
their middle. Claws divaricate. Prosternal process rather
feebly arched and narrow between the coxe, gradually dilated
behind up to its posterior border. Mesosternal process nar-
rowing posteriorly, bearing a feeble tubercle between the
coxe.
—
Srom the Indian Region. 385
The shape of the head seems to determine the position of
this genus in the group of the Agniides. The absence of
spines or tubercles from the sides of the prothorax and the
incomplete margin to the cicatrice of the antennal scape are
two characters which will distinguish the genus from the other
members of the group, and which at the same time seem
to point to an affinity with the Mesosides.
Paragnia fulvomaculata, sp.n. (Pl. XIX. fig. 5.)
Ferrugineus ; capite fronte crebre minuteque punctulato, lateraliter
et supra minus dense punctato ; prothorace sat sparse punctato et
dense minuteque rugosulo ; elytris a basi ad medium grosse cre-
breque punctatis, subrugosis, pone medium minus fortiter sat
denseque punctatis, utrisque maculis 12-14 fulvyo-auratis ; an-
tennis quam corpore longioribus, articulis 4°-11"™ apice infuscatis,
basi grisescentibus.
Long. 14, lat. 5 mm.
Hab. Mungphu, in Sikkim (A¢tkinson).
Ferruginous red. Head minutely and very closely punctured
in front, more sparingly and strongly punctured on the sides
and vertex. Prothorax rather thickly punctured, and with
numerous short and minute ridges which have a general
transverse direction. Elytra with very large and closel
placed punctures occupying almost the whole of the basal half ;
behind the middle, and for a short distance along the suture
in front of the middle, the punctures are much smaller and
less dense, while towards the apex they become still smaller
and sparser; each elytron has about twelve or fourteen
irregularly arranged spots of golden-tawny pubescence, nearly
halt of these spots being very small and punctiform. The
underside of the body is coloured like the upperside, but the
sides of the meso- and metathorax are somewhat blackish.
KUSEBOIDES, gen. nov.
Elongate. Head with the front slightly transverse and
somewhat widened towards the base. Eyes emarginate, their
lower lobes rather small. Antenne about half as long again
as the body; first joint subcylindrical, reaching to a littie
beyond the middle of the prothorax ; third joint about equal
in length to the first, the fourth a little longer; the fifth and
following joints gradually decreasing in length. Prothorax
cylindrical, parallel-sided, its length scarcely appreciably
greater than its breadth. LElytra elongate, gradually nar-
rowing from the base backwards; their apices narrowly
386 =Mr. C. J. Gahan on new Longicorn Coleoptera
truncate near the suture and prolonged at the outer angles
into a rather strong spine; each elytron with a slightly raised
and obtuse dorsal carina which runs at a short distance from
the suture, and is separated from it by a shallow channel-like
depression. Intermediate tibie with an oblique groove just
below the middle of their length. Prosternal process feebly
arched in the middle and rather widely dilated behind.
Mesosternum with its intercoxal process rather narrow and
almost parallel-sided.
This genus is allied to Nyctimene, Thoms., but may be
distinguished by its shorter prothorax, the shorter scape of its
antenne, and the relatively longer fourth joint. Its legs also
are longer, the hind femora being almost equal in length to
the two first abdominal segments, while in Nyctimene they
are not longer than the first segment.
Luseboides plagiatus, sp.n. (Pl. XIX. fig. 6.)
Fuscus, fulvo-brunneo haud dense pubescens ; elytris utrisque plagis
tribus pallidioribus—una prope basin, secunda vix pone medium
nonnihil albescente, tertia prope apicem quoque albescente.
Long. 16-18 mm.
Hab. Mungphu, in Sikkim (Atkinson Collection).
- Dark brown, with a not very dense fulvous-brown pubes-
cence. Head thickly punctured. Prothorax somewhat more
strongly punctured. Elytra rather thickly punctured, each
with three rather ill-defined and somewhat broken patches of
paler pubescence separated from one another by two areas
covered with dark brown pubescence. ‘Ihe first paler patch,
somewhat fulvous in colour, is near the base; the second, of
a dirty white colour, is just behind the middle; the third,
similar in colour to the second, is near the apex. Breast
rather strongly and sparsely, the abdomen more feebly, punc-
tured, each of the punctures being marked in position by a
minute rounded dark brown spot.
Enispia bella, sp.n. (Pl. XIX. fig. 7.)
Q . Capite fusco, crebre punctato; antennis 12-articulatis, ferrugineo-
fuscis, griseo pubescentibus et sat longe ciliatis, basi dense punc-
tatis, articulo quarto testaceo, albido pubescente et ciliato; pro-
thorace dense cinereo-hirsuto ; elytris fuscis, basi et vitta longi-
tudinali, leviter arcuata, rufo-brunnescentibus, griseo pubescen-
tibus et setosis, fascia transversa paullo pone basin et fasciis
duabus leviter arcuatis paullo ante apicem albidis ; corpore subtus
pedibusque testaceis, cinereo tenuiter pubescentibus, his longe
Srom the Indian Region. 387
setosis ; lateribus metathoracis et femoribus quatuor anticis paullo
infuscatis,
Long. 8°5 mm,
Hab. Burma (G. Q. Corbett).
Head dark reddish brown, very closely punctured and with
a thin greyish pubescence. Antenne longer than the body,
twelve-jointed, with the first joint about equal in length to
the third and distinctly longer than the fourth, with the joints
from the fifth to the eleventh subequal or very slightly
diminishing in length, each shorter than the fourth; the
twelfth joint distinctly shorter than the eleventh; the fourth
joint with its proximal half testaceous and clothed with a
whitish pubescence. Prothorax slightly constricted near the
base and still more slightly near the apex, clothed with a
dense yellowish-cinereous pubescence, which is raised in the
form of two small tufts on the middle of the disk; the sides
and two small patches near the base are less densely pubes-
cent, are darker in colour, and are seen to be rather thickly
punctured. ‘The elytra are mostly of a dark brown colour,
with the base, a dorsal (slightly arcuate) vitta on each side,
and a narrow lateral border reddish brown; they are rather
strongly and thickly punctured on the basal two thirds, with
the punctures arranged, some irregularly, others in tolerably
definite rows, one of which accompanies the dorsal reddish-
brown vitta ; the pubescence is mostly greyish in colour, but
forms three narrow white bands—one transverse and slightly
zigzag, forming a border behind to the basal reddish-brown
portion, two posteriorly, which, as they approach the suture,
separate from one another and again converge, so as to
enclose a somewhat rounded space with a dark spot in its
centre,
Eunidia lateralis, sp. n.
Supra pube fulvescente dense vestita, vitta lata utrinque a capite ad
apicem elytrorum extensa nigro-fusca ; corpore subtus pedibusque
cinereo tenuiter pubescentibus ; antennis nigro-fuscis.
Long. 6°5 mm.
Hab. Canara, in South India (A. Bell).
This species resembles Hunidia simplex, Gahan, but may
be readily distinguished by the rather broad and distinct dark
brown vitta which runs along each side from the head to the
apex of the elytra: this vitta has a nearly straight upper
margin ; it is narrowed posteriorly in correspondence with the
narrowing of the elytra towards the apex, and it ends at the
angle formed by the slightly curved postero-external margin
388 Mr. C. J. Gahan on new Longicorn Coleoptera.
of the elytron with the oblique inner or apical margin. The
body underneath and the legs are black, with a faint ashy-
grey pubescence. ‘The front of the head is grey in its lower
portion, fulvous towards the vertex. The antenne are almost
black and are about half as long again as the body.
Glenea Andrewest, sp. 0.
Testacea, pube griseo-cervina sat denso vestita; capite medio nigro-
lineato; prothorace vittis sex interruptis aut maculis duodecim
nigris ornato ; elytris utrisque maculis duabus—una basali, altera
paullo pone medium—et postice vittis tribus angustis, nigro-
fuscis; apicibus truncatis, extus brevissime dentatis.
Long. (¢) 11, (2) 15-16°5 mm.
Hab. Canara, in South India (A. Bell).
This species is allied to and rather closely resembles
G. spilota, Thoms., but may be distinguished by the different
position and arrangement of the dark markings on the elytra.
One rounded black spot is just behind the humeral depres-
sion at the base of each elytron ; a second rounded black spot
is placed nearer the suture just behind the middle of the
elytron, and from near its outer border two narrow dark
brown vitte proceed backwards for a short distance, while a
third brownish vitta arises a little behind its inner border and
runs close alongside the suture to within a very short distance
of the apex. ‘The apices of the elytra are very briefly and
scarcely perceptibly toothed at the outer angles.
Glenea Belli, sp. n.
Nigra, dense punctata, vittis maculisque argenteo-viridi-squamosis ;
elytris lateraliter utrinque unicarinatis, apicibus oblique truncatis
angulis dentatis.
Long. 9°5 mm.
Hab, Canara, in South India (A. Bell).
Head rather thickly punctured, black, with the sides and
two longitudinal bands in front, united across the base, silvery
green. Prothorax more thickly punctured, with a median
longitudinal silvery-green band and with some band-like
spots of a similar colour on each side. Scutellum silvery
green. Elytra with a distinct carina on each side separating
the vertical lateral portion of the elytron from the slightly
convex or nearly horizontal disk. The vertical lateral portion
bears two rows of punctures divided by a slightly raised line ;
the disk is closely and irregularly punctured, with the punc-
fe SO pein Ramee 4? 5-4 Cop re
On some allied Pentatomidex. 389
tures becoming sparser posteriorly and disappearing near the
apex. ach elytron has six silvery-grey spots, of which two
are in the same transverse line at a short distance behind the
base, the third is just in front of the middle and is placed
transversely, with its inner extremity rather close to the
suture ; the fourth and fifth are smaller and more remote from
the suture, the fourth just behind the middle, the fifth about
midway between this and the apex ; the sixth is at the apex,
and from its inner edge a narrow sutural vitta passes forwards
for about half the length of the elytron, The body under-
neath and legs are greyish, with the anterior portion of each
of the metathoracic episterna, an anterior band on each of
the first four abdominal segments, and the whole of the
fifth segment so faintly pubescent as to appear black and
subnitid.
EXPLANATION OF PLATE XIX. Fies. 4-7.
. Polytretus cribripennis (3).
Fig. 5. Paragnia fulvomaculata.
Fig. 6. Euseboides plagiatus.
7. Enispia bella.
~
LXII.—On some allied Pentatomide, with Synonymical
Notes. By W. L. Disrant.
Dalpada mirabilis, sp. n.
Olivaceous green ; a central elongate spot at base of head,
lateral margins, a central fascia (not extending beyond centre),
and a narrow transverse fascia on anterior area of pronotum,
five small basal spots and an obscure posterior central fascia
to scutellum, lateral margins, apex, and the whole inner area
of corium castaneous. Membrane very dark castaneous, its
apex paler. Body beneath and legs ochraceous; lateral
margins of head, sternum, and abdomen somewhat broadly
olivaceous green; sternum with transverse spots of the same
colour and with two large black spots between the anterior
and intermediate coxe. Intermediate and posterior femora
spotted with castaneous, posterior tibiee with the base and
apex castaneous. Anterior legs, intermediate tibic, and the
antenne mutilated. Abdomen beneath with a central black
spot on the pate segment. Lateral angles of the
pronotum moderately prominent and nodulose; lateral lobes
of the head slightly longer than the central lobe.
Ann. & Mag. N. Hist. Ser. 6. Vol. xi. 28
390 Mr. W. L. Distant on
Long. 25 millim. ; exp. pronot. angl. 13 millim.
Hab. India, Naga Hills.
I have for some years possessed a single mutilated specimen
of this extremely fine species, and hoped to have procured a
more perfect example. As I have not succeeded in doing so,
I now describe sans antenne and anterior legs. It cannot
be mistaken, and there is no other species of the genus which
resembles it either in size or colour.
Dalpada insularis, sp. n.
Ochraceous or reddish ochraceous, coarsely punctured with
greenish black. Head with the punctures thickest on the
lateral Jobes and on the margins of the central lobe ; antennz
pale castaneous, bases of the fourth and fifth joints broadly
ochraceous. Pronotum with the lateral margins strongly
serrate, the lateral angles, which are moderately prominent
and robust, black, with their apices ochraceous. Scutellum
with three obscure and ill-defined pale basal spots. Head
and corium more finely punctate than the pronotum and
scutellum. Membrane cupreous. Connexivum reddish
ochraceous, greenish black at the incisures. Body beneath
and legs reddish ochraceous; head and sternum ornamented
with fascie of greenish-black punctures. Abdomen with a
broad submarginal fascia of green punctures. Anterior tibiz
dilated, with their apical halves sometimes darker in hue.
Long. 15-19 millim. ; exp. pronot. angl. 83-10 millim.
Hab. Malayan Archipelago: Ombay, Putar.
Coctoteris exiguus, sp. n.
Ochraceous. Head coarsely punctate; antenne with the
first and second joints ochraceous, outwardly margined with
black, third, fourth, and fifth joints black, the base of the
third joint brownish ; pronotum (excluding the anterior area)
with exceedingly coarse black punctures, and with two central
black spots on anterior margin; scutellum with the anterior
half with exceedingly coarse black punctures, the central
basal area impunctate, posterior half more finely and sparsely
punctate ; an elongate black fovea at each basal angle; corium
somewhat thickly and finely punctate; membrane pale
fuscous with the margins paler, or uniformly brownish ochra-
ceous with the veins darker. Body beneath and legs ochra-
ceous; lateral areas of the head and sternum coarsely but
sparingly punctured with black; two small black spots on
the lateral areas of the pro-, meso-, and metasternum. Abdo-
minal spiracles black.
some allied Pentatomide. 391
The central lobe of the head is very little longer than the
lateral lobes; the pronotal angles are moderately and sub-
acutely produced ; the lateral pronotal margins are somewhat
obscurely denticulated; the posterior angles of the sixth
abdominal segment are only moderately produced.
Long. 13 millim.; exp. pronot. angl. 74 millim.
Hab, New Guinea (Goldie) ; Duke of York Island.
This species is rather aberrant in its generic characters, but
the position of the ocelli alone will show that it is a Coctoteris
and not a Spudeus.
Halys neelgiriensis, sp. n.
Ochraceous, thickly punctured with brassy black; head,
anterior area and lateral angles of pronotum metallic green ;
antenne black, the joints very narrowly ochraceous at base,
fourth joint annulated with ochraceous near base (fifth joint
mutilated). Head with a spot at apex, an oblique spot on
each side near eyes, and a reversely directed oblique spot on
each side near base ochraceous; eyes castaneous, with their
inner margins ochraceous; ocelli red. Pronotum with the
anterior and lateral margins ochraceous, and with a dull
ochraceous spot on the green lateral angles. Scutellum with
five small ochraceous spots at base and the apex ochraceous.
Connexivum ochraceous, broadly greenish black at the inci-
sures. Body beneath and legs ochraceous, lateral margins of
head and sternum broadly metallic green ; femora and sub-
lateral areas of abdomen darkly punctate. Abdomen with
small marginal spots at incisures and a spot on the three
ultimate segments of the abdomen ; tibiz with outer spots at
base, centre, and apex, and the apices of the tarsi black.
Long. 20 millim.
Hab. India, Neelgiri Hills (Hampson).
This second Indian species of Halys is distinguished from
H. dentata, Fabr., not only by the colour differences, but by
the head being much broader, especially at the apex, and its
lateral margins more strongly toothed; the lateral margins
of the pronotum more finely erenulate ; the apical third of the
scutellum much more elongate and narrow; second joint of
the antenne much shorter than the third, &c.
NEVISANUS, gen. nov.
Head about as long as central portion of the pronotum,
broad, the lateral margins moderately concavely sinuate, but
not toothed, the anterior margin truncate, the central and
28*
392 Mr. W. L. Distant on
lateral lobes equally long. Antenne a little more than half
the length of the body, five-jointed, inserted about halfway
between the eyes and apex ; the second and third joints rather
longly pilose, the basal joint reaching the apex of the head,
second and third joints subequal in length and a little shorter
than the fourth joint. Rostrum slightly passing the posterior
coxe, third joint longest, fourth a little shorter than the first.
Body ovately elongate, depressed. Pronotum with the ante-
rior margin strongly concave, the lateral margins entire or
dentate, sinuate near centre, the lateral angles subprominent.
Scutellum subtriangular, passing the centre of the abdomen,
moderately tumid at base and gradually narrowing to about
one fourth before apex, which is convexly rounded. Corium
with its apical. margin moderately sinuate. Membrane with
the veins robust and longitudinal. Connexivum slightly
notched or toothed at the incisures. Abdomen beneath with
a broad basal furrow. Legs pilose ; tibiz suleated.
Allied to the African genera Canomorpha and Scribonia.
Nevisanus orientalis, sp. n.
Ochraceous, thickly punctured with brassy black, Head
with the lateral lobes, the margins of the central lobe, and
the base very thickly punctate. Antenne black, the base of
the third joint narrowly and the bases of the fourth and fifth
joints broadly ochraceous. Pronotum rugulose and thickly
punctate, with the extreme anterior and lateral margins
ochraceous, the last with the margins at lateral angles black.
Scutellum with a levigate spot (variable in size) at basal
angles, and a small central levigate basal spot ochraceous, apex
and acentral fascia from about centre very sparingly punc-
tate. Corium thickly punctate, especially on disk and near
base. Membrane black, with some apical pale spots. Con-
nexivum ochraceous, broadly brassy black at the incisures.
Body beneath and legs ochraceous, the lateral areas of the
sternum and abdomen sparingly punctate ; longitudinal fascize
to head, transverse fascia to sternum, abdominal segmental
margins, duplex marginal spots at the incisures, and a trans-
verse streak near spiracles black. Legs with the femora
punctured with black; tibie outwardly black at base and
apex ; apex of the tarsi black.
Long. 18-20 millim.
Hab. India, Sikkim, Khasia Hills.
I Ee
some allied Pentatomide, 393
1aeey y
Nevisanus nagaensis, sp. n.
Ochraceous ; head, pronotum, and scutellum thickly covered
with greenish-black punctures, the corium more sparingly so.
Pronotum with the lateral margins coarsely dentate, the lateral
angles subacutely prominent. Scutellum strongly trans-
versely wrinkled at base, and with three pale spots on basal
margin. Membrane cupreous, with the apex paler and the
veins black, with subapical black spots between them. Con-
nexivum ochraceous, greenish black at the incisures. Body
beneath as above, but with the punctures much scarcer on
the disk of the abdomen. Legs ochraceous, spotted with
black. Antenne mutilated.
Long. 20 millim.; lat. pronot. angl. 10 millim.
Hab. India, Naga Hills.
Nevisanus tectus.
Dalpada tecta, Walk. Cat. Het. i. p. 224. n. 17 (1867).
Hab. India, Silhet. Brit. Mus.
Apodiphus integriceps.
Apodiphus integriceps, Horvath, Rey. d’Ent. vii. p. 172 (1888).
This species was described from Turkestan. I possess two
specimens from Silgeet, in Dardestan of the Kashmeer State,
which agree fairly well with Dr. Horvdth’s description and
are certainly quite distinct from A. amygdali, Germ. The
anterior margin of the head is less cleft between the apices of
the lateral lobes than in Germar’s species.
SynonyMICAL NOTES.
Dalpada collocata, Walk. Cat. Het. i. p. 221. n. 13 (1867),
belongs to the genus Halyomorpha and is very closely
allied to H. picus, Fabr., if not but a variety of that
species.
Dalpada apicifera, Walk. Cat. Het. 1. p. 222. n. 14 (1867),
=WDalpada varia, Dall. List Hem. i. p. 185. n. 8
(1851).
Walker’s specimens are from Hong Kong and have the
appearance of being discoloured by immersion in spirit.
Dalpada bulbifera, Walk. Cat. Het. i. p. 223. n. 16 (1867),
= Dalpada clavata, Fabr.
Walker’s type! is in far too mutilated a condition to have
YP
394 Mr. G. Lewis on a new Beetle from Japan.
warranted description; and, although -he allies it with his
D. nodifera, the shape and structure of the head is quite
different.
Dalpada brevivitta, Walk. Cat. Het. i. p. 224, n. 18 (1867).
This species does not, as stated by Walker, agree “in
structure”? with his D. tecta, which, as already mentioned,
belongs to my genus Nevisanus. The shape of the head is
totally different, and locates it rightly in the genus Dalpada.
Dalpada consobrina, Walk. Cat. Het. i. p. 225. n. 20 (1867),
= Dalpada clavata, Fabr.
Dalpada brevis, Walk. Cat. Het. i. p. 226, n. 21 (1867),=
Dalpada remota, Walk. ibid. p. 227. n. 22,=Dalpada
proxima, Walk. ibid. p. 227. n. 23,= Halyomorpha
ptcus, Fabr.
Dalpada japonica, Walk. Cat. Het. i. p. 228. n, 24 (1867),
= Lrthesina fullo, 'Vhunb.
LXIII.—On a new Beetle from Japan (Omaliide).
By G. Lewis, F.L.8.
THE curious insect to which this note refers was not submitted
to Dr. Sharp while he was writing his second memoir on the
Staphylinidee of Japan in 1889, for by an error in grouping it
was set aside as belonging to the Silphide; and it was only
on my coming to write a description of it as such that I
discovered my mistake. I then sent an example to Dr. Sharp,
who kindly returned it to me with this memorandum :—“ You
may say in your description that it is the only species in the
Staphylinide yet known in which the elytra are entire, not
truncate, and completely cover the abdomen. In Trigono-
desmus and Lathrimeum the wing-cases are truncate and
leave the tip of the body exposed. It has a pair of ocelli,
which brings it into the Omaliide.”
CAMIOLEUM, gen. nov.
Resembles Lathrimaum and Olophrum in many of its
characters, especially in the antenne, palpi, and mouth-
organs as they appear without dissection. ‘orm depressed ;
the head somewhat transverse ; eyes prominent and circular in
outline ; two ocelli before the neck; the thorax also a little
transverse and widening out laterally, the expanding portions
Mr. G. Lewis on a new Beetle from Japan. 395
being equal to one third of the whole breadth; the elytra
straight at the base, but otherwise completely oval in outline,
sides much dilated and formed, especially in the epipleure,
like Silpha atrata, L.; viewed from beneath they are widest
behind the humeral angle, and behind the posterior cox the
marginal expansion narrows and gradually ceases before the
apex. The legs are slender and doubtless vary in structure
in the sexes.
Je ‘
Camioleum loripes, sp. n.
Alutaceum, parum nitidum, depressum ; elytris ovatis haud abbre-
Viatis, striato-punctatis, marginaliter dilatatis.
L. 4 mill.
Alutaceous ; disks of the head, thorax, and dorsum, with
five terminal joints of the antenne, darker; the antenne are
slender, with the joints relatively longer than in Olophrum
simplex, Sh. (to compare them with a very common Japanese
species), but otherwise the structure of the articulations is
sim.lar; the head uneven between
the eyes, with irregular and _scat-
tered punctures ; punctures smaller
and fewer between the antenne;
epistoma smooth; the thorax mar-
gined on all sides with a faint stria,
anterior margin bisinuous; angles
rounded off, with the edge behind
them feebly crenulate, posterior angles
‘rectangular, sides dilated and slightly
narrowed behind, punctured some-
what more coarsely than the head,
surface uneven, with a fovea before
the scutellum and two others, less
distinct, placed longitudinally on each
side of it near the margin (or they are merged in one, as in
the figure); the scutellum triangular, with a few small
punctures ; the elytra complete, dilated along the outer edges,
ovate, except at the base ; dorsal area striate-punctate, sutural
row of punctures and those next to it most distinct, outside
the fifth row the punctures become denser and lose their linear
arrangement ; the basal segments of the abdomen are infus-
cate. Male: anterior and intermediate tibiae widely bent;
in the anterior pair the inner edge of the tibia is swollen and
somewhat angular at the point where the bending ceases,
Hab. Main Island, near Kashiwagi, 19th June, 1881.
I found two males under some marshi-refuse ; the female is
not known to me,
396 Mr. A. G. Butler on the Genus Acronycta.
LXIV.—Notes on the Genus Acronycta of Authors and its
Position in the Classification of Heterocerous Lepidoptera.
By A. G. Burver, F.LS., F.Z.8., &e.
In the ‘ Transactions of the Entomological Society’ for 1879
I proposed (pp. 313-317) to break up the old genus Acro-
nycta and distribute it, chiefly on account of its very distinctive
larval characters, amongst the various families of Bombyces
&e. to which the peculiarities of the larvee seemed to show
affinity. It must, however, be borne in mind that my con-
clusions were based chiefly upon the larve of the British
species and the published illustrations of Abbot ; nevertheless
J examined the neuration of the species in each group, com-
paring it with that of each of the families to which the larval
types seemed to point.
Unfortunately, at the time when I wrote my paper the
importance of the position of certain veins in the wings of
moths, as having more than generic value, was not so fully
appreciated as at the present time; therefore, although I
correctly represented the venation of the Notodontide, I failed
to see that the position of the median branches and lower
radial of the primaries in that family precluded the possi-
bility of any Acronycta being regarded as even remotely
allied to it. On the other hand, it is now known that the
Liparide and Arctiidz are much more nearly related to the
Noctuz than was formerly supposed; so that the similarity
in neuration is by no means surprising, nor is there any
reason why larval characters found in the former should not
also occur in a genus of Noctue.
A recent study of the transformations of Acronycta in
Abbot’s unpublished volumes has now clearly shown that the
differences in the clothing and outline of the larve charac-
teristic of the Bombycid families already referred to occur in
various species of the same group in Acronycta, and so are
valueless as indicative of their affinities. I am therefore
obliged to renounce my former opinion and admit that
Acronycta is in truth a genus of Noctue, probably nearest
allied to Polia.
The sections into which Acronycta has been divided may
still be used (as having the value of numerals only) to indicate
the near affinities of the species; and, as the Museum series
is represented by nearly eighty species, I propose to indicate
them below. I must, however, first point out that Acronycta
simplex of Walker is Xylomyges crucialis of Harvey, and
Mr. A. G. Butler on the Genus Acronycta. 397
that Hadena externa of Walker (which I formerly placed in
Triena) is probably a very worn and stained specimen of
Bryophila muralis, Forst.; its neuration certainly corre-
sponds closely with that of Bryophila, and such markings as
can be traced are also found in BL. muralis. I have failed to
recognize Acronycta declarata, Walk., A. telum, interrupta,
and pachycephala, Guen. ; though the description of A. cnter-
rupta is rather suggestive of A. vinnula, Grote, the absence
of the illustration upon which M. Guenée based it from the
Museum collection of drawings prevents its being satisfactorily
identified. Possibly A. telwm and pachycephala may be
known to our American friends; but we possess no specimens
so labelled.
Grote’s group erolonche seems to me to be far more
distinct than the other sections of Acronycta and may perhaps
prove to have generic value, the thicker antenne of the males
and the well-developed lateral anal tufts, combined with the
regular fasciated character of the markings, should, I think,
have some significance. A. lithospila, Grote, appears to me
to be closely allied to the European Scotochrosta pulla, and
eannot, I think, be an Acronycta ; nor can I believe that
A, lanceolaria and insolita, Grote, should be widely separated
from the latter or from Cucullia. I therefore admit Hulonche,
for these two species, as a genus.
One or two of the American species which have been
wrongly identified I shall be obliged to rename.
AcronyctTa (Acronicta, sic, Ochs.).
Typical Section.
1. A. leporina, Linn., and var. bradyporina, Treit.
Europe.
*2. A. felina, Grote. United States.
3. A. lepusculina, Guen., and var. populi, Riley.
Hudson’s Bay &e. -
*4, A. insita, Walk. United States and Canada.
*5, A. innotata, Guen. New York, Hudson’s Bay,
Canada, &c.
Of the above we possess the types of 2, 4, and 5. To save
trouble I shall indicate all species of which we possess the
type specimens by an asterisk.
398 Mr, A. G. Butler on the Genus Acronycta.
Section MrGacronycra, Grote.
1. A. americana, Harris.
A. acericola?, Guen. New York and Foo-
chow.
*2. A. dactylina, Grote. United States.
3. A. hastulifera, Sm. Abb.
* g. A. contacta, Walk. New York and
Hudson’s Bay.
In the Grote collection an example of A. americana was
labelled A. hastulifera; but the figure corresponds far more
closely with a female recognized as the latter and labelled by
M. Guenée. We have a fourth species of this group from
Ichang fT.
Section Arncromyscis, Hiibn.
A. acerts, Linn. Switzerland, Christiania, Li-
vonia, &e.
2. A. abscondita, Treit. North Germany, Hungary.
3. A. euphrasiw, Esper. Sicily, Russia.
4. A, euphorbiw, Gmel. Switzerland, Germany, &c.
A, sperata, Grote. New York.
*6, A. tota, Grote. Texas.
No. 6 is so much nearer to the dark variety of A. aceris
than to any species of the section Zriana, that 1 have without
hesitation placed it here.
Section ApaTELa, Grote (APATEL®, part., Hiibn.).
1. A. hercules, Feld. Yokohama and Tokio.
2. A. rubricoma, Guen. Philadelphia and Texas.
3. A. luteicoma, Grote. Kansas,
*4, A. pallidicoma, Grote. Illinois.
*5. A. impressa, Walk.
A. fasciata, Walk., and Verrillii, Grote.
A. brumosa, Grote (not Guen.). Hudson’s
Bay Xe.
+ I hesitate to describe the latter, Messrs. Leech and Oberthiir having
done so much work in the Chinese fauna of late years as to render such a
course risky.
Mr, A. G. Butler on the Genus Acronycta. 399
A, distans, Grote. United States.
A. megacephala, Schiff... Zurich, Frankfort, &e.
8. A. noctivaga, Grote. New York and Canada.
A, superans, Guen. New York and Canada.
?
. A, brumosa, Guen.
3. A. longa?, Guen.
* 9, A. persuasa, Harv. Florida and Yoko-
hama.
*11. A. perdita, Grote. Sanzalito.
A
. afilicta, Grote.
A. brumosa, var., Guen. Florida.
13. A. xyliniformis,Guen. Rhode Island, Florida, &e.
*14. A. extricata, Grote. Texas? (locality not on
labels).
15. A. oblinita, Sm. Abb. New York, Ohio, Nova
Scotia, Canada, &c,
Section LeprrorEumMa, Grote.
1. A. rumicis, Linn.
A. diffusa, Walk. Zurich, Brussa, Turkey,
Ichang, Japan.
2. A. leucoptera, Butl. Yokohama,
3. A. impleta, Walk.
A. subochrea, Grote (on label and in Check-
List, p. 23. n. 66). New York.
In Grote’s Revised Check-List the author (p. 4) states that
there is no such species as A. subochrea. If this is the case,
how did he come to label one specimen as his type of that
species, a second specimen as also “‘Apatela subochrea, Grote,”
and to include it in his Check-List of 1882? It may not be
a distinct species from the following, but it is certain that at
one time it was so regarded by Grote.
*4, A. hamamelis,Guen. United States.
Our example is evidently a co-type, though not mentioned
by Guenée, since it still bears his label; probably as it has
no abdomen he thought it not worth recording under the
description.
*5. A. hesitata, Grote. United States.
400 Mr. A. G. Butler on the Genus Acronycta.
. dentata, Grote. United States.
. increta, Grote. New York.
. dissecta, Grote. Philadelphia, New York,
Kansas.
. clarescens, Guen. Trenton Falls, New York,
Nova Scotia.
. modica, Walk.
*A. exilis, Grote. New York and Texas.
ra
ees : oi
Be pe fa Se
*11. A. spinigera, Guen.
*A. Harveyana, Grote. New York.
*12,. A. ovata, Grote. Texas.
*13. A. alborufa, Grote. New York.
*14, A. grisea, Walk. Hudson’s Bay.
Section PHareEtra, Hiibn.
1. A. auricoma, Schiff. Zurich, Livonia, &e.
. ortentalis, Mann. Bithynia.
aie
bb
. menyanthidis, View. St. Petersburg, Berlin, &e.
Section Trrana, Hiibn.
A, pst, Linn. Regensburg &c.
A, increta, Butl. Yokohama.
3. A. tridens, Gmel. Frankfort &e.
A. leucocuspis, Butl. Yokohama and N. China.
3
. A, lobelia, Guen.
* 9. A. thoracica, Grote. New York.
6. A. Grote, sp. n.
A. lobelizw, Grote (not Guen.).
Half as large again as A. lobeliv, the longitudinal black
streaks on the primaries slightly more elongated, but all the
markings very similar: secondaries decidedly darker, with
more golden gloss, the grey markings also darker. Expanse
of wings, ¢ 2 56 millim. (one dwarfed male 47 millim.).
New York and Kansas.
It is possible that this may be no more than a large dark
form of M. Guenée’s species; but it is clear that it is not so
regarded in America, and therefore I name it.
Mr. A. G. Butler on the Genus Acronycta. 401
7. A. furcifera, Guen. New York.
8. A. cuspis, Treit. Germany.
9. A. maxima, Moore. Kilt and Murree.
*10. A. anedina, Butl. Hakodaté and Tokio.
11. A. vinnula, Grote. New Jersey.
12. A. Smithit, sp. n.
A. clarescens, Grote (not Guen.).
Primaries above chalky white, irrorated and clouded with
sandy greyish or yellowish and grey mixed, somewhat after
the manner of A. morula; the submedian basal black streak
has somewhat the character of that of the true A. clarescens,
only it is more elongated, sharply defined, and its upper
margin is regularly tridentate; the reniform and orbicular
spots are narrowly black-edged; the twin discal or post-
median lines are sigmoidal and dentate-sinuate, the outer line
being partly black ; the subapical longitudinal dash is vague
and diffused, and the dagger-mark has a brown background,
which gives it a blurred appearance ; in other respects the
species 1s like A. furcifera, the male having whitish and the
female more or less golden-brown secondaries with the usual
markings. Expanse of wings 40-45 millim.
New York and Kansas.
*13. A. pauperata, Grote. United States.
14. A. occidentalis, Grote. Rhode Island, New York,
Buffalo, Kansas.
“A. furcifera, Pack.” (not Guen.), on Zeller
specimen.
15. A. morula, Grote. New York and Canada.
*16. A. faleula, Grote. United States.
*17, A. parallela, Grote. Colorado.
*18. A. Radcliffe’, Grote. New York and Vancouver.
19. A. tritona, Hiibn. New York and Florida.
20. A. hasta, Guen. ‘‘ Amherst”? (United States or
Canada ?).
*21. A. quadrata, Grote. Kansas.
Section Hysoma, Hiibn.
1. A. strigosa, Fabr. N. Russia, Livonia, &c.
402 Mr. O. Thomas on some Mexican. Oryzomys.
*2. A. nigrivitta, Hamps. Nilgiris.
3. A. divisa, Moore. Dharmsala,
Section Jocur=ra, Hiibn.
1. A. alni, Linn. Ausbach, Livonia, &e.
*2. A. connecta, Grote. New York.
We also have a pretty species from Madagascar. It has
probably been named by Herr Saalmiiller or M. Mabille ;
but at present I cannot spare the time to look it up. Grote’s
A. funeralis is unknown to me, but (if confounded with
A, americana) can hardly belong to the section Jocheera,
Section MastreHanes, Grote.
*1, A. denticulata, Moore. Dharmsala and Sabathu.
*2. A. edolata, Grote. Arizona.
Section MERoLoncHE, Grote.
*1, A, spinea, Grote. California.
2. A. Lupint, Behr. Mendocino.
It is possible that one or two of the Japanese and Chinese
forms enumerated above may be synonymous with species
described from East Siberia; but the illustrations to the
‘ Lepidopteren Ost-Sibiriens’ and ‘ Reisen und Forschungen
im Amur-Lande’ are so poor that, without seeing examples
from the Amur, it would not be possible to be certain of the
identity of the species, in a genus containing so many closely
related forms.
LXV.—Notes on some Mexican Oryzomys.
By OLDFIELD THOMAS.
Dr. A. C. Butter has lately sent to the British Museum a
specimen of an Oryzomys from Jalisco which appears to be
new, and in working this out I find that some confusion
exists as to Alston’s Hesperomys Couest, in clearing up which
a second species in the Museum collection proves to need
description.
Mr. O. Thomas on some Mexican Oryzomys. 403
Hesperomys Couest, Alston.
Hesperomys Couesi, Alston, P, Z. 8. 1876, p. 756.
This species, described by Alston from three specimens now
in the British Museum, has proved rather a stumbling-block
to later workers, and this is not surprising, as an examination
of his specimen shows that the three examples represent
certainly two, and perhaps three, different species. The
question therefore that at once arises is as to which of the
three should be taken as the type, a question which, on the
principle of the selection of the first-named species as the
type of a genus, may best be settled by taking Mr. Alston’s
specimen @ as the type of //. Couest?. This selection is
supported both by the fact that the first five lines of the
description itself are solely based on the spirit-specimen and
also by Mr. Alston’s known partiality to working on alcoholic
material ; so that no one who knew his methods would doubt
as to which he himself would have chosen. I propose there-
fore for the future to look upon Mr. Salvin’s spirit-specimen
from Coban, Guatemala, whose measurements are given in
Mr. Alston’s first column, as the type of his H. Coues?.
Whether c, from Mexico (Verreauz), is or is not the same
species | am at present unable definitely to decide ; but that d
is distinct from either is perfectly clear, its distinctive cha-
racters, both cranial and external, being more definite than
are those separating any other two members of this difficult
group.
The true O. Couesi, as represented by the type specimen, is
a small species with slender limbs and long tail. Its colour
is fulvous, darker along the centre of the back and paler, but
not white, on the belly. The fur is fairly soft, but not thick
and woolly as Alston stated, that remark evidently resting on
specimen b (my O. fulgens). The measurements of the
typical skull are given below and also those of the skull of an
old male from the Hacienda Cubilguitz, about 25 miles north
of Coban, which with some hesitation I refer to this species.
Oryzomys fulgens, sp. n.
Size large. Fur very thick, coarse, and woolly. General
colour above bright fulvous, brighter than in any other
Central-American species ; anterior half of the body, including
the head, rather paler and duller than the posterior half.
Ears decidedly small, broadly rounded, thinly hairy, their
hairs practically the same colour as those of the head in
general, so that they are not distinguishable by colour at a
404 Mr. O. Thomas on some Mexican Oryzomys.
distance. Lips, chin, throat, and inguinal region whitish,
belly with a strong suffusion of fawn, which reaches a maxi-
mum on the breast between the fore legs; passage of upper
colour into lower quite gradual. Outer sides of limbs like
back, inner sides whitish ; upper surfaces of hands and feet
thinly clothed with pale silvery-fawn hairs. ‘Tail long, thinly
haired, the scales not hidden by the hairs; above blackish,
below yellowish, darkening towards the tip.
Skull readily distinguishable from all allied species by its
great breadth, the bold expansion of the zygomata, and
especially by the evenly incurved outline of the supraorbital
edges; in all other species these edges form two approxi-
mately straight lines diverging from the narrowest interorbital
point, but in O. fulgens the whole inner wall of the orbit forms
one even curve, the breadth at the posterior end of the olfac-
tory chamber being scarcely greater than at the anterior end.
Nasals broad and flattened. Frontal premaxillary processes
very narrow and barely attaining to the same level as the
back of the nasals. Anterior palatine foramina large, widely
open, their posterior margin just level with the front of ™1.
Measurements of the type (an adult male in skin) :—
Head and body 160 millim. ; tail 151; hind foot, without
claw 35, with claw 37°53 ear (approximate) from notch 13:3.
Skull (see below).
Hab, Mexico. Coll. A. Boucard.
Type B.M. 70. 6. 20. 3. Purchased of Geale.
As already noted, this species is founded on specimen 6 of
Mr. Alston’s description of Hesperomys Couest. Its exact
locality unfortunately must remain unknown until further
specimens of it are found; but its characters, both external
and cranial, are so striking that I have no hesitation in
describing it as a distinct species.
Oryzomys melanotis, sp. n.
Size small, form more slender and delicate than in the
allied species. Fur straight, close and crisp. General colour
grizzled rufous, brighter and clearer on the sides and rump.
Lips, inner sides of limbs, and whole under surface white, the
line of demarcation on sides well defined ; the belly-hairs are,
however, as usual slaty grey basally. ars large, projecting
far beyond the fur, closely covered with short black hairs,
their black colour contrasting conspicuously with the general
rufous colour of the head. Upper surfaces of hands and feet
whitish. ‘Tail long, very scantily haired, blackish above,
whitish below.
Mr. O. Thomas on a new Species of Perognathus. 405
Skull unusually broad considering its small size. Nasals
very broad behind, surpassing. in length the premaxillary
processes. Interorbital space very broad. Supraorbital beads
widely divergent. Anterior and posterior edges of inter-
parietal forming two nearly equal curves. Anterior palatine
foramina ending in front of ™- a distance nearly equal to the
length of ™%, Posterior nares widely open, rounded.
‘Teeth small and delicate.
ise og of the type (B.M. 93. 3. 6. 25), an old male in
skin :—
Head and body 97 millim.; tail 127; hind foot, without
claws (damped) 27, with claws 28 ; ear from notch 18.
Skull (see below).
Hab. Mineral San Sebastian, Jalisco, Mexico. Coll.
Dr. A. C. Buller, Jan. 25, 1893.
The type specimen of this species being quite old, with the
molar teeth much worn down, its small size will readily
distinguish it from all its allies, except the still smaller
O. Alfari, Allen, from Costa Rica.
Measurements of the Skulls of the above Species.
Oryzomys Couesi.
ay esl lls
Hacienda O. fulgens. O. melanotts.
Type, 2. Cubilguitz,$. Type,d. Type,d.
—
millim. millim, millim. miullim.
Basal length ........ (c.) 26°6 30°4 Ges 2571
Upper length* ...... (c.) 80° 34:8 (81°8t) 29°5
Zygomatic breadth .. .... 17'8 (c.)17°8 = (e.) 152
Nasals, length ...... 1b Aa! 14°] 13:2 12:0
Interorbital breadth .. (c.) 5:2 59 48 51
Breadth of brain-case . 12°7 13°2 were 128
Interparietal, length .. 35 33 or
- breadth. . 79 7°8 10°0
Palate-length ........ Stas Gls sats 155
PHBBEANIG cos sce 55784 86 9°3 91 8]
Length of palatine fo-
POMIM AL rc oe oid ks 6:2 7:0 7:2 58
Length of upper molar
SNE stra a sx 48 48 52 4°3
LXVI.—Description of a new Species of Perognathus from
Colorado. By OLprirLp THOMAS.
AMONG a series of what appear, from Dr. Merriam’s able
monograph, to be Peregnathus flavus, Bd., collected by
Mr. W. G. Smith in Colorado and acquired by the British
Museum, there is a single specimen obviously different from
* To back of interparietal, excluding supraoccipital.
+ Interparietal lost ; from back of parietal suture only.
Ann. & Mag. N. Hist. Ser. 6. Vol. xi. 29
406 Mr. O. Thomas on a new Species of Perognathus.
the others, and, so far as I can make out, not referable to any
hitherto known species. It may be termed
Perognathus infraluteus, sp. n.
Size small, about as in P. longimembris. Pelage soft.
General colour above tending towards greyish olivaceous,
decidedly more so than in P. flavus, but less than in P. Lordi,
Lighter patches behind ears much reduced, scarcely percep-
tible ; ears very small (but doubtfully quite perfect in the
type), their infolded edges yellowish. Fulvous lateral band
strongly defined from the dark upper surface, but extending
beneath the body so as to cover the whole under surface
except just the throat and the centre of the neck and breast ;
the limbs also wholly fulvous; the hairs of this fulvous
underside everywhere slaty grey for their basal halves. This
coloration, strikingly different as it is from the usual pure
white under surface, seems to be only paralleled by that of
Dr. Merriam’s P. olivaceus amenus, a very much larger
animal. Soles hairy for their posterior half. ‘Tail short-
haired throughout, yellowish above, whitish below.
Skull of about the size and proportions of that of P. longi-
membris, but the mastoids are markedly less swollen, do not
approach each other so much, and do not project backwards
beyond the level of the occiput. The interparietal is conse-
quently much broader, although of the usual length ; it is in
fact “ broadly pentagonal,” ratio of length to breadth 53 per
cent. Frontal, median, and mastoid sides of parietals sub-
equal and longest, about 4 millim. each.
Teeth_™8 very small, scarcely larger in section than the
rounded anterior cusp of the premolar. Lower premolar
rather larger than 5.3, rounded quadrangular, its posterior
side the longest. |
Dimensions of the type (an adult female) :—
Head and body (measured in flesh by collector) 60 millim. ;
tail 57; hind foot, without claw (camped and measured in
skin) 14°9, with claw 16; hairy part of sole in centre 6°6.
Skull: basal length 17°3; basilar length of Hensel 15°3 ;
greatest median length 21:2; greatest mastoid breadth 111 ;
nasals, length 7°7; interorbital breadth 4°95; interparietal,
length 2°7, breadth 5:1; least distance between mastoids
above 5°2; combined lengths of three upper molars 19;
ditto, three lower molars, 2°2.
Hab. Loveland, Larimer County, Colorado, 5000 feet.
Coll. W. G. Smith, April 4, 1592.
This species is perhaps most nearly allied to P. fasciatus,
Wied, from Montana, but it is readily distinguishable by its
different coloration and somewhat smaller size.
ea ee
-ee-,
~
On a new Spectes of Tree Trap-door Spider. 407
LXVII.—Description of a new Species of Tree Trap-door
Spider from Trinidad. By R. I. Pocock.
[Plate XIX. figs. 1-3.]
Mr. J. H. Hart has recently sent for determination to the
British Museum a small series of insects and spiders from
Trinidad. Three species of spiders were represented in the
series: one of them is the well-known Argiope argentata; a
second appears to be Actinopus scalops of Simon; the third,
however, a species of Psewdidiops, appears to be undescribed.
I propose therefore to name it in honour of its discoverer. It
may be diagnosed as follows :—
Pseudidiops Hartii, sp. n.
Colour.—Carapace olivaceo-piceous, with black postero-
lateral border and black ocular tubercles; abdomen purplish
brown ; legs mostly testaceous, but the whole of the patella
and the distal end of the tibia black ; tarsi reddish black ;
sternum aud coxe flavous, maxille and labium olivaceo-
castaneous,
Carapace smooth, high, the posterior portion sloping
upwards to the deep crescentic fovea; the area immediately
in front of this fovea abruptly elevated, deeply longitudinally
sulcate, and furnished on each side with a single large seti-
ferous puncture; the area of the carapace between these
punctures and the anterior border nearly flat, but bearing the
two elevated ocular tubercles, the area between the two
tubercles only a little larger than the diameter of the anterior
eyes of the posterior tubercle, which is furnished with a
strong seta between these eyes.
Mandibles weak, smooth above, furnished in front with
long stiff sete, the inner angle above the base of the fang
produced into a short spicular prominence; fangs short but
robust.
Labium separated from the sternum by a deep depression,
narrowed in front, its distal border rounded, beset with long
stiff sete and armed with a transverse row of four short
conical spines, behind which are two less conspicuous but
similar spines.
Mawille covered with stout conical spines. The femoral
segment of the palp furnished internally with a few irregu-
larly arranged long sete, which distally increase in stoutness
and become spiniform ; the patella armed internally with two
408 Mr. R. I. Pocock on a new Species of
long spines and furnished with a distinct tubercle externally ;
the tb¢a hairy beneath, but armed externally and internally
with about two rows of strong spines; the tarsus similarly
armed, the claw with a large basal tooth, which is itself
also armed. The legs of the anterior pair armed like the
palpi, except that the inner surfaces of the femur and patella
are not armed with either spines or spiniform hairs and the
anterior spines on the tibia are fewer. The second leg is,
like the first, stout, the spines on the tibia still more reduced,
those on the anterior surface becoming shorter and those on
the posterior surface setiform and fewer. In the third leg
there are a few small spines interspersed amongst the hairs on
the anterior aspect of the patella and on the anterior and
posterior aspects of the tibia; a few, too, only remain upon the
proximal segment of the tarsus, which is, however, armed
beneath with two strong long spurs. In the fourth leg the
patella is armed with a few short spines in front and one
behind ; the tibia is furnished with setiform spines beneath
and the proximal tarsal segment with a few long spines
beneath. The claws of all the legs are armed with a large
basal tooth, behind which, except in the posterior claws of
the first and second leg, there is a second minute tooth.
The abdomen high and rounded. The anterior spinners
shorter than the basal segment of the posterior, which are
stout, three-jointed, and conical.
Length of carapace 6°5 millim., width 6; length of abdo-
men 8, width 6; length of maxillipede 12°8, of first leg 15-2,
of secoud and third 13°8, of fourth 20.
Of this interesting genus Pseudidiops, with which Den-
dricon of Cambridge is synonymous, two species have been
described *, and these are very probably identical.
Simon’s original specimens were from Cayenne, and he
has subsequently obtained the species from Venezuela. Un-
fortunately his description is very brief and is not accom-
panied by figures ; so it is only possible to point out that this
torm from Trinidad appears to differ from his op/fex in having
the legs flavous and ringed with black. By the same
character it may be recognized from the British-Museum
example from Labia, which Mr. Cambridge has described as
Dendricon rastratum in the Proc. Zool. Soc. 1890, p. 623.
* Pseudidiops opifer, Simon, Ann, Soc. Ent. Fr. (6) ix. p. 215 (1889).
Dendricon rastratum, Cambridge, Proc. Zool. Soc. 1889, p. 250.
(The description of Pseudidiops appeared on Sept. 11th, that of Den-
dricon on Oct. Ist. The former tirerefore has the priority.)
Tree Trap-door Spider from Trinidad. 409
But it further differs from this last-named specimen in having
the cephalothorax less elevated, a smaller distance between
the two ocular tubercles, the eyes on the posterior tubercle
set more closely together, and in possessing fewer spines on
the labium.
The nest of this species, which was sent with the specimen,
appears to resemble that of P. rastratus from Bahia. It is a
short tube permanently closed at the bottom, with a hinged
door at the entrance formed of closely woven silk threads, and
perfectly smooth inside. The length of the tube from the
hinge-margin to the bottom is oniy about twice the greatest
diameter of the aperture. The surface by which it was
attached to the tree-trunk is tolerably flat; the free surface,
on the contrary, is strongly convex and thickly covered, as
also is the door, with particles of bark. The shape of the
cavity corresponds to that of the outside and of the door.
The door, which is slender and with upturned edges, is wider
one way than the other, 7@. e. its width from the hinge to the
margin opposite to it is less than the width taken along a
line at right angles to this measurement. It opens outwards
and closes by the elasticity of its hinge. When closed the
plane of its outer surface meets that of the trunk of the tree
at an angle of about 45°.
Note 1.—In the nest of Actinopus scalops, which Mr. Hart
sent with the specimens of this species, the hinge of the door
is strengthened and protected by thick tough layers of
greenish silk, so that the area above the hinge projects far
above the rest of the surface of the door (Pl. XIX. fig. 3).
Note 2.—Mr. Hart forwarded with the specimens of
Argiope argentata examples of the cocoons of that species.
These cocoons show an interesting variation in colouring,
being either bright yellow or green on both sides or green on
one side and yellow on the other.
EXPLANATION OF PLATE XIX. Fries. 1-3,
Fig. 1. Pseudidiops Hartii, sp. n., nat. size. la. Lateral view of cara-
pace. 16. Lateral view of upper part of carapace, to show
arrangement of eyes and disposition of sete. 1c. Labium,
1d. Pseudidiops rastratus (Cambr.), for comparison with fig. 1 b.
Fig. 2. Nest of Pseudidiops Hartii, sp. n.
Fig. 3. Nest of Actinopus scalops, Simon.
410 Bibliographical Notice.
BIBLIOGRAPHICAL NOTICE.
The Fauna of British India, including Ceylon and Burma.—Moths.
Vol. I. By G. F. Hampson. Edited by W. T. Branrorp.
Royal 8vo. With numerous Illustrations and 527 pages of
letterpress. Published under the authority of the Secretary of
State for India in Council. London: Taylor and Francis, 1892.
Wirnovr doubt Mr. Hampson’s work is one of the most important
contributions to entomological literature which has _ hitherto
appeared—valuable alike to the student and collector of Indian
moths, to whom it will be a priceless boon; to the cabinet worker
who has to deal with the Heterocera of the world it will be a
necessary text-book.
The classification of the families of Butterflies was studied in 1864
by the late Mr. H. W. Bates, and with such satisfactory results that
his arrangement commended itself to all lepidopterists who took the
trouble to test its accuracy; indeed, Bates’s classification, with very
slight modifications, is generally adopted at the present day. On
the other hand, to form a key to the many families of Moths seemed
such a stupendous task, that few men ventured to attempt it. The
arrangements proposed by Messrs. Boisduval and Guenée were gene-
rally followed, and the blunders of these pioneers were copied and
multiplied by their successors until the chaos into which the Hete-
rocera were brought looked almost hopeless.
At length the study of the Tineina by Stainton and others and of
the Noctue and Pyrales by Lederer began to throw a little light
upon the obscurity ; but entomologists still needed a guide to point
out how, by the use of a simple pocket-lens and a little benzine, to
decide at once whether a moth was a Geometer, a Noctuid, a Pyrale,
and so forth.
In his ‘ Vlinders van Nederland’ Heer P. C. T. Snellen eventually
produced an admirable key to the families and genera of European
Moths—a work unfortunately overlooked by most students of
Exotic Lepidoptera ; the confusion which therefore existed in public
and private collections became year by year more confounded.
Happily Mr. Hampson, when seeking a basis on which to found a
general classification, discovered Snellen’s key, and upon this, with
slight modifications and many additions, he formed his classification
of the Moths of the World.
In his Introduction Mr. Hampson has for the first time pointed
out a character by which Moths can be distinguished from Butter-
flies, namely—all which resemble Butterflies in the possession of
clubbed or dilated antennz, also possess a frenulum, a character
invariably wanting in the so-called Rhopalocera.
The descriptive matter and illustrations in the first volume of the
Moths of India leave nothing to be desired, the former being terse
and to the point, whilst in every genus one species is admirably
figured, usually with accurate structural details, andin many instances
a typical larva is represented. When absolutely necessary the
Miscellaneous. 411
synonymy of a species is given, but otherwise a reference to Cotes
and Swinhoe’s Catalogue of the Moths of India is substituted.
Touching the sinking of many described forms to the rank of
synonyms, doubtless considerable differences of opinion will exist
amongst lepidopterists ; Mr. Hampson has used his private judgment
in the matter, and, as an experienced collector of Indian Moths, his
opinion must be allowed to have some weight. Without doubt the
reduction of spurious species was much needed; but nevertheless
the decision of no one man, however trustworthy, can be considered
as final, until the life-history of many more species has been studied ;
because it is an acknowledged fact that, whereas some families of
Moths are remarkable for their variability, others are almost as
constant in all their characters, A. G. Burrer.
MISCELLANEOUS.
New Observations on the Affinities of the different Groups of Gastro-
pods (Expeditions of the Yacht ‘ Hirondelle’). By M. E.-L.
Bouvier.
Gastropops are divided into two groups according as they are
unisexual or hermaphrodite; the former (Prosobranchia) are further
characterized by their decussating visceral commissure in the form
of a figure of eight, while the latter (Opisthobranchia, Pulmonata,
Pteropoda) are distinguished by their visceral commissure being
more or less free from torsion. After a previous study * I had
succeeded in partially removing the abnormal hiatus which an
incomplete investigation had allowed to exist between these two
groups, which I remained convinced must formerly have been
united by a transitional form. If this form still existed it could
only be found among the oldest Opisthobranchia, the Actzeonide,
which made their appearance in the Carboniferous period, and
which are represented at the present time by the genus Acton, of
Triassic origin. Investigations which I have made upon specimens
of Acton solidulus, kindly handed over to me by M. Jousseaume,
show that this Gastropod is, as a matter of fact, an ideal tran-
sitional form not only between the Prosobranchia and the Opistho-
branchia, but also between the latter and the Pulmonata. As I
have already published a succinct résumé of the organization and
affinities of Actaon ft, I shall here confine myself to an exposition of
the general considerations to which the organization of this animal
gives rise.
* “Quelques observations anatomiques sur les Mollusques gastéro-
podes,”” Comptes rendus de la Société de Biologie, December 17, 1892.
+ Société philomathique, séance du 24 décembre, 1892, and Société de
Biologie, séance du 7 janvier, 1893,
412 Miscellaneous.
The nervous system of Acton * is decidedly chiastoneurous, like
that of the Prosobranchia. The commissural ganglion on each side
is fused with the corresponding cerebral ganglion. One of the
branches of the visceral commissure starts from the left cerebro-
commissural ganglion, travels obliquely from left to right and from
front to rear, passing beneath the long buccal mass, and ends in the
subintestinal ganglion, which is situated to the right close to the
body-wall; the other branch is detached from the right cerebro-
commissural ganglion, travels from right to left and from front to
rear above the buccal mass, and ends in the supra-intestinal
ganglion. Starting from this ganglion, which is situated upon the
body-wall to the left, the commissural branch passes backwards,
and, shortly before reaching the anus, inclines to the right above
the cesophagus, and terminates in the visceral ganglion, which lies
between the latter and the oviduct. In thissame ganglion likewise
terminates the prolongation of the subintestinal branch.
The supra-intestinal ganglion innervates the gill and the left
portion of the mantle; the subintestinal ganglion emits a nerve
which proceeds to the right portion of the latter organ. The inner-
vation of the mantle is, however, a little further complicated, owing
to the presence of two small accessory ganglia, which we may term
secondary pallial ganglia, and which are the more important since
they will enable the chiastoneurous nervous system of