Skip to main content

Full text of "Transactions of the Linnean Society."

See other formats 


THE A 
TRANSACTIONS 


OF 


THE LINNEAN SOCIETY 


lon TN 


OF 


LONDON. 


VOLUME XXII. E 


LONDON: 


PRINTED BY TAYLOR AND FRANCIS, RED LION COURT, FLEET STREET: 


SOLD AT THE SOCIETY’S APARTMENTS, BURLINGTON- HOUSE ; 
AND BY LONGMAN, GREEN, LONGMANS, AND ROBERTS, PATERNOSTER-ROW. 


M.BOCO.LIX. 
MS 


MISSOURI 
BOTANICAL 
GARDEN. 


CONTENTS.. 


PART I.—1856. 


I. On the Structure and Affinities of Balanophoreæ. By Josera Daron Hooker, Esg., 
MED le ELN Qc E. . page 1 


IL. On the Development of the Ovule of Santalum album ; with some Remarks on the 
Phenomena of Impregnation in Plants generally. By ARTHUR HENFREY, Esq., 
F.R.S., F.L.S., Prof. of Botany in King's Dum LEM . ... ... À 


III. Remarks on the Nature of the outer fleshy covering of the Seed in the Clusiaceæ, 
Magnoliaceæ, §c., and on the Development of the Raphe in general, under its 
various circumstances. By JOHN Miers, Esq., F.R.S., F.L.S. ges. MM 


IV. On several instances of the Anomalous Development of the Raphe in Seeds, and the 
probable causes of such deviations from the usual course of structure, especially in 
reference to Stemonurus (Urandra of Thwaites), with some Prefatory Remarks on 
that Genus. By Jon Miers, Esq., RS, FLS. &e. . . . . . . . 91 


PART II.—1857. 
V. On a new form of Corynoid Polypes. By Pure Henry Gosse, Esq., F.R.S., 
A. L.S. vo. vw o s o. P RU. re . 113 
VI. Description of a new Species of Euplectella (Euplectella Cucumer, O.). By Professor 


EE NT 
VII. On Brachynema and Phoxanthus, two new Genera of Brazilian Plants. By 
GEORGE BENTHAM, Esg., F.L.S. . ee > us «à RED 
VIII. On some new Fungi. By the Rev. M. J. BERKELEY, M.A., FL.S.. . . . 199 


IX. On the Growth and Composition of the Ovarium of Siphonodon celastrineus, Griffith, 
especially with reference to the subject of its Placentation. By Josern Daron 
Hooxne, 440. DG KES FLEST pre | ag 


X. Further Remarks on the Organs found on the bases of the Halteres and Wings of 
Jue. by J.D, Mo D, MD FELS =: V v... > |, 141 


vi CONTENTS. 


XI. On a new Structure in the Antenne of Insects. By J. B. Hicks, Esq., M.D., 
BI... s; 84 9 US os Ppåge 147 


PART 111.—1858. 


XII. Observations on Entozoa, with notices of several new species, including am account 
of two experiments in regard to the breeding of Teenia serrata and T. cucumerina. 
Tee, erences COBRBOID: Rs MD, LLS. o o V. -— s. a B6 


XIII. On the Arrangement of the Cutaneous Muscles of the Larva of Pygæra bucephala. 
By Joux LuBBock, Esq., F.R.S., F.L.S. $c. . es 173 


XIV. On the Agamic Reproduction and Morphology of Aphis.—Part I. By Tuomas 
H. Huxzey, F.R.S., Professor of Natural History, Government School of Mines. 
Communicated by G. Busk, Esg., FR SF LS... . + ap … os. 198 


XV. On the Agamic Reproduction and Morphology of Aphis.—Part II. By Tuomas 
H. Huxzey, F.R.S., Professor of Natural History, Government School of Mines. 
Communicated by G. Busk, Esq., ERS, FLS. 2.020. . . : 221 


XVI. On the external Anatomy and Natural History of the Genus of Annelida named 
Palolo by the Samoans and Tonguese, and Mbalolo by the Fijians. By JOHN 
Denis Macponatn, Assistunt-Surgeon of H.M.S. * Herald, employed on Survey- 
ing Service in the South-western Pacific, under the command of Captain H. M. 
Dennam, R.N., F.R.S. Communicated by G. BUSK, Beg; ERD ELS. 987 


XVII. On the probable Metamorphosis of Pedicularia and other Jorms ; affording pre- 
sumptive evidence that the Pelagic Gasteropoda, so called, are not adult forms, 
but, as it were, the Larve of well-known genera, and perhaps confined to species 
living in deep water. By Joux DENIS MACDONALD, Assistant-Surgeon of H.M.S. 
* Herald,’ employed on Surveying Service in the South-western Pacific, under the 
command of Captain H. M. DENHAM, R.N., F.R.S. Communicated dy G. Busk, 

H, ERS BLS. à. ee NN NE pe *41 


XVII. On the Anatomy of Eurybia Gaudichaudi, as bearing upon ils Position amongst 
the Pteropoda. By Joux Denis Macponaup, Assistant-Surgeon of HMS. 

‘ Herald, employed on Surveying Service in the South-western Pacific, under the 
command of Captain H. M. DENHAM, R.N., F.R.S. Communicated by G. Busk, 

Es, PRS, PLS. eee silts me 228 50 24b 


XIX. Notes on Phoronis hippocrepia. By F. D. DYSTER, Dogs MISC. | oH. 251 


XX. Synopsis of the Fructification of the Compound Sphæriæ of the Hookerian Herba- 
rium, By FREDERICK CURREY, EN MA ERS RLE L o 2, 257 


CONTENTS. vii 


PART IV.—1859. 


XXI. Observations on the Structure of the Stem in certain Species of the Natural Orders 
Caryophylleæ and Plumbagineæ. By DANIEL OLIVER, Jun., F.L.S. . page 289 


XXII. On the Genus Henriquezia of Spruce. By GEORGE BENTHAM, Esq., V.P.L.S. 295 


XXIII. Remarks on Gnetum. By the late WILLIAM GRIFFITH, Esq., F.L.S., Madras 
Medical Service. Communicated by A. HExrREY, Esq., F.R.S., F.L.S., Professor 
Ofolfolany; King's College. 4a) SS a ØP SØR ID 


XXIV. Synopsis of the Fructification of the Simple Sphæriæ of the Hookerian Her- 
barium. By FREDERICK CURREY, Esq. M.A., PBS. FOS. . . . . . 313 


XXV. Synopsis of the Genera Camellia and Thea. By BERTHOLD SEEMANN, Esq., 
PED, BELS . . 249 2124 0  . . ee 


XXVI. On Tomopteris onisciformis, Eschscholtz. By WıLLıam B. CARPENTER, Esq., 
UBD SERE ERES. . |. 4 9 o 5» s ws. JM 


XXVII. On some new forms of Entozoa. By T. SPENCER Cossoi, Esq., M.D., 


XXVIII. On the Anatomical Characters of three Australian species of Tunicata referable 
to Savigny’s subgenus Cæsira. By Joux DENIS MACDONALD, Esq., F.R.S., Assistant 
Surgeon of H.M.S. * Herald, commanded by Captain H. M. DENHAM, R.N., 

FRS. Communicated by Guorce Busk, Esq., F.R.S., F.L.S. . . . . . 367 


XXIX. On the Anatomical Characters of a remarkable form of Compound Tunicata. 
By Joux Denis MACDONALD, Esq., F.R.S., Assistant-Surgeon of H.M.S. * Herald,’ 
commanded by Captain H. M. DENHAM, R.N., F.R.S. Communicated by GEORGE 
BUE ERN BER... . . 4 oS o so ru . . c D 


XXX. On the Anatomical Characters of an Australian species of Perophora. By Joun 
Denis MacpoNALD, Esq., F.R.S., Assistant-Surgeon of H.M.S. * Herald, com- 
manded by Captain H. M. Denmam, R.N., F.R.S. Communicated by GEORGE 
LE NE . . 971 


XXXI. Observations on the ** Bitentaculate Slug” of New Zealand (Limax bitentacu- 
latus, Quoy and Gaimard; Janella antipodarum, Gray; * Aneiteum | Slug” £ 
Macdonald). By CHARLES Knient, Seg; PDS. . . . . . . . . 881 


viii CONTENTS. 


XXXII. Further Remarks on the Organs of the Antenne of Insects, described in a 
Paper published in the * Transactions of the Linnean Society, vol. xxii. pp. 147-154. 
By J. Braxton Hicks, Esq. M.D., F.L.S. &e. . . . . . . . . page 383 


XXXIII. O» the Embryos of Endogens and their Germination. By BENJAMIN CLARKE, 
ENN TE 


XX XIV. Notes on the Structure and Affinities of Batideæ, Callitrichaceæ, Vochysiaceæ, 
and Cassytheæ. By B. CLARKE, RD ee . 9. 


XXXV. On the Origin and Development of the Pitchers of Nepenthes, with an Account 
of some new Bornean Plants of that Genus. By J. D. Hooker, Esq., M.D., 
ENN NNN 


XXXVI. On å New Genus of Balanophoreæ from New Zealand, and Two New Species 
of Balanophora. By J. D. HOOKER, Esq., M.D., ERS., F.L.S.&c. . . . 425 


THE 


TRANSACTIONS 


DE. 


THE LINNEAN SOCIETY 


OF 


LONDON. 


VOLUME XXI. 
PART THE FIRST. 


LONDON: 


PRINTED BY TAYLOR AND FRANCIS, RED LION COURT, FLEET STREET: 


SOLD AT THE SOCIETY’S HOUSE, SOHO-SQUARE ; 
AND BY LONGMAN, BROWN, GREEN, AND LONGMANS, PATERNOSTER-ROW. 


M.DCCC.LVI. 


CONTENTS. 


PART I.—1856. 


I. On the Structure and Affinities of Balanophoreæ. By Josern Darron Hooker, Bsq., 
DS EBR Ko FL GC: ere PT Se je QO 


IL On the Development of the Ovule of Santalum album; with some Remarks on the 
Phenomena of Impregnation in Plants generally. By Awrmug HENFREY, Esq., 
F.R.S., F.L.S., Prof. of Botany in King’s College, London . . . . . . 69 


III. Remarks on the Nature of the outer fleshy covering of the Seed in the Clusiaceæ, 
Magnoliaceæ, &c., and on the Development of the Raphe in general, under its 
various circumstances. By JOHN Miers, Esq., F.R.S., F.L.S. $c. . . . 81 


IV. On several instances of the Anomalous Development of the Raphe in Seeds, and the 
probable causes of such deviations from the usual course of structure, especially in 
reference to Stemonurus (Urandra of Thwaites), with some Prefatory Remarks on. 
that Genus. By Joux Mimes, Esq., PRS. ELS &e. . ..... . 97 


TRANSACTIONS 


OF 


THE LINNEAN SOCIETY. 


I. On the Structure and/Affinities of Balanophoreæ. 
By Josera Dauron Hookék, Esq., M.D., F.R.S., F.L.S. $c. 


Read February 6th, February 20th and June 19th, 1855. 


THE materials from which this Essay has been drawn up, have been accumulating on 
my hands for a considerable period. They consist principally of—1. A very complete set 
of the American species, especially of the Helosidee, formed at the desire of Sir William 
Hooker, in New Grenada, Jamaica, and Trinidad, in 1846 and 1847, by Mr. William 
Purdie, at that time collector for the Royal Gardens at Kew. He gathered nine species, 
including several new genera, and preserved many specimens of most, both dried and in 
spirits. 2. I am indebted to Prof. Liebmann of Copenhagen: for the species collected by 
him in Mexico, together with drawings of them; a translation of his paper on Thonningia 
and Helosis mexicana, read before the Society of Scandinavian Naturalists at Christiania ; 
and the loan of the original specimens of Thonmingia, from the herbaria of Vahl and 
Schumacher. 3. Mr. Miers has placed at my disposal his Brazilian specimens of Zangs- 
dorffia hypogea and Helosis guyanensis, from Rio and the La Plata district, together with 
his sketches of them made on the spot. 4. Sir Robert Schomburgk has given me his 
. drawings of the same genera, made in Guiana; and I have also received from other 
travellers numerous specimens of them. 5. For the South African genera Sarcophyte 
and Mystropetalon I am indebted to Dr. Harvey, who, during his residence at the Cape, 
communicated beautiful specimens of them to Sir William Hooker. 6. Of the Indian 
Balanophoreæ Y have very extensive suites of specimens indeed; having had, in the Hima- 
laya and Khasia mountains, the opportunity of studying several species in many stages 
of growth. I have also examined most of the specimens collected by Mr. Griffith himself, 
from which he described the species for the Society's Transactions; and I have received 
the Peninsular and Ceylon B. indica from Wight, Gardner, and Thwaites. 7. Sir William 
Hooker has procured the Javanese species from Mr. Thomas Lobb, Prof. de Vriese of 
Leyden, and others. 8. For specimens of the original species of Balanophora (B. fun- 
gosa of Forster), I am indebted to Mr. M‘Gillivray, who found it on the N.E. coast of 
VOL. XXII. B 


-^ 
f 


2 DR. J. D. HOOKER ON THE STRUCTURE AND 


Australia, during the voyage of H.M.S. “Rattlesnake ;” and I have also received it from 
Tanna (the place of its original discovery by Forster), where it was collected by Mr. Hinds, 
during the visit of H.M.S. “Sulphur” to that island :—these I have compared with 
Forster’s original specimens in the British Museum. 9. For other species I have been 
indebted t6 various sources, including the herbarium of the British Museum, which con- 
tains the original specimen of Lophophytum mirabile*. 

The total number of species thus brought together is about twenty-eight, of which I 
have examined both sexes of twenty-six. 

I have not considered it necessary to give a detailed list of the authorities who have 
written upon this Order, nor a history of the successive additions that have been made to 
our knowledge of it; these subjects having been minutely and well detailed by Richard, 
Griffith, and Endlicher; and a réswmé of them by Dr. Lindley will be found in his 
valuable ‘ Vegetable Kingdom.’ I may however mention that, in their several ways, the 
original Essay of the great Linnæus upon Cynomorium, in the fourth volume of the 
‘Ameenitates Academicee,’—the admirable one of Richard on Cynomoriwm, Helosis, and 
Langsdorffia, in the * Mémoires du Muséum,’—Gceppert’s very valuable remarks on the 
anatomy of the Javanese species, in the * Nova Acta, —Mr. Griffith's observations in the 
19th and 20th volumes of our Society’s Transactions,—Schott and Endlicher’s paper in 
the * Meletemata,’ and Weddell’s paper in the fourteenth volume of Ser. 3. of * Ann. Sci. 
Nat.,’ are by far the most important. A very complete summary of other authors will be 
found in Unger’s paper upon parasites in the Annals of the Vienna Museum; since which 
period, however, Goeppert’s, Griffith’s and Weddell’s papers have appeared, as also 
Junghuhn’s in the * Nova Acta,’ all of which are accompanied by valuable plates. 


4. Parasitism and Structure of the Rhizome. 
I shall employ the term rhizome for the principal axis of Balanophoreæ: it was 


* Since the above was read before the Linnean Society I have examined several other collections, of which the most 
important are—10. The original specimens of Langsdorfia and Helosis, collected by Von Martius, and preserved at 
Munich; 11. those of Seybalium (which are to this day unique), in the Vienna Herbarium; 12. the valuable collec- 
tion in the Jardin des Plantes at Paris. I have also to record my great obligations to my friend M. Weddell of Paris, - 
. who has already contributed so much to our knowledge of the plants of this Order, and through his good offices to the , 
Museum of the Jardin, for specimens of Lophophytum in several stages of growth, of Ombrophytum peruvianum, of 
Corynæa Weddellii, and of Langsdorfia rubiginosa ; all collected by Weddell in Bolivia, Peru and Brazil; for 
drawings of these made on the spot by himself; for dissections of Sarcophyte sanguinea, showing the central 
embryo which he discovered and figured, and for others of Langsdorfia hypogæa with the fruit fully formed, and 
which confirm Liebmann’s drawings and descriptions of the fruit of that genus. 

The result of the materials thus added has been to strengthen the views I have adopted of the structure and 
Mini of the Order, to enable me to classify Sarcophyte with Monostyli, and to reduce the subgenus Lepidophytum, 
which I had proposed, to the previously imperfectly known Lophophytum, with which I had doubtfully associated it. 

I nave aliy to express my obligations to our ingenious and accomplished foreign member M. Hofmeister of Leipsic, 
for showing a m drawings of the impregnated ovule of Cynomorium, with the pollen-tube in the foramen of the 
ovule: this, n is the most important discovery in favour of my view of the normal condition of the nucleus of the 
ovule and function of impregnation in the embryonate species, is also a most remarkable instance of skilful dissection. 


. lam encouraged to hope that M. Hofmeister will take up the subject of the embryogeny of the Balanophoreæ, and 


need hardly add, that from his unrivalled skill as a phytotomist, and extensive acquirements in embryogeny, the 


subject will receive the fullest illustration at his hands.— December 4th, 1855, 


AFFINITIES OF BALANOPHOREÆ. 3 


suggested doubtfully by Richard, who however adopted those of * radix," * tuber," and 
* caulis :” it is the “axis” of Griffith; “rhizome” and “caudex” of Geeppert; “ basilar 
receptacle” of Junghuhn, &c. In mode of origin and development it sufficiently accords 
with the definition of a rhizome, as usually employed in descriptive botany. In speaking 
of the root, I wish always to be understood as referring to that organ of the plant upon 
which the parasite grows. : 

The simplest and at the same time most frequent form assumed by the rhizome of Bala- 
nophoreæ, is that of a simple or branched tuber, sessile on the root from which the plant 
derives its nourishment, and giving off one or more flower-bearing peduncles. In the 
earliest stage at which I have examined any of the species, the young plant appears as a 
cellular mass, nidulating in the bark of the root (but partially exposed), with whose cellular 
tissue its own is in organic adhesion, though easily distinguishable. It offers at first no 
trace of a vascular system, nor any distinction of parts; but before it has reached the 
cambium layer of the bark, and before its upper extremity has attained any considerable 
size, an opaque line of white cellular tissue, different from the rest, may be found in the 
centre of the mass or beneath each of its lobes, in which vascular tissue makes its appear- 
ance. Shortly afterwards, the wood of the root upon which the parasite grows appears 
to become affected; its annual layers are displaced, and at a still later period vascular 
bundles, enclosed in a cellular sheath, are found in the axis of the rhizome, and are 
continuous with those already formed in it. For illustrations of these stages of deve- 
lopment see Plates IV. & VI. våg 
. Some genera do not present the appearance of any vascular bundles communicating 
with those of the root-stock; but their own vascular bundles may be traced descending 
to the line of union between the root and the parasite, where they become closely applied 
_ to the vascular system of the former, without, however, forming any interlacement or 
organic union. Of this, Lophophytum and perhaps Scybalium are examples. 

The fully formed rhizomes are roughly divisible into the simple, or merely forked or 
lobed, and those which are cylindrical, elongated horizontally and branched; forms which, 
though exceedingly dissimilar, and associated with very important anatomical details, are 
not accompanied by such modifications of the floral organs as would afford sectional 
characters in the Order; as a comparison of Helosis with its very near allies, Seybaliwm, 
&c., proves. The elongated rhizomes of some species form attachments by their ramifi- 
cations to the various roots they encounter; and such never have foliaceous appendages, 
except at the bases of the peduncles or flowering branches. The amorphous or simple 
rhizomes again are often provided with scales (as in Cynomorium, Lophophytum, and 
Spherorhizon) , or with cellular papillæ (as some species of Balanophora). These papillæ 
consist of simple or divided extruded masses of cellular tissue traversed by a furrow ; they 
are very numerous and cruciate in B. dioica, and are probably intimately connected with 
the aération or respiration of the plant; they present nothing remarkable in structure, 
and resemble the rimæ with swollen lips on the spongy bark of some Venispermee, Vitis, 
and many other plants*. 

_ * Junghuhn says (Nova Acta, xviii. Suppl. p. 223) that B. globosa is a species which he never found bearing these 
papillæ, except when it grew on the same root with B. elongata, when, like that species, it was always provided with them. 
| B2 


å DR. J. D. HOOKER ON THE STRUCTURE AND 


The rhizomes in many species attain a considerable age; but it is difficult to ascertain 
their duration after they have commenced flowering. Helosis seems to be capable of in- 
definite increase; the individual patches of the plant flowering at all or most seasons of the 
year, and the old branches of the rhizome dying as new ones are formed. In Phyllocoryne 
also, the large many-lobed rhizome seems perennial, and to flower at various seasons. In 
Rhopalocnemis and several species of Balanophora, it appears to me as if the rhizome 
continues to increase for several years; and then, after throwing up many peduncles in 
one season, to die the following autumn. Others however, as B. involucrata, which causes 
great knots to form on the roots of trees, either live many years and flower perennially, or 
else a perennial succession of young plants germinate upon the swollen root; a mode of 
increase suggested by the germinating specimen represented in Plate VI. fig. 8. In 
Lophophytum each tuber-like rhizome gives off only one or two peduncles, and the root on 
which it grows forms a shallow cup round its base; which I have found to be of many 
years’ growth. Cynomoriwm appears to be decidedly annual, but I have not examined a 
sufficient number of specimens with the attachment preserved, to decide this point. 
Langsdorffia has certainly a perennial branching rhizome, and Sarcophyte a perennial 
tuberous one. Most of the Balanophoræ have lobed or branched rhizomes, which perhaps 
die after flowering. i 
-. In no case is the vegetation of the rhizome very rapid, in comparison with that of many 
plants; and especially of Fungi, with which some authors have compared them. On the 
contrary, I believe that the growth of all the parts is very slow ; and with regard to Rho- 
palocnemis and Balanophora especially, I have had many opportunities of observing that 
the peduncle did not flower for several weeks after its protrusion from the rhizome. 

The modes of attachment above indicated suggest another division of Balanophoree, 

namely, into—1. those in which the vascular tissue of the parasite is continuous with 
that of the root; 2. those in which the attachment is by means of the cellular ‘system 
only; and 3. those in which bundles of vessels from the root terminate definitely in the 
parasite, a short distance from the point of attachment; the vascular systems of the two | 
plants forming no evident confluence. 
_ Of the first of these classes Balanophora and Rhopalocnemis are the best examples, from’ 
the great development of their vascular systems (which in some species present woody 
zones, à cortical system, and medullary rays), and from the fact that in many instances 
bundles of vessels appear to run in unbroken continuity from the woody system of the 
root to the very flowers of the parasite. 

In the species of this first group, the appearance of the parasite having derived all its 
vascular tissue from the root has given rise to the hypothesis, that the whole production 
is an abnormal development from the root of the plant on which it grows :—thus Junghuhn 
quotes Trattinick (Linnæa, iii. p. 194) as saying of Sarcophyte, * hasce parasitas degene- 
rationes plantarum specificas, sine seminum aditu creatas, modo spontaneo genitas ; ” 
and adds (Nor. Act. Acad. xviii. Suppl. p. 205), * Mihi Balanophorarum vegetatio 
fungosa est, originaria, Succi arborum, e quarum radicibus vivis sanisque Balano- 
phore progerminant, nimis copiosi, cursu consueto perturbati, morphosin arboris redun- 
dantis, ut ita dicam, RETROGRADAM provocant, atque (directione vegetationis mutata) in 


AFFINITIES OF BALANOPHOREÆ. 5 


novam et matricis nature alienam prolem consumuntur * * *." These theories have 
been well combated by Gæppert, who adduces the fact of the same species of Balanophora 
growing indifferently on various plants of very different natural families, as being quite 
opposed to them; to which may be added, that they have an independently developed 
vascular system of their own, which only in some species blends with that of the root; 
and that they are propagated by seeds. 

Griffith does not seem to have traced the vascular bundles of the root into the peduncle 
of the parasite; for in his valuable paper on .Balanophora (Linn. Soc. Trans. xx. p. 96), he 
describes them all as rising from the root into the rhizome, and terminating abruptly in 
the axis, towards its periphery : this well describes the appearance of those bundles which 
form the main body of the parasite; and they may be seen in the vertical section given 
in Plate IV. fig. 20, radiating in a fan-like manner from the root, and terminating in 
broad truncate masses towards the circumference of the rhizome. In a transverse section 
again (fig. 19) of a young, symmetrically formed, unbranched rhizome, with one peduncle, 
the vascular bundles will be found to be much more regularly disposed round a cellular 
axis, and separated by broad rays of cellular tissue. 

Gæppert and Unger both consider that there is a double vascular system in the parasite ; 
the one given off by the root on which it grows, and the other confined exclusively to the 
peduncle and its appendages, though passing downwards through the axis of the rhizome 
to within a very short distance of the base of the parasite, and there terminating abruptly. 

The result of my own observations on live plants of Rhopalocnemis (and which were 
verified by Dr. Thomson), is that the vascular bundles of the peduncle are so intimately 
united with those of the rhizome towards the base of the latter, that they are organically 
one and the same tissue. In illustration of this I will refer to Plate IV. fig. 22, as being 
taken from one of the simplest and most symmetrical forms presented by a Balanophora : 
in this the letter a indicates the union of the vascular bundles of the peduncle and rhizome. 
Of Rhopalocnemis and Balanophora dioica Y macerated many specimens in all stages of 
growth, some being in ripe fruit, when the vascular bundles have most consistence; and I 
never failed in dissecting them out in continuous masses from the bases of the apparent 
root-branches in the rhizome to the capitulum itself. 

The vascular branches that connect the root with the rhizome of the parasite, are 
altogether analogous to those found in the exostoses of DeCandolle on the roots of various 
. Leguminous plants; and especially such as have been pointed out to me by Prof. Henslow 
as being frequent on the roots of Laburnum*. 

The root itself of the plant on which B. fungosa grows, has no pith (Plate VIII. fig. 15); 
but the branches which it appears to send into the parasite, enclose a pith (figs. 10 & 11 a), 
and the wedges of wood of which these branches are composed become broken up at a 
distance from the base of the rhizome (fig. 11 55) ; the branches terminate in cylindrical 
masses of cellular tissue, enclosing a few imperfect spiral or barred vessels in their axis. 


* These latter are coralloid masses, consisting of a cortical and woody system, the latter provided with obscure 
medullary rays: as their distance from the root is increased, their branches become simpler in structure, being merely 
cellular cylinders with a vascular axis or core, the latter consisting of a little pleurenchyma and very imperfectly 
developed annular and other vessels. 


- 


6 DR. J. D. HOOKER ON THE STRUCTURE AND 


Two forms of attachment are found amongst the genera with branched and much- 
-elongated rhizomes. In Helosis the rhizome forms a tuber at each point of its attachment 
to the various roots it meets with in its subterranean course, and a few vascular bundles 
from the root are rarely sent into it at these points; but these do not appear to commu- 
nicate directly with the previously existing vascular tissue of the rhizome, nor to become 
blended with it: possibly, however, they may have been given off by it, or have been 
independently formed in the rhizome; a point which can only be determined by exami- 
ning the nature of the attachment at its first formation, and which I shall hereafter 
discuss. In Langsdorffia the branch of the rhizome corrodes the bark of the roots it 
encounters; the first contact in the case of L. rubiginosa being by means of woolly hairs. 
Both the rhizome and the root generally swell considerably, but often do not, and the 
root sends long vascular branches, apparently covered with the cellular bark of the root, 
right and left into the axis of the rhizome; with whose vascular system, however, I have 
never found them to form an organic adhesion (see Plate IT. figs. 10, 12, 13, 16, 17). In 
this genus two or more species of dicotyledonous plants sometimes send their roots into 
one tuber of an old rhizome, each penetrating at several points. 

In the Annals of the Vienna Museum (ii. 53), I find Balanophoree arranged by Unger 
under three of the divisions, into which all parasites are separated by that author according 
to the nature of their parasitism ; they are the following :—1. Parasites which form a _ 
rhizome by which they adhere to the roots of plants, and from which the flower-buds rise. _ 
Example, Scybalium. 2. Parasites which exercise a powerful specific action upon the root, 
causing it to send vascular bundles into the rhizome, which hence becomes an organ in- 
termediate in nature between the stock and the parasite. Examples, Balanophora, Sarco- 
phyte, Cynomorium, Lophophytum ? Ombrophytum ? 3. Parasites which form a rhizome 
intimately attached by its vascular tissue to the root. Examples, Helosis, Langsdorffia. — 

It appears to me that the above are rather distinctions of words than of facts; and that — 
in so far as they are correct, any one of the three definitions is more or less applicable to 
all the species : for all form rhizomes, all owe their adhesion to their power of exerting à 
specific action upon the roots from which they derive their nourishment, and except inthe 
case of Lophophytum (and perhaps of Ombrophytum, which I assume to. have the same 
mode of parasitism as Lophophytum), all more or less present the appearance of the vas- 
cular bundles of the root being enclosed in the cellular tissue of the parasite. Further, if 
my observations are correct, both Helosis and Langsdorffia should be transferred to the 
first class; for there is certainly no distinct union of their vascular bundles with those of 
the root, nor do their rhizomes appear to send any bundles towards the root; on the 
contrary, the appearance is perfectly distinct of the root sending its branches into the 
on Langsdorffia indeed is described both by Richard and (apparently following 
a 5 ling forth root-fibres from its rhizome ; but I not only fail to discover 
ok ES : e very numerous specimens I have examined, but I find this appearance 

produced by fibres being given off from the roots of the plant on which the parasite 


grows, which fibres become included within the rhizome (Plate II. fig. 11) 


The differences therefore that prevail amongst the modes of parasitism of Balanophoreæ, 


are of degree only: the power of erosion and of forming an organic adhesion is the main 


AFFINITIES OF BALANOPHOREÆ. 7 


point; that of inducing such a diseased action on the root as gives the appearance of 
the latter forming growths within the rhizome is a secondary one, and varies in amount; 
from Lophophytum, in which it seems to be none, to Rhopalocnemis, in which the bulk of 
the vessels in the rhizome are confluent with those of the root. As a general rule, the 
older the root attacked by the parasite, the fewer are the branches which it appears to 
send into the parasite; and as all my specimens of Lophophytum are on very much older 
and larger roots than are those of any other species, and indeed on wood of many years’ 
growth, it is quite possible that in the case of its attacking younger and feebler roots it 
may develope the same power. 

Under this view, the propriety of considering the rhizome of Balanophora and its con- 
geners to be an intermediate body, as suggested by Unger, seems, as Goeppert has pointed 
out, to be erroneous; indeed, there are stronger objections to it than have hitherto been 
urged, derived from the development of that body. 

I am unable to confirm Gceppert’s observations on one extremely difficult point, namely 
the presence of the two wholly independent and unconnected systems of vascular tissue. 
This author maintains, 1. that no free vascular bundles originate in the rhizome previous 
to the formation of flower-buds, but that the root gives off bundles to the rhizome, within 
which they ascend, prolonging, increasing in diameter, and branching, with the corre- 
sponding development of the cellular system of the rhizome. 2. That on the formation 
of the peduncles (floral organs), free and independent vascular bundles are developed in 
them, which ascend as the peduncles elongate, and also descend into the rhizome, occu- 
. pying a position between the vascular bundles of the latter, with which they do not unite. 

3. That these independent vascular systems present anatomical characters by which they 
may constantly be recognized, at any rate in the individual species. These positions I 
shall examine consecutively, premising that it is with considerable diffidence that I venture 
to dissent from the conclusions of this eminent author, since though I possess the advantage 
of having repeated my observations, both on living and dead plants of several species, I 
cannot regard these as entitled to more consideration than M. Gæppert's known skill 
and accuracy *. ; 
1. With regard to M. Goeppert’s first observation, it must be remembered that he nev 
had the opportunity of examining very young specimens, the importance of which 
desideratum he fully admits. In the section in Plate VI. figs. 7 & 8, which represents the 
independent formation of vascular tissue in a germinated Balanophora involuerata (and 
in other similar cases), I find in the axis of the rhizome pale transparent lines consisting 
of elongated cells, which contain no wax or cytoblasts, surrounding rudimentary vascular 
bundles. I have never examined a very young specimen in which these bundles were 
. found to have descended to the vascular system of the root, but I infer that they do so, 
‘and, becoming incorporated with the vascular bundles of the root, present the appearance 


* The difficulty of investigating these points is further far greater in living than in dead specimens : this is owing 
to the rapid sphacelation of the parts when cut, and the quantity of viscid Balanophorine (the term applied by Gæppert 
to the peculiar waxy secretion of Balanophora) contained in their cellular tissue, which prevents dissection with any 
approach to nicety : impediments so great, that I have no hesitation in saying, that in many cases better results may 
be obtained from specimens preserved in acid or spirits, than from living ones. 


8 DR. J. D. HOOKER ON THE STRUCTURE AND 


of having ascended from it, instead of having descended to it. This union once established, 
the difficulty of regarding the vascular bundles as originating in the parasite and drawing 
their nourishment from the root, appears to me less than that of regarding them as 
dependent both for origin and increase upon a reversed and diseased action of the root. 
The great theoretical objection to this view is, that the anatomical characters of the vas- 
cular bundles of the parasite precisely resemble those of the root, and that in some species 
they are even found to arrange themselves in the forms of woody plates and medullary - 
rays, enclosing a pith axis, and to be surrounded by a cortical layer (Plate VIII. figs. 10, 3 
llo) It must however be borne in mind, that there is no law more universal in the 1 
vegetable kingdom than that vascular tissue is developed according to the requirements | 
of the plant, both as to abundance and kind ; and that the formation of a perfect organic _ 
cohesion between the walls of the individual cells of the cellular systems of the parasite - 
and root, is in no respect less anomalous than the similar perfect and intimate organie 
cohesion between their respective vascular systems. As the rhizome increases, the 
organic cellular cohesion extends with the increased surfaces of the parasite and root, by 
the merismatie subdivision of the cells of both; and the vascular system increases by the 
development of pleurenchyma, ducts, &e. from those nucleated cells which are found in 
the positions in which vessels are required. 

- In a case of parasitism like that of Balanophora, which involves perfect organic cohesion « 
between the individual cells of different plants, it must obviously in many instances be 1 
impossible to draw the line between the tissues of the parasite and those of the root on 1 
which it grows. With regard to the cellular tissues, however, there is generally no diffi- 3 
culty; for, that of the Balamophora containing organie compounds (wax), the line of 1 
union is evident; but it is different with the vascular systems, which consist in both cases 4 
of tubes of indefinite length, containing no solid organized contents, and presenting an q 
extreme simplicity of form. Again, granting (as we must) that in Lophophytum (and in | 
Scybalium, according to Unger’s observations) the vascular tissue of the rhizome never - 
descends to that of the root, and hence cannot form an organic cohesion with the latter, we 
must assume an independent origin for it in these genera, at any rate; the application of | 
which to Goeppert’s views involves the necessity of concluding that there are two funda- 1 
mentally distinct principles of development amongst very closely allied species ; namely, that 3 
the germinating plant of some does form. independent vascular bundles (in common with all . 
Phænogamic plants), but that that of others does not. To me it appears more in accord- ' 
ance with the known laws of development, to suppose that the origin of the vascular 
system is the same in both, but that its after-development is modified in different cases. 
In Langsdorfia, where the rhizome has certainly a highly developed vascular tissue of 
its own, and where the root also appears to send branches into the rhizome, although I 
have never found the vascular system of the latter to unite with that of the root, I can- 
not but admit that such a union may exist, for the difficulty of dissecting the mixed brittle, 
woody, and flaccid tissues of this plant is very great. : 

The last argument which I shall bring forward in favour of considering the vascular 


system of the rhizome as in its origin proper to the parasite, is derived from the fact of 
free vascular bundles being formed in the flower-buds or nascent peduncles; which is 


ea da ne inc 


AFFINITIES OF BALANOPHOREÆ. 9 


conceded by Gæppert and all observers, and which is easily demonstrated. Hence, 
besides the difficulty of reconciling the ‘theory of two origins for the vascular tissue in 
one plant to any known law, we must also break through the well-established law, that 
the formation of buds is a repetition of the process of germination. 

2. M.Gœppert accurately describes the vascular bundles of the inflorescence as origina- 
ting in the buds whilst still enclosed within the rhizome; but whereas he figures and 
describes them as having free terminations, I find them to become confluent with the 
vascular bundles of the rhizome. To any one versed in the dissection of vegetable tissues 
it can be no wonder that this is a point almost incapable of demonstrative proof in the solid, 
opaque tubers of Balanophoreæ, which generally turn of a deep brown when first cut, and 
become black in spirits; whose tissues cannot be torn; and in which the vascular bundles 
of the peduncle are so delicate, and run in such sinuous courses, that it is impossible so to 
bisect a plant that these bundles shall be traced continuously from the inflorescence to 
the base of the rhizome: I have, however, repeatedly found that the appearance of a free 
termination to the bundles is produced by cutting them obliquely across. A long mace- 
ration of the parts, and a careful picking away of the cellular tissues, are the only means 
I have found available for proving their confluence by direct observation; but at the same 
time I must confess that, whilst carrying on these dissections in various species of Bala- 
nophora, and in Rhopalocnemis, I have repeatedly changed my opinion, and indeed have 
on some occasions been almost convinced of the truth of the contrary view to that I have 
finally adopted, so deceptive are appearances. _ 

In the tuberous Helosideæ, and in Cynomorium and Sarcophyte, the bundles of the 
rhizome are so unsymmetrically arranged, so much smaller in diameter, and so much 
more tortuous, that I have hitherto been unable to trace this confluence in them; whereas 
in both species of Helosis, and in Langsdorffia, which present the most perfect develop- 
ment of a cylindrical rhizome, the origin of the vessels of the peduncle in those of the 
rhizome is perfectly evident, and requires little skill in dissection to demonstrate. 

+ 8. With regard to the anatomical differences stated by M. Geeppert to exist between 
the vessels forming the vascular system of the rhizome and of the peduncle, they 
- certainly do not exist in all the species. This is however quite consistent with Goeppert’s 
analysis being perfectly accurate, for it is to be expected both that the vessels of the 
perennial rhizome should differ from those of the annual peduncle, and that from the 
form and direction of development of these organs being essentially different (the one 
chiefly increasing in breadth and the other in length), their vessels would be different also. 
In Balanophora the cellular sheath enclosing the vascular bundles is the same in the 
rhizome and peduncle (except that the individual utricles are longer in the latter); in 
both cases the cells are colourless, void of solid contents, and with few dots or markings 
on their walls; thus always contrasting strongly in appearance with the adjacent paren- 
chyma, which abounds in wax. (See Plates IV., VI. &e.) j 
- In the fully-formed flowering specimen of B. involucrata (Plate IV. fig. 1), I find no 
greater differences between the vessels in the rhizome and those in the peduncle than 
might be expected in organs so dissimilar in age and proportions. Plate IV. fig. 14. is a 
transverse, and 15. a vertical section of å vascular bundle from below the capitulum, com: 
VOL. XXII. C 


10 DR. J. D. HOOKER ON THE STRUCTURE AND 


posed of elongated cells which are more opaque towards the centre of the bundle; lower 
down in the peduncle they present the same appearance as is represented in figs. 26 and 27 f 
which show vessels from the apices of the bundles in the rhizome. In fig. 22 some vessels 
of the rhizome, and on the left the base of one of those of the peduncle, are seen : at this 
part of their course both are regularly banded; which is better seen in figs. 23 and 24, 
where they are more highly magnified. Every intermediate form of vessel may be found 
between those represented at figs. 15 and 27; and occasionally in both organs the form 
of cellular tissue, seen at fig. 25, is found; which appears to be another modification; 
intermediate between the vascular and cellular, and which, in fig. 23, is placed outsidé 
the barred vessels. | "n; 
In older specimens of B. involucrata much larger cylindrical vessels are found in thé 
rhizome, mixed with hexagonal tubes with barred or otherwise marked sides, and bundles 
of pleurenchyma, which also occur in the peduncle, but in a much less perfectly deve- 

 loped state. 
I find considerable uniformity in the microscopical character of the vessels amongst 


different specimens of B. dioica, although these have grown on widely different genera of l 


plants (and the same remark applies to Rhopalocnemis) ; and I do not in any case find å 


more highly developed tissue in the peduncle than in the rhizome. Ona comparison of : 
my dissections of young specimens of B. involuerata with Gæppert's of old ones af 1 


B. elongata, the differences between them are perfectly reconcilable. Gceppert figures 
barred cylindrical vessels of cellular tissue as occurring only in the peduncle, and larger 


vessels with short transverse bars as occurring only in the rhizome; this I also find to be _ 
the case in old specimens ; but in younger ones the barred cylindrical vessels are abundant - 


in the rhizome, and comparatively rare in the peduncle; from which it may be inferred; 
that the said vessels are an imperfectly developed tissue. 


In full-grown specimens of B. fungosa (Plate VIII. fig. 12), the same forms, relations, 4 
and modifications of vascular tissue prevail to a considerable extent; and the same may |. 


be said of other species which I have examined, though less in detail; whence I conclude 
that the anatomical differences between the vessels of the rhizome and those of the peduncle 
are dependent on position and degree of development. i 
The rhizome of the most perfect species of Balanophoreæ is decidedly exogenous. Ifa 
transverse section of the elongated one of Helosis mexicana be taken, the mass will be 
found to be composed of cellular tissue, enclosing (in the specimen given at Plate XV. 
fig. 14) a vascular system consisting of seven wedges, which surround a narrow cylindrical 
axis. Each of these wedges is (on a transverse section) narrow and oblong, and consists 
_of many rows of annulate or transversely barred cylindrical or angular ducts, which 
occupy the position of the pleurenchyma of ordinary exogenous plants; outside of these is 
a reniform mass of stout, elongated liber-cells, into whose concave faces the outer ends of 
. the wedges are thrust. Beyond the vascular system is a very thick spongy cellular mass 


are smaller and denser: this cellular 


asses of thick-walled sclerogen-cells, round 
which the cells of parenchyma radiate, and which, in a transverse section, resemble scat- 


å seven-lobed zone of stout sclerogen-tubes, 


él oc L 


AFFINITIES OF BALANOPHOREÆ. 11 


the lobes of which project outwards as the bases of the medullary rays ; and between these 
lobes lie the axial ends of the vascular wedges. The following is a summary of these 
characters :—1. The axis is occupied by hexagonal cells, which become vertically elon- 
gated and woody (see 2.) towards the vascular wedges, and then radially elongated in the 
medullary rays, and pass insensibly into the membranous hexagonal tissue of the cortical - 
portion: these cells contain grains of starch, and chlorophyll in abundance. 2. The 
woody tubes forming the outer zone of the axis (which is in many respects analogous to 
a medullary sheath) consist of long and strong cylindrical pleurenchyma, with much- 
elongated angular sclerogen-cells: these are all extremely hard, and their walls are per- 
forated by innumerable canals. In old specimens the pith passes gradually into these 
tissues; its utricles becoming first cubical, with thick dotted or perforated walls; then 
becoming tubes elongated vertically ; which are succeeded by tubes with blunt ends and 
narrow cavities. 3. The wood consists wholly of scalariform vessels which are cylin- 
drical in young rhizomes, but polygonal with transversely barred or gashed walls in older 
ones; intermixed in every instance with smaller, more irregular and variously marked 
or perforated cells and tubes. 4. The liber-bundles consist of large, stout-walled, woody, 
hexagonal tubes, of great density ; their walls everywhere perforated by canals. 5. The 
isolated sclerogen-cells in the cortical portion in no respect but shortness differ from liber. 

Both in arrangement and in anatomical characters this description of the rhizome re- 
sembles in most particulars that of the stems of many Menispermee ; and a more close 
examination bears out this resemblance. 

- In a transverse section of the peduncle of Helosis mexicana (Plate XV. fig. 12), eight 
symmetrically disposed vascular bundles are seen, and outside of these a few smaller 
irregularly scattered ones: and, as in Balanophora, the anatomical structure of the ves- 
sels composing these differs from that of the rhizome only in degree. The bundles consist 
of a sheath of elongated cellular tissue, enclosing a few fusiform vessels, some scalariform, 
others with spiral bands or transverse bars, with a few woody tubes and sclerogen-cells ; 
and these may be traced up to the scales of the capitulum, to which scales much stronger 
bundles are given off than to the flowers. 

- In the rhizome of Helosis guyanensis (Plate XVI. fig. 80) I find—1. The whole pith 
torpid of the same woody vessels as surround the pith of H. mexicana; and this both in 
New Grenada, Trinidad, and Rio de Janeiro specimens; these pass into a muriform tissue 
of woody tabular cells, which occupy the broad medullary rays, and of polyhedral cells 
still with very thick walls, in the circumference of the rhizome. 2. The wood is seen in 
a transverse section to be formed of seven lanceolate wedges of soft, white, scalariform, 
or spirally unrollable tubes. 3. A very large reniform mass of liber-cells or short tubes _ 
is placed outside each wood-bundle, and in contact with it. This does not seem to in- 
crease annually, but other and equally large liber-bundles form a zone exterior to these, 
and alternate with them; as in many Menispermeæ. 4. Isolated masses of sclerogen- 
cells and long liber-vessels are scattered throughout the parenchyma of the periphery. — — 

The peduncle of Helosis guyanensis presents innumerable bundles of vascular tissue, 
composed of sclerogen-cells, spirally marked and scalariform vessels, and a few woody tubes, 


generally occupying definite relative pe In a specimen of H. guyanensis from 
c2 


' ^ . ‘ DR. J.D. HOOKER ON THE STRUCTURE AND 


Columbia the medullary system is much more utricular, lax, and membranous; but there « 
are so many modifications of all these tissues in different specimens of the same species 
and parts of the same specimen, that it would be useless to multiply descriptions of them. — 
In all the other Helosideæ the same vessels are very conspicuous; but owing to the - 
form of the rhizome they are confused in arrangement and variable in amount, frequently 1 
presenting no system whatever. i 
Langsdorffia presents the same exogenous arrangement in its rhizome as Helosis, but | 
its axis (pith) is formed wholly of long wood-tubes (Plate II. figs. 5 & 6): its tissues are — 
more particularly described under the remarks on the genus itself; where also its resem- 1 
blance to the Indian Balanophoreæ in its waxy cell-contents is noticed. 
Cynomorium has a rhizome which I have never seen to branch, though luxuriant speci- - 
mens probably do so. The fusiform axis at the base of the peduncle, which is probably — 
not the rhizome, but only the base of the peduncle, presents in a transverse section - 
many small, unsymmetrieally disposed vascular fascicles: each of these is composed of— 
l. towards the axis a bundle of delicate, white, cylindrical and angular, barred or scala- — 
riform vessels, or long polygonal cells with variously marked faces:—2. externally to - 
this is a rather broad mass of vertically elongated oblong cells, of equal length; with — 
blunt superimposed extremities, which all meet at the same height; giving this tissue a 
transversely marked appearance. | 
The tissues of Sarcophyte and Mystropetalon present nothing remarkable. EG 
Cellular tissue.—This has been extremely well described in the Java species, by Geeppert, | 

of whose remarks the figures of B. involucrata (Plate IV. figs. 7, 8, &c.) are illustrative. — 
The walls of the cells are almost invariably dotted; in some cases owing to pores, and in | 
others to deposits of wax and chlorophyll. Very frequently (and always in young speci- 1 
mens) each cell presents a conspicuous cytoblast, firmly adherent to a discoid spot. At - 
Plate IV. fig. 11. are seen some of the waxy contents of the cells, in the shape of spherical. 1 
or rounded nuclei of various sizes; full of utricles, which appear to burst, and scatter their - 
granular contents within the cell, which is seen ruptured in fig. 13. b 
The wax of Langsdorffia and Balanophora is replaced in most of the other genera by : 
starch-grains: these are especially abundant in Sarcophyte, Oynomorium and Lopho- | 
phytum, which are in consequence eaten, as are other species occasionally*. The fluids . 
of most of the species are colourless or pale yellow; those of the Indian Balanophore are | 
quite white, and often very viscid. b 
I have never observed the appearance of the red cortical layer of the bark of the root, - 
which Gæppert describes as ascending with and surrounding the vascular bundles of the | 
rhizome in Balanophora, and which, he adds, contains tannin: it is, however, very con- | 
spicuous in Langsdorfia, and probably developed more or less in many other species. I 
have not found the raphides which he describes in the J avanese B. alutacea. | 
Unger calls the elongated parenchyma-cells with cytoblasts, « 
and notices their similarity to vessels that occur in Filices ; and 
affinity between Balanophoreæ and Acrogens. Gæppert also, 


‘-* A chemical analysis of this wax is 
the wax of Ceroæylon andicola. . 


pseudo-pleurenchyma,” - 

he hence alludes to an 
considering that the cellular — 
given by Gæppert, who ealls it Balanophorine, and observes that it resembles — 


AFFINITIES OF BALANOPHOREÆ. - E 


tissues of Balanophoreæ are more uniform throughout the whole plant than in any other 
vegetables in which so abundant and high a development of cytoblasts occurs, is inclined 
to refer all Rhizanths to one class, which he would place amongst Acrogens, and near 
Filices. I need scarcely say that these feeble analogies do not appear to me to be of the 
smallest systematic value; so long as they are unsupported by definite characters, and 
that any such affinity is negatived by every other point in their structure and deve- 
lopment. 

The cuticle of Balanophoreæ never presents stomata, but is very simple in its structure, 
and formed of small cells, sometimes however of large vesicular ones, either isolated or 
in groups; as in B. involucrata (Plate IV. figs. 7 & 16): in other specimens clusters of 
bladdery cells form warts on the rhizome (fig. 16), which are arranged in lobed masses in 
B. elongata and others. 

Hairs rarely occur on plants of this Order, though they are abundant on Langsdorffia 
tomentosa, and found on the flowering stem of Thonmingia: in both genera they are un- 
branched, cylindrical, rather blunt tubes, with swollen and often bulbous bases, and more 
or less rough surfaces, and have thin walls and a large continuous empty cavity. 

Sclerogen-cells, or clostera, abound in most of the species, and always present very 
thick, woody, perforated walls: they are especially conspicuous in the rhizome of Langs- 
dorffia, in the cortical layer of that of Lophophytum, and in the leaf-scales of the mel 
plant, in which they pass into the form of tubular vessels. 

Foliar organs.—No species is wholly deprived of these, though in some they are almost 
absent, and in others represented by scales on various parts of the plant; rarely on the 
rhizome, and most frequently on the capitulum, where they form more or less perfect 
bracts. There is, however, no obvious law for their development. In Cynomorium and 
Lophophytum they occur on all parts, from the rhizome to the apex of the capitulum. In 
Langsdorfia, Thonningia, Balanophora, Scybalium, and Mystropetalon, they are more 
or less highly developed on the peduncle, and very much reduced on the capitulum: in 
Phyllocoryne, they clothe both the peduncle and capitulum : in Sphærorhizon, they oceur 
only at the base of the peduncle, on it, and on the capitulum : in Helosis guyanensis they 
cover the capitulum, but on the peduncle and at its base are reduced to a few small scales ; 
whilst in H. mexicana, Rhopalocnemis, and Corynæa, they are almost er to the 
former organ. 

It hence appears that their chief development is upwards ; the most rudimentary forms 
occurring on the rhizome at the base of the peduncle, where they compose the bud-scales ; 
the most perfect on the capitulum, where they appear as bracts. 

The bud-scales are numerous and imbricating in Spherorhizon, and probably also in 
Scybalium; valvate in Langsdorffia and Helosis guyanensis; reduced to a volva or ring 
in Balanophora and Rhopalocnemis; and absolutely wanting in Corynea. In most of 
. the species the foliar organs are alternate; but in several Balanophore, in Langsdorffia, 
and Helosis guyanensis, those of the peduncle are whorled, and together form a cup; 
while they are reduced to an obsolete ring in Helosis mexicana. 

Inflorescence.—The flowers are arranged in a uni- or bi-sexual spherical, oblong, cylin- 
drical, or ovoid capitulum, in all the genera except in those of Lophophyteæ and in Sar- 


14 DR. J. D. HOOKER ON THE STRUCTURE AND 


cophyte, in which they occur in compound spikes or panicles. However simple these 
capitula appear, they are invariably found to be compound if examined at an early period 
of growth, when the bracts or scales imbricate completely over them, and cover definite 
masses of flowers, representing branches of the inflorescence. Sarcophyte presents 
the most perfect inflorescence, and the only one with a fully branched panicle; it has 
general bracts on the main axis below each ramification, but no partial ones. Lophophy- 
tum presents the next degree of perfection in inflorescence: each bract is a very highly 
developed peltate organ, subtending a cylindrical branch of the main axis, which is covered 
with flowers :—a modification of this arrangement is found in all the Helosidee, and in 
Cynomorium, where the bracts are peltate and imbricate in a young state, and either 
peltate and attached by their margins, or scattered, in the older state. 
In Ombrophytum the flowers are whorled round the pedicel of a very complete peltate 
bract, and in most Balanophore the female flowers are similarly arranged round a very 
-rudimentary clavate one. In Thonningia and Langsdorffia the female flowers have no 
bracts whatever, and the male flowers very rudimentary ones. Mystropetalon, the most 
highly developed genus in many other respects, has a trifid bract under each flower, and 
no general bracts on the capitulum. : 
Articulated filaments occur abundantly over the whole ‘surface of the capitula of most 
of the Helosideæ, and are probably rudimentary female flowers: their similarity to the: 
paraphyses of Musci has been dwelt upon by Griffith, who (with some other authors) 
attaches great systematic value to this resemblance. These anomalous organs will be 
described under the respective species: analogous ones may be seen in the capitulum of . 
Langsdorfia, and between the male flowers of some Balanophore. For further structural 
particulars respecting the inflorescence, the individual genera must be consulted. | 
The periods of inflorescence present some remarkable anomalies in Balanophore, and — 
especially in the Helosideæ with bisexual capitula; a curious phenomenon, first observed 
by Richard (fully described by him under Z. guyanensis), which necessitates the agency 
of dichogamy, or the fertilization of the ovaria of one capitulum or plant by the males of 
another. | 
Some genera are constantly diecious; as Langsdorffia, Thonningia, Rhopaloenemis, — 
Sarcophyte, Lophophytum, and some Balanophoræ; though in B. dioica, which is one 
of the most constantly so of that genus, I have occasionally found male capitula on 
the same rhizomes with female ones. The inflorescence is bisexual or monæcious, with 
the male flowers below, in some Balanophore : the male flowers are above in Lophophy- 
tum and Mystropetalon, and the two sexes are irregularly mixed in Helosidee and Cyno- 
morwm, which latter occasionally presents also hermaphrodite flowers, 
Flowers.—These present many gradations of perfection, both in the male and female. _ 


They are most fully developed in Mystropetalon, and the least so in the female of Bala- 
nophora and the male of Lophophytum. 


The perianth, when present, is almost invariably 
male flowers : in those of Lophophytum it is who 


AFFINITIES OF BALANOPHOREÆ. 15 


scales; in Rhopalocnemis it is tubular; in Spherorhizon and Corynea it is tubular 
below and campanulate above; in Helosis, Scybalium, and Sarcophyte it is tubular below, 
with three valvate segments; in Balanophora solid below, with three to eight valvate 
segments; in Mystropetalon it is irregular and oblique, of one free and two combined 
pieces, all valvate, and forming a tube below. The æstivation of the perianth is inva- 
riably valvate. 

In the female flowers of all the genera but Cynomorium, the perianth differs very 
widely indeed from that of the male; as much soas in any Natural Order of plants. It is 
generally far less highly developed than the male, though more so in Lophophytee and 
Thonningia. It is assumed to be more or less adherent with the ovary in all the genera, but 
is perhaps totally suppressed in Balanophora, which presents the simplest possible form of 
female flower. In Sarcophyte the ovaries are immersed in a fleshy perianth, and all cohere 

into a solid capitulum. All the Distyli have two confluent ovaria, forming a one- or rarely 

two-celled pistil, and crowned by a two-lipped perianth ; except in the case of Lophophytee, 
in which the limb is truncate or suppressed. In Cynomorium the six pieces of the 
perianth adhere to the ovary at irregular heights, being rarely wholly superior or wholly 
inferior. Thonningia and Langsdorffia have slender tubular perianths, which are solid 
below, and bear at their very base a small ovule, which is sunk in the fleshy capitulum : 
in the latter of these genera the female perianth much resembles the male, and its mouth 
is sometimes swollen and obscurely three-lobed. The female flower of Mystropetalon 
departs widely from the general type of the Order: the spherical ovary is seated on an 
oblate disc, and crowned by a small, campanulate, three-lobed, deciduous perianth, which 
may either be considered as the articulate free limb of the adherent calyx, or as a corolla. 
Analogy with Haloragee, Rubiacee, Composite, &c., suggests the latter explanation, 
which however is opposed to the fact of there being no double perianth in the male 
flowers of this genus, or in any other plant of the Order. 

Stamina.—The diversity of form so conspicuous in the perianth of different genera of 
Balanophoree is shared by the male organs, which agree in no point save the production 
of pollen. In Lophophyteæ the stamen is of the normal form, but only two in each 
flower, and without any other perianth than two mamillæ: the stamens have a very short 
filament and a long linear anther. In Cynomorium the stamen is solitary and of the usual 
form, but surrounded by a perianth, and subtended by a rudimentary style: in herm- 
aphrodite flowers it is epigynous, and the filament is stout, attached to the anther by a 
very small point, and thé anther is introrse. Mystropetalon presents the next modification, 
having three free stamens, each opposite a division of the perianth, and being similar 
to that of Cynomorium, but with an extrorse anther. Sarcophyte has three free stamens, 

opposite the valves of the perianth; they have fleshy filaments and adnate subspherical 
capitate anthers, full of polliniferous cavities. In all the remaining genera, viz. those of 
Helosideæ, Thonningia, Langsdorffia, and Balanophora, the stamens are opposite the lobes 
of the perianth; they are usually three in number, but vary in this respect; and are more 
or less confluent, both by their filaments and anthers. The dehiscence varies extremely, as 
does the number of cells. In all the species of Balanophora, Langsdorffia, and Thonningia, 
the anthers burst extrorsely, and have two or more loculi, which are confluent or anfrac- 


16 DR. J.D. HOOKER ON THE STRUCTURE AND 


tuose in some species of Balanophora: in all the Helosidee they burst introrsely, and 
also by their apices, which decay away, and thus allow the pollen to escape. : 4 

The tissues of the anthers present little peculiarity ; that of the connective is simply 
cellular, without any of the spiral vessels or beautiful modifications of banded or annulate 
cells, so conspicuous in the endothecium of many plants. It is frequently lined with a . 
pulpy mass of mucilaginous filaments of excessive tenuity, which appears to be the | 
remains of the tissue amongst which the pollen-cells were elaborated. - = 

The pollen of various species has been carefully described by Gæppert, Griffith, and — 
others, and presents nothing remarkable: it is generally spherical, often 3-lobed or 3- 
nucleate; in Mystropetalon it is polygonal. The surface of the extine is occasionally 
minutely granulated. Impregnation is probably mainly effected by insect agency, and at 
night; for during the day there is a singular want of insect life in the still, humid forests 
frequented by the species of Balanophora. I have, however, failed in many attempts to 
trace insect action; and a small Acarus feeding on the pollen of the monæcious B. invo- 
lucrata is the.only one I have found to be concerned in the operation, and that no doubt 
quite accidentally. The necessity of cross impregnation is manifest in the Helosidee, as 
indicated by Richard, and elsewhere explained in this Essay (under the genus). The fact 
of insects forming a nidus in the fleshy plants of this Order, has been used as an argument 
in favour of insect action assisting in impregnation; but the same might be applied to — 
any fleshy fungus or fruit. 

Ovaries.—These vary in number, from one in the Monostyli and Sarcophyte, to two in 
the Distyli ; and, according to Endlicher, to sometimes three in Helosis and Scybalium. 
When there are more than one, they are congenitally coherent, enclosed within the ad- 
herent perianth, and all the cavities but one are suppressed (being rarely present, according b 
to Endlicher and Schott, in some flowers of Helosis and Scybaliwm), whilst the styles — 
invariably remain, and are equal, and symmetrically disposed (right and left to the axis) 
at the summit of the perfectly symmetrical one-celled ovary. In those species which 
have a perianth, it may be traced surrounding the ovary, if examined before the latter 
begins to swell, and at all periods in some species; in most, however, the walls of the 
ovary become indurated, and blend insensibly with the adherent perianth, whose limb 
ee on à rud lea ae o the EHE in the Lophophyteæ it 
ture has been alread ax lai ed en ig Mats ee SASE ep Ade dte ric 
a y explain uud 'ongst the Monostyli the ovary is always one-celled; 

gsdorffia and Thonningia it is enclosed in a very evident perianth. In Bala- 

nophora there are not even rudimentary traces of a perianth. 
sit pre, ll, d Maw 
few oblong cells surrounding pei pulpy en en d Aes 
distinct stigma, and the termination of the style WR fr did. 
fiogu M a ee J'e scarcely differs from any other part of 
I have found a the ks t gi Sag ee = horis die Nig $i å 
- capitate, and rather more ek in th ina å er ho; De gr. 
globular cells. After ee Be så cras Eies par larger, oen 

| 'gnation, ese cells, when very highly magnified, 


_ AFFINITIES OF BALANOPHOREÆ. 17 


appear minutely wrinkled on the surface. The long single style of Mystropetalon termi- 
nates in a clavate or capitate, and evidently 3-lobed stigma. Sarcophyte has a sessile, 
broad, discoid stigma. The style of Cynomorium is more complex than in any other 
species, and terminates in a 2-lobed stigma; it is provided with two vascular cords and a 
central groove occupied by stigmatic tissue :—a detailed account of it will be found in the 
remarks upon C. coceineum. : 

In all the above-mentioned plants the cellular tissue of the ovary is very loose, consist- 
ing of oblong utricles, usually furnished with cytoblasts, and without any vascular tissue 
in its walls (except in the style of Cynomorium) : there is, however, a manifest approach 
to vascular tissue in the woody cells of the superior perianth of Thonningia, and perhaps 
also of Langsdorffia. 

Ovule.—This is invariably solitary, and pendulous from the summit of the cavity of the - 
ovary. In both Monostyli and Distyli its insertion is so near the very centre of the cavity, 
that I cannot detect any deviation in its position from the axis of the ovary; nor in the 
Distyli do I find it to be placed nearer to one of the styles than to the other. 

The earliest appearance of the ovule of Balanophora is as a solitary cell, protruded from 
the wall of the ovary: its subsequent stages I have followed to some extent in B. involu- 
erata, though, owing to the rapid sphacelation of the cellular tissue of the ovary imme- 
diately after opening it, and the extreme minuteness of all the parts, the analysis is one of 
great delicacy, and proportionately liable to error. 

Plate V. figs. 11 & 12. represent an opened ovary of B. involucrata, showing a very 
young ovule, consisting of a delicate hyaline sac suspended almost immediately below the 
insertion of the style, and containing two free spherical cells, each full of fluid and covered 
with opaque spots, which are probably cytoblasts. I found it impossible to detach the 
ovule, or to view it, except im situ, and by transmitted light. The formation of cells pro- 
ceeds with great rapidity within the sac, but I was unable to trace their evolution. The 
resemblance between the cells thus developed, and those in the embryo-sac of ordinary 
ovules, is obvious, and it suggests the possibility of the ovule being reduced to an embryo- 
sac. I could obtain no clue to the period at which impregnation is effected, nor to the 
particular action of the pollen-tubes, which I never found within the cavity of the ovary 
or ovule* : nor could I trace on any part of the surface of the ovule, any indication of a 
chalaza, raphe, or foramen, at which impregnation is probably effected. After the ovule . 
has swelled, so as to fill the cavity of the ovary, it adheres by means of its membranous 
coat to the walls of the ovary ; at which time it consists of a dense opaque mass of 

eohering hexagonal cells. 

. "The ovule, as thus described, does not materially differ from that of Viscum, as described 
in Decaisne’s admirable memoir on that plant (Mémoires de l’Académie de Bruxelles), 
except in being more simple; the ovule of Viscum consisting of an embryo-sac covered by a 
delicate cellular membrane (the tercine of Mirbel), and the greater portion of the substance - 
of its nucleus being undeveloped. Regarding Balanophora as presenting the most reduced 
form of ovule, Loranthacee are a step higher, and from these the passage is direct to the 

. naked nuclei of Santalacee and their allies, of Cornea, Caprifoliaceæ, Rubiacee, Umbelli- 
* But which M. Hofmeister has observed in the ovule itself of Cynomorium. . 


VOL. XXII. D 


18 Å DR. J. D. HOOKER ON THE STRUCTURE AND 


Jere, and some others; and from these, through the single-coated ovules of Menispermeæ, — 
many Monocotyledones, and other Orders, to the double-coated ovule of most Phænogams, « 
and lastly to the three ovular integuments of Gnetum. | 

This rudimentary ovule can by no means be compared to the archegonium of a moss; | 
nor does the reduction of the ovule to its simplest form argue any alliance between Bala- « 
nophoree and €ryptogamic plants. The affinity which Griffith endeavoured to establish, « 
is in this respect founded upon erroneous views of the origin and development of the ovule 1 
of Balanophora, with which he was not acquainted; and the development of this reduces 1 
the grounds of the argument to a casual external resemblance, or rude analogy, between - 
two organs which are not homologous, and have no similarity of origin, structure, or — 
function. ; 1 

 lvery much regret my having been unable to trace the development of the ovule in | 
any of the three embryonate and albuminous genera, Sarcophyte, Mystropetalon, and « 
Cynomorium; and can only suggest that in them the albumen is endospermic, or deve- - 
loped within the embryo-sac, and not in the substance of the nucleus. The position of — 
the radicle in Mystropetalon being close to the hilum of the seed, suggests the probability j 
of the ovule being anatropous, and hence somewhat more complex than its congeners; 1 
whilst the lateral position of the embryo of Cynomorium is consistent with an obliquely - 
pendulous ovule. Until, however, we become acquainted with the process of impregna- 1 
tion or the development of the ovule or the albumen, we have no materials for forming 
an opinion on the real nature either of the ovule or the seed. I have repeatedly dissected 

- half-grown ovules of Cynomorium preserved in spirits, but never found a trace of any coats 
to the ovule, which always appeared as a membranous sac, full of cells as in Balanophora, 
and amongst which cells one is free, and from it the embryo is developed. 

M. Weddell (Ann. Sc. Nat. sér.-3. v. 14) has also considered the ovule to be an embryo- 
sac, but he makes this opinion depend upon views of the nature of the ovary, style, stigma, 
and perianth, so different from my own, as to render our accordance upon this individual 
point purely accidental. I shall return to this subject after describing the seed. 

Seed.—There are two types of seed in this order; the embryonate, and what has been 
called by various authors the exembryonate, and which has been described as consisting: 
of a homogeneous or sporuliferous mass. The only known embryonate genera are Cyno- 
moriwm, Sarcophyte, and Mystropetalon. 

The seed is always pendulous from the summit of the cavity of the pericarp. The 
excessively thin testa contracts an intimate but not organie adhesion with the walls of the 
generally erustaceous endocarp, and is always so closely applied to the surface of the seed 
that it cannot be detached. "This structure is very frequent in various Orders of Exogens, 
as in Gunnera, whilst there is a manifest tendency to it in Araliaceæ, Boldoa (a South 
American genus of Monimiaceæ), and other plants. In the exalbuminous species the 
substance of the seed is uniformly cellular; the cells, which are loose in the ovule, and — 
fill the cavity of the pericarp previous to the swelling of the seed, become densely packed, 
probably from the cavity being limited in size, and its walls indurated before the seed has 
con E fall eaim When ripe, the seeds of most are densely corneous, especially 

periphery. The comparison of the seed to a loose cellular mass, so frequently 


AFFINITIES OF BALANOPHOREÆ. 19 


made by authors, probably in all cases arises from their having only examined immature 
specimens. 

The individual cells of the homogeneous embryo are angular, with very thick transparent 
walls, and small cavities filled with a few chlorophyll-granules. I have never found starch 
in the embryo of any species; the contents of their cells being browned by iodine. Oil 
abounds in the exalbuminous species, and in the embryos of Cynomorium and Mystropetalon. 

Hitherto the true nature of the exalbuminous, so-called exembryonate seed of Balano- 
phoræ has eluded all research; mor till its germination has been traced*, is it probable 
that this point will be satisfactorily cleared up. In the mean time it may involve less of 
hypothesis to assume that the embryo is a homogeneous mass, in so far as any evident 
distinction of cotyledons and radicle is concerned, than to regard it as an albumen in which 
the embryo has not yet been discovered. Much may be said on both sides of this ques- 
tion; for instance, analogy with Oyaomorium, in which the embryo is oily and the albu- 
men not so, is in favour of the seed of Balanophora being considered to be embryo; on 
the other hand, if the cellular ovule of Balanophora so perfectly resembles the embryo of 
Cynomorium, it appears reasonable to conclude that the albumen of Cynomorium is endo- 
spermic, and developed in the same delicate sac with the embryo itself; against which view 
there appears no theoretical objection t. 


* Impressed with the great importance of this point, I endeavoured, when in India, but uniformly in vain, to induce 
the seeds of Balanophora and Rhopalocnemis to germinate. 

t Amongst the many Natural Orders whose homogeneous seeds or embryos present more or less analogy with those 
of Balanophoreæ, none have so close a similarity as those of Triurideæ. I have examined a species of this Order in 
a living state in the Khasia Mountains (East Bengal) : its ovule (which has not hitherto been described) is manifestly 
anatropous, and consists of one integument and nucleus; offering one out of many proofs that the structure and posi- 
tion of the ovule in no degree influence the after-development of the embryo: in other words, that the development 
of the embryo, so far as its form and structure are concerned, is in a great measure irrespective of the presence or 
absence of envelopes to the embryonary sac. 

Mr. Miers, in his valuable and elaborate paper on Triurideæ (Linn. Soc. Trans. xxi. p. 51) considers that it is con- 
sistent with the simplicity of the stracture of other parts of the plants belonging to that family, “to expect a nucleus 
equally simple in its nature, formed merely of an aggregation of cytoblasts, which, under farourably-exeiting eircum- 
stances, are endowed with the faculty of self-development.” The true nucleus of the ovule in Triurideæ is however in 
no way different in structure or position from that of ordinary Phænogamic plants, from which it follows that although 
the embryo appears amorphous, its radicular extremity must be a determinate point with relation to the seed, and that in 
germination that end will elongate, and perform the function of the roots. The term * Protoblastus,” therefore, as indi- 
cating an embryo that germinates from no determinate point, cannot under this view be adopted for that of Triurideæ, 
though, if it were proved that the germination of Balanophore (the structure of whose ovules does not reveal the posi- 
tion of the radicle) were from an indeterminate point, it might be more applicable to them. In both Cynomorium and 
Mystropetalon however, the radicular end of the embryo is very evident, and as there can be no doubt that the embryos of 
most or all Balanophoree germinate whilst still within the pericarp, it may be inferred that the radicle will protrude 
from a given ruptured point of the latter, and not indifferently through any part of its walls. Under these circum- 
stances, I hesitate to adopt a term, which, in the present state of the inquiry, and as far as regards this order, implies, 
not that the germinating point is indeterminate, but that this, and the whole process of germination, are absolutely 
unknown. 

With regard to other embryos which would come under the definition of a Protoblastus, that of Orchidee evidently 
germinates from a given point; and Caspari’s beautiful observations on Orobanche show that the same is the case 
in that genus. Blume describes the embryo of Amorphophallus as throwing out plumulary leaves from several 
points at once, which probably indicates a development of. several much-reduced internodes crowded together, 


D2 


20 DR.J. D. HOOKER ON THE STRUCTURE AND 


Richard (Mém. du Muséum, viii. p. 429), and latterly Lindley (Veg. Kingd. p. 85), have | 
assumed the seed to be embryonate in all Balanophoree; arguing from that of Cymo- « 
moriwm, which both well understood; and I at one time adopted the same opinion, | 
being much influenced by the fact that in certain plants with densely fleshy albumens, | 
formed of large coherent cells, the embryo scarcely exceeds one such cell in bulk, and — 
often eludes a very careful search; as that of Mystropetalon escaped Harvey, Griffith, and 
others. Yielding, however, to the mass of evidence in favour of the absence of any visible — 
embryo within the seed of Balanophora and of all the Distyli, I am now inclined to agree M 
with Griffith (Linn. Trans. xx. p. 93) in considering, the embryo as a homogeneous mass, — 
or “ indivisus albuminiformis.” : | 

Endlicher (Meletemata, p. 9; and Gen. PI. p. 73) describes the seed as a nucleus, * nucleo | 
e tela cellulosa, massa sporacea farcta, conglobato,” and adds that the testa is coriaceous, - 
hard or subosseous, evidently mistaking the endocarp for a testa. Blume (En. PI. Jav.i. 
p. 87) seems to have taken a similar view of the contents of the seed. J unghuhn, an inge- 
nious and acute observer, says (Act. Acad. xviii. Suppl. p. 205), * Semina nulla adsunt; — 
quod (supra) ovaria salutavi, vix nisi analoga sunt germinum plantarum perfectarum que 
nunquam maturescunt, sed more fungorum putredine pereunt." "Trattinick also (Linnea, | 
iii. p. 194) says, under Sarcophyte, “that these plants are not developed from seeds, but | 
are specifie degenerations of the plants on which they grow." l 

Gœppert (Nov. Act. l. c. p. 257) considers that Balanophore grow from seeds, and | 

_ describes these as “nuda exembryonata;” and Nees von Esenbeck (Nov. Act. J. c. p. 225) 
calls them acotyledones of a high class. | 

Liebmann (Proceedings of Assembly of Scandinavian Naturalists at Christiania) says 1 
of the seed of Langsdorffia hypogea, that it is intermediate in character between a crypto- — 
gamic spore and a naked seed. 1 

Martius (Nov. Gen. et Sp. iii. p. 186) regards the seed as an embryo, and states that he — 
has seen small fibres given off from its basilar end, like rootlets; an observation not I 
hitherto confirmed. 3 

I have reserved to the last the discussion of M. Weddell's views, because they differ 4 
from those of other authors, and are based upon å comparison of an extensive range of . 
organs, which cannot be considered separately; they are published in a paper read before — 
the Société Philomatique of Paris, and more at length in the * Annales des Sciences Natu- _ 
relles* (ser. 3. xiv. p. 166): they especially refer to the relationship between Balano- 
phoreæ and Rafflesiacee, and may be thus summed up :— 

1. "The so-called fruit of Balanophoreæ is constructed on the same plan as the seed — 
of Rafflesiacee ; the so-called styles, which are almost always observable on that organ A 
before its maturity, are appendages of one of the essential parts of the ovule. The fruit of - 
Balanophoree must hence be regarded as a naked seed.” 


but all belonging to one axis. In Griffith’s admirable paper on Ambrosinia (Linn. Trans. xx.), an extremely ano- : 
malous embryo is reduced to the ordinary type b 


1 y a careful study of development and germination, and it is shown 
that though its parts are undistinguishable at first sight, each has its functions defined. It is remarkable that Griffith 


has not alluded to the strong resemblance between the embryo of Ambrosinia and the bulbils formed on the deformed 
inflorescence of several species of its near ally, Remusatia. 


AFFINITIES OF BALANOPHOREÆ. 21 


2. “The so-called flower of Raflesiaceæ may be regarded as an inflorescence; the 
pericarp of the fruit is a receptacle, of which the folds form the placentæ.” 

It is not my intention to discuss the second of these propositions*, and I therefore 
confine myself to the first. In his descriptions M. Weddell states that the female repro- 
ductive organ of Balanophora is the nucleus of an ovule, and that of Cynomorium, Helosis, 
Ombrophytum, and Sarcophyte, a nucleus surrounded by a peculiar envelope, which is not 
a perianth, but is formed from the axis, and is to a certain extent analogous to the inte- 
guments (pericarp) of ordinary seeds: he was led to this conclusion by a comparison of 
the fruit of Balanophora with the seed of Rafflesia; and adds, that there is no more 
fundamental difference between the pericarp with its anfractuous cavity, of Rafflesia or 
Hydnora, and the convex or peltate receptacles of Balanophora or Ombrophytum, than 
there is between the receptacle of a fig and that of a mulberry. 

In support of these views, Weddell contrasts the fruits of Rafflesia and Balanophora, 
and of Hydnora and Sarcophyte; but the comparison being maintained by the employment 
of the same terms for organs that do not appear to me to be homologous, the similarity 
becomes one of words, and not of facts. The term * styliform processes,” for what other 
authors consider the styles of the ovary, and which analogy suggests to be such, appears 
to be the most anomalous; and by describing them as almost always present in Bala- 
nophora, it is implied that they are sometimes absent, which I have never found to be the 
case. : 

In commenting upon Griffith’s theory that there is an analogy between the pistil of 
Balanophora and the pistillidium of Musci, Weddell points out that Griffith is in error in 
describing the styles as perforate, and adds in a note, * On reaching the periphery of the 
capitulum, this styliform process becomes eroded at the apex, when its internal cavity 
communicates with the external air: consequently the styliform process, being bathed in 
a mucous fluid that surrounds the capitulum, is exposed to the action of the fovilla of the 
pollen, which is mingled with that fluid, and fecundation is thus effected.” In proof of 
this it is added, that in some dicecious species of the Order, which do not secrete this fluid, 
the ovules remain sterile; such at least being the case with Langsdorffia and ‚Helosis. 

Weddell is right with regard to the imperforation of the style, for at no period do I 
find an open canal in the style either of the American or Indian species; but neither do 
I find any erosion, or other arrangement of the organs by which I can conceive an 
erosion to-be effected. The only fluid exudation I have seen on Balanophoree was a 
limpid watery one, on old capitula of Rhopalocnemis, after they had been removed from 
the ground; and this is a diœcious species, which was then in ripe fruit. 

Lastly, the structure of the hermaphrodite flowers of Cynomorium (which M. Weddell 
was not acquainted with) is conclusive against the pistil being regarded as a naked ovule. _ 


Affinities of BALANOPHOREZ. 
Polymorphism and an extreme simplicity in every organ are the prominent features of 
* No explanation of the staminal apparatus in Raffesia is given that is at all consistent with this view; and this 


therefore, as well as the presence of a discoid stigma, is adverse to the theory. Also, it is not shown how, if the 
seeds of Raflesia are truly naked, the pollen is applied to the nuclei. 


22 DR. J. D. HOOKER ON THE STRUCTURE AND 


this Order, when surveyed in a structural point of view; and were value to be attached to 
the fact of every organ appearing in a most degraded state in one or more of the species, _ 
Balanophoreæ would rank low in the system of Phænogamic plants. If however we 
disregard imperfection, and inquire what organs are wanting in the Order, we shall find | 
that, with the exception of terrestrial roots, all are present which are necessary to justify | 
their being placed amongst Phaenogamie plants. 3 
The arguments which have been used to exclude Balanophoreæ from Phænogams, all « 
appear to have originated on the one hand in mistaking feeble analogies between the forms 1 
of organs that are not homologous, for affinities; and on the other, in overlooking a mul- « 
titude of positive characters. These arguments may be summed up as being :— | 
1. An erroneous view of the nature of the seeds, by Endlicher, Martius, Blume, and | 
others, who describe them as a sporuliferous mass ; a term which, even were it applicable, - 
has no meaning. ; ; 
2. An erroneous view of their origin being in a diseased state of the plants they grow … 
upon; adopted by Junghuhn and Trattinick. i 
3. A supposed similarity in appearance to Fungi*, and an erroneous idea that their 
appearance is meteoric, and their growth rapid; a theory advocated by Endlicher, who — 
(Meletemata, p. 5) says of the horizontal rhizome of Helosis and Langsdorffia, “ mycelio 
Fungorum quam maxime analogum.” ER : 4 
4. The resemblance between the articulated filaments on the capitula of the Helosideæ, 
and the paraphyses of Musci; and between the pistilla of Balanophora, and the pistillidia 
of Musci; strongly advocated by Griffith and Lindley. 1 
5. The resemblance of the cellular and vascular tissues in some of their characters to — 
some of those of Filices; as indicated by Unger and Gæppert. 1 
6. A very peculiar view of the nature and relations of the parts of the female flower, | 
entertained by Weddell; who hence considers Balanophoreæ (together with Raflesiaceæ) — 
to approach nearer to Gymnosperms than to any other group of plants. 2: 
It would be fruitless to discuss these opinions at length; for it cannot be doubted 
that, had the authors who advocate them been sufficiently furnished with specimens and « 
facts, they would never have been entertained. On the other hand, it is not easy to 
account for the little importance attached by so many good botanists to the positive evi- 
dence afforded by the presence of sexes, the perfection of the essential organs of the male 
flower in all the species, the total dissimilarity in structure and function between the 
female organs of all the species and those of Cryptogams, and their identity of structure | 
in all essential points with those of other Phænogams. 
With regard to the union of Balanophoreæ with Rafflesiaceæ, into one great class of 
Phænogams, equivalent to Monocotyledones or Dicotyledones, the arguments brought | 
against it by Brown and Griffith are conclusive. Not only have these Orders no cha- 


either physiological or structural, except para 
of widely different affinities; which in the case 


» Brongniart, and Griffith have all placed Rafflesiacee in close 


* H : 
I may mention here that the species I have examined never became putrescent, 


AFFINITIES OF BALANOPHOREÆ. 23 


proximity to Aristolochiæ, an Order with which Balanophoreæ have not the slightest affi- 
nity. The arguments employed by Blume, Endlicher, Lindley, &c., for combining these 
Orders into one group, are also employed for removing both from Phænogams; the 
strongest reason for allying them being, not that they present characters in common, 
but that neither of them is considered to be allied to any other known Order of Phæ- 
nogams. 

Of the authors who consider Balanophoree to be Phænogamie, the majority refer them 
to Monocotyledones; Richard, and others following him, placing them in the neigh- 
bourhood of Aroideæ. 

Griffith places them in Dicotyledones, and suggests their being the homogeneous em- 
bryonate group of Urticee; to which the structure of the Distyli is quite opposed: nor 
indeed does he endeavour to support this hypothesis by any arguments, but merely 
throws it out as a suggestion. 

In my opinion, the arrangement of the vascular bundles in the rhizomes of Helosis and 
Langsdorffia is sufficient evidence of these plants being exogenous; for these, as I have 
elsewhere shown, are altogether exogenous, differ little from the stems of Menispermeæ 
and other anomalous, but still undoubted, Dicotyledons, and resemble no known Endogen 
in structure or arrangement. 

In endeavouring to determine the affinities of Balanophoreæ, I shall disregard the nega- 
tive characters, as those may be termed which are founded on the imperfection of organs; 
and I shall take the most perfectly developed species, as the best expositors of the typical 
structure of the Order. In so doing, I believe I am obeying a maxim supported by an 
attentive study of the natural system ; for there are few Natural Orders, however perfect, 
that do not present structurally incomplete genera or species, many of which in point of 
development of their organs might rank below many Balanophoreæ and some Cryptogams ; 
but which, nevertheless, are not departures from the type of their Order, but simply less 
developed forms of it. : 

Balanophoreæ have an adherent perianth in all the genera where this organ is deve- 
loped, and an epigynous stamen in Cynomoriwm, the only species in which hermaphrodite 
flowers occur. These characters indicate a position amongst the epigynous Calyciflore ; 
a group which, though far from being well limited as a natural class, is in our present 
state of knowledge one of considerable value, as comprehending many nearly allied natural 
families. Amongst them, the most direct relation of Balanophoreæ is certainly with 
Halorageæ, and especially Gunnera; with which it presents many important characters 
in common, especially the valvate perianth and stamens opposite its lobes, and near which 
I would place it in the linear series. 

A detailed comparison of the individual organs of a family so eminently polymorphous, 
with those of its allies, can alone establish its affinities: the more conspicuous of them 
are :— 

1. Between Cynomorium and Hippuris, in the one epigynous stamen, on an ovarium 
consisting of one carpel with a solitary pendulous ovule and simple style. 

2. Between the Distyli and Halorageæ a relation is established through the Australian 
genus Loudonia, which is a very peculiar form of the latter Order, having four styles, and 


24 DR. J. D. HOOKER ON THE STRUCTURE AND 


only one cell to an ovarium containing one or two pendulous ovules. This tendency to — 
suppression of the ovaria, combined with the constant presence of their styles, and the 
styles of the suppressed ovaries being in all respects similar to that of the developed ovary, - 
and equally perfect, is a very peculiar character, frequent in the Haloragee, though — 
not absolutely confined to them : it is very conspicuous in Gunnera. The greater tendency . 
to imperfection in the female than in the male flowers of Halorageæ, is also a marked . 
feature shared by Balanophoree. | 

3. Between Gunnera and Lophophytum the affinity is so close that the female flowers — 
of these genera might be mistaken for one another; and the male flowers of Lophophytum — 
in their two stamens, linear anthers and basal short filaments, are absolutely identical — 
with those of several species of Gunnera: in the subgenus Misandra especially, the male | 
flower often consists of two small sessile calyx-lobes, with two alternating stamens. 

4. If the female inflorescence of Gunnera and Lophophytum be compared, the affinity 
may be very easily pursued: in each, short conical branches of the flower-head project 
laterally from a stout axis, and are subtended by a large bract, and studded with a dense 
mass of flowers, which consist of an adherent perianth, no trace of rudimentary stamens, 
two styles, and a one-celled ovary, with a pendulous ovule, whose integument, in ripening, 
contracts an adhesion to the inner wall of the cavity. i 
. 5 The tendency to a dimerous or tetramerous arrangement of the parts of the flower, 
so conspicuous in all Halorageæ, and in Gunnera, is common to the Helosideæ and Lopho- 
phyteæ. 

Griffith has suggested an affinity between Mystropetalon and Loranthaceæ, founded on 
the form of the male perianth, and the opposition of the stamens to its lobes; but this i 
not borne out by the female flowers, which must be considered of the highest importance | 
in establishing affinities. Griffith further was ignorant of the true structure of the seed 
of Mystropetalon, and supposed that the genus had no relationship with Balanophoreæ 
After much consideration, however, I have included that genus in this Order, for reasons 
appended to some notes upon its structure; and in which view I am likewise followin 
that taken by Mr. Brown (Linn. Soc. Trans. xix. p- 233, in note). 

I have not dwelt upon the character afforded by the extreme dissimilarity of the sex 
of Balanophoreæ, and which is also conspicuous in Gunnera and Haloragee ; because i 
1s common to many other Orders, and indeed is perhaps a very constant accompaniment of 
reduction in structure, or of a normally imperfect development of the floral whorls. 
Amongst the objections that may be urged against associating Balanophoree with th 
epigynous Calyciflore, the strongest will probably be considered to be derived from the 
habit, and the imperfection of the foliar organs: with regard to the former, it appears 
wholly valueless, as will be proved by a cursory inspection of many Orders; and of these 
none are so conspicuous as Halorageæ, which, for its extent, is one of the most polymor- 


phous in the vegetable kingdom, and further, one consisting for the most part of reduced 
forms of Onagrariee. 


_ The extreme simplicity of the 
importance, 
there is a 


| structure of the seed and ovule is another point of some 
and may be used as an argument against the alliance I have proposed; bu 
manifest tendeney to such imperfection in the epigynous Calyciflore, especially 


AFFINITIES OF BALANOPHOREÆ. 25 


amongst Corneæ, Loranthaceæ, and several other Orders in which the ovule is reduced to 
a nucleus. The homogeneous embryo is (as indicated by Brown and Griffith) a form of 
that organ which, if taken alone, appears to be of little value in a systematic point of 
view; for it occurs in various genera belonging to natural families which have, typically, 
highly developed embryos; and this argument is further weakened by the fact of Cyno- 
morium, Sarcophyte and Mystropetalon having very obvious embryos immersed in 
| albumen. $ 

To parasitism, as a character of systematic value, I need scarcely allude; its invalidity 
being universally conceded. 

As an Order, Balanophoreæ may in one sense be considered a strictly limited one, not 
passing directly into any other, except perhaps through Gunnera into Halorageæ; and 
forming a sufficiently natural assemblage of species, though, owing to causes I have 
repeatedly dwelt upon, not easy of exact definition. Putting aside any consideration of 
. its relationship with other Orders, and regarding it per se, it is not easy to say whether 
it should abstractedly be considered as ranking high, or the contrary. Assuming that 
the conventional definition of perfection in use amongst zoologists is applicable to the 
vegetable kingdom, and which argues that a high degree of specification of organs and 
morphological differentiation of them for the performance of the highest functions, indi- 
cates a high rank, Balanophoreæ may in some respects be considered to hold a very bigh 
one. Thus:— : 

1. The moncecious and generally dicecious flowers show that either a whole plant, or a 
eonsiderable portion of it, is specialized for each sex. 

2. The great difference between the perianths of the sexes indicates a very high degree 
- of morphological differentiation for each special function or sex. 

3. The deviation of the parts composing the perianth of all the species from the com- 
mon leaf-type indicates great differentiation. 

4. The cohesion of the parts of the perianth in the male flower is a further deviation 
from that theoretical simplieity which assumes the leaves composing the floral whorls to 
be developed free from one another. 

5. The cohesion of the anthers and filaments in most of the species is a further —Z 
of specialization of the same nature. 

6. The adhesion of the perianth of the female flower to the ovary shows its special 
adaptation as a protecting organ to be carried to the highest degree. 

7. If it is safe to assume that either of the two perfectly equal and similar styles of the - 
group Distyli is capable of conveying pollen-tubes to the solitary ovule, we have here 
a very remarkable case of specialization; for these two styles undoubtedly belonged 
originally to as many ovaries, though finally specialized for the use of one only. 

As however is the case in all theoretical inquiries which are not based upon fixed prin- 
ciples, so in this as to the comparative rank of Balanophoree, there is much to be said on 
both sides. The reduced axis, the imperfect vascular system, the absence of leaves in many 
of the species, and their reduction to scales without stomata in the remainder, together 
with the invariable absence of a corolla, and of integuments to the ovule, are all evidences 
of a very low development. Whatever difference of opinion there may be as to the posi- 


VOL. XXII, E 


26 DR. J. D. HOOKER ON THE STRUCTURE AND 


tion of Halorageæ, of which I regard them to be reduced forms, it cannot be disputed that 
amongst Phænogams there are few groups so uniformly incomplete as regards the normal … 
complement of organs, or the arrested development of those organs which are present, - 
The value of this consideration is however much diminished by the fact, that there are no | 
limits to the suppression of organs in the individual genera of Orders which are, never- - 
theless, typically highly developed. 3 

In a systematic point of view, the value of these suppressions in the Vegetable Kingdom — 
diminishes to a great extent in ascending from the root towards the ovary: thus, the - 
absence of a root of the ordinary structure, and the adaptation of the lower portion of the : 
stem to a parasitic attachment, occur in six or seven natural families of Exogens which — 
are normally terrestrial, and perhaps in many more. A total absence of leaves, or a — 
reduction of them to minute scales, occurs in many natural families. A reduction of the — 
whole plant to a leafless, single- or few-flowered stem, is found in many parasites, and in — 
Orchidee, Burmanniacee, Ericeæ, Scrophularine, Triuridee, Rafflesiace@, Gentianee, « 
and other families which have no mutual affinities; whilst the reduction of the inflo- | 
rescence to a single flower, and the parts of the latter to its essential organs, is too . 
frequent to need specification. That of the embryo to a homogeneous mass is found in — 
various genera, as indicated by Brown and Griffith: the reduction of the ovule to an 1 
embryo-sac is however, in the present state of our knowledge, almost peculiar to Bala- | 
nophoreæ. (3 

As regards the including the Monostyli and Distyli under one Natural Order, these 1 
are so manifestly different, that it is a theoretical question how far, were there more | 
genera of each, or had they a widely different geographical distribution, they would E 
by common consent have been united into one natural family: and the same argument 1 
might indeed be applied with equal force to the removing Mystropetalon and even Cy- 

It is difficult to indicate any particular genus of Balanophoreæ which can be considered 
typical of the Order, though Cynomoriwm may be taken as such for the Monostyli, and 
Helosis for the Distyli. Mystropetalon, though in many respects the most perfect 
genus of the Order, cannot in any degree be considered typieal of it; for it departs far 
more widely from the prevalent structure of its allies than any other genus does. Our 
es of aet IS or 1s not typical, are, however, vague and arbitrary; the ideal type 
ae = prevalent form of the group, or that which unites most of the pecu- - 
stinguish it, or that which possesses the fullest. complement of organs 


united in one individual, or that in which these are most compl ially 
* ex, as well as specially 
adapted to the functions they perform. : E 


N Classification of BALANOPHORE.E. 


In the following arrangement of Balanophoreæ, I have been chiefly guided by the 


structure of the female flowers which are sen: 
3 ’ erally f. . t 
characters for systematic en g y ound to afford the most important 


The primary division into Monostyli and Distyli was proposed by Griffith (Linn. Soc. | 


AFFINITIES OF BALANOPHOREÆ. 27 


Trans. xx. p. 103); who however erroneously refers Rhopalocnemis (Phæocordylis, Griff.), 
of which his specimens appear to have been imperfect, to the Monostyli. The genera of 
each group are all more nearly related to one another than to any of those of the other 
group; and such a division is therefore perfectly natural: but there are, notwithstanding, 
such very great differences between the members of each group, that the genera of Mono- 
styli especially have all characters of far more than generie value, and may well be 
conceived to be types of very distinct assemblages of genera. This is not so much the 
case among the Distyli, for the two genera of Lophophyteæ are very nearly related, and 
- those of the Helosideæ so much so, that it may be doubted how far Corynæa, Scybalium, 
and Sphærorhizon are distinct from Helosis. j 

Sarcophyte must at present be considered as in many points a doubtful member of the 

Monostyli, from my inability to discover whether the ovarium is simple or compound, at 
any period of growth. The absolutely sessile, discoid stigma is found neither in Mono- 
- styli nor Distyli, and in some other characters it partakes as much of one group as of 
the other; thus in its three stamens it agrees with Balanophora and all the Helosideæ, 
but it differs from them in its stamens being free, and in its anomalous anthers (which 
however suggest an affinity with B. polyandra). In its branched inflorescence it resembles 
Lophophytee, but differs totally in the structure of the male flowers. 
- For characters of secondary importance, I have availed myself of the male flowers, and - 
especially of the number and form of the stamens and their cohesion. A remarkable 
analogy between two of the sections into which each primary group. is divided, is esta- 
blished by these organs ; the Lophophytee (of Distyli) having free stamens of the ordinary 
type of Phænogamic plants, and being hence analogous to Cynomorium and Mystrope- 
talon (among Monostyli), while all the other genera have anomalous anthers. 

In framing the genera, characters of the third degree of importance have been employed ; 
such as the presence or absence of the perianth, and its structure; aided occasionally by 
the stamina, the cohesion of the flowers, the nature of the inflorescence, and lastly, the 
habit; this term implying in these plants not merely differences in the outline of organs, 
but also in their development and mode of evolution. * : 

Besides Griffith's arrangement of Balanophoreæ, that of Endlicher (Meletemata, p. 4) is 
the only one of any importance hitherto proposed. His division is founded on the stamens 
being free or combined : this however not only assumes the position and arrangement of 
the stamina to be of more importance than those of the ovaria, but brings together genera 
which have otherwise little in common. * 


Geographical Distribution, and Variation. 


The greater number of Balanophorec belong to the tropical and subtropical mountains 
of Asia and South America, where they probably occur in nearly equal proportions. In 
both hemispheres certain species ascend to 10,000 feet ; comparatively few being found in 
low tropical forests, almost the only ones being Balanophora fungosa in the eastern, 
Helosis guyanensis in the western hemisphere, and Thonningia in Africa. A considerable 

E 2 


' 


28 DR. J.D. HOOKER ON THE STRUCTURE AND 


number aïé extra-tropical; as Cynomorium, which attains lat. 41° N. in Europe; the two 
Mystropetala and Sarcophyte, which inhabit South Africa; Helosis guyanensis, which — 
extends to the La Plata district; and the North Indian species of Balanophora and Rho- — 
palocnemis. 1 
The genus Balanophora is confined to India, and the Malay and western Polynesian 1 
islands: it extends from the N.W. Himalaya at Simla throughout that mountain-range to 
the eastward, thence to the Khasia Mountains, Burma, and the Malay Peninsula, Sumatra, [ 
Java, the N.E. coast of New Holland, as far east as the New Hebrides; it is also found in | 
the Indian Peninsula, Ceylon, and the Philippine Islands, and no doubt occurs in Borneo — 
and New Guinea. The Helosideæ, with the exception of the Indian Rhopalocnemis, are 1 
confined to the American continent and islands, where they extend from Jamaica and | 
Mexico to the Pampas. Of the Langsdorfie, two species are American, and one (Thon- — 
ningia) is found on the west coast of tropical Africa. The Lophophyteæ, as far as is | 
known, all inhabit tropical South America, and are chiefly confined to South Brazil, Peru, - 
and New Grenada. 
The individual species of this Order have often exceedingly wide ranges, though some 
are extremely local. The most conspicuous examples of extensive distribution are: Cyno- 
morium coccineum, which ranges from the Canary Islands to the mouths of the N ile, viz. 
through 3000 miles of longitude ; Rhopalocnemis is found in lat. 27° N. in East Nepal and 
Sikkim, in the Khasia Mountains of East Bengal, and in Java, under the Equator, places 
no less than 3000 miles apart; Balanophora dioica, which has probably a still wider 
range; and B. fungosa, which is found both in East Australia and Tanna, places separated 
by 1500 miles of ocean. In the new world, Langsdorffia hypogea has been found in the | 
province of Oaxaca in Mexico, lat. 18° N., by Prof. Liebmann, in thé mountains of New 
Grenada by Mr. Purdie, at Rio de Janeiro by many collectors, and in the Pampas by 
Mr. Miers (lat. 84^ 8.) ; having thus a range of 52 degrees of latitude, and 4000 miles in 
a straight line. 1 
I do not find that the widely distributed species vary much according to the distance 
they spread; specimens from the most distant localities often being absolutely identical 
and all being very constant to one form : on the other hand, some of the most local species - 
as well as some of the most widely spread are excessively variable. As a general rule, 
the most imperfect forms vary most in general characters, especially Balanophora, th 
individual species of which differ in the size of their parts, in the form of their scales, their 
rhizome and their capitula, in the capitula being unisexual or bisexual, and in the size, 
form and number of the parts of the flower. Helosideæ again vary extremely in size an 


habit, but much less in the capitulum. Langsdorffie are rather variable, and Cynomo- 
rum 18 conspicuously so. , 


` 


— AFFINITIES OF BALANOPHOREA. 29 


BALANOPHOREARUM TABULA SYNOPTICA. 


Div. I. Monosryur (Griff). Stylus 1. 
§ I. Stamina libera, Semen embryone et albumine instructum. 


A. Mystropetaleæ. 


Gen. I. MysrRoPeTALON (Harv.). Perianthium fl. & 3-partitum, 2-labiatum, segmentis valvatis, 
2 anticis connatis; fl. ? epigynum, campanulatum, 3-lobum. Stamina 3, segmentis perianthii 
opposita, iisque inserta; antheris extrorsis. Embryo hilo proximus.—Pedunculus solitarius, 
squamosus. Capitulum oblongum, biseæuale; floribus 9 inferioribus, 3 3-bracteatis. 

1. M. Polemanni, Harv.; bracteå anticå spathulatå, perianthio 9 tubuloso. 
Hab. Africå Australi. 

2. M. Thomii (Harv.); bracteå anticá latè oblongå, perianthio 9 subgloboso. 
Hab. Africá Australi. 


B. Cynomorieæ. 


Gen. II. Cynomorium (Mich.). Perianthium utriusque sexus 6-phyllum. Stamen 1, in fl. $ epi- 
gynum; filamentum in fl. 4 basi stylo deformato suffultum ; antherá introrså. Embryo 
lateralis, hilo remotus.—Pedunculus solitarius, squamosus. Capitulum cylindricum. Flores 
unisexuales, rarius bisexuales, 8 et 2 immizti; bracteis sparsis remotis. 

1. C. coccineum (Mich.). 
Hab. Regione Mediterraneá, et Insulis Fortunatis. 


C. Sarcophyteæ. 


Gen. III. SARCOPHYTE (Sparrm.). Flores dioici, 4 paniculati; perianthii lobis 3, valvatis. 
Stamina 3, antheris multilocularibus, liberis. Fl. 9 in capitulis globosis arctè cohærentes. 
Stigma discoideum, sessile.—Rhizoma simplez, lobatum. Pedunculus nudus, ramis inflores- 
centie primariis basi bracteatis. 

1. S. sanguinea (Sparrm.). 
Hab. Africå Australi. 


$ II. Stamina connata. Semen homogeneum ? 


D. Langsdorffieæ (Endl). Perianthium fl. 9 tubulosum. 

Gen. IV. Langsporrrıa (Mart). Staminum columna cava. Perianthii fl. 3 lobi 3, valvati, 
prefloratione genitalia includentes. Anthere breves.—Rhizoma horizontale, ramosum. 
Pedunculi terminales, squamis imbricatis tecti. Capitula uniseæualia. 

1. L. hypogæa (Rich.); rhizomate glaberrimo v. glabrato. 
Hab. Americá tropicá. 

2. L. rubiginosa (Wedd. MSS.) ; rhizomate tomentoso v. lanato. 
Hab. Brasiliå et Guianá. 


Gen. V. THONNINGIA (Vahl) Staminum columna solida, infra medium squamis 2-6 aucta, 
Antheræ lineares.— Habitus e£ vegetatio Langsdorfie. . 
1. T. sanguinea (Vahl). 
Hab. Africå tropicá occidentali. 


30 DR. J. D. HOOKER ON THE GENERA AND SPECIES 


E. Balanophoreæ. Perianthium fl. 9 0. 

Gen. VI. Bauanopuora (Forst.). Perianthium fl. å 3-6-phyllum. Anthere extrorsæ. Fl. 9 
pistilla bracteolis elavatis immixta v. pedicellis bractearum inserta.—Rhizoma £uberosum-v. 
ramosum. Pedunculi nudi v. squamosi. Capitula unisexualia v. bisexualia. Flores uni- 
sexuales. 

æ. Pedunculi squamis in cupulam v. involucrum connatis. 

l. B. involucrata (n. sp.). 
Hab. Himalayå temperatå. 
B. Pedunculi squamis alternis v. imbricatis; antheris 3-6 2-locularibus. | 
2. B. dioica (Brown); capitulis unisexualibus cylindricis, rhizomate tuberoso lobato pustulis 
lobulatis instructo. i , 
Hab. Sylvis montosis subtropicis Indiæ borealis, Bengaliæ et Birmæ. 
3. B. elongata (Blume) ; capitulis unisexualibus subcylindricis, rhizomate elongato ramoso ramis 
cylindricis pustulis lobulatis instructis. 
Hab. Montibus temperatis Javæ et Peninsulæ Indiæ orientalis. 
4. B. Indica (Wall.) ; capitulis unisexualibus, ? obovoideis globosisve, pedunculis elongatis, rhi- 
zomate tuberoso lobato.—(An B. globose var. ?) 
Hab. Sylvis montosis subtropicis Peninsulæ Indiæ orientalis et Ceyloniæ. | 
5. B. globosa (Jungh.); capitulis unisexualibus, 9 globosis, pedunculis brevibus, rhizomate 
tuberoso lobato.—(An B. abbreviata, Blume?) 
Hab. Sylvis temperatis Javæ. 2 
6. B.fungosa (Forst.) ; capitulis bisexualibus ovoideis subglobosisve, rhizomate tuberoso lobato _ 
granulato. 
Hab. Ora orientali Australiæ tropicæ, et Ins. Novæ Hebrides dietis. E 
7. B. alutacea (Jungh.) ; capitulis bisexualibus cylindricis, rhizomate lobato levi, pedunculi — 
Squamis paucis vaginantibus. ; 
Hab. Sylvis tropicis Javæ et Ins. Philippinis. | 
y- Pedunculi squamis alternis v. imbricatis ; antheris multilocularibus. 
‚8. B. polyandra (Griff.). 
Hab. Sylvis montosis subtropicis Himalayæ orientalis et Khasiæ. 
Div. II. Disryzr (Griff). Styli 2. : 
F. Lophophyteæ (Endl.). Stamina libera. 


Gen. VII. Lornornyrum (Schott & Endl.). Flores secus ramulos pedunculi mamillæformes — 
apice obtusos congesti, mamillis basi bracteis deciduis suffulti.Rhizoma crassum, superne — 
squamis imbricatis tectum. Pedunculus basi nudus. 


1. L. mirabile (Schott & Endl.) ; paleis cum floribus immixtis v. 0 
usculisve. 


Hab. Brasilia. 
2. L. Bolivianum (Wedd.) ; paleis inter flores 0, stylis 0 (an delapsis ?). 
Hab. Cordillerå Boliviæ. 


3. L. Weddellii (n. sp.) ; floribus dioicis, paleis inter flores 0. 
Hab. Cordillera Nove Granade. 


Gen. VIII. OMBROPHYTUM (Pæpp. & Endl.). Flores secus pedicellos bractearum peltatarum | 


congesti— Vegetatio ef habitus Lophophyti, sed squamæ nulle. Pedunculus basi volva v. 
annulo circumdatus. 


1. O, Peruvianum (Pæpp. & Endl.). 
Hab. Cordillerä Boliviæ et Peruviæ. 


3 stylis 2 elongatis brevi- 


OF BALANOPHOREÆ. __ 31 


G. Helosideæ (Endl.). Stamina connata. 

Gen. IX. Scysauıum (Schott & Endl.). Rhizoma tuberosum, lobatum. Flores pedunculis 
distinctis squamis imbricatis tectis monoici. Capitula convexa v. planiuscula. Perianthium å 
3-lobum. 

A. S. fungiforme (Schott & Endl.). 


Hab. Brasiliä. : 
Gen. X. SPHÆRORHIZON (n. g.. Rhizoma tuberosum, indivisum. Pedunculus solitarius, 


squamis deciduis tectus. Capitula sphærica v. oblonga. Perianthium 3 3-lobum. 
1. S. curvatum (n. sp.). 
Hab. Novå Granadá. 
Gen. XI. PHyLLOCORYNE (n. g.). Rhizoma lobatum v. ramosum, Pedunculi plurimi squamis 
persistentibus subhexastiché imbricatis tectum. Capitula cylindracea v. oblonga. Perian- 
_thium 3 3-lobum. 
1. P. Jamaicensis (Cynomorium Jamaicense, Sw.). 


Hab. Ins. Jamaicå. 
Gen. XII. RnoraLocnemis (Jungh.). Rhizoma tuberosum, simplex v. lobatum. Pedunculi 


pauci v. solitarii, basi annulo v. volvå instructi. Capitulum oblongo-cylindraceum, Perian- 
thium 3 tubulosum. : 

_ 1. R. phalloides (Jungh.). 

: Hab. Sylvis montosis Himalayæ temperate orientalis, Khasiæ, et ins. Javæ. 

Gen. XIII. Corynæa (n.g.). Rhizoma tuberosum, simplex v. lobatum. Pedunculi solitarii v. 
pauci, basi annulo v. volvå obscurá instructi. Capitula sphærica v. oblongo-cylindracea. 
Perianthium 3 campanulatum. 

1. C. crassa (n. sp.) ; capitulo cylindraceo. 
Hab. Montibus temperatis Novæ Granadæ. 

2. C. spherica (n. sp.) capitulo rhizomatis fossá sessili sphærico. 
Hab. Montibus temperatis Novæ Granadæ. 

3. C. Purdiei (n. sp.) ; capitulo sphærico pedunculato. t 
Hab. Cordillerá Peruvianå. : ; 
Gen. XIV. Herosıs (Rich). Rhizoma cylindraceum, ramosum. Pedunculi plurimi, nudi v. 
basi v. medio annulati. Capitula ovoidea, oblonga v. globosa. Perianthium 4 3-partitum. 
1. H. Guyanensis (Rich.); pedunculis basi volvå v. involucro 5-7-phyllo instructis medio 

nudis. > ; 


Hab. Americå tropicå. 
2. H. Mexicana (Liebm.); pedunculis medio v. supra medium annulo finas instructis. 


Hab. Americá tropicá. 


I. MxsrRoPETALON, Harv. 
(Tas. I. B.) - 


1. MysTROPETALON Tuomi, Harv. in Ann. Nat. Hist. 1839, vol. i. p 386. t. 19; Griff. 
in Linn. Soc. Trans. vol. xix. p. 336. 


_I have little to add to the excellent descriptions of Harvey and Griffith, except with 
regard to the embryo, which I have found to be constantly present in the ripe fruit; it is 
clavate, placed at the upper part of the seed, lying quite loose in its axis, with the radicle 
pointed to the hilum, and close to it. "The albumen-grains are oleaginous, large, loosely 
coherent, and enclosed in a delicate cellular membrane. I find no membrane enclosing 


32 DR. J. D. HOOKER ON THE GENERA AND SPECIES 


the embryo, which is pale, transparent, and formed of minute cohering cells, and - 
is not oleaginous. The apex of the stigma is obscurely 3-lobed: there is no very | 
distinet stigmatic canal; but the cells of the stigma and centre of the style are soft and | 
pulpy. | | | 

Griffith, who did not detect the embryo of this plant, considered that this albumen 
(which he describes doubtfully as an embryo) suggests a greater resemblance to a sporu- å 

_liferous mass than is shown by any other so-called Rhizanth : it is remarkable that this « 
observation should refer to one of the only three known plants of the Order, of which the — 
truly albuminous nature of the great mass of the seed is indisputable; and is an example 
of the inexpediency of assuming an organ to be anomalous because its structure is 
unexplained. 

The inflorescence of Mystropetalon differs from that of other Balanophoree in the male | 
flowers being at the summit of the capitulum, and the female below. In the monæcious - 
Qynomorium and most monccious Helosideæ they are promiscuously mixed, with the 

males generally lowest; in moncecious Balanophoree the males are normally at the — 

bottom of the capitulum; though I have seen traces of male flowers at the apex of å — 

capitulum of B. involucrata. 1 

Mystropetalon is certainly in all respects the most highly developed genus of the Order, » 

both from the complexity of the floral envelopes, and from the presence of a perfect embryo, 1 

placed in the usual position of that organ in Phænogamous plants. It was referred to — 

Balanophoreæ by Harvey and by Mr. Brown (Linn. Soc. Trans. xix. p. 233, in note*); and — 

though very anomalous in its greater perfection, must remain as a section of that Order, - 
or in close proximity to it. Griffith considered it as sui ordinis, but with great sagacity — 
indicates its affinity with Cynomorium (a plant he had not examined), and finally described ; 
it (doubtfully) as the homogeneous-embryo form of a group of plants including Loran- | 

thacee, Proteacee, Santalaceæ, and other Orders comprehended in Lindley’s alliance à 

Tubiferose. Considering the very great structural and morphological differences pre- - 

sented by Balanophoree, it becomes impossible to exclude Mystropetalon from the Order; 
of its affinity with which there are many positive evidences, and the curious negative one | 

of extreme dissimilarity between the perianths of the sexes. Of positive characters, 1 

the most conspicuous besides habit, are, in the male flowers, the valvate 3-partite 1 

perianth, tubular below, and enclosing a rudimentary ovarium, the stamens opposite the - 
segments of the perianth, and the extrorse anthers; in the female flowers, the epigynous 
monosepalous perianth, the deciduous style, the structure of the walls of the ovary, the : 
extreme simplicity of the ovule, and the adherent membranous coat of the seed. It - 
departs from all the rest of the Order in its male flowers being normally above the female; 
in its very unequal male perianth, its frequently imperfect odd stamen, and angular pollen; | 
in the ovarian disc, deciduous perfect female perianth, 3-lobed stigma, presence of an im- - 
perfect pistil in the male flower, and axile clavate embryo. Considering its monostylous - 
ovary, with a tubular perianth, its nearest ally would appear to be Langsdorffia, between — 
Which and. Cynomorium it will rank in a linear series. ; 


* The species there referred to does not appear to me to be specifically distinct from M. Thomii. 


OF BALANOPHOREZÆ. 33 


II. Cynomortium, Mich. 
(Tas. I. A.) 


1. CYNOMORIUM COCCINEUM, Mich.—Ad citationes evulgatas adde, Linn. Amœn. Acad. 
iv. 351. t. 9; Webb, Flor. Ins. Canar. iii. 431; Weddell in Ann. Sc. Nat. sér. 3. 
xiv. p. 176. t. 11. 


Though this curious plant has received so much illustration from many able botanists, 
there are still some points in its structure which are little known; and there are points 
in Linnæus’s description in the * Amcenitates Academicæ,* which have, I believe, escaped 
most subsequent observers. 

The geographical range is very remarkable: it extends from the Canary Islands to the 
Levant; i.e. over fully fifty degrees, or 3000 miles of longitude. I have examined speci- 
mens from its extreme eastern and western limits ; namely, from Lancerotte and from 
the delta of the Nile; and I have compared these with others from various intermediaté 
localities, as Oran, Malta, Sardinia, S. Spain, and Sicily; and I have found no traces of any 
differences that suggest the propriety of establishing even varieties. It is also found in 
Etruria, lat. 41° N. (its northern limit), Lampedusa, and Tunis according to Linnæus. 

Cynomorium is not singular amongst Balanophoreæ in this wide distribution; though 
it is more local and scarce than any of its congeners which occupy an equally extensive 
area. It is the only species known to inhabit a dry climate and soil, and is no less 
remarkable for delighting in the immediate neighbourhood of the sea, and growing in 
salinas, and often on saline plants (Linn.). Mr. Webb informs us (Hist. Ins. Canar. Bot. 
vol. iii. p. 431) that it is eaten in the island of Lancerotte. | 

In a young state the lower part of the peduncle is remarkably distinct from the upper; 
it is broader, fusiform, and covered with short, broad, acuminate, imbricating, spirally 
arranged scales (well shown in Micheli’s plate), giving it a polygonal appearance: these 
scales are much less conspicuous in the old plant, and are probably very deciduous; 
whence the discrepancy that Richard remarks between his Egyptian specimens and 
Micheli’s figure. The lower portion of this contracts suddenly at the point of union with 
the root-stock, and there is, I believe, a well-developed rhizome; but I have no complete 
specimens of it. The parasitism consists in an intimate organic adhesion between a small 
surface of the cellular tissue of the Cynomorium, and the wood of the root on which it 
grows; but no vascular tissue (at any rate in the old state) unites the latter with the 
parasite. 

The vascular system (represented by Unger, Ann. Wien. Mus. ii. t. 5. f. 32) consists 
of many bundles irregularly scattered through the peduncle and capitulum; form- 
ing waving lines, but never crossing or resembling the endogenous type in structure or 
arrangement. These, in a transverse section, are seen to consist of two kinds of vessels ; 
namely, 1. internally of a small bundle of delicate cylindrical or angular white tubes with 
transverse marks or bands; 2. a broader dark external tissue which in a transverse 
section appears cellular, and in a longitudinal one is found to consist of many series of 
` linear, superimposed, oblong cells, regularly placed, and all terminating at the same plane, 
thus giving a barred appearance to the tissue ; the contents of these are all highly coloured. 

VOL. XXII. d 


34 DR. J. D. HOOKER ON THE GENERA AND SPECIES 


Besides these there are thick-walled pleurenchyma-cells. The cellular system, which M 
forms by far the greater mass of the plant, consists of large polygonal utricles, with thin — 
transparent walls, full of starch and chlorophyll granules. 1 

The scales upon the upper part of the peduncle are broader than those at the base; blunt, ` 
often transversely oblong, peltately attached, and, on the capitulum, gradually assume the 
character of peltate bracts. In the youngest specimens I have examined (Tab. I. À. fig. 1) | 
long before the flowers are discernible, except as lobed papillæ, the appearance of all the M 
scales from the base to the top of the plant is remarkably uniform ; they are broadly ovate, — 
acuminate, and imbricated at the fusiform swollen base, more scattered and broader, and 
blunter or truncate, on the upper contracted portion of the peduncle, and again acute 
and densely imbricated from the base to the summit of the capitulum, the surface of 
which they wholly conceal. At this early period the scales are very obliquely peltate; 
in a vertical section each is seen to curve upwards and cover the lower part of the peltate 
scale immediately above it, whilst the lower part is produced into a long incurved lobe. 
The lower lobe presents a semi-lunar curve towards the capitulum, and arches over a 
mamilla of the capitulum covered with nascent flowers. The concave upper surface of 
the next scale below is closely applied to the dorsum of the lower lobe of that above it. 
A strong vascular cord enters each scale, and is united at a little distance from its base, 
within the body of the capitulum, with an equally stout cord from the mamilla above it. - 
It will thus be seen that each scale forms the protecting organ to a definite mass of . 
flowers below its point of insertion, but is connected by its vascular system with the 
mamilla of flowers above it ; an arrangement similar to that which occurs in Lophophytum 
and some Lycopodiacee, but which can in Cynomoriwn only be discovered at a very early 
period. : 

Owing to the much more rapid growth of the capitulum than of the scales, these even- 
tually become scattered, at the same time losing their bracteal form, and becoming 
broader and fleshy. These changes are precisely the same in their nature as occur in the 
bracteal scales of Helosis and its allies. The paleæ which occur abundantly amongst the 
flowers, and vary extremely in form, consist of rudimentary flowers, both males and 
females, and of perigonial leaves, removed from their flowers by unequal growth. There 
is a disposition in some of the floral scales of larger size than the rest, to assume the 
position of a bractlet under each flower, or group of flowers. 

The male and female flowers appear promiscuously in succession for a considerable 
period: in this respect Cynomoriwm presents a remarkable contrast to Balanophora and 
Helosis, &c.; as in these the evolution of the sexes occurs at different times: and it is 
worthy of remark that this phænomenon is perhaps confined to this genus, and is there- - 
fore peculiar to the only plant of the Order which exhibits a strong tendency to herm- 
aphroditism. 

In Webb's ‘Flora of the Canary Islands’ (iii. 431), I have described hermaphrodite 
flowers of Cynomoriwm; which were pointed out to me by Mr. Brown in a drawing of 
Bauer's: they were originally discovered by Linnæus (Gen. Plant. ed. 5, 1754), and 
described by him; as indicated by Richard, who, however, failed to find them himself. - 

- The palea of Richard, surrounding and half enveloping the stamen of the male flower, 


OF BALANOPHOREÆ. 35 


is å remarkable organ : from analogy with Balanophora, Helosis, and indeed the majority 
of the Order, it might be considered an imperfect perianth ; whereas in the hermaphrodite 
flowers it is seen to be a well-developed style, the ovary of which is usually suppressed. In 
these flowers the stamen rises from near the summit of the ovary, above the insertion of 
any of the perigonial scales, and the base of the filament is lodged in a narrow vertical 
canal immediately opposite the concave face of the style (figs. 3, 4 & 5). I have not 
been able to ascertain the position of the style and stamen with reference to the axis of 
the plant; but, considering the simple nature of the pistil, and that the flowers are 
collected in groups, representing theoretically ramifications of the infloreseence, I assume 
that the concave face of the style is opposite to the ideal axis of each such ramification. 

* In the youngest state of the male (fig. 6), the floral envelopes are often symmetrically 
disposed. A spathuläte palea subtends each flower, and within it are six perigonial scales ; 
these are frequently perfectly regular, and form a verticil round the rudimentary ovarium 
and filament: the whole flower afterwards grows with great rapidity, and some of the 
perigonial scales are left at its base, while others are carried up on the elongating imper- 
fect ovarium, which resembles a pedicel, and branching bundles of vessels are developed 
in their axes. The style at the same time elongates rapidly, but increases also in breadth 
upwards, so as to resemble a perigonial scale much more than the style of the female 
flower does; it has however no vascular tissue. In some male flowers all the perigonial 

Scales remain symmetrically disposed round the filament and style, till the flower has 
attained its full development. 

At the earliest period the female flowers (fig. 2) appear as compressed, pedicelled bodies, 
subtended by a bract similar to that of the males. The pedicel dilates, and divides into 
two (rarely three) linear perigonial leaves, and again contracts, forming the base of a com- 
pressed obovate ovary, on whose summit is the broad linear style, somewhat contracted 
immediately below the papillose stigma; and at its base are three other linear blunt peri- 
gonial leaves of equal size, which are always symmetrically disposed, two being placed 
right and left in the same plane as the style and the two lower perigonial leaves, and one 
at the dorsum of the style. Two vascular cords are seen, one on each side of the ovarium, 
meeting at its base in the pedicel, and again at its summit immediately at the base of the 
style, below which an opaque circular spot marks the cavity of the ovary; these vascular 
cords again separate in the style, and are free to its apex. The base of the style is con- 
tracted, and of a very dark colour at the contraction: I am not aware what this indicates. 
The opacity is also seen at the base of the filament in hermaphrodite flowers, and often 
upon all the perigonial scales, at a point exactly corresponding to the base of the style. 

During the growth of the female flowers, the perigonial scales do not attain the deve- 
lopment they do in the males, nor do they generally contain vascular bundles. They 
become variously displaced; some remaining on the summit of the ovary, and others 
being more or less basal, or adnate to its surface. In the adult female flower a small 

- cellular protuberance may often be seen on the summit of the ovary (fig. 7), fronting the 
concave face of the style: this may be a rudimentary stamen. = s 

The hermaphrodite flowers present both the pistil and stamen in as perfect a condition 


as they attain in the unisexual flowers. The situation of the perigonial scales varies as 
F2 


36 DR. J. D. HOOKER ON THE GENERA AND SPECIES 


much in these as in the female flowers, but though they never attain the same develop- ; 
ment as they do in the males, they approach it, and often contain vascular cords. In. 
one hermaphrodite flower (fig. 9) I found six perigonial scales, symmetrically disposed 
round the summit of the ovary. 4 
There is thus in a complete flower of Oynomorium a superior perigonium, an epigynous . 
stamen, a pistil consisting of one carpel with one simple style, and one pendulous ovule, - 
succeeded by an albuminous seed :—characters common also to Hippuris, and indicating | 
an affinity I have elsewhere noticed, and endeavoured to support by the structure of other 1 
Balanophoree. y ; 1 
I may here repeat what I have stated in the * Flora of the Canary Islands' concerning 
the pistil of Cynomorium; namely, that the simple concave style with two parallel vas- 
cular cords terminating an ovary which undoubtedly consists of one carpellary leaf, is à. 
strong evidence of the compound nature of a style of the simplest type; and that the two 1 
lateral stigmata are here perfectly obvious: which agrees with Mr. Brown's remarks on 
the composition, &c. of the pistil* (Plant. Jav. Rar. p. 110 in note). The stigmatic tissue - 
runs down the mesial line of the style, occupying the canal, and is covered by a very - 
delicate epidermis. 4 
I have never succeeded in tracing the development of the ovule in Oynomorium. The 
structure of the ripe seed has been determined by Richard, Lindley, and Weddell; but | 
admits of some little further illustration. In the first place, the embryo when fully ripe 
is considerably larger than is figured by any of these authors, and is never exactl 
globular, but sometimes broadly conical, the narrow end being placed next to the firm 
cellular integument of the seed. It consists of large cells with dark nuclei, full of o 
and presents no integuments whatever: it lies in a cavity of the albumen, nearer the base 


albumen as mucilaginous, and the cells 
either microscopically, or by the iodine test, whi en applied to the most delicat 
slices of fully formed albumen, turned its granular cell-contents brown. Mr. Weddell 
states that the albumen as well as the embryo contains oil (Ann. Se. Nat. 7. c. 178) 


“dans les Balanophora, dans le Oynomorium, et dans le Sarcophyte, j'ai rencontré un 
albumen charneux ou huileux, et un embryon de méme nature.” — 


Å On showing Mr. Brown my analysis of the style of Cynomorium, he informed me that this was a case he had had 
m view, and that he considered it strongly confirmatory of his theory. 


f 


t^ 


OF BALANOPHOREÆ. 37 


III. SARCOPHYTE, Sparrmann. 
(Tas. I. C.) 


1. SARCOPHYTE SANGUINEA, Sparrm. in Act. Holm. xxxvii. p. 300. t. 7; Schott & Endl. 
Melet. Bot. p.11; Griffith, Linn. Soc. Trans. xix. p. 339; Unger, Ann. Wien. Mus. 
ii. t. 5. f. 28; Weddell in Ann. Sc. Nat. ser. 3. p. 14. t. 10. f. 34-38. 


Ichthyosma Wehdemanni, Schlecht. in Linnæa, ii. 671. t. 8. 
Hab. Africå Australi, ad radices Ekebergie prope Grahams-town (Wehdemann, &c.). Ad radices Acaciæ 
Capensis, Quagga's Flat, Uitenhage (Zeyher /). 

My observations on this remarkable plant chiefly refer to two points: the structure of 
the anther, and the relation of the genus to other Balanophoreæ, in both which I differ 
from Mr. Griffith. 

The anther is rightly described by Endlicher as consisting of a solid capitate body, 
containing many loculi filled with pollen. The contracted persistent septa between these 
loculi have been mistaken by Griffith for pedicelled anthers, which he describes as forming 
_ together a “ caput antherarum,” crowning a common peduncle, which rises from the axil 
ofa bract. On the contrary, I find that the anther* contains about fifteen to twenty cells 
of very variable size, radiating from a cellular axis: a transverse section shows about twelve 
such cells symmetrically disposed round the apex of the filament; and a vertical one 
exhibits about eight, which radiate from the blunt summit of the filament, and of which 
the outermost are very small. At a very early period the septa consist of three tissues : 
an inner cellular not materially different from that of the filament, from which, however, 
it is separated by a broad dark line; this is lined by a delicate hyaline endothecial coat, 
and upon this is a mass of matted filaments of excessive tenuity and pulpy nature pre- 
cisely similar to the anther-lining of PAyllocoryne, &c., amongst which the pollen-grains 
nestle. Each cell is distended with spherical pollen-grains. 

I have not observed the circumscissal dehiscence of the outer membrane described by 
Endlicher, and which was apparently suggested to that author by the appearance of an 
annulus at the base of the dehisced anther; as on the contrary the anther dehisces by 
the disruption of the membrane over each loculus, in a manner quite analogous to that of 
the apices of the anthers of Rhopalocnemis, Helosis, and Corynæa, on the one hand, and 
of Balanophora polyandra, &c., on the other; and is an instance of the general tendency 
to dehiscence by irregular disruption of the anther-wall, which prevails throughout the 
Order. I have not been able at any period of its growth to reduce the anther to the 
ordinary 4-locular type; the pollen being developed, as in Viscum and various Rhizo- 
phoræ, simultaneously in many independent points of the epithelium: that these points 
originated along definite lines, answering to the position of anther-cells of the ordinary 
type, can therefore only be assumed. 

From the above it is evident that Griffith is perfectly correct in insisting (J. c. p. 339) 
that a continuous solid tissue must exist between the cells of the anther, if it be assumed 
that these cells are not separate anthers. 

In assuming that the filaments of Sarcophyte are axillary to the lobes of the perianth 


* A correct section of the anther is given in Unger's paper; Ann. Wien. Mus. l c. t. 7. fig. 48. 


38 . DR. J. D. HOOKER ON THE GENERA AND SPECIES 


(which he hence considers bracts), rather than bodies forming a verticillus on an inner ] 
and different plane, Griffith overlooks the fact, that their position in no way differs from . 
that of the stamen of other Balanophoree, and that all stamens opposite to and seated at , 
the base of perigonial leaves, are in the same category. On the other hand, his argument | 
against the stamens being axillary, because they do not appear to form an inner whorl, 
may be equally applied against considering the perigonial leaves as being bracts, for the M 
latter decidedly do form an outer whorl, and are all on one plane; a fact which, as well | 
as that of their decidedly valvate æstivation, is opposed to their bracteal origin. E 
Another remark of Griffith's is to the effect, that * the analogies of Balanophora are in M 
favour of Endlicher's generie character; but that it requires a very exalted idea to be 1 
held of the value of parasitism, to conceive any affinity between Sarcophyte and Balano- 3 
phora” (p. 340). If, however, the homologies in the structure of the flowers are ad: - 
mitted, it cannot be said that systematists have depended on an undue value attached to — 
parasitism, for the supposed affinity; and in the second place, the argument derived from 
parasitism, if of any value, does not rest upon the mere fact of parasitism, but on that of — 
the root appearing to send vascular bundles into the rhizome of Sarcophyte as it does 
into that of other Balanophoree, a kind of parasitism not hitherto detected in any other — 
Natural Order*. ; E 
The male flower of Sarcophyte differs in no essential particulars from that of Balano- : 
phora; the pedicel (tube of perianth) and three valvate perigonial leaves being identical, | 
and both having the stamens opposite to the latter. The chief difference is, that in Bala- 3 

à; nophora the stamens are united by their filaments and connectives, whilst in Sarcophyte 4 
they are free. The suspected analogy between the structure of the stamen of Sarcophyte — 
and the sorus of Cyathea and Spheropteris, suggested by Griffith, is under any view quite 3 
untenable. 4 
Griffith’s description of the ovarium and its contents does not accord with my obser- 4 
vations; nor could I suggest any explanation of his “brown central nuclei, containing — 
one, or not unfrequently two, other brown nuclei ;” but Weddell points out that Griffith — 
examined an abnormal state of the fruit, which he has frequently observed himself, 
and in which the embryo is abortive, and the albumen and integuments become con- 
founded into an ossified mass. I find, in the ovarian cavity of specimens preserved in 
acid, an immature ovulum, consisting of loose white cells, enclosed in a delicate mem- 
brane as in Balanophora. This albumen and its erustaceous coat (formed of the peri- 
carp) are well illustrated by Weddell (Ann. Se. Nat. l. c), as is the central embryo, - 
discovered by himself. i : j 
Griffith goes too far in stating that the female flowers of Sarcophyte are widely different 
from those of Balanophoree, in their greater general perfection, the union of the ovaria, 
and the obvious stigmatic surfaces : for, in the structure of the female flower, and of the — 
Seed, except in the development of the embryo, they are identical : the more highly organized 
stigmata attain a greater degree of perfection in Sarcophyte than in those genera with which 


* Except Orobanche, 
1854, p. 577, t. 3). 
of Orobanche, 


the germination of which has been so admirably illustrated by Caspary (Regensburg Flora, 1 
It appears most probable that the germination of Balanophoreæ will prove very similar to that I 


OF BALANOPHOREÆ. 39 


Griffith was best acquainted, but not so great as in Cynomorium and others; whilst the 
greater general perfection in other respects, to which he alludes only, I do not appreciate. 
The union of the ovaria does not indicate a difference in the female flowers, but a different 
inflorescence ; a point of much less systematic value, and indeed of none in Balanophoreæ, 
as is proved by a comparison of Balanophora, Cynomorium, Lophophytum, and their allies. 
In this point the analogy is perhaps complete with Thonningia and Langsdorffia, whose 
ovaria are entirely united; as are the perianths of Langsdorffia, in some states at any rate. 
Ås an indication, however, of the female flowers of Sarcophyte being furnished with a 
perianth, this point is of considerable importance. 

Another fact connected with the inflorescence of Sarcophyte is the sudden suppression 
of bracteal scales; these, which are very conspicuous at the base of the primary branches 
of the male and female inflorescence, are not developed at the base of the capitula, or of 
the individual male flowers. This appears, further, opposed to Griffiths supposition that 
the perigonial leaves of the male flower are bracts, though perhaps not conclusive 
against it. < 

The tissues of the stem of Sarcophyte are full of starch-granules, but do not differ 
otherwise from others of the Order, though in the peduncle the vascular bundles are very 
irregularly deposited. The roots of the plant upon which it grows are connected by stout 
woody branches with the rhizome of the parasite; and there seems to be a complete 
fusion of the vascular tissues of both. 


IV. LANGSDORFFLA, Rich. 
(Tas. II.) 


1. LANGSDORFFIA HYPOGÆA, Martius in Eschwege’s Journal von Brasilien, ii. p. 179; Nov. 
Gen. et Sp. iii. 181. t. 199; Unger in Ann. Wien. Mus. ii. t. 4. figs. 21 & 22, & — 


t. 7. fig. 40. 


Langsdorffia Janeirensis, Rich. M&m. Mus. viii. p. 412. t. 19; Endl. & Schott. Meletem. p. 12. 
Thonningia Mexicana, Liebmann in Proceedings of Assembly of Scandinavian Naturalists, 1844. 


Sendfenbergia Moritziana, Klotzsch & Karsten, Herb. Mus. Berol. i 
` Hab. Americå tropicå, a Mexico ad Brasiliam meridionalem. Sylvis montis Serra d’Estrella (Martius) ; 


Rio de Janeiro (Miers, Gardner, Stephan). Sylvis montosis provinciae Oaxaca, Mexico (Liebmann) ; 
f. Nov. Dec. Ad radices arborum ad Tucouroma, provincie Ocanæ (W. Purdie); Colombia 
(Karsten). 


After a very careful examination of specimens from all the localities quoted above, 
* except the male and fruiting specimens of Martius and Prof. Liebmann, I have referred 
all to one species. As however this is a very important point, involving the question of 
the range of the species extending throughout the tropics of both Americas, I feel that it 
is necessary to dwell at length upon it. From Mr. Purdie I have large suites of speci- 
mens, which show that the capitula on the same rhizome vary quite as much as those of 
Mexican specimens do from Brazilian ones. Prof. Liebmann, in his résumé of the cha- 
racters distinguishing the Mexican plant, seems chiefly to rely upon a comparison of it 
with the drawings and descriptions of that from Brazil; but I find that none of my 
Brazilian specimens agree in all their details with Martius' and Richard's drawings; nor 


40 DR. J. D. HOOKER ON THE GENERA AND SPECIES 


do these quite accord with one another, but only in what I assume to be mere individual, 
and not specific characters. 

Prof. Liebmann sums up the differences between the Mexican and the Brazilian indivi 
duals, as residing in the more globose female capitulum, shorter stem, more shortly pedi: | 
celled perianth, twisted style, binate paleæ of the male receptacle, which are also clavate 3 
and dilated at the base, white papillose (male) perianth, longer filament, globose synema - 
and globose pollen. It is also added, that the anthers are 2-celled, and dehisce differently, 
leaving a triangular opening between them, and that the filaments are free immediatel 
below the anthers. | 

With regard to these points, I find the capitulum if anything more depressed in the 
Mexican plant and in Liebmann’s accurate figure of it, than in Richard’s drawing of the 
Brazilian, or than in most of my specimens either of the Brazilian or Colombian plant. 
The stem (peduncle) varies extremely in length, from 4 an inch to 8 inches, and con- 
siderably even on the same rhizome. The perianths of Mexican specimens are much 
longer than those of Liebmann's figure, and they are of the same length as those of my 
Brazilian specimens, though shorter than in Richard’s or Martius’ figures. The styles of 
the Mexican plant are very slightly twisted, and that from left to right, not the opposite 
way, as represented in Prof. Liebmann's figure; and there is the same twist in Mr. Purdie’s 
and in some of the Brazilian specimens. The perianths seem constantly papillose, though 
varying in degree with age, drying, and other less obvious causes, Globose pollen is the 


so extremely short, that it appears impossible to draw a character from it; the synema. 
varies in form, according to its age, and that represented in the figure of 7. Mexicana 
. entirely agrees with Brazilian individuals; and finally, the anthers of all, though 4-celled 

in their early and perfect state, become 2-celled previous to dehiscence, by the contraction 
of the septum. I therefore feel justified in referring the Thonningia Mexicana to 
Langsdorffia hypogæa. 

. The parasitism of Langsdorffia is remarkable: the dichotomously branching rhizom 
appear most frequently to corrode, as it were, the bark of the roots they encounter, which 
they even sever, and then enclose the end that remains attached to the parent plant: the 
root swells considerably at the junction, and appears to send prolongations of wood into 
the rhizome of the parasite, which run along its axis for several inches; but though 
there is an intimate union between the wood of the root and the cellular tissue of the 
parasite, there seems to be no blending of their vascular systems. The rhizome also 
invariably swells at the junction, but does not branch from that point, as is often the case 
with Helosis. Both Richard and Martius represent rootlets as given off from the rhizome 
at a considerable distance from any parasitic union; but I do not find such in any of my 
Ru — have any other Balanophoreæ rootlets, though at the junction of root and 
parasite similar rootlets to those figured by Martius are often given off by the root, and 
these being partially enveloped by the parasite, appear to proceed from it. Martius an 
Langsdorff further say that the plant grasps other roots by means of these fibres, and 


OF BALANOPHOREÆ. 41 


that it does not appear to be really parasitical; but in both these points I think these 
authors are mistaken. 

In Purdie's specimens of L. hypogea, which are immature, the female perianths 
adhere firmly, so that their tissues are torn in*sundering them. I have not found this to 
be the case in the Brazilian or Mexican specimens. The style varies exceedingly in 
length, as does the perianth of the female flower and its pedicel; the mouth of the 
perianth is obscurely lobed, and a few slender, straight, rigid, woody tubes traverse it 
longitudinally, as in Thonningia. 

In common with Richard and Von Martius, I have sought in vain for any traces of the 
cavity of the ovarium. For a knowledge of the fruit I am indebted to Prof. Liebmann’s 
invaluable communications, and to specimens given me by M. Weddell. The two 
former authors indeed hazard the supposition that all the specimens they examined were 
of a barren state of the species; to which, if such were the case, would have to be added 
upwards of fifty specimens examined by myself. Considering the extraordinary minute- 
ness of the seed, it seems safer to conclude that the cavity of the ovary being still smaller, 
sunk in the substance of the fleshy receptacle, and probably filled with the ovule, which 
adheres to the cavity, it has escaped observation*. The fruit-bearing receptacle is quite 
similar to that of Thonningia; it dilates greatly after flowering, causing the surrounding 
scales to spread horizontally; its surface is covered with the persistent fleshy conical 
perigonia, which adhere so closely that they may be removed in a body as a fleshy covering 
to the receptacle. Beneath each flower is a minute oblong seed, nidulating in the fleshy 
receptacle, and pendulous from the apex of a unilocular crustaceous putamen. The seed - 
is compressed, oblong, covered with a membranous coat, and has an evident raphe down 
one of the edges. The specimens I have examined are immature and had no discernible 
embryo, as were probably those examined by Liebmann, who describes the contents of the 
seed as a pulpy mass of globular cells. The clavate scales of the male capitulum appear 
to me to be undeveloped female flowers; and the small, hard, prominent, imbricating 
scales which surround the base of the female capitulum, are connate articulate filaments, 
analogous to those of Helosidee. ; 

This species yields so large a quantity of wax, that candles are made of it in New 
Grenada. The secretion is contained entirely in the cellular tissue, where it appears as a 
large opaque mass in every utricle. Mr. Purdie informs me that near Bogota the stems 
are collected, and sold in the markets under the name of Siejos, and used as candles on - 


saints’ days. 


* Since writing the above, I have examined some excellent flowering specimens of L. hypogea, which, though 
originally preserved in spirits and afterwards transferred to acid, have not turned brown; and in these I find unim- 
pregnated ovules. The perianths in these specimens adhere firmly throughout the upper two-thirds of their length, 
but their cylindrical bases, though densely packed, are quite free. Their substance is very loosely cellular and 
diaphanous, and a dark spot immediately above the insertion of the flower on to the fleshy capitulum, marks the 
position of the excessively minute ovarian cayity and ovule. Owing to the extreme minuteness and laxity of the 
cellular tissue of the ovary, I had great difficulty in opening it and dissecting out the ovule, which forms a pen- 
dulous globular transparent sac, consisting of a few loosely packed nucleated cells, enclosed in a membrane of ex- 
cessive tenuity. This ovule is the most minute that I have met with in the vegetable kingdom. I further found 
pollen-tubes in the style of some flowers, traversing a conducting tissue formed of long, soft, lax, tubular cells in the 
axis of the style.—Kew, March 4, 1856. 


VOL. XXII. G 


42 ' DR. J. D. HOOKER ON THE GENERA AND SPECIES 


, 


On the Tolima range it is called “ Belacha;” and * Melousita” on the mountains 
around Bogota, where its soft receptacle is eaten when ripe, and considered stimulating 
and refreshing. It is remarkable that Langsdorffia, the only monostylous American 
genus, should resemble Balanophora in the abundance of its waxy secretion; whilst 
Rhopalocnemis, the only distylous Asiatic species, resembles the other American Helo- . 
sideæ in the absence of wax. 


2. LANGSDORFFIA RUBIGINOSA, Weddell (Ann. Sc. Nat. ser. 3. xiv. t. 11. f. 48-51) 
rhizomate densè tomentoso. (Taz. II. figs. 1-19.) 
Hab. Sylvis montosis regionibus superioribus fluminis Orinoco, alt. 3500 ad 4000 ped. (Schomburg, 


Ic. Pict.): locis umbrosis humidis inter Quebradas de San Juan et Panones, Parana de Ruiz (Purdie) 
provincia Goyaz Brasiliz (Weddell). 


This species does not seem to have been distinguished from L. hypogea by Mr. Purdie, 
- from whom I first received it, and who mentions its also being used for candles. It in every 
respect resembles L. hypogea, except in the dense, matted, woolly vesture of the rhizome, 
which appears as if wrapped in wool. This indumentum is formed of long, simple, very 
sparingly articulated transparent hairs, with broad bases and blunt apices, and walls 
covered, especially towards the base, with minute granulations ; they are quite hollow, and 
very flaccid. All my specimens are males, and have very short peduncles, clothed wi 
rather narrower scales than in L. hypogea. The small clavate bodies situated on the male 
capitulum, and placed at the angle where four flowers meet, are often connate: they are 
quite analogous to the similarly placed organs in dicecious Balanophore. Wedd 
figures the female perianths as connate throughout their length, as is sometimes the ca 


with L. hypogea; his specimens appear to be immature, and I doubt its proving ev 
tually distinct from the last-mentioned species. 


V. THONNINGIA, Vahl. 
(Tas. III.) 
Conophyta, Schum. Hæmatostrobus, Endl. MSS. (Gen. p. 76). 


FL. 3. Perianthium incompletum, e squamis 2-3 subulatis inæqualibus infra medium column 
antheriferæ insertis. Anthere 3-5, longissimè lineares, in columnam conico-fusiformem elong 
solidam coadunatæ, 12-20 ?-loculares, loculis linearibus, extrorsis. FL. 9 lineares. Periant 
superum, tubulosum, inæqualiter 3-5-dentatum. Ovarium lineare, perianthio zequilongum imá 
ovuliferum. Stylus cylindricus, filiformis, perianthio dupld longior, supra medium papillosus s 
matiferus, Ovulum 1, pendulum. Achenia receptaculo spongioso immersa.—Rhizoma r 
ramosum. Pedunculi erecti, 1-6 unciales, basi nudi, squamis coccineis circa inflorescentiam elon, 
tecti, inter squamas pubescentes. Squamæ dens? imbricate, late ovato-acuminate, inferiores 4 
superiores 3-1 unc. longe. Receptaculum spongiosum lat? conicum v. hemisphericum, convexum, 
bus obtectum squamisque omnind velatum. Fila articulata nulla. Flores rubri. 


Dioica. 


1. THONNINGIA SANGUINEA, Vahl! (Act. Soc. Hist. Nat. Hafn. vi. p. 124. t. 6, 181 
Schumacher et Thonning, Dansk. Selsk. Skrivt. vi. p. 124. t.6. Liebmann in P 
ceedings of Assembly of Scandinavian Naturalists at Christiania, 1844, p. 177. 
Conophyta purpurea, Isert, Reise nach Guinea, p. 283. 
Hab. Guinea ad Aquapim, Thonning. Abeokuta, Beat. Dom. Irving (fid. Ic. Pict.). 


OF BALANOPHOREÆ. 43 


I am indebted to Prof. Liebmann of Copenhagen for the opportunity of deseribing this 
rare plant; he having forwarded to me the original specimens from Vahl’s and Schuma- 
cher’s herbaria, which are deposited in the museum at Copenhagen. In his paper read 
before the Association of Scandinavian Naturalists at Christiania in 1844, M. Liebmann 
discusses the propriety of restoring Vahl’s name of Thonningia to the American Langs- 
dorffiæ of Martius and others, under the impression that they are all congeneric. As his 
information is very curious, and as I am obliged to dissent from his conclusions, I shall 
give the substance of his communication here, the paper being little likely to become 
generally accessible in England. 

Thonningia was brought to Europe by Thonning in 1804, and described, and named 
after its discoverer by Vahl in the same year, in a paper read before the Natural History 
Society of Copenhagen, and accompanied by a plate. Whether the paper was printed 
does not appear; but Vahl died in 1804, and the Society was dissolved immediately 
afterwards. The volume, of which Vahl’s paper formed a part, was not completed till 
. 1810, when a few copies were distributed, and the rest retained by Prof. Viborg, on account 
of an obnoxious preface by M. Ratke, detailing a controversy between Professors Viborg . 
and Vahl, and which was suppressed on the ultimate publication of the volume in 1818. 

During the same year (1818) Von Martius published the Brazilian Langsdorffia in 
Eschwege’s Journal; and the question brought forward by Liebmann is: supposing it 
to be congeneric with Thonningia, which name should be retained? Prof. Liebmann 
advocates Vahl’s, on the ground of priority, and because his plate enables the genus to be 
identified, though he considers his description to be faulty*. As far as priority of 
publication is concerned, the claims of the names are on a par; but it appears to me 
impossible to include the Brazilian plants in the same genus with the African, on account 
of the great differences between their male flowers. 

My description of Thonningia is drawn up from Vahl’s and Schumacher’s specimens 
and drawings. The male flower consists of a very long spindle-shaped synema, curved at 
the base, broadest in the middle, and tapering to a sharp point: a little below the middle 
it bears two or three subulate narrow fleshy scales, which are the rudiments of a perianth 
that is never further developed. The upper half appears from Vahl’s drawing to be 
covered with pollen; and according to his specimens this is perfectly correct, and further 
agrees with Schumacher’s description. In Schumacher’s specimens I find no traces of 
anthers or pollen. In Vahl’s specimens, however, I find four or five vascular bundles, 
and as many very long linear connate anthers, each 4-valved, bursting Jongitndipally, and 
containing globose hyaline pollen-grains with transparent borders. - 

The female flower of Thonningia only « differs from that of Langsdorffia i in a more complete 
tubular 3-5-toothed perianth. The parenchyma of-this organ is much inflated, and is 

formed of very lax cellular tissue, traversed by four to six remarkable nerves. These consist 


* Prof. Liebmann says that Vahl must have been in error in describing both male and female flowers, as from the 
plate accompanying his paper it appears that he had only female specimens : but Vahl is here right, for he certainly 
figures both the male plant and its flowers, t. 6. figs. a, 5, c, d; and though not very intelligible, they are accurate, 
and accord perfectly with the description,of Schumacher and Thonning, whose specimens Vahl examined, and which I 


have also examined and described here. 
62 


44 DR. J. D. HOOKER ON THE GENERA AND SPECIES 


of one or two rigid, stout, cylindrical, yellow tubes of sclerogen, with blunt apices; their | 
walls are transparent, but extremely thick, and they are sometimes solid in places; they , 
entirely resemble the woody tissue commonly developed in other parts of Langsdorffia | 
and Helosis. The style is very long, filiform, and continuously papillose along the 
exserted portion. I have not seen the seeds, which occupy a very minute cavity in the 
base of the columnar ovary, and are said to be sunk in the receptacle. The latter expands 
considerably after flowering, when the scales fall away from the flowering branch, and the 
latter turns black, and probably decays. | 

I know nothing of the parasitism of Thonmingia; the rhizome is brown, slender, — 
smooth, and sparingly branched, and rises into an obscure cup round the base of the 
peduncle, which is clothed with bright red scales. I find no hairs upon the rhizome, as - 
is the case with Langsdorffia, but there are small woolly tufts at the bases of the leaves, 4 
on the stem. The hairs are simple, long, inarticulate, flexuose, broad at the very base, | 
rough on the surface, and with a very large continuous cavity. 


VI. BALANOPHORA, Forst. 
1. BALANOPHORA INVOLUCRATA (supra, p. 30). 


- Var. a. rubra, pedunculis et capitulis rubris, capitulis ovoideis bisexualibus (Tas. IV., V. & VL). 

Var. B. flava, pedunculis et capitulis stramineis v. flavis, capitulis unisexualibus rariüs bisexualibus. 
Var. y. gracilis, pedunculis elongatis gracilibus capitulisque flavis, capitulis unisexualibus parvis 

- (Tas. VII. A.). 

Var. 8. Cathcartii, pedunculis robustis capitulisque albis roseisve, capitulis unisexualibus (Tas. VII. B.). 
Hab. In Himalayz temperatæ sylvis humidis ; Sikkim, alt. 7-9000 ped. (J. D. H.) Simla, alt. 6000 ped. 
- (Thomson) (fl. Jul.). ; ; 
Rhizoma 2-6 unc. latum, pustulis parvis cellulosis asperum, variè lobatum, nodos 3-4 poll. diam. radicibus 
Aceris et Quercds efficiens. Pedunculi graciles v. crassi, breves v. elongati, medio involucrati, interdüm — 
compressi v. fasciati. Capitulum ovoideum v. globosum, rariùs depresso-globosum; ¢ profunde 
alveolatum. Flores å 2-5-meri, plerumque 3-meri. Anthere tot quot lobi perianthii synemate brevi 


sessiles, transversè oblongæ, supernè rimå transverså dehiscentes, FZ. 9 capitulo sessiles v. circa 
basin bracteolæ clavatæ aggregati. 


The extreme varieties which I have here included under one species are so very dis- 
"similar, that no one who had not seen large suites of specimens, presenting every inter- 
mediate form between them, could venture to unite them under one: as it is, I found that - 
neither colour, form, nor the sexuality ofthe capitula are constant characters. In the same 
woods wherein I gathered the var. gracilis growing upon roots of oak, I also gathered var. 
flava growing on those of an Araliaceous shrub, and differing from the var. gracilis only 
In its more robust habit. In general there is a greater tendency in the female capitula | 
to bear male flowers than in the males to produce female; for though I often met with 


female capitula bearing male flowers at their base 


» and sometimes at their summit, and 
occupying a considerable portion of the surface, I never found male capitula to bear any - 


but very rudimentary female flowers scattered along the edges of the alveoli in which the - 
lower part of the male perianth is sunk. 


. The present is the most alpine species of the genus known to me, and is common in 


Sikkim at 8000 to 10,000 feet elevation. I have found it on the exposed aërial rootlets of | 


OF BALANOPHOREÆ. 45 


oaks in very humid forests, but, like the rest of the species, it generally grows at the foot 
of the trees immersed in the spongy soil. It causes large knots 2-4 inches in diameter to 
form on the roots of oaks and maples, and these are much sought by the natives for the 
manufacture of the wooden cups in general use throughout the Himalaya and Tibet. 


2. BALANOPHORA DIOICA, Brown in Wall. Cat. 7246; Linn. Trans. xiii. 207 in note. 
Royle, Ill. Plant. Himal. p. 330. t. 99 or 78 a. Schott & Endl. Melét. p. 13 (sub 
B. elongata, BL). 

B. typhina, Wall. Cat. 7248. ; 

B. picta, B. alveolata, B. Burmanica et B. affinis, Griff. in Linn. Trans. xx. pp. 94, 95. t. 3, 4, 5 & 6. 

Variat insigniter rhizomate plus minüsve lobato v. ramoso, lobis crebré v. laxe pustulatis, pustulis simpli- 
cibus stellatim lobulatisve, pedunculis brevibus elongatisve, flavis albis rubrisve, squamis arctè v. 
laxè imbricatis, capitulis omninà unisexualibus v. fæmineis basi androgynis, cylindricis ovoideis 
conoideisve, columnå stamineå brevi v. elongatä, antheris 3-5 arcte v. lax& compactis, floribus 9 ee 
v. longè pedicellatis, capitulo v. pedicello bracteolæ clavatæ insertis. 

Hab. In Himalayæ orientalis, centralis et mont. Khasiæ sylvis subtropicis vulgatissima, alt. 3-7000 ped. 
(T. Thomson et J. D. H.) ; Nepal (Wallich); Birma (Wallich et Griffith); Mont. Mishmee (Griffith). 

- Fl. Aug.-Decembr. (v.v.) 


This is an extremely common species in the Eastern Himalaya and Khasia, and so 
variable, that I am quite unable to define its varieties. Specimens of all sizes may be 
found, from an inch to a foot high, of all degrees of robustness, and of all colours between 
blood-red, yellow and white, or brown. Though usually strictly dicecious, I have found 
capitula bearing only male flowers on female plants, and more frequently male flowers 
towards the base of the female capitula. Schott and Endlicher, and latterly Junghuhn, 
have united this species with Blume’s B. elongata, and I should not be at all surprised 
that they proved the same; but I have never found the Indian to have the long branching 
rhizomes of the Javanese species, and there is much less wax in the plant. 

. B. dioica grows indifferently on the roots of many species of shrubs and trees, but I 
have never found that it produces knots on these, as B. involucrata does. 


3. BALANOPHORA ELONGATA, Blume, En. Pl. Jav. i. 87; Schott & Endl. Melet. 13; Unger, 
Ueb. d. Paras. pp. 26 & 33. t. 2. f. 1, 2; Junghuhn in Nov. Act. Acad. Ces. Nat. 


Cur. xviii. Suppl. 207. t. 1. 


Cynopsole elongata, Endl. Gen. PI. 74. es 
Var. maxima. B. maxima, Jungh. in Nov. Act. Acad. Cæs. Nat. Cur. xviii. Suppl. 209. t. 1. 
Hab. Montibus Javæ alt. 5-9000 ped. (Blume, Junghuhn, Lobb). Fl. Mart., Maio et August. Montibus 


` Peninsula Indiz orientalis (Wight) et Ceyloniæ? (Gardner, Thwaites). 


I have, under B. dioica, stated what seem to me the only differences between this species 
and B. dioica, and these are rather modified in the Peninsular specimens figured by Dr. 
Wight in a drawing he has had the goodness to give me, and in the Ceylon specimens. 

Junghuhn's B. maxima seems to differ only in size from B. elongata, the difference 
between these two forms being exactly Serge to ta between B. dioica and B. typhina, 


Wall. (picta, Griff.). 
This species produces wax in great abtsideno which i is used for making candles in Java. 


46 DR. J. D. HOOKER ON THE GENERA AND SPECIES 


4. item INDICA, Wall. Cat. 7247; Weddell in Ann. Sc. Nat. sér. 3. xiv. p. 167. 1 
1.9. f. 11-22. 3 : 1 


Cynomorium, Herb. Wight. e 
Langsdorfia Indica, Arnott in Hook. Ic. Plant. t. 205, 206, et in Ann. Nat. Hist. ii. 36. 


Hab. In montibus Peninsulæ Indiæ orientalis (Wight, Gardner) et Ceyloniæ (Gardner, Thwaites). 


This much resembles a large state of B. globosa, differing chiefly in the longer peduncles f 
of the female capitula, which also have many more scales. 


9. BALANOPHORA GLOBOSA, Junghuhn, Nov. Act. Acad. Cæs. Nat. Cur. xviii. Suppl. p. 210.t.2. 1 


Bal. gigantea, Wall. Cat. 7249, nov. gen. Sarcocordylis (fid. Bennett in Linn. Soc. Trans. xx. p- 94, in note). 1 
Hab. Sylvis montosis Javæ alt. 3-5000 ped. (Junghuhn, Lobb). Birma (Wallich). Fl. April. : 


Junghuhn makes a very curious observation, that when growing with B. elongata (on | 
the same root) he found this species to have the lobed pustules on -its rhizome which 
distinguish that species, but not when it was solitary. E 

According to Wallich, this species is sold for medicinal purposes in the bazars of Burma. : 


E 


6. BALANOPHORA FUNGOSA, Forster, Gen. t. 50; Richard, Elem. de Bot. (1833) t. xv. 1 
(Tag. VIII.) | 
Cynomorium australe, Willd. Sp. PL v. 177. 


Hab. Insulå Tanna Novæ Hebrides ad radices Paritii tiliacei (Forster, Hinds) : ad * Goold Island” in Sinu 
Rockingham, orå orientali Novæ Hollandiæ, fruticetis densissimis (M*Gillivray). Fl. Mai. 


All the specimens of this plant which I have examined have bisexual capitula with the 
female flowers at the base. The surface of the rhizome is minutely granular, and not 
pustular; the peduncles short, stout and leafy. The male flowers have 4—5-lobed peri- 
anths, the lobes grooved inside from pressure against the anther-lobes in the bud. | 

I am not aware upon what plant the Australian specimens were found, but the root is 
very woody, as thick as a crow-quill, and consists of wood and bark with no pith, but 
obscure medullary rays. The wood-fibres are slender and intermixed with large cylin- 
drical ducts and long hexagonal cells whose walls are marked with numerous short 
transverse bars. The vascular bundles in the rhizome are large and stout, branch in 
the usual manner from the root radiating outwards to the lobes of the rhizome, and 

consist (as in B. involucrata) of a thick cylinder of soft colourless parenchyma distin- 
guished from that surrounding it by the absence of chlorophyll or wax, and in this 
respect resembling the bark of the root; its cells are also smaller than the other cells of 
the rhizome, and have rather more numerous punctuations on their walls. The individual 
wood-bundles form a more or less complete zone of wedges, separated by masses of the 
surrounding parenchyma, which also forms a broad cylinder of pith in the interior. The 
wood-wedges are traversed by large ducts, quite similar to those of the root ; these are 
most abundant near the root, and become smaller and inconspicuous at a distance from 


> and A the extremity of the bundles are found as elongated hexagonal cells with 


OF BALANOPHOREÆ. 47 


stem, the wood forming a zone of wedges round a central pith (Tab. VIII. fig. 11) enclosed 
by a cellular zone that communicates with the pith by broad medullary rays: the total 
absence of pith in the root, with whose wood these bundles communicate, would thus seem 
to indicate that the wood of the rhizome belongs to itself, though it has all the appear- 
ance of being solely produced by the root; the root, in short, supplies the nutriment from 
its own vascular tissue, but the parasite organizes it. 


7. BALANOPHORA ALUTACEA, Junghuhn in Nov. Act. Acad. Cæs. Nat. Cur. xviii. Suppl. 
205; Goeppert, ibid. p. 230. t.3. An B. abbreviata, Blume, En. Pl. Jav. i. 87? 


Hab. Sylvis tropicis Javæ (Junghuhn). Ins. Philippinis (Cuming). Fl. Aprili. 


A very much smaller species than any of the preceding, according to Junghuhn's 
description and plate, but probably, like its congeners, extremely variable in size. Its 
prominent characters are the tuberous rhizome, like that of B. dioica and B. involucrata; 

-its few, short, broad, subvaginate scales in the peduncle, and its cylindrical capitula with a 
few male flowers at the base, in which character it resembles B. fungosa and certain states 
of B. involucrata. 


8. BALANOPHORA (POLYPLETHIA) POLYANDRA, Griff. in Linn. Soc. Trans. xx. p. 94. t. 7. 


Hab. Sylvis subtropicis Mont. Khasiæ (Griffith) et Himalayæ provincia Sikkim, alt. 4-6000 ped. (J. D. H.). 
Fl. August.-Novemb. (v.v.) : 

This species is very abundant in the localities enumerated above, and varies in height 
(from 2 to 6 inches), in robustness, in colour, and in the form of the capitula, which are 
however always short and subeylindrie or conical. I have frequently not been able to 
distinguish female specimens of this from those of B. dioica, nor indeed, except by the 
alternate scales, from those of B. involucrata. The numerous anthers of the male flower 
and usually larger perianth of that sex distinguish it from its congeners. 

I have made many detailed analyses of the anatomy of this species at all stages of - 
growth (except the germinating), both in the Khasia Mountains and Himalaya, but do 
not find any point of importance except the anthers in which it differs from B. dioica, 
fungosa and involucrata. The male flowers are well figured and described by Griffith.” 


VII. LormornyTUM, Schott & Endl. 


In habit this genus approaches to Cynomorium more nearly than to any other of 
the Order, as may be seen by comparing their very young states; in each the upper 
part of the rhizome is clothed with spirally arranged imbricating scales, which pass into 
the bracteal scales of the inflorescence. In both the flowers are aggregated into definite 
masses, which masses are immediately covered by the dependent portion of the peltate 
bracteal scales; but whereas in Cynomorium any further tendency to a branched inflores- _ 
cence is arrested at a very early stage, in Lophophytum the development of the branches 
proceeds with that of the whole plant. The paleæ observed by Weddell amongst the 
female flowers of L. mirabile are a further point of resemblance, as are the irregular dis- 
position of the vascular bundles in the rhizome and great abundance of starch-granules 


in the parenchyma. 


48 DR. J. D. HOOKER ON THE GENERA AND SPECIES 


It is a remarkable fact that impregnation appears to be effected in this genus when the 
greater part of the inflorescence is completely clothed with the imbricating bracteæ, and in 
some cases when the plant is still under the surface of the soil. Thus, Weddell's drawings — 
of L. Bolivianum show that even after the fruit is mature it is wholly subterranean, the 
upper part of the male portion of the inflorescence alone being above ground. In my 
specimens of Z. Weddellii, which have all the appearance of being subterranean, and 
which are very young, the pollen is partially shed and the fruit fully formed (though 
abortive), and yet the peduncle would have to lengthen to three times its present length 
before the scales fall away and the plant appears to be in perfection. With regard to 
Weddell's drawings and specimens, they suggest the idea that the upper or male part of 
the spike in elongating under ground sheds so much pollen in the soil, that the female 
capitula on the lower part of the spike, which are afterwards carried up through the same 
soil, are impregnated by the pollen-grains remaining imbedded in it. On the other hand, 
L. Weddellii being diœcious, suggests insect-agency as absolutely essential to the process. 

I have already (p. 24) alluded to the remarkable similarity between the inflorescence 
and flower of this genus and of Gunnera. : ; 


1. LoPHOPHYTUM MIRABILE, Schott & Endl. Melet. i. t. 1 ; Weddell in Ann. Se. Nat. 
ser. 3. xiv. p. 185. t. 10. f. 31-33. 

Archimedea, Leandro. 

Hab. Sylvis tropicis Brasiliæ meridionalis prov. Sebastianopol (Schott): Leandro (in Hb. Mus. Paris.). 


This very remarkable plant is well figured by Endlicher, who however represents it in 
a very advanced state, when the peltate scales have fallen away from the base of the 
capitula of flowers; the younger specimens preserved in the Paris Herbarium altogether 
resemble those of L. Weddellii, but are much smaller. According to a drawing of M. — 
Weddell's, apparently of this species (and which was copied from one by M. Descourtils), 
the whole plant appears immersed in the soil with the exception of the inflorescence, — 
which rises up like the upper part of a long pine-cone with sharp erect brown scales that — 
conceal the red male flowers. 

The scales observed and figured by Weddell amongst the female flowers much resemble 


those of Cynomorium; they are apparently bracteolæ, subtending the female flowers, to 
which they are very obscurely attached. 


2. Lornornytum Botivianum, Weddell in Ann. Sc. Nat. ser. 3. xiv. p. 185. t. 10. f. 29, 30. 
Hab. In Boliviz prov. de Cordillera, sylvis humidis. Fl. Novembri (Weddell). 


I am indebted to Dr. Weddell for a fine drawing of this species and for specimens. In 
general appearance it closely resembles L. mirabile ; but the rhizome is lobed, the lobes 
. rounded, and it appears to be buried up to far beyond the middle of the 
the soil, the whole of the female flowers and the lower half of the male 
of the spike being subterranean: as however the Balanophore and R 
Sometimes wholly exposed (even the rhizome), and at others 
so may these plants be; to which it may be added, that in 


inflorescence in 


hopalocnemis are 
almost immersed in mould, 
the tropical forests wherein 


or upper portion — 


OF BALANOPHORE2. 49 


these plants grow, sudden accessions of rain may often alter the relative level of a plant 
and the soil in which it grows. | 

M. Weddell remarks the absence of styles in this species as constant; but all his speci- 
mens being advanced, and these organs being extremely caducous, I am not disposed to 
lay much stress upon the fact of none of the specimens presenting them. The form of 
the fruit differs from that of L. mirabile, being much less contracted at the base. This 
however is a variable character, and I am inclined to agree with M. Weddell in suspect- 
ing the possibility of its being only a variety of the Brazilian species. 

The arrangement of the stamina upon the lobes or mamillæ of the peduncle appears to 
be definite, though at first sight they appear irregularly clustered. M. Weddell's figure 
and specimens show that the whole surface of each mamilla is covered with lobed conical 
masses, which are connate male flowers, each consisting of two mamillæ placed right and 
left to the axis, and two stamens, one opposite each mamilla. 


3. LOPHOPHYTUM WEDDELLII, nob. (supra, p.30). (Tas. IX.) 


Hab. In sylvis humidis montium prov. Ocaniæ, alt. 3-4000 ped., Novæ Granadæ (Purdie). (Nom. vern. 
* Cardon de la Cordillera.”) | 


I have very fine specimens of this plant sent by Mr. Purdie, but all in a young state, 
with the peduncle wholly covered by the imbricating peltate bracts; but the stamens and 
pollen fully developed, the anther-cells dehisced, and the fruit, though empty, fully formed, 
and its walls crustaceous or bony. It differs from L. mirabile in the much larger size, 
and in being apparently invariably dicecious : I find no paleæ amongst the female flowers. 

The root upon which my specimens grew is as thick as the wrist; it is of considerable 

age, is not much swollen at the surface of contact, has no pith or annual rings, but very 
evident medullary rays. The rhizome of the largest specimens is sunk 3-1 inch ina 
shallow obconical cup in the root. I find no traces of vascular bundles uniting those of 
the rhizome and the root. ; 

The tissue of the rhizome consists of a very dense cortical layer of crustaceous sclerogen- 
cells, which sometimes separates like a bark and encloses a loose cellular mass full of 
starch-granules and vascular bundles traversing the whole in sinuous courses. The 
vascular bundles consist of fusiform ducts and tubes with barred or ringed cylindrical 
or angular walls, surrounded by a laxer tissue of more elongated cells. The arrange- 

^. mení of the peltate scales on the peduncle, and their position relatively to the masses of 
the flowers in the bud, are exactly as in Cynomoriwm ; the lower or dependent porson of 
each bract immediately covers the mamilla of flowers below it, its upper or ascending 
portion covering the dependent base of the scale above it. The vascular bundle of each 
mamilla of flowers does not, however, unite with that of the bract above it, but with that 
below it. — i 


VIII. OmBROPHYTUM, Popp. & Endl. 


This genus bears the same relation to Lophophytum that Balanophora does to Cyno- 
morium, inasmuch as the flowers, instead of being inserted upon the capitulum, are 
Whorled round the pedicel of the bract. In the structure of its female flower it hardly 


VOL. XXII. : 2 2 


50 DR. J. D. HOOKER ON THE GENERA AND SPECIES 


differs from Lophophytwm; the male flowers, according to Weddell’s drawing, appear to - 
consist of a solitary stamen with no trace of a perianth. 3 

The figure given in Pæppig and Endlicher's * Nov. Gen. et Sp. Plant.’ is probably made 4 
up in part from notes or memory, for it can hardly be doubted that the plant they in- 
tended is specifically the same as Weddell's O. zamioides, as he himself informs me. The 3 
rhizome is solitary, tuberous, not lobed, and bears one pedunele, which is surrounded at 4 
the base by a large volva. The inflorescence is cylindrical, and there is no interruption 
between the male flowers which occupy the upper half of the peduncle and the females. 
The bracts are quite similar in each, are persistent, and have a broad orbicular peltate dise. 


1, OMBROPHYTUM PERUVIANUM, Pæpp. & Endl. Nov..Gen. et Sp. Plant. ii. p. 60. t. 155. 
O. zamioides, Weddell, in Ann. Sc. Nat. ser. 3. xiv. t. 10. f. 23-28. 


Hab. In sylvis densis Peruviæ subandinæ ad Cuchero (Peppig); et prov. de las Cordillera (Weddell). 
Fl. Sept.-Nov. (Nom. vern: “ Mays del Monte,” Pæpp.) 


IX. ScyBaLıum, Schott & Endl. 


1. SCYBALIUM FUNGIFORME, Schott & Endl. Melet. p. 3. t.2. Unger in Ann. Wien. 
Mus. ii. t. 2. f. 4, t. 4. f. 19, 20. 


Hab. In sylvis Brasilize ad Sierra d'Estrella (Schott). 


I have nothing to add to the excellent description cited above. In habit the genus - 
approaches Langsdorffia more than any of the Distyli. In the crowded and often connate 
peduncles and unisexual capitula, as well as in the plane figure of the latter, it differs | 


from its allies; also in the filiform hairs which occur amongst the male flowers and 


= are exserted far beyond the articulated threads, and which are probably undeveloped 1 
females. The 2-celled ovar 


y invariably (as it is described) followed by a one-seeded fruit, | 
is unique in the Order. ] 


The rhizome resembles that of Corynea, the scaly peduncle that of Sphærorhizon. The 


fact of one male peduncle being always surrounded by several females on the same rhizome 
is unique, and does not necessitate the ope | 


ration of dichogamy which occurs in Helosis. 
From fig. 13 of the plate quoted, it appears that the male flowers are developed in succes- 
sion for a considerable period. 


X. SPHÆRORHIZON, Hook. fil. 


Rhizoma napiforme seu depresso-globosum, solitarium, Pedunculus solitarius, curvus, squamatus, basi 


annulo brevi rhizomatis circumdatus, junior squamis densissimå imbricatis velatus, demüm elongatus, 
squamis oblongis obtusis subpeltatis denique deciduis. 


E Capitulum ovoideum, monoicum, squamis 
deciduis peltatis tectum. Flores et fila 


| articulata Helosis, sed synemate 3-loculari apice dehiscente. 
1. SPHÆRORHIZON CURVATUM, Hook. fil. (Tan. X.) 
Hab. Sylvis alpinis Novæ Granadæ inter vicum Ni 


Rhizoma 3-3 unc, diametr., obscurè lobatum, 


validus, I une. diametr., interdåm brevi 
catis tectus, 


laxè velatus, 


va et montes Paramo de Ruiz dictis (Purdie). Fl. Jul. 


radicibus validis sessile. Pedunculus flexuosus 1-6 uncialis, 


x : ssimus et squamis multiseriatis patentibus densissimé imbri- 
interdüm elongatus squamis longioribus å unc. longis obtusis latà basi subpeltatim affixis 


Squamz infra capitulum magis peltatæ latè adnate basibus apicibusque truncatis sub- - 


- 


OF BALANOPHOREÆ. 51 


recurvis. Capitula laté oblongo-cylindracea v. ovoidea, filis articulatis densissimè operta, e squamis 
delapsis areolata. Flores masculi synemate solido triloculari (nempe antheris unilocularibus) cylin- 
draceo, apice dehiscente, septis e strato duplici conflatis, exteriore celluloso, interiore floccoso albido 
e fibris minutissimis implexis. Pollen globosum, immaturum 3-nucleatum, maturum peripheriá 
hyalinå, tuberculis 3 notatá. Ovarium lineari-obovatum compressum, perianthii labiis latè ovatis, 
stylis breviusculis recurvis. Fila articulata lineari-clavata simplicia v. varié coadunata. 


Å very curious and peculiar-looking plant, of which Mr. Purdie has sent several spe- 
cimens belonging possibly to two species, and differing remarkably in the amount and 
disposition of the scales of the peduncle. These in small short specimens form a dense 
mass between the rhizome and capitulum, are very numerous, closely packed, broad, 
short and patent: in specimens 3-5 inches long, the scales are more loosely placed, linear- 
oblong, adnate by a broad base, and those towards the capitulum are completely peltate 
or sessile by the middle and lower part of the face of the scale; the upper parts and base 
being truncate, free, and slightly recurved: still larger and apparently old specimens 
appear to have shed these scales altogether. The bracts on the capitula are much larger 
than in Helosis, but are in an imperfect state upon the only specimen retaining them. 

The plants are evidently of annual duration, penetrating the last year's wood only, and 
producing no injury in the layers below that. 


XI. PHYLLOCORYNE, Hook. fil. 


Rhizoma crassum, deforme, ramosum. Pedunculi crassi squamis hexastichè imbricatis tecti, supernè tuber- 
culis conicis densissimè obsiti, | Capitula androgyna, cylindracea, squamis imbricatis deciduis tecta : 
filis articulatis densissim operta. Fr. 4. Perianthium 3-lobum. Anthere 3 in capitulum 6-loculare 
apice dehiscens connatæ. Fr. 9. Ovarium obovatum, compressum, calycis limbo bilabiato coronatum. 
Fructus turgidus, utrinque truncatus. 


l. PHYLLOCORYNE JAMAICENSIS. (Tas. XI) 


Cynomorium Jamaicense, Swartz, Fl. Ind. Occ. i. p.11; Browne, Jam. p. 334. 

Helosis Jamaicensis, A. Richard, Mém. Mus. viii. 432. 

Scybalium? Jamaicense, Schott & Endl. Melet. p. 12. 

Hab. In sylvis savannisque Jamaicæ (Swartz, Purdie, Wilson) Fl Jan.-Jul. (Nom. vern. * John 


Crow's nose.") 

Richard suggested the separation of this from Helosis, with which genus however it 
entirely agrees in the structure of the flowers of both sexes, but differs remarkably in 
habit, in the leafy peduncles, and imbricated bracts of the capitulum. Like its congeners, 
it varies extremely in stature and in the relative size of its organs; old specimens form 
subterranean masses a foot in diameter. 

The rhizome on a transverse section presents a thick, brown, cellular cortical layer, 
formed of hexagonal cells full of starch-granules and chlorophyll, with occasionally masses 
of sclerogen-cells. "The axis is occupied by a slender column of cellular tissue forming a 
true pith; it is surrounded by a layer of long woody sclerogen-cells or tubes ‚that pass 
between the wood-wedges as medullary rays, and are there shorter and cubical. The 
wood-wedges are about twenty, lanceolate (on the transverse section), symmetrically 


disposed round the axis, and are composed of pale slender tubes, which are scalariform 
H2 


MISSOURI 
BOTANICAL 
GARDEN. 


52 DR. J. D. HOOKER ON THE GENERA AND SPECIES 


or subspirally marked. Outside of each wedge is a bundle of liber-cells, which are - 
long sclerogen-tubes. This arrangement accords with that of Helosis, and is essentiålly 
exogenous. | ; 

The young peduncle and capitulum are wholly concealed by the imbricating scales and 
bracts, which are hexastichously arranged in my specimens, but pentastichously as — 
described by Swartz. As the peduncle elongates the bracts fall away from the lower 
part of the capitulum, leaving an areolated surface; the uppermost are persistent for a 
considerable period. The upper part of the peduncle or base of the capitulum is rough 
with conical papillæ, which become fusiform and slenderer upwards as they mix with the 
articulated threads and female flowers, of which they appear to be arrested states. 4 

The female flowers protrude their styles, which are bent down under the bracts, as - 
soon as the latter fall away, and the evolution of the flowers follows the same law as in 
-Helosis. The ovary resembles that of Scybaliwm more than of Helosis, being broader, with 
larger semicircular lips to the calyx and short stout diverging styles. The fruit is broadly 
ovoid, truncate at both ends, striate, sulcate towards the apex, and more turgid than usual 
amongst the Helosideæ. On a transverse section the style is found to consist of about - 
eight large cells surrounding a few smaller ones that enclose a soft conducting tissue. 
The seed is broadly oblong, compressed, very oily, of the same structure as Helosis. 

The articulated threads of the capitulum are (like the ovaries) much broader in this 
genus than in its allies, and are often fusiform and geminate. The apical cells turn black, 
and their cell-walls become minutely wrinkled soon after the bracts fall away. 

The male flowers have often a 4-lobed perianth, and the odd lobe is as often the lower 
. as the upper; there being no constant arrangement of the lobes with reference to the 
axis of the capitulum. The anthers are 2-celled, narrow oblong, and the pollen escapes 
through an irregular opening at the apices of the loculi: the filaments are free just below 
the insertion of the anthers. Pollen globose, with four minute papillze on the surface. 


XII. RuoParooxEuis, Junghuhn. 


CHAR. EMEND.—Rhizoma deforme, globosum, lobatum. Pedunculi basi volvå carnoså irregulariter fisså 
circumdati. Capitula elongata, bracteis peltatis deciduis velata, unisexualia, filis articulatis densè 
obtecta. Fu. 3. Perianthium campanulatum, integrum, basi columnæ stamineæ adnatum. Filamenta 
coadunata, longè exserta; antheræ 3, 2-4-loculares, arctè cohærentes, apice dehiscentes. FL. 9 ob- 


longæ, compressæ; perianthio bilabiato; stylis 2 elongatis; ovulo 1 pendulo, Fructus lineari- V. | 
ovato-oblongus, turgidus. Semen Heloseos. 


1, RHOPALOONEMIS PHALLOIDES, Junghuhn, in Nov. Act. Acad. Cees. Nat. Cur. vol. xviii. | 
Suppl. p. 215. (Tan. XII.) 


Pheocordylis areolata, Griff. in Linn. Soc. Trans. xx. p- 100. t. 8, descript. incompleta. 
Hab. Sylvis Acacie montibus Ins. Javæ, alt. 7000 ped. (Junghuhn)! Sylvis densis montium Khasiæ, et 


Himalayz orientali iæ orientali ikki | 
e Duc. 2n entalis, Nepaliæ orientalis et Sikkim, alt. 6-8000 ped. (Griffith, Thomson, J. D. Hm | 


| This ne remarkable plant has been much misunderstood, both by the author of the | 
genus, .J unghuhn, and more recently by Griffith. From J unghuhn' s des criptio n it is P 


OF BALANOPHOREÆ. 53 


probable that his specimens were most imperfect, and perhaps covered with mould. The 
museum at Kew is indebted to Prof. de Vriese of Leyden for beautiful Javanese specimens 
of both sexes, which being authentically named, enabled me to recognize Junghuhn’s 
plant as identical with the Himalayan and Khasian one; and which, making allowance 
for the absence of male flowers, and for his erroneous description of the females, is also 
identical with the Phæocordylis of Griffith, gathered at the same spot in the Khasia 
Mountains where Dr. Thomson and I procured an abundant supply of specimens. 

Rhopalocnemis is by far the largest and the handsomest of the Helosideæ, and it is the 
only one which I have had an opportunity of examining in a living state; it is most 
closely allied to Corynæa, differing in the presence of a volva and in the unisexual capi- 
tula. It grows gregariously, in shady mountain woods, its large heads of a pale yellow- 
brown colour alone appearing above ground: it is of a firm, fleshy consistence, perfectly 
inodorous even when decaying. I have vainly tried to induce the ripe seeds to germinate, 
and have examined many hundreds in the fruitless attempt to discover any embryo in the 
mass filling the whole cavity of the seed. During the shedding of the fruit, the capitula 
(of gathered specimens) copiously exuded a transparent sugary fluid, but I have never 
observed this on the living plant: it is no doubt analogous to the fluid described by 
Weddell as bathing the capitula of some Balanophoreæ, and supposed by that author to 
be of use in the operation of fecundation. , 

The rhizome varies from the size of an egg to that of the human head, and is supplied 
internally with many stout woody branches, which appear continuous with the wood of 
. the rootstock, and which upon maceration are found to send continuous bundles to the 
top of the capitulum. The peduncles are solitary, or many together on large rhizomes, 
and are enveloped at the base by a hard, fleshy, erect, cylindrical volva, 3-2 inches high ; 
they vary in length from 2 to 6 inches, and in ‘diameter from } to 2 inches; they are 
altogether naked below, but in the upper part are covered to a greater or less distance 
below the capitulum with fleshy, patent, and somewhat recurved scales, 4 inch long, 
which appear to be persistent, and to occur chiefly on the male plants. The hexagonal 
fleshy scales which cover the whole capitulum are altogether similar to those of Helosis ; 
as are the female perianths to those of Corynæa, and the males to those of Helosis. —— 

In flowering, both males and females expand at the same time, throwing off their 
cohering bracteal scales in large masses, and exposing a velvety pile of styles, and a 
dense mass of subjacent articulate threads. There are several crops of male flowers, 
which expand successively ; and in the dense humid calm woods in which this genus grows, 


insect agency is probably necessary to impregnation. During the ripening of the fruit, 
d from the swelling of the masses answering 


the surface of the capitulum becomes areolate 
to an obscure lobing of that organ, and at first externally defined by one of the fallen 


bract-scales, and internally by a vascular bundle from the- plexus of vessels within the 
capitulum. ; ee: 

My examination of living specimens, both in the Khasia Mountains E ii on , ied 
to no results which may not as well be obtained from those preserved in spirits, for the 
sphacelation and browning of the cut surfaces were SO instantaneous, that I had to put 
the sections in spirits as soon as made. Å careful study of the ovule and seed at all 


54 DR. J. D. HOOKER ON THE GENERA AND SPECIES 


stages of growth led to the same conclusions as were obtained from Balanophora involu- 
erata. After flowering, the capitulum and peduncle wither, dry, and fall away from the 
base of the volva, but do not deliquescé nor turn putrid, nor are they attacked by insects 
to any extent. Each peduncle seems to become detached from the rhizome by a very 
obscure articulation, which may be seen as å dark transverse line, on å longitudinal section 
of the base of the peduncle, some time before its falling away. The rhizome is perennial, 
but does not give forth its buds until July, and the plant ripens its fruit in September. 3 

The apex of the stigma, described and figured by Griffith as an opaque rounded body, - à 
is when young, and even after impregnation, perfectly simple. 

The apices of the articulated hairs consist of several series of cells, which are (as figured — 
by Griffith) much darker than the rest; and the outer cell-membrane becomes transversely 
wrinkled, as in the South American Helosideæ. 


XIII. Corynæa, Hook. fil. 


Rhizoma deforme, lobatum. Pedunculi nudi. Capitula androgyna, squamis hexagonis peltatis velata. | 
Fu. 3. Perianthium infundibuliformi-campanulatum, margine crenatum. Anthere 3 v. synema — 
` 6-loculare. Fr. 9. Heloseos. : 


1. Coryn#a CRASSA, Hook. fil.; capitulis clavatis cylindraceis. (TAB. XIII.) 


Hab. Sylvis densis Crategi, Thibaudie et Befarie, prope S Fé de Bogota, Nove Granade, alt. 8000 ped. 
(Purdie, 1846). : 

Rhizoma lobatum, a magnitudine nucis Juglandis ad capitis humani. Pedunculi plurimi, juniores basi E 
annulo cincti, 1-8 unc. longi, 1-12 unc. crassi, nudi, cylindracei. Capitulum obovato-oblongum v. 
clavatum, in pedunculum attenuatum, flores perplurimos utriusque sexus gerens, filis articulatis 
immixtis. Fr. 4 ad basin capituli plurimi. Perianthium basi cum tubo stamineo connatum, 
supernè ampliatum, filis paulló longius, margine crenatum, rariüs fissum. Columna staminea longè 
exserta, filamentis omnind coalitis. Anthere 3, 2-loculares, arctè cohærentes, introrsüm et ad apices 
dehiscentes. Pollen 3-4-nucleatum. FL. 9. Ovarium obovato-oblongum, compressum, limbo peri- 


anthii breviter bilabiato coronatum ; styli graciles, elongati. Fructus latè oblongus v. orbicalaris, 
compressus. - 


This fine species often weighs many pounds, and is so abundant that the roots of å 
plant attacked by it resemble a mass of potatoes. The rhizome encloses the roots of the 
plant on which it grows parasitically. In the evolution of the flowers it follows 
the same law as Helosis. The male flower entirely resembles that of Rhopalocnemis. 
After the fall of the scales from the capitula, both the stigmata and the tips of the arti- 
culate threads covering the head sphacelate; the latter wrinkling, and the stigma, which | 
consists of two series of globular cells (enclosing a mucilaginous passage down the style), 
becomes minutely punctulate. The body of the style consists of about eight large peri- 
pherical cells, enclosing the cellular conducting tissue. The cavity of the ovarium is 
small, compared with its congeners; and the seed * is also small and broad. | 

* In Tab. XIII. figs. 11, 12, 13, I have figured a large embryo in i : thi i | 
fruits (now ten years ago) ; but having meer fed since das in me ine" prem dad g | 


in the body of this memoir. Since Mr. Weddell has informed me of the very rare occurrence of the calyx in Sar- | 


cophyte, I have been induced to reconsider the propriety of introducing these drawings, and now do so with the more 


confidence from the embryo of Sarcophyte being so entirely similar to this.—Kew, Aug. 2, 1856. ft 


OF BALANOPHOREÆ. ` 55 


At the earliest period each female flower often appears as two flagon-shaped bodies, con- 
nate from the middle downwards; and at that stage is evidently composed of two ovaria : 
when fully formed, and the scales are still attached to the capitulum, the styles are 
exserted far beyond the articulate threads, and are sharply bent down, so that the stigmata 
are in contact with the apices of the threads. "The utricular tissue of the capitulum is 
composed of hexagonal cells, each containing a loose inner coat, full of starch and 
endochrome. 3 


2. CORYNEA SPHJERICA (Hook. fiL); capitulo solitario globoso cavitate rhizomatis semi- 
immerso. (Tas. XIV.) 


Hab. Sylvis montanis Novæ Granadæ, alt. 5-8000 ped. (Purdie). 


Rhizoma deforme, lobatum, diametro 2-4 unciali, infra capitulum in cupulam hemisphæricam marginibus 
crassis obscuré lobatis ampliatum. Pedunculus brevis, crassus, nudus. Capitulum exactè globosum, 
2 unc. diametr., squamis peltatis hexagonis velatum. Flores masculi ut in C. crasso, sed perianthio 
breviore latiüs campanulato truncato obscuré trilobo. 


A very different species from C. crassa. The rhizome completely surrounds the root of 
the stock; and in a transverse section I find that the latter is sometimes so completely 
broken up, that the end which enters seems cut off from that which leaves the rhizome. 
(See Tas. XIV. fig. 14.) 


3. Conyx xA Purpret (Hook. fil.) ; rhizomate depresso horizontali lato lobato, pedunculis 
brevibus crassis, capitulis oblongis globosisve. 

Hab. Sylvis montosis Novæ Granadæ (Purdie), radicibus Cinchone : sylvis prope St. Juan del Oro, 
Peru (Weddell). 

I am indebted to M. Weddell for an excellent drawing and specimen of this very distinct 
little species, of which I had previously received a small example from Mr. Purdie. The 
rhizome forms a continuous broad flattened lobed mass enveloping large roots of Cinchona 
several inches to half a foot in extent, and is about 2 inch in average thickness. The 
lobes are but little elevated, and give off short white peduncles, 4-1 inch high, that are 
conical below and about 4 inch in diameter. The capitula are nearly globose or ovoid, 
about 1} inch diameter, are quite white at first, but covered with red-brown hexagonal 
peltate seales. "The flowers entirely resemble those of C. sphærica. 

There is sometimes a very obscure volva at the base of the peduncle. 


XIV. HELOSIS, Rich. 
(Tas. XV. & XVI.) 


Caldasia, Mutis, Sem. Nov. Granad. Lathræophila, Leandro de Sacram, 


Rhizoma horizontale, gracile, teres, ramosum, hic illic nodosum, ad nodos pedunculos (ramos floriferos) 
olucello annulari donati, rariùs 


emittens. Pedunculi erecti, basi v. medio v. infra capitulum inv Å 
omnind nudi. Capitula androgyna, filis articulatis operta, juniora Serie Pen herpa — 
Fr. å. Perianthium tubulosum, limbi lobis 3 valvatis, tubo intüs basi rudimento ovarn conico in- 
structo. Synema fauce perianthii adnatum. Anthere 3, introrsæ, in massam 6-12-locularem coadu- 


nate.  Fr.9 subsessiles. Ovarium: elliptico-oblongum, utrinque obtusum, compressum, limbo 


56 DR. J. D. HOOKER ON THE GENERA AND SPECIES 


perianthii breviter bilabiato coronatum, 1- (rarius 2-3) loculare. Styli 2, rariüs plures, filiformes, 
Stigmata subcapitata, papillosa. Ovulum 1, pendulum. Fructus subcrustaceus. Semen 1, achenio 


conforme, testá tenuissimá hyaliná reticulatá. 

Helosis appears to be the commonest American genus of the Order, inhabiting both — 
sides of the Andes, and extending from Mexico to the river la Plata. The species much 4 
resemble one another in general characters, and are of a whitish colour tinged with red, 
and become red-brown when dry: they are said to inhabit moist grounds, where their 
rhizomes spread annually by innovations to a considerable distance, seeking nourishment 
from various roots in their progress, and seeming to have the power of attacking such as 
they come in contact with. Each year’s rhizome is probably annual, and it gives off an 2 
innovation before dying, as described by Richard; the whole mass sometimes perishes … 
at once. 

The parasitism is simply that of adhesion by the contact of the tissues of the Helosis 
with those of the root-stock ; in the older specimens there are no vascular bundles uniting 
both, and the roots attacked do not swell up to any remarkable size at the point of union; 
though the parasite often penetrates deeply into the wood by a conical protuberance. In 
very young plants, however, the wood of the root-stock ramifies extensively through the 
tubers of the parasite. A transverse section of the rhizome shows a most distinctly | 
exogenous structure, very curiously modified, and varying considerably in the different — 3 
species, under which the details will be given which have been already referred to in the | 
general remarks on the anatomy of the Order. å 4 

The peduncles are always erect, and rise from a swelling on the rhizome, whence they — 
receive many vascular bundles. The bundles in the peduncle are, however, simple, and 4 
either promiscuously scattered, or arranged in a circle; each resembles in structure that : 
of a monocotyledonous stem, having its own liber, wood, and vascular portions; but the E 
bundles do not follow the course that they do in endogenous stems, and are not to be — 
regarded as indicating any affinity between Helosis and Monocotyledons: they are, in 
fact, solitary bundles such as occur in the leaves, and often in the annual flowering | 
branches of other Exogens. E å = 

An incomplete involucre, generally divided into 3-6 broadly ovate segments, is frequently 
present in this genus; in H. Guyanensis it is placed at the base of the peduncle, in the 
Andes variety of that species it is carried up towards the apex, while in H. Mexicana it 
is either reduced to an elevated ridge round the centre, or entirely absent. When fully 
developed, this involuere never encloses the young capitulum. : 
Se peltate, fleshy scales cover the whole capitulum, as in the Indian Rhopa- 
48. na very young state these will be found to be developed as imbricating, 
ascending, bracteal leaves, each covering a definite portion of the inflorescence, which is 
indicated by a vascular bundle, given off from a plexus in the bod 


thei en. À y of the capitulum; 
_ their position is hence analogous to bracts subtending branches of a flowering axis. As 


the inflorescence grows, they become peltate, hexagonal from mutual pressure, and ad- 


hering by their contiguous edges, fall away i : 
R , y in large masses, 1 ; 
faintly marked on the capitulum, : 18» PaVIDG corresponding areola 


The male flowers haye usually a conical body at the base of the tube of the perianth, 


OF BALANOPHOREÆ. 57 


which probably represents a rudimentary ovary. The filaments are free just below 
the anthers, to a greater or shorter distance, which varies in the individual species, 
as does the length of the filaments. The anthers burst introrsely; they are firmly 
united into an obtusely trigonous mass enclosing a central cavity; each is 4-celled, the 
mass consequently being originally 12-celled: the cells are shown in a transverse section 
to be disposed in two concentric series, of which the inner has much the smallest cells ; 
generally the two rows become confluent. E 

The female flower offers little worthy of notice, except the occasionally 3-lobed young 
flowers, indicating three ovaria, as figured in Tag. XVI. figs. 8,9, 10; and the anomalous 
membrane enveloping the terminal cells of the articulated threads in 77. Mexicana, which 
is probably mucous, and may be the source of the fluid which is said to bathe the capitula 
of some species during flowering, and thus to facilitate the dispersion of the pollen. 


1. Hetosis Guyaneysts, Rich. Mém. Mus. viii. p. 416. t. 20; Martius, Nov. Gen. et Sp. 
Plant. Bras. iii. p. 184, t. 300 & 208. fig. 2. 


Caldasia Cayennensis, Mutis, fid. Steud. 

Cynomorium Cayennense, Swartz, Fl. Ind. Occ. i. 13. 

Var. a. pedunculo elongato gracili, volva v. involucro ad basin pedunculi. 

Var. 8. pedunculo abbreviato, volva v. involucro ad basin pedunculi. H. Brasiliensis, Schott & Endl. 


Meletem. p. 12. 

Var. y. andicola, pedunculo brevi, volva v. involucro 4-6-fido infra capitulum sito. 

Hab. Sylvis humidis Guiana, Richard; Para, Martius, Spruce (Aug.-Dec.) ; Jamaica et Trinidad, Purdie 
(May 1848); Berbice, Schomburgk ; Pampas, Buenos Ayres, Miers.—Var. 8. Rio de Janeiro, Miers ; 
Serra d’Estrella, Brasiliæ, ScAott.—Var. y. Vegas de Rio Quindiu, Goudot (in Herb. Webb, No. 140). 


This remarkable plant has been well described by Swartz, and again (with illustrations) 
by the elder Richard, and by Von Martius. It appears to be common in damp woods, 
on the east coast of South America, ranging from Trinidad to south of the Equator. It 
varies extremely in size, being from an inch to nearly a foot in height; with slender or 
robust peduncles and rhizomes, and ovoid or subcylindrical capitula, which (according to 
à drawing by Sir Robert Schomburgk) are sometimes lobed or even deeply bifid at the 
summit. Schott and Endlicher have made a species of the var. Brasiliensis, because of 
its 3-lobed involucre and small size; but the involucre is generally 3-lobed, and is 
described as such in the Guiana species by Richard and Martius, and Miers's Brazilian 
specimens have 5-6-lobed involucres. 

The rhizome creeps to great distances in spongy soil, forming adhesions with the roots 
it encounters. A transverse section of Trinidad specimens displays an arrangement of 
the tissues in several respects closely resembling that of many Menispermous plants. 

The axis, or position of the pith, is occupied by a cylinder of elongated, hard, woody, 
cylindrical tubes, with very narrow, often interrupted cavities, and this sometimes sur- 
rounds a central pith of loose hexagonal cells*. These tubes become broader and shorter 

* The occasional presence of a cellular pith within this woody axis is important, as it reduces the type - which 
Langsdorfia belongs, which has no cellular pith, to that of most other Balanophoreæ. It is a curious fact, that in many 


Balanophoreæ the relation of the vascular system to the cellular is reversed, in respect of the latter being excessively 
dense, hard and rigid, whilst the true woody system is composed of extremely lax, soft, thin-walled vessels,—of ducts, 


‚in fact, with little or no pleurenchyma. 


VOL. XXII. r 


58 DR. J. D. HOOKER ON THE GENERA AND SPECIES 


between the wood-wedges, where they appear as medullary rays, lastly passing into the — 
loose hexagonal tissue of the circumference of the rhizome. 

Seven narrow, elliptical, pale white wedges succeed the woody axis that occupies 
the position of the pith, then radiate, are equal in length to about half the radius of the 
rhizome, and placed midway between the periphery and circumference; they are separated 
from one another by the broad medullary rays, which assume the character of cubical, 
hard, brittle wood-cells consolidated into one dense firm mass, protecting the softer tissue 
between them. The wedges themselves are formed of delicate, white, large tubes, placed 
end to end, and transversely marked with short lines, annular or spiral bands. 

A broadly semilunar or kidney-shaped mass of wood-cells (liber) is placed externally . : 
to each vascular wedge and curves round its outer extremity, and is either placed closely — 
in apposition to the vascular wedge, or is separated from it by a little cellular tissue. … 
These wood-cells are very large and thick-walled, are vertically elongated, and form long 
parallelograms placed end to end, and adhering firmly one with another, become of a 
dense yellow, almost crustaceous or osseous consistence; their walls are everywhere per- 
forated by minute canals, giving them a punctate appearance. 

The cortical portion or cellular tissue of the periphery is formed of hexagonal thick- 
walled, almost woody cells, with perforated faces, and there are scattered irregularly 
through it very large sclerogen-cells and liber-bundles. "This cortical portion is spongy 
in consistence, and its hexagonal cells gradually pass into the cubical ones of the me- 
dullary rays. ; 

Such appears to be the arrangement in the first year; in the second, more woody liber- | 
bundles are formed outside the semilunar ones, and alternating with them. The wedges 
of vascular tissue do not appear to be added-to much, but there is an appearance of incom- 
pleteness towards their circumference, as if a cambium-layer existed there. Strictly 
speaking there are only two well-defined kinds of tissue in the rhizome :—1. the delicate 
vascular wedges, and 2. the coarse, hard, hexagonal cellular tissue of the periphery, which 
becomes indurated between the vaseular wedges and passes into the slender woody tubes 
of the pith: the other tissues that are so conspicuous on a transverse section are not so 
on a vertical one, the broad sclerogen-tubes of the semilunar bundles of liber differing 
little from the cubical cells surrounding them, and the liber-bundles of the periphery 
altogether resembling the long woody tubes of the pith. 

Many deviations may be found in different specimens from the above-described 
arrangements of the cellular and vascular systems of the rhizome; but all, I think, may 
be easily reduced to this type. 

Red sas sears ee en consists of scattered bundles that run free and 
a plexus within the danh ice — where they partially anastomose, i 
erence, from which bundles are given off with great regularity 


towards the base of each scale. I do not find the tissues of these bundles to be more 


than rudimentary ; but traces of their each consisting of a bundle of woody tubes towards 


the axis, followed by delicate transversely b 
tubes : arred vessels, an À n- 
» may, I think, be detected. å s, and these again by sclerogen- 


nn the styles as occasionally united at their bases, which I have never 
3 that he has never observed the plants to be truly parasitical, though 


OF BALANOPHOREÆ. 59 


their roots intertwine with those of other plants: my specimens undoubtedly contract 
broad organic adhesions with the roots they encounter, and in the young state receive 
woody bundles from them. Richard's admirable account of the epoch of fecundation 
agrees with what I have observed in the monecious Balanophore of India. The capitula 
are never self-fertilized; the styles of the female flowers are protruded immediately after 
the fall of the scales, and fertilized by the pollen of a neighbouring capitulum ; the styles 
then fall away, and during the maturation of the fruit, the male flowers are protruded 
and shed their pollen to fertilize another capitulum ; by the time that the latter opera- 
tion is performed the fruits have ripened, are shed, and the peduncle and capitulum 
perish, though the latter still contains an abundant crop of young male flowers, apparently 
destined never to perform their functions. This apparent superfluity of male blossom is 
a very remarkable phænomenon, and not at all comparable with the common one of 
numerous male flowers on one inflorescence never becoming perfected except under 
favourable conditions, for in this case there appears to be a second crop of males after the 
first have performed their office, and after the females of the same and all the other 
capitula are fertilized, and it is difficult to conceive any circumstances arising at all likely 
to call for the operation of these complementary males. 

Martius mentions that a beetle of the family of Curculionide, or its larva, possibly 
assists in the fecundation, as it is found nidulating in the capitula; judging however from 
the fact of one capitulum being fecundated by another, the larvæ could be of little use, 
nor can the beetles themselves be of much, under ordinary circumstances. | 

Martius mentions delicate thread-like radicles as proceeding sometimes from the base 
of the seed (embryo): that author also states that the disposition of the vascular system, 
both in its nature and arrangement, is monocotyledonous, an error to which I have else- 
where alluded in my general remarks on the Order. 

Schott and Endlicher (Meletem. p. 8) observe, that in their H. Brasiliensis there are 
sometimes two and even three cavities in the ovarium, accompanied in the latter case by 
three styles. I have never seen such an arrangement in any specimens of this species, 
but indications of it will be shown to occur in the lobed young flowers of H. Mewicana. 
Swartz (Fl. Ind. Occ.) describes the styles as sometimes solitary, probably from one 
having fallen away, as he did not examine living specimens. | 

In the variety +, for which I am indebted to the late P. B. Webb, Esq., the cellular 
tissue of the periphery consists of vertically elongated and much more delicate utricles, 
often filled with starch and chlorophyll grains: there is also a slender central column of 
true cellular pith surrounded by those woody tubes that are often seen to be the only pith 


of the varieties a. and ß. 


2. Herosts Mexicana, Liebmann, Proceedings of the Scandinavian Meeting of Natural- 
ists, p. 181. 


H. aquatica, Mutis MSS. in Herb. Hook. : vo 
Hab. Mexico montibus ditionis Vera Cruz et Oajaca, alt. 3-5000 ped., Liebmann (v. ic. pict. a cl. auct.). 
Mirador (Linden), Jul. Convallibus humidis Novæ Granadæ ad Melgar (Purdie), Febr. 1846. 


Less variable in form and more so in robust or slender habit than the €— The 
! 12 


60 DR. J. D. HOOKER ON THE GENERA AND SPECIES 


earliest stage at which I have examined this plant is that of an amorphous cellular hemi- | 
spherical mass nidulating within and almost enclosed by the bark of the roots, into the 
wood of which it had penetrated, its cellular tissue being in immediate contact with the 
wood-fibres: in this early stage it contained no vascular tissue and no traces of lobing or — 
of peduncle. When fully developed, this species so entirely resembles H. Guyanensis in - 
all but the nature and position of the volva or involuere (here reduced to an oblique ring 
about half-way up the peduncle), as to preclude the necessity of a detailed description, 
and I shall proceed at once to some remarks on its structure and development. 

The structure of the wood of the rhizome is essentially the same as that of the preceding 
species. The vascular bundles of the peduncle are, however, more regularly disposed. 
The young peduncles arise as buds from circular depressions in the rhizome, and do not 
exhibit any volva or involucre. 

The scales on the young capitulum are all ascending and imbricating, and do not 
assume the peltate and hexagonal form till a subsequent period. 

At the base of the capitulum are seen several rows of small conical protuberances that 
appear to be undeveloped bracts. The annular projection which represents an involucre 
never to be developed, is situated close under the capitulum for some time after the 
lengthening of the lower part of the peduncle; finally, however, the upper part of the 
peduncle elongates most, and the annulus hence occupies a middle position upon it. 

The articulated threads of the surface of the capitula are stout, of several collateral cells 
in breadth, and the upper eight or ten cells are smaller and become sphacelate very soon. 
In a young state these terminal cells are seen to contain several nuclei, and they are all 
enclosed in a membrane of excessive delicacy, forming a balloon around them ; they appear 
much whiter than the other cells at this early period, but afterwards are darker and 
become minutely wrinkled on the surface. 

At the earliest period at which the female flowers can be recognized, they appear as 
minute 2-3-lobed cellular papillæ, broader than long, upon the surface of the capitulum. 
When 2-lobed, the lobes (which lengthen into styles) are widely separated at their bases - 
and diverge. As they increase, the lobes lengthen and approximate; and if a third one be - 
present, it is the middle one of the three that is suppressed. 
At this period the perianth is not distinguishable, but appears after the flowers have 
assumed their complete form, as a broadly campanulate 2- (sometimes 3-4- ?) lobed 
superior calyx, much larger in proportion to the ovary than at any future period; the - 
styles at the same time have approximated, and appear united at their bases into a conical 


body surmounting the ovary. The full-grown ovary differs in no important particular 
from that of the preceding species. 


OF BALANOPHOREÆ. ; 61 


EXPLANATION OF THE PLATES. 
TAB 4. 


A. Cynomorium coccineum, Mich. 


Fig. 1. Vertical section of a portion of very young capitulum, showing two bracts and their included 
masses of flowers ;—though the lower limb of each bract immediately covers only the mass of 
flowers below it, the vascular cord of each bract unites with that of the mass of flowers 
above it. 

Fig. 2. A very undeveloped female flower, showing the vascular cords of the ovary. 

“ Figs. 3, 4 & 5. Hermaphrodite flowers. 

Fig. 6. A male.flower with a regular 6-leaved perianth. 

Fig. 7. A female flower showing at «. a mamilla, which may be an undeveloped stamen. 

Fig. 8. Portion of a male flower showing an undeveloped ovary a, and a wholly inferior perianth. 

Fig. 9. Another male flower with undeveloped ovary and wholly superior perianth. 

Fig. 10. Vertical section of immature fruit. 

Fig. 11. Ripe seed and embryo cut vertically. 

Fig. 12. Embryo very highly magnified. 

Fig. 13. Apex of style and stigma, showing the two vascular cords and intermediate groove filled with 
-conducting tissue. 


All the figures very highly magnified. 


B. Mystropetalon Thomii, Harv. 
Fig. 1. Female flower. 


Fig. 2. Apex of style and stigmata. 
Fig. 3. Vertical section of unripe fruit. 
Fig. 4. Ditto of ripe fruit. 
Fig. 5. Transverse section of the same. 
Fig. 6. Seed. : 
Fig. 7. Vertical section of ditto. 
Fig. 8. Embryo. — 
All very highly magnified. 
C. Sarcophyte sanguinea, Sparrm. 
Fig. 1. Male flower. 
Fig. 2. Segment of perianth with stamen with undehisced anther. 
Fig. 3. Vertical section of the same. 


Fig. 4. Transverse section of the anther. á 
Fig. 5. Vertical section of portion of anther with all the cells burst and pollen discharged. 


Fig. 6. Tissues of epithelium and pollen-grains. 
All very highly magnified. 


Tas. IL. 


Langsdorfia. 
a k. 
Fig. 1. Portion of plant of L. rubiginosa (Weddell), nat. size, from a drawing by Sir R. Schomburg 


Fig. 2. Female flower. 
Fig. 3. The same in a very young state. 


62 DR. J. D. HOOKER ON THE GENERA AND SPECIES 


Fig. 4. Rudimentary female flowers that occur on the male capitulum, 
Fig. 5. Unexpanded male flower with rudimentary female at its base. 
ig. 6. anded male flower. 
* 3 vr. section of the rhizome showing the five regularly disposed vascular bundles. 
Fig. 8. a. Tubular barred vessels that form the woody system; 5. thick-walled elongated cells that 
form the liber system. 

Fig. 9. Surface of rhizome with hairs. 

Fig. 10. Portion of a hair. 

Fig. 11. Rudimentary condition of a Langsdorfia on a rootlet, cut vertically. 

Fig. 12. Portion of a rhizome horizontally sliced, showing the intrusion of the root. 

Fig. 13. Swollen portion of a rhizome at the point of junction with a root, showing the rootlets of. 
the latter given off apparently by the rhizome, and partially enclosed in it. 

Fig. 14. Transverse section of the same, showing the abbreviated termination of the divisions of the 
root. 

Fig. 15, Apex of a rootlet enclosed within the cellular system of the parasite. 

Fig. 16. Vertical section of a portion of a rhizome corroding the bark of a rootlet which it has 
attacked. : 

Fig.17. Another vertical section of a root and rhizome, the former sending vascular prolongations 
into the latter. 

Fig. 18. Another section showing the first contact of a rhizome with a rootlet. 

Fig. 19. A portion of a rhizome enveloping the branch of a rootlet. 

Fig. 20. Ripe fruit of L. hypogea, Rich. 

Fig. 21. Vertical section of the same, showing the pendulous seed. 

Fig. 22. Seed removed, showing at a. the hilum. 

All but figs. 1, 12, 13 & 14, very highly magnified. 


Tas. III. 


Thonningia sanguinea, Vahl. 


Fig. 1. Female branch from VahPs herbarium, 

Fig. 2. Male capitulum from Schumacher’s herbarium. 

Fig. 3. Vertical section of the same. 

Fig. 4. Female capitulum from the same herbarium. 

Fig. 5. Vertical section of the same. 

Fig. 6. Imperfect male flower from Vahls specimen : 
perianth. 

Fig. 7. Male flower from fig. 2, —the anthers dehisced and partially separated at the apex. 

Fig. 8. Transverse section of the synema,—the anther-cells all dehisced. 

Fig. 9. Pollen-grains. 

Fig. 10. Female flower. 

Fig. 11. The same with the free tubular limb of the perianth laid open. 

Fig. 12. Another female flower with a more developed perianth. 


Fig. 13. diei of apex of perianth, showing the strong sclerogen-tubes that form its woody 
system. 


Fig. 14. Hair from the peduncle. 
Fig. 15. Cellular tissue and vessels of the perianth. 
All but figs. 1-5 highly magnified. - 


a. the bract at its base; 4. two scales of the 


Fig. 


1. 


. 2. 
. 3. 


4 
45, 
6 


OF BALANOPHOREÆ. 63 


Tan. IV: 


Balanophora involucrata, Hook. fil. 


Fully-formed individual, consisting of a simple tuberous rhizome, and one branch or peduncle 
with a hermaphrodite capitulum. 

Vertical section of the same. 

Transverse section of peduncle. 

Ditto of capitulum. 

Cellular tissue of surface of peduncle. 


. 6. Transverse, and fig. 7. vertical sections of cellular tissue at the circumference of peduncle, where - 


«8. 


= 5 
. 10: 


“LE. 
12. 
. 18. 
. 14. 
„15. 
. 16: 
SH 
“is 
19. 
. 20. 
» 21. 


. 23. 


. 24. 
. 25. 
. 26. 
27 
. 28. 


there is a little deposit of wax. 

Cellular tissue of young peduncle, the cells each with a globule of wax. 

A single cell detached, showing its dotted walls and wax-globule with its nucel. 
Another cell more advanced, with the wax-secreting cell burst and its contents discharged in 
the cavity of the cell. 
Wax-secreting cells at different stages of growth. 
Cellular tissue of rhizome. 
A single cell from the same. 
Transverse section of portion of peduncle and vascular bundle. 
Vertical section of the same, showing the vascular bundle to consist of simple delicate tubes. 
Cuticle of rhizome with empty bladdery cells that give it a granular appearance. 
Vertical section of a portion of the sheathing upper part of the rhizome. 
Transverse section of ditto. 
Transverse section of the rhizome, showing the symmetrical arrangement of the vascular system. 
Vertical section of the same. 
Transverse, and fig. 22. vertical sections parallel to the radius of a portion of the rhizome, 
showing the termination of the vascular system. Å 
Very highly magnified view of longitudinal section of a vascular cord and its surrounding 
tissues ; it consists of barred tubes, a, enclosed in a cellular tissue whose cells are filled with 
viscous fluid, 5: at c, some of the cellular tissue of the rhizome is seen containing wax-deposits. 
Barred vessels from 23 a. 
Cells surrounding the same with reticulated walls. : 
Imperfectly developed spirally marked tubes from the apices of the vascular bundles of rhizome. 
The same much more highly magnified. 
Another fully formed individual. 


All but figs. 1; 2, 3, 4 & 98, very highly magnified. 


Tas. V. 


Inflorescence of Balanophora involucrata, Hook. fil. 


Fig. 1. Female flowers intermixed with clavate bracts. 

Fig. 2. The same with sessile female flowers and pedicelled bracts. 
Figs. 3 & 4. Bracts. 

Fig. 5. Two adnate female flowers. 

Figs. 6, 7 & 8. Female flowers. 

Fig. 9. Ovary with base of style. 

Fig. 10. Transverse section of cavity of ovary. 


Fig. 11. Vertical section of ovary and ovule. 
Fig. 12. Ovule at earliest stage examined, supposed to consist 0 


f a simple embryo-sac. 


Sue 


A. Male plants of Balanophora involucrata 


64 DR. J. D. HOOKER ON THE GENERA AND SPECIES 


Fig. 13. Vertical section of more advanced ovary. 
Fig. 14. Immature seed. * 
Fig. 15. Apex of style, and pollen-grain ; 15 a, cells of apex of style before impregnation. 
Fig. 16. Base, & fig. 17. apex of style, with pollen-tube traversing it. 
Figs. 18 & 19. Male flowers with 2-lobed perianths. 
Figs. 20 & 21. Male flowers with 3-lobed perianths. 
Fig. 22. Imperfect male flowers showing their reduction to bracts, 
Fig. 23. Vertical section of male flower. 
Fig. 24. Pollen-grains. 
Fig. 25. Acarus found on the capitulum. 
All very highly magnified. 


Tas. VI. 


Balanophora involucrata, Hook. fil. 

Fig. 1. A young plant which has already caused the root upon which it grows to enlarge very much. 

Fig. 2. Vertical section of the same. 

Fig. 3. Very highly magnified view of the root and portion of the Balanophora, showing the mode of 
attachment, displacement of the woody system of the root, and apparent interlacement of the 
vascular systems of the root and parasite: a. indicates the bundles of the peduncle, 5. of the 
rhizome. 

Fig. 4. Very highly magnified slice of the root and parasite at the point of attachment of the latter. 


Fig. 5. Transverse section of the root from near the swollen portion, showing the separation of the bark 
and wood. 


Fig. 6. Woody system of the root. 


Fig. 7. Vertical section of an exceedingly young Balanophora, nidulating in the cellular bark of a root: 
a. spongioles ; 5. position of the nascent vascular bundle in its axis. 


Fig. 8. 


Vertical section of a more advanced, but still very young Balanophora, which has apparently ger- 
minated on one of the large tubers that this species eventually forms on the roots of maples, &c., 
and which tuber consists of a confused mass of the tissues of the parasite and root: a. spongioles; — 
b. nascent vascular bundles in its axis; c. root traversing the mass; d. cellular tissue of parasite. 


. Fig. 9. Vessels and cells of the vascular bundle taken from fig. 85. 


Fig. 10. Section at point of union of root, a, and parasite, b. 
Fig. 11. Vascular tissue of the wood of the root taken from fig. 8c. 
Fig. 12. Very young vascular tissue from the axis of fig. 7. 

All but figs. 1 & 2 very highly magnified. 


Tas. VII. 


4, var. gracilis, growing on the roots of an Oak. 1. male, and - 
2. female plant. : 


B. Male and female plants of B. involucrata, var. Catheartii. 
Fig. 1. Male flower magnified. 
Tas. VIII. 


B Balanophora fungosa, Forst., from N.E. Australia. 
Fig. 1. Plant of the natural size, but the flowers not fully developed. 

Fig. 2. Male flower. 

Fig. 3. Synema. 

Fig. 4. The same with the anthers dehisced. 


OF BALANOPHOREÆ. 


. Pollen-grains. 
. Female flowers on the pedicel of the bract. 
Female flower. 


= 
3 LI 
© Dr wc 


. Apex of style. 


- Transverse section of column of synema, showing four vascular bundles and anther-cells. 


65 


Fig. 10. Transverse section of rhizome from near the root, showing that the arrangement of the woody 
systems of the vascular branches that radiate outwards from the root in the mass of the rhizome 


is the same as that of the root, but dislocated. 


Fig. 11. The same from a point further removed from the root, showing a further dislocation of the 


woody system. 


Fig. 12. Very highly magnified vertical section of vascular bundle of rhizome, showing a. the cellular 
tissue of the Balanophora; b. the cellular tissue surrounding the woody system; c. the woody 


system. 
Fig. 13. Cellular tissue of circumference and cuticle of the rhizome. 


Fig. 14. Cellular and vascular tissue of the termination of the bundles. 


Fig. 15. Transverse section of root of the stock. 
Fig. 16. Woody tissue of the root of the stock. 
All highly magnified. 


Tas. IX. 
Lophophytum Weddellii, Hook. fil. 


Fig. 1. Portion of section of male inflorescence. 
Fig. 2. Ditto of female. 
Both magnified. 


Tas. X. 
Spherorhizon depressum, Hook. fil. 


Figs. 1-4. Individuals at various stages of growth. 
Fig. 5. Very young male flower. 
Fig. 6. Mature bud of male flower with articulate filaments. 
Fig. 7. The same with the perianth partly removed. * 
Fig. 8. 'Transverse section of anther-cells. ; 
Fig. 9. Ditto of one anther-cell. 
Fig. 10. Tissue of anther-cells. 
Fig. 11. Pollen-grains. 
Fig. 12. Very young female flower and articulated filament. 
Fig. 13. Mature female flower. 
Fig. 14. Vertical section of ditto. 
Figs. 15 & 16. Articulated filaments. 
All but 1-4 very highly magnified. 
Tap. XE 
Phyllocoryne Jamaicensis, Hook. fil. 
Figs. 1, 2. Male individuals in different stages of growth. 
Figs. 3, 4. Female individuals. 


Fig. 5. Articulated filaments ; 5a, apex of ditto. 
Fig. 6. Male flower. 


VOL: XXIL 


66 DR. J.D. HOOKER ON THE GENERA AND SPECIES 


Fig. 7. Male flower with portion of perianth removed. 
Fig. 8. Synema with anthers burst at their apices. 
Fig. 9. Transverse section of anthers. 
Fig. 10. Pollen-grains. 
Fig. 11. Female flower. 
Fig. 12. Longitudinal section of ditto. 
Fig. 13. Apex of style and stigma. 
Fig. 14. Transverse section of styie. 
Fig. 15. Fruit, in situ. 
Fig. 16. The same removed. 
Fig. 17. Vertical section of ditto. 
Fig. 18. Seed. 
Fig. 19. Grains of albumen. 
Fig. 20. Rudimentary flowers at base of male capitulum. 
Fig. 21. One of the same detached. 
Fig. 22. Another of the same, more developed. 
All but figs. 1-4 very highly magnified. 


Tas. XII. 


Rhopalocnemis phalloides, Jungh. 
Fig. 1. Male plant. 
Fig. 2. Female ditto. 
Fig. 3. Male flower and articulated filaments. 
Fig. 4. Transverse section of anthers. 
Fig. 5. Female flower and articulated filaments. 
All but figs. 1 & 2 highly magnified. 


Tas. XIII. 

Corynea crassa, Hook. fil. V 
Fig. 1. Plant of the natural size. 
Fig. 2. Very young plants attached to the branches of a root. 
Fig. 3. Bud of male flower. | 
Fig, 4. Expanded male flower. 
Fig. 5. The same far advanced, the pedicel of the synema having lengthened greatly. 
Fig. 6. Transverse section of anthers. 
Fig. 7. Pollen-grains. 
Fig. 8. Very young female flowers, showing the ovary to be 2-lobed at that age. 


Fig. 9. Articulated filament and female flower before the falling away of the bracts, showing the position 
of the styles. 


Fig. 10. Mature female flower. 
Fig. 11. Vertical section of nearly ripe fruit exposing the seed and embryo*. 
Fig. 12. Section of ripe seed and embryo. 
Fig. 13. Embryo removed from the seed. 
Fig. 14. Apex of style and stigma. 
. Fig. 15. Transverse section of style. 
Fig. 16. Young articulated filament, its cells still nucleated. 


* See foot-note at p: 54, 


OF BALANOPHOREÆ. 


Fig. 17. Cells from fig. 16, showing the cell-contents. 

Fig. 18. Fully formed articulated filaments after the fall of the scales of the capitulum. 

Fig. 19. The same, showing the appearance of the terminal cells. 

Fig. 20. Old terminal cells of the articulated filaments, showing their wrinkled walls. 
All but figs. 1 & 2 very highly magnified. 


b. | 


Tas. XIV. 
Corynea spherica, Hook. fil. 


Fig. 
Fig. 


o 


Fig. 


Mature, and fig. 2. Immature plants. 
Vertical section of fig. 2. 

. Peltate bract. 

Fig. 5. Immature male flower and articulated filaments. 
Fig. 6. Ditto more advanced. 

Fig. 7. Ditto expanded. 

Fig. 8. Transverse section of anthers. 

Fig. 9. Articulated filaments. 

Figs. 10 & 11. Mature female flowers. 

Fig. 12. Vertical section of fruit. 

Fig. 13. Seed. 


67 


Fig. 14. Section of portion of rhizome showing the two portions of the root to be completely severed in 


the axis of the rhizome. 
All but figs. 1, 2 & 3, highly magnified, 


Tas. XV. 


Helosis Mexicana, Liebm. 
Fig. 1. Plant of the natural size. 
. Vertical section of capitulum and upper part of peduncle. 
. Very young peduncle on a branch of the rhizome. 
. Vertical section of the same magnified. 


. Fully developed bract. 
. Vertical section of ditto. 
Fig. 8. Nascent Helosis on a rootlet. ; : 
Fig. 9. Vertical section of a very young Helosis which has fully established itself on a nid : 
Fig. 10. Vertical section of a full-grown tuber of the rhizome, showing its attachment to the root. 
Fig. 11. Similar section of another and larger one. 
Fig. 12. Transverse section of the peduncle, showin 
Fig. 13. Vascular tissue of the same. eot in 
Fig. 14. Transverse section of the rhizome, showing—a. 

or pith; 5. wedges of vascular tissue; c. liber ; d. detached masses - A T 
Fig. 15. Cells of liber. 
All but figs. 1, 2, 3, 5 & 10, very highly magnified. 


2 

3 

4 en P 
Fig. 5. Portion of capitulum vertically cut, showing the imbrication of the very young bracts. — 

6 

7 


g eight regularly placed vascular radiating bundles. 


edullary sheath surrounding the axis 


68 DR. J. D. HOOKER ON THE GENERA AND SPECIES OF BALANOPHOREÆ. 


Tas. XVI. 


Helosis Mexicana, Liebm. 
Fig. 1. Bud of male flower. 


Fig. 2. More advanced male flower. 
Fig. 3. Fully formed male flower with the lobes of the perianth spreading. 
Fig. 4. Young synema. 
Fig. 5. Fully formed synema. 
Fig. 6. Pollen-grains. These are probably incorrectly represented; the appearance of pollen-tubes may 
be due to the presence of the mycelium of a fungus: these and the pollen-grains were black. 
Fig. 7. Tube of the male perianth laid open, showing the conical rudiment of an ovarium at its base. ; 
Figs. 8, 9 & 10. Very young state of female flowers with articulated filaments, showing the compound. 
nature of the ovarium. ; 
Fig. 11. Fully formed but immature female flower, with the limb of the perianth laid open. à 
Fig. 12. Another female flower of a different form. 
Fig. 13. Vertical section of young female flower showing the albumen of the seed to be already formed, 
apparently before fecundation has taken place. 
Fig. 14. Mature female flower. , 
Fig. 15. Vertical section of the same, showing the pendulous ovule. 
Fig. 16. Apex of style and stigma. 
Fig. 17. Vertical section of nearly ripe fruit. 
Fig. 18. Transverse section of ditto with the seed removed. 
Fig. 19. Seed. 
Fig. 20. Transverse section of ditto. 
Figs. 21 & 22. Articulated filaments, showing the bladdery 
Fig. 23. Apex of the same more highly magnified. 
Fig. 24. Apex of another articulated filament, 
Fig. 25. Transverse section of portion of rhizome: a. vascular 
' wedge; d. liber; e. sclerogen-cells. - 
Fig. 26. Vertical sections of tissues forming the vascular system from the axis to the circumference: på 
a. pleurenchyma that occupies the axis; 0. medullary sheath; c. vascular wedge of wood - 
formed of angular thin-walled vessels; d. liber-cells; e. cellular tissue. | : 
Fig. 27. More highly magnified vessels of axis or pith. 
Fig. 28. Ditto of wood. 


Fig. 29. Transverse section of vascular system ;—the letters refer to the same tissues as in fig. 26. 


Fig. 30. Section of stem of Helosis Guyanensis from Brazil (Mr. Miers), showing the supplementary 


membrane enclosing the uppermost cells. 


axis; d. medullary sheath; c. vascular | 


liber-bundles alternating with those first formed. 
Al very highly magnified. 


På 
3 


LEP D), ay 
- 
m ie 


FR) 
ug 


titi, 


X 


Langsdorffia rubiginosa | 
en. g 


Brooks i 


Vincent 


Os À 
Samui ER 


NN | 


ANS [ É 
NUE 


LA 


phora. involucrata; Hf 


Dalano 


Re 


286 a a NR 


PRE : 
" 


SN iii He 


u. 
U 


Vincent Brooke dm. 


Balanophora involucrata Hf 


ORT Ty 

yl? 

NU 
SN US 


TUTTI 
HELLE 


Ji Hd HE 


Trans Linn. Soc. Vor XXILP 
Plate VI. 


i eh 2 ia 


200 
BOOT 

ROOTED: 
M 


Ned 


Balanophor | fangosa , Fors 


[rans Linn Soc Vor AXIN 


T F 
Plate IX 


ncent Brooks Imp 


Lophophytum Weddeln, HA 


ae ren iet 


à N 
e NE 
Ar PIPE 


SLR tak m» e 


* 


Junge 


] 


aa 
Fe) 


illoide 


phe 


RNS RAI owe 


e Seien TET 


w 


Vincent Brooks imp 


FNS 
DAN NS S 
X) NS 


s 
SS 
o DR 


Sat 


IB 


CX 


Å 


t — 


je t | 
ant mh, 


Mars 


so? Li 
Have 


ON 
4 


À 
i 
$^ 


CR, 


e. 
per^ 


+ 


SN LL 
TT] v EE ny 
GITT TT TITTEN 


E ORE 
: v OPEP GK 
ERBE DICH Denn 


bolt ae er 


CT 


II 

OG ER > 
^ 

Bergs 


y 
qi 


Meudon Lim (Hos 1-29) Hguyanensis, ich (fg. 30 


[3 


IL. On the Development of the Ovule of Santalum album; with some Remarks oh the 
Phenomena of Impregnation in Plants generally. By ARTHUR sa 
F.R.S., F.L.S., Prof. of Botany in King's College, London. 


Read March 4th, 1856. 


IN the 21st volume of this Society’s “Transactions” are published the results of some of 
my earliest observations on the phenomena of fertilization and development of embryos 
in flowering plants. To myself, these were at that time conclusive, and they were in 
accordance with those published shortly before by Amici, von Mohl, and Miller, sub- 
sequently confirmed, with far more extensive illustration, by Hofmeister and Tulasne. In 
spite of all the adverse evidence, however, Schleiden still continues to maintain the doc- 
trines he formerly propounded, and during the last few years he has found a most active 
supporter in Schacht. : 

Those who have not directed their labours to the practical examination of these delicate 
points, may feel surprise at the discrepancy existing between the statements of different 
observers. "Those who have been engaged in the dissection of ovules in the earlier stages 
of development, will not much wonder at doubts arising in my mind whenever I have 
met with statements directly contradicting those made in my former paper. The number 
of instances in which a satisfactory observation is made, is very small in comparison with 
the number of specimens examined; and notwithstanding that I have year after year 
devoted much time to the dissection of ovules, these time-devouring researches have 
yielded me a far smaller number of facts than I could wish. 

Å certain amount of discrepancy exists even between the statements of the disciples of 
Amici, in regard to the origin of the germinal vesicle (Keimbläschen) ; and to whether it 
exists before, or is formed after fecundation. Hofmeister* says before; Tulasnet says 
he never could find it anterior to the fertilization. The latter, in the memoir just referred 
to (page 115), adds, * This delicate question no longer (1849) perhaps possesses all the 
interest which was accorded to it by MM. Mirbel and Brongniart, and eF recently by 
Mr. A. Henfrey, in his * Report on the Progress of Physiological Botany f It is true, 
the existence of the embryonary vesicle at a period anterior to the arrival of the pollen- 
tube would, if placed beyond doubt, prove invincibly that this vesicle could not owe its 
origin to the latter organ; even now that the error of the pollinists on — Boe 
tain, the question seems to me worthy of the attention of botanists, especially on accoun 


of the theoretical consequences involved in its solution." : Re 
Confidently as Tulasne expressed himself as to the origin of the germinal vesicle inde- 


: i . 9"* ser, xii. p. 114. 
* Entstehung der Embryo. Leipsie, 1849. fn du Dámon sat, Svr e p 


t Ann. Nat. History, ser. 2. i. p. 49 (1848). 


70 PROFESSOR HENFREY ON THE DEVELOPMENT OF THE OVULE 


pendently of the apex of the pollen-tube, this very point is most warmly contested by 
Schacht*, and notwithstanding that I have tolerably positive opinions derived from å | 
certain number of cases, where the end of the pollen-tube and the germinal vesicle (the 
summit of the suspensor) were seen together, but distinct, in one preparation, the objects 
are so delicate, and the cause producing obscurity of the preparation so difficult to guard 
against, that I cannot but think the demonstration of the pre-existence of the germinal 
vesicle in the embryo-sac, the most important fact that can be brought forward in oppo- 
sition to the views of Schleiden. As remarked by Tulasne, it has a great philosophical 
importance in reference to speculations as to the source of the vitality of the new being; 
and, as will be shown below, it is of no less importance for the establishment of the rela- 
tions of the processes of embryogeny in the various classes of plants, and of the analogies 
which these present to phænomena attending the reproduction of animals, — 4 
As in my former paper I selected one well-established series of observations for the 
support of the doctrine I advocated, passing over without notice numerous fragmentary — 
researches which, although corroborative, had in themselves nothing absolutely decisive, : 
I shall here confine myself to the course of development of the ovule of one plant, in | 
which the complete series of observations have been repeated many times. I feel the less : 
hesitation in laying before the Society a paper thus restricted, from the circumstance that — 
the example brought forward possesses features of great interest in other respects besides » 
the main point of the fertilization, The principal facts are not indeed now brought for — 
ward for the first time, even before this Society, but they are such as few botanists have 
the opportunity of testing for themselves, and hence may be admitted as supplementary, 
partly confirmatory, partly emendatory, of the memoirs on the same subject by the late | 
William Griffith f. pr 
Å melancholy interest is attached to the investigations now presented; for the mate- 
rials which have enabled me to repeat and control the observations of Griffith, were 
furnished by my lamented friend Dr. Stocks, another Fellow of this Society, suddenly cut 
off from us in the midst of active labours for the advancement of Indian botany. — 
A bottle containing a large quantity of blossoms and fruits of Santalum album, m 


have acquired a firmer consistence than I have usually found in fresh objects. nr 
_ When very young buds are examined in vertical sections (Tab. XVII. fig. 1) the ov 
Is found superior, forming a conical body arising in the centre of the flower; as 
advances in age, the adherent tube of the calyx and the side-walls of the ovary. 
rapidly (Tab. XVII. figs. 3-5), so as at length to render the ovary altogether m 

(Tab. XVIL fig. 14); the original conical summit (continued into the style) b 
gradually flattened by the lateral elevation (Tab. XVIII. figs. 22, 27, 28). 
The Ovary never exhibits any open cavity; the centre is occupied by an e 
* Flora, 1855, p. 145; Botanische Zeitung, 1855, p. 641. ; 

t Linn. Trans. xviii. pp. 59 & 71; xix. pp. 171 & 487. 


OF SANTALUM ALBUM. 71 


conical, or rather spindle-shaped pillar, the free placenta of authors, arising from the 
centre of the base; the internal surface of the walls of the ovary are closely applied to 
the placenta, without however contracting any adherence either at the sides or above. 
In the youngest buds I examined (Tab. XVII. fig. 1) I found the central placenta with 
three cellular papillæ projecting downwards from the thickest portion, near the base, that 
part of the placenta below their origin being narrowed into a kind of peduncle (Tab. XVII. 
fig. 2). These papillæ, of cylindrical or slightly conical form, with blunt ends, pointed 
downward and a little outwards; they were composed of tolerably compact cellular tissue, 
and according to the view commonly taken of the structure of this genus, they are the 
nuclei of three ovules (fig. 2. a,a). In the youngest specimens examined, no trace of an 
embryo-sac could be seen, but a lighter streak (as seen by transmitted light) soon appears 
in the axis of each nucleus, indicating a cavity, and in a little farther advanced state a 
closed tubular process was found projecting from the point of each nucleus, the free 
extremity of a tubular embryo-sac extending internally up to the organic base of the 
nucleus. . Examined in successive stages, the tubular embryo-sac is seen to extend down- 
wards at first ; then its external (or anterior) end (organic summit) turns outwards, next 
suddenly upwards over the side of the nucleus; it becomes then more and more elongated 
until it has grown up upon the side of the placenta so far as nearly to reach its free sum- 
mit. It lies upon the outside of the placenta in the form of a slender filamentous tube 
(Tab. XVII. fig. 4, 5). At the same time the posterior or internal end of the tube elon- 
gates in the substance of the placenta until it makes its way nearly to the apex. These 
phænomena occur in all the other (occasionally four) ovules, alike. The tubular embryo- 
sacs contain protoplasmic matter with a few granules, but are comparatively clear as 
contrasted with the cells of the nucleus, which are filled with dense protoplasm and 
(starch-) granules. 
. The next stage is the formation of a bulbous enlargement of the embryo-sac at the 
point where it turns up to rise over the placenta (Tab. XVII. fig. 6). The free points of 
the tubular sacs also contract a slight adherence to the side of the placenta near the sum- 
mit, remaining free however in the greater part of their length, although closely applied 
to the side of the column. The protoplasmic pre ær to increase in quantity 
and consistence, especially at the apex and below the bulb. A septum is soon 
formed at the bot of the bulb, ital at first flat, becomes convex (looking upwards), 
and finally appears as an inner bulb projecting up into the cavity of the primary bulb 
(Tab. XVII. fig. 7). The septum divides the embryo-sac into two distinct portions ; I ar 
never discovered any cell-formation within the cavity behind (or below) the septum, either 
externally or internally to the nucleus. The only change this posterior (or oo cape Å 
inferior) part of the embryo-sac undergoes, is a tubular ramification in the su en 
the placenta, to be described presently. get n ae | 
About the time of the formation of the septum just nere - di ceva: 


period of the opening of the flower, the apex of the embryo-sae 
(clavate), and protoplasm accumulates on it. Soon after this, before any pollen-tubes 


i ular cell-nucleus, becoming gradually better 
reach the placenta, from the stigma, a granula of the clavate end, not quite at the extre- 


defined, makes its appearance in the protop 


72 PROFESSOR HENFREY ON THE DEVELOPMENT OF THE OVULE 


mity (Tab. XVII. figs. 8 &9,e). A portion of the protoplasm in the absolute extremity 
(adherent to the placenta) collects into two granular masses, which become much darker. 
coloured than the surrounding substance, and apparently almost solid, while the closed 
end of the embryo-sac becomes moulded as it were on these so as to present a kind of 
notch or depression between them (figs. 8 & 9, d, d). They lie nearly in contact, occupying 
(like a plug) the summit of the embryo-sac; the nucleus before mentioned being quite 
below them. At this period the nucleus is devoid of a cell-membrane. 

The summits of embryo-sacs, examined soon after the above period, present the ends of 
one or more tubes adherent to them; these tubes extend down to the embryo-sacs from 
the summit of the placental column; I have never traced them up the style, for this 
structure was so hard and resisting in my preserved specimens as not to allow of my 
examining the canal minutely; but there can be no doubt as to their nature. They are 
the ends of pollen-tubes. Usually one applies itself upon the very apex of the embryo. 
sac (Tab. XVII. figs. 10 & 11 J). The tubes appear to creep down between the papillose 
projections of the cells of the surface of the placenta, being moulded in some degree on 
them; and these grooves appear to conduct the tubes to the points of the embryo-saes, 
which themselves adhere to the superficial cells of the placenta; sometimes so firmly as 
to carry away fragments of their walls when dissected out free (Tab. XVII. figs. 9-11). 

I have directed my utmost efforts to the accurate observation of the ends of the 
embryo-sacs with the pollen-tubes adherent, They are tolerably easily extracted free 
from the ovary, with needles under a low doublet; I have examined at least five-and- | 


_ tions, have applied every means to make the structures clear; mounting in water and then 
in glycerine, between very thin glass, so as to observe both sides; boiling in nitric acid; 
treating with dilute sulphurie acid alone, and with this and solution of iodine; examining 
the objects with a4 and } object-glass under the compound microscope, with and without 


embryo-sac, in the situation of the ‘notch’ above mentioned (figs. 10 & 11). But I di 
incline to believe that a phænomenon analogous to conjugation takes place. For asl —— 
have said, the adhesion is intimate, but the nucleus before spoken of (Tab. XVIL 
figs. 8 & 9e) lies away from the pollen-tube, separated from it by the two coagula (d, d); 


the fissure, however, between these leads exactly from the end of the pollen-tube to the de 
nucleus (figs. 10 & 11). Moreover, very soon after the pollen-tube becomes adherent, the ee 
nucleus acquires a Proper coat of cell-membrane,—becomes a real cell, the germinal — 


vesicle, from which the Suspensor is developed. This cell is slightly pyriform, with an —— 


obtuse projection directed toward the fissure between the coagula (Tab. XVII. fig. 119)» — 


I think that the pollen-tube, after becoming adherent to the summit of the embry0-S26, 
contents pass into the embryo-sac, reach the nucleus, and 


OF SANTALUM ALBUM. 73 


determine its conversion into a cell. The nucleus becomes perfectly defined, as separate 
from the surrounding protoplasm, just at the period of fertilization. At the same time 
this surrounding protoplasm (contained in a * primordial utricle * lining the entire embryo- 
sac, down to the septum of the bulb (Tab. XVII. fig. 6)), becomes more dense and granular, 
sometimes exhibiting largish starch-granules. 

Little further change occurs at the apex of the embyro-sac for some time; merely the 
cell-membrane enclosing the nucleus becomes thicker, and the cells assume å more elliptical 
form. The next step is the formation of endosperm-cells from the protoplasm of the 
embryo-sac. This takes place by segmentation, or free cell-formation from the whole 
abundant mucilaginous protoplasm, commencing always in the bulb, often advancing 
from them before it proceeds up the tubular part of the embryo-sac, which at first presents 
only a single row of ‘primordial utricles’ (Tab. XVII. fig. 12); the latter then divide 
perpendicularly (Tab. XVII. fig. 13), and the cells produced, appear to apply themselves 
to the side-walls and multiply for some time by free cell-formation in the centre 
(Tab. XVII. fig.15). Ultimately cell-division occurs, and the whole cavity of the swollen 
embryo-sac being filled up with cellular tissue analogous in character to that of the 
nucleus,—namely, composed of squarish cells with thin walls, filled with dense protoplasm, 
containing increasing quantities of minute starch-granules,—the albumen of the end 
becomes a distinct structure, on the surface of which all trace of the originally bounding 
embryo-sac is soon lost (Tab. XVII. fig. 16). 

The first change in the germinal vesicle is its elongation downwards into a cylindrical 
form; then cross septa appear (Tab. XVII. fig. 15 g, Tab. XVIII. fig. 19 g) one after an- 
other, so that it is converted into a short row of cells. The uppermost remains appressed 
to the coagula in the apex of the embryo-sac, and does not appear to become developed 
further ; forming a kind of suspensor. The lower cells multiply greatly (Tab. XV ITI. fig. 23) 
and form an elongated, clavate cellular body, the embryo, in which at first no trace of 
regions can be detected,—only a greater density of the tissue and abundance of granular 
contents at the cotyledonary (inferior) extremity (Tab. XVIII. figs. 22 & 24 g, fig. 25). 

The conditions of the walls of the ovary during these changes deserve some attention; 
the outer substance of the young fruit is formed of a firm layer of tolerably equal thick- 
ness all over, constituting what we may call the epicarp together with the mesocarp 
(Tab, XVIII. figs. 27 & 288). These define and correspond to the outward form of ". 
fruit. The portion immediately beneath the epidermis is composed of oblong d pe 
their longest dimension in the direction of the axis of the fruit; these pass insensibly In 
a denser layer of closely-packed polygonal cells with thick walls. These ea pen ires 
coloured, while the endocarp is nearly colourless (Tab. XVIII. figs. 27 = ) sone 
of the mesocarp (well defined internally to the naked eye) appear, under the microscope, 


å å nice .27 & 28 r), composed of membranous, 
to pass again insensibly into the soft endocarp pico je hate ihe siget; 


globular, and polygonal cells, loosely packed, of m 

than those of the mesocarp. The endocarp (figs. 27 & 287) - veste et 
the dark rind of the fruit (s) and the placenta (and ovules) (0, p), å Å li, i6 is of 
expansion of the ovary. Where in contact with the 2... se. dee A perpendi- 
denser texture and of darker colour, having a definite, separa i i 

VOL, XXII. 


74 PROFESSOR HENFREY ON THE DEVELOPMENT OF THE OVULE 


cular line is seen leading from the apex of the placenta to the point corresponding to the 
base of the style; the cells of the endocarp exhibit a radiating linear arrangement on the 
upper half, the lines running upwards and outwards from the placenta (fig. 27). 

Å remarkable phænomenon occurs meanwhile in the interior of the placenta. It has 
been stated that there is never any appearance of development of cells in that part of the 
embryo-sac, behind (or below) the septum formed a little way outside the nucleus 
Tab. XVII. figs. 7-12). The inner bulbous expansion, formed by the septum itself, which 
becomes convex, and protruded upwards into the centre of the larger bulb, is found in the 
same condition, as long as the structures are traceable, and it finally forms a kind of stalk 
or “funiculus” to the seed, inserted into the substance of the albumen (Tab. XVII. 
figs. 12, 17, and Tab. X VIII. fig. 26), and connecting this with the remains of the placental 
structure. I have already mentioned, that while the tubular prolongation of the embryo- 
sac, outside the nucleus, is growing up over the placenta, the posterior end, inside the 
nucleus, also grows up, breaking down the tissue before it, into the substance of the placenta 
(Tab. XVII. fig. 4. Within this organ it proceeds nearly to the summit, and then turns 
round somewhat suddenly, and grows down again, with various ramifications, in the 
centre of the placenta, and even into the receptacle below where these arise (Tab. XVIII. 
figs. 20, 21): I have never seen anything like cell-formation, or even production of septa 
in these posterior branches of the embryo-sac; they contain a granular protoplasmic 
substance, which, in my preserved specimens, is of a red colour. The ends of the barren 
embryo-sacs undergo the same kind of development within the placenta, although the 
changes in the external portion cease at the period of the formation of the germinal 
yesicle. The ramifications of the three distinct embryo-sacs become somewhat inter- 
laced, but I have never certainly detected any conjunction or adhesion of them, as 
suspected by Griffith. 

It remains only to notice the further changes exhibited by the ovule. The endosperm 
or albumen of the embryo-sac increases enormously in quantity, so as to expand the sac 
in all directions (Tab. XVIII. figs. 16, 20, 26); the placenta is broken off just below the 
point of origin of the ovules (fig. 26), and pushed outwards and upwards by the enlarge- 
ment of the albumen (Tab. XVIII. figs. 27, 28 p); in the ripe fruit it is found lying upon 
the surface of the latter (Tab. XVIII. figs. 30 & 82 p), which, through the displacement 


and destruction of the endocarp, is finally in immediate contact with the woody mesocarp. - 


On examining the remnant of the placenta, even in the ripe fruit, it is found to be connected — 
with the endosperm by the bulb. When the placenta is then broken away from the 
receptacle, and pushed up, the receptacle upon which it was seated is also carried away 
towards the same side of the albumen, on the outside of which, near the base, it is 
ultimately found (Tab. XVIII. figs. 28, 30, 32 v) as a little mass of sphacelated tissue 
overlying a pit or foramen, which leads to an internal dark line, running through the — 
endosperm to the cotyledonary extremity of the embryo (Tab. XVIII. fig. 300). In he 
line, and in the sphacelated mass, are found remains of the posterior ramified processes 0" — 
the embryo-sac, some of which are also found attached to the upper fragment or placenta. — 
In the ripe seed, the embryo is found lying a little out of the axis of the albumen, na — 

i * This seems to resemble what Mr. Bentham describes in Olacaceæ, Linn. Trans. xviii, p- 675- 


OF SANTALUM ALBUM. 75 


vertical position (fig. 33), or with the cotyledons curved a little towards the lateral 
basilar scar just mentioned (fig. 35). The radicle is at the upper end of the seed and 
terminates in an acute apiculus (Tab. XVIII. figs. 34 & 36). 

In the mature fruit the mesocarp forms a hard shell, outside which the epicarp forms a 
thin layer of pulpy substance (Tab. XVIII. fig. 22). The woody shell is slightly pointed 
and trigonal above, presenting three converging ridges (Tab. XVIII. fig. 81) when the 
epicarp is removed. Within the woody mesocarp the albumen or endosperm of the 
seed lies free; the coat formed by the embryo-sac is no longer distinguishable, but the 
endosperm is covered with brownish membranous scales (Tab. XVIII. fig. 82), composed 
of compressed withered fragments of the lax cellular tissue of the obliterated parenchy- 
matous endocarp. 

The above observations confirm in almost every respect those published by Griffith in the 
Transactions of the Linnean Society, the main point of difference lying in the statements 
made with respect to the phenomena presented at the summit of the embryo-sac at the 
time of fertilization; in which my account is strongly opposed to those given by that 
author, not only in Santalum, but in Osyris and the Loranthacee. Notwithstanding 
the high value I attribute to Griffith’s labours, increased and confirmed by the researches 
now brought forward, I feel very confident of the correctness of the account I have given 
of the origin of the embryo from a pre-existing germ, and I have little doubt that the 
process of fecundation is such as I have described in Osyris, and the other cases. The 
importance attaching to the truth of the view I have propounded will be farther illustrated 
below. 

Every one who has studied the development of the ovules of Santalum, and the allied 
genera, has been struck by the remarkable anomalies which present themselves, The 
entire protrusion of all the (apparently) essential part of the embryo-sac from the apex 
of the nucleus, the development of the endosperm in the external compartment of the 
sac, altogether independently of the nucleus, are very remarkable; while the posterior 
development of the embryo-sac is no less singular. The idea has been suggested that the 
entire central body here described as a free placenta with the ovules reduced to nuclei, 
might be one ovule with three embryo-sacs; and also that the central body is all placenta, 
the nuclei being merely the funiculi of ovules reduced to embryo-sacs. There does not 
seem to be sufficient ground for either of these assumptions, although the former is in 
some respects plausible. : : 

The i reason which could be advanced in favour of the second idea, is the nn 
rently abnormal position of the embryo, the coty. ledonary extremity of nee "rs m 
apex of the nucleus, and the radicle end pointing in the direction caeli a n 
nucleus. But this is rather an apparent than a real irregularity. It wo - a senum 
insistence upon terms to call that end of the embryo-sac engaged "M e ede ifr. 
nucleus, the micropyle-end; it is really the middle, and the erhbryo-sie is in ampy 

: : ine in the interior of the nucleus, while the 
lotropous, its organic base or chalazal end being m ed up so as to lie (out- 
micropyle-end is prolonged out beyond the micropyle, and turned up 


side) against the chalazal end. e i 
In regard to the first of the two views above referred to, the only | which 


76 PROFESSOR HENFREY ON THE DEVELOPMENT OF THE OVULE 


it appears to me can be urged in favour of the idea that the central body is not a placenta, 
but a compound nucleus with three embryo-sacs, are as follows d : 

The central body is stated by Griffith to form at first, in Osyris, but a slightly -elevated 
cone with three (or four) papillæ (or nuclei) at its sides. In the youngest specimens of 
Santalum I examined, the placenta was highly developed as a pillar in the centre of the 
ovary (Tab. XVII. figs. 1 & 2). The papillæ might be either three points of one ovule, 
or three nuclei with three chalazal ends blended in the centre, and the growing-up of the 
central column, which goes on for a certain time after the embryo-sacs are distinguish- 
able, might be regarded as a conversion of the compound ovule, or the three conjoined 
ovules, into an anatropous structure, the chalazal end being at the apex of the conical 
mass, the micropyles free and turned down next the “ funiculus.” The subsequent retro- 
version of the embryo-saes would still be anomalous, connected however in some degree, 
through Osyris, Avicennia* and Myzodendron+ (in which the embryo-sac is not extended 
unless fertilized) with the ordinary conditions. Then the growth of the posterior ends of 
the embryo-sacs would seem to represent a kind of chalazal structure, connecting them 
below in a kind of central raphe with the peduncular placenta. The relative positions of 
the remains of the nuclei and the central body, and of the remains of the peduncle of the 
placenta, in the ripe fruit, would then indicate a rupture at the umbilicus, which was 
carried up to near the top of one side of the seed, while the funiculus was driven outwards 
on the same side, and almost obliterated f. 

Some degree of likelihood attaches to the explanation just given when the Santalacee 
are compared with the Loranthacee, with which they appear to be nearly connected 
through Myzodendron (which genus is nearer the former than the latter family as regards 
ovulary structure$). In the Loranthaceæ the supposed compound, triple ovule, being 
atropous instead of anatropous, would naturally be still more completely combined into 
one piece; it would here be almost a question of words whether there were confluent 
nuclei or one nucleus with three parallel embryo-sacs. Then the phenomena observed in 
Loranthus might afford a still farther confirmation, as indicating analogy with the Coniferæ, 
in which the existence of a number of (secondary) embryo-sacs is the rule. 

These points deserve further attention, and can only be elucidated by the study of the 
development in more of the genera of these remarkable families. I should not omit to 
mention here the observations of DeCaisne on Thesium ||, which I have not yet repeated, 


but which agree essentially with the statements respecting Santalum made by Griffith 
and myself. 


The reason which perhaps most of all induced me to present these observations to the 


—- of the Society, is the remarkable analogy which is shown to exist between the 
. phenomena of fecundation above described, and those which have recently been demon- 


isti ished in the conical plac > aan, ine to Griffith, 
(I. e.) the vascular cord eme mecal placenta ; in Avicennia, according to 
reaches nearly to the base of the embryo-sac, so that the chalazal end of the ovule is marked. 


OF SANTALUM ALBUM. 77 


strated to take place in some of the lower plants. I pass over the numerous discoveries 
that have been made of late years in the higher Cryptogamia, from the Marsileaceæ to 
the Liverworts, showing the pre-existence of a germ, and its fertilization by spermatozoids. 
I may refer to my own publications on this subject elsewhere*. The cases most imme- 
diately interesting to me in this instance are those described by Thuret in the Fucaceæ, 
already repeated by Pringsheim, and the researches of the latter and of Cohn on certain 
filamentous Confervoids. According to the elaborate investigations of Thuret+ the 
spores of Fucus are discharged from the Spore-sacs as globules of protoplasmic substance, 
bounded by the structure denominated by Von Mohl the primordial utricle, without a 
cellulose coat. While swimming free in the water, spermatozoids come in contact with 
them in large numbers, and after a certain time a cellulose coat is developed upon the 
surface of the spore; the latter thus becomes encysted, and forms a true cell, which then 
germinates, to produce a new plant. Pringsheimt describes essentially analogous pheno- 
mena as occurring (inside the parent-cells) in Vaucheria, and Cohn’s$ account of the 
fecundation of the spores of Sphæroplea also agrees with these. 

These facts, together with those I have brought forward in this paper, tend to prove 
that the process of impregnation in plants consists in the absolute admixture of the pro- 
toplasmic substance of two cells (*male” and * female"); of which the female (or 
germinal) substance or body always pre-exists in the form of a nucleus or * protoplast,” 
while the male (or spermatic) substance exists in the form of a granulose fluid. In the 
Flowering Plants the spermatic fluid is conveyed directly into the embryo-sac by the 
channel of the pollen-tube; a similar process appears to exist in the conjugation of the 
lower Algæ; in other cases the spermatic fluid is conveyed from organs situated at a 
distance from the parent-cell of the germinal vesicle, by the agency of locomotive struc- 
tures (spermatozoids) developed in the spermatic cells, bathed in and discharged with 
their contents, and themselves composed of the nitrogenous protoplasmic matter of cell- 
contents. 

In my Memoir on Orchis Morio||, I described the nascent germinal vesicles as cells. 
Hofmeister and others in like manner call them cells; but comparison of my older draw- 
ings and those of Hofmeister with new observations, leads me to believe that, on careful 
examination, these bodies will be found to consist of nuclei or “ protoplasts” before fertili- 
zation. I may note in reference to this, that I have already some confirmation from 
another case besides Santalum, and I trust to bring forward hereafter more complete 


evidence on the subject. 
Jan. 30, 1856. 


; : ay tions, 
* Report of the British Association, 1851; Annals of Natural History, 2 ser. ix. p. 441 ; Linnean Transactions 
xxi. p.117. 
| ; . March 1855. 
+ Ann. des Sciences nat. 4 * ser. ii. p. 197. t Een er en å 
$ Bericht Berlin Akad. May 1855. : | . Trans. xxi. p. 7. ES 
: for inqui 
Y Certain circumstances which I observed in the archegonia of the F ue uni onis gn Rd: Rr de 
‘whether the parent-cell of the germinal vesicle is not open at a certam period eide ser rd dé the Devo- 
Spermatozoids to a naked nucleus. See especially the figures 56, 57, 63-70 of plate = r 
lopment of Ferns (Linn. Trans. xxi. p. 117). 


78 PROFESSOR HENFREY ON THE DEVELOPMENT OF THE OVULE 


EXPLANATION OF THE PLATES. 
The figures maguified 50 diameters and upward were drawn by the aid of the camera lucida. 


Tas. XVII. 


Fig. 1. Vertical section of the ovary of an unopened bud. 

Fig. 2. Placenta with two nuclei (a a) from the same. 

Fig. 3. Vertical section of the ovary, &c., of a flower just opened. 

Fig. 4. Placenta of the same with two nuclei (a, a) and one embryo-sac (b). 

Fig. 5. Section of an ovary somewhat older. 

Fig. 6. Nucleus (a), embryo-sac (ö) with germinal vesicle and pollen-tubes at the apex; from fig. 5. 
septum of the bulb not yet developed. 

Fig. 7. Point of a nucleus and the embryo-sac (b), with a fully-developed septum in the bulb (c). 

Fig. 8. Apex and unimpregnated embryo-sac, with terminal coagula (dd) and a nucleus (e). 

Fig. 9. Another more advanced. 

Fig. 10. Another with a pollen-tube (f) applied. : 

Fig. 11. Another when the nucleus (e) is converted into a cell with a cellulose wall (g). 

Fig. 12. Part of an impregnated embryo-sac (6) attached to its nucleus (a), with endosperm-cells formed 
in it. 

Fig. 13. Upper end of another one with the endosperm more developed. 

Fig. 14. Vertical section of an ovary some time after impregnation, 

Fig. 15. Upper end of the endosperm of an ovule from fig. 14, with the germinal vesicle (g) undergoing 
division. 

Fig. 16. A less advanced ovule, with the endosperm considerably developed; the dotted line runs to the 

- point of insertion of the inner bulb (fig. 17) formed by the septum. 


Fig. 17. The inner bulb (similar to that in figs. 7 & 12) drawn out from the endosperm of fig. 16; a, 
nucleus. : 


Fig. 18. Inner bulb drawn out from another endosperm. 


Tas. XVIII. 


å à Apex of a young ovule with the embryo (g) dividing into four cells; i, endosperm cells. 
1g. 20. Placenta and ovules, one fertile, the other barren, of a more advanced ovary; the future prolon- 


gations of the embryo-sacs are seen in the placenta: lettering as before. 
Sa 21. Posterior prolongations of an embryo-sac partly dissected out. 
1g. 22. Section of an imperfect fruit, 


Fig. 23. Apex of the ovule of fig. 22, with embryo 
the germinal vesicle (suspensor). 
Fig. 24. Upper point of a more advanced ovule, 
section of the ovary from which it was 
Fig. 25. Embryo from fig. 24. 
Fig. 26. Placenta and ovule of a fruit ne 


. Teceptacle : lettering as before. 


(9)5 dd. coagula; k. upper cell of the row formed from 


with the embryo (y). The small figure on the left is å 
taken, nat. size. 


arly as in fig. 27, the placenta p. already torn away from the 


OF SANTALUM ALBUM. 79 


Fig. 27. An immature fruit; 0. ovule, on the side of which lies the placenta p; r. endocarp; s. rind com- 
posed of the blerided mesocarp and epicarp ; v. peduncle of the placenta. 

Fig. 28. Another, more advanced ; lettering as before. 

Fig. 29. Å fruit nearly ripe; part of the succulent epicarp removed, laying bare the shell or mesocarp s. 

Fig. 30. Section of fig. 29: letters as before; p. & s. are the sphacelated placenta (p), and its base (v) 
removed from the points indicated by the dotted lines. The endocarp is now obliterated. 

Fig. 31. Ripe fruit with the pulp removed from the shell. 

Fig. 32. The seed of the same with the shell or mesocarp removed; 7. scaly fragments of the obliterated 
endocarp ; p. placenta ; v. its original base. 

Fig. 33. Section of fig. 32 with the embryo (g) in the albumen ». 

Fig. 34. Embryo extracted. 

Fig. 35. Section of another seed. 

Fig. 36. Its embryo extracted, 


Trans Linn Soc Vol XXL tab! 1 p 78 


[tab I9 p78 


GJarman Se. : 


Trans LinnSoc Vol 


[ 8] 


III. Remarks on the Nature of the outer Jleshy covering of the Seed in the Clusiaceæ, 
Magnoliaceæ, &c., and on the Deøelopment of the Raphe in general, under its various 
circumstances. By Joux Mss, Esq., F.R.S., F.L.S., $c. 


Read March 18th, 1856. 


IN a notice read before the Linnean Society on the structure of the seed of the Clusiacee, 
I described the external fleshy envelope of the seed of the Clusiee, and offered evidence 
to prove that this coating is a product of extraneous placentary growth, subsequent to the 
development of the primine, and therefore a kind of arillus. It was of some importance to 
ascertain this point, as in the tribe Tovomiteæ of the same family, the outer coating, similar 
in substance and colour, is unquestionably an aril: this is manifest from its peculiar form ; 
it is free from the testa, but may be opened out like a flat plate; itis folded round the seed 
which it envelopes and conceals, its margins being free and overlapping one another: in 
the other tribe of the Garciniee, this covering is also entire, is soft, and assumes the 
character of an enveloping pulp. 1f, therefore, in the two latter tribes, the outer coating 
be unquestionably an aril, it was fair to conclude that the analogous envelope in the 
Clusiacee is of a similar nature. This inference was still farther confirmed by the 
presence of a distinct simple raphe, which extends from the base to the apex of the seed 
_ beneath the inner pellicle of the aril; it lies within a groove of the testa from which it is 
wholly free, the apex of which it perforates, and becomes lost in the chalaza of the inner 
integument. Under the evidence of such demonstrative proof, as far as regards the, 
Clusieæ, I was led to institute a comparison of similar facts observable in the Magnoliaceæ, 
because, if that coating be considered an aril in the one family, it must be of the ume. 
nature in the other. I then referred to the admirable work of Dr. Asa Gray (‘The 
Genera of the Plants of the United States’), where a different view is entertained : = 
the external fleshy coat of the seeds of Magnolia is described as the testa, and er 4 
bony shell as the tegmen, or inner integument, the true tegmen having estaper Å 5 
notice of that excellent botanist. In opposition to this view, I referred to tha eu 
had made many : years before, of the seed of Talauma, à er re ibed by 
Magnolia: the evidence then collected, convinced me that the ‘testa’ på - 
Dr. Gray is arilliform, and that his * tegmen ’ is the true testa. If we ng c this i z 
coating in Talauma, where it is entire, we find it easily detached from må pi Is gå 
shell; and if we begin to pull it away from the summit, the raphe, as å adis en 

à : 5 ae er rresponding groove which 
be seen quite free from it, as in the Clusieæ, and to lie in a correspo : Mn 
extends from the base to the apex: the upper end of this raphe is rn scm ied 
aperture near the summit of this osseous shell, and to become lost in ies ms 
chalaza of a membranaceous inner integument. To all appearance, the raphe 


; : 1 
quite free; but if we examine it more attentively, a corresponding portion of the extremely 


VOL. XXII. 


M 


82 MR. MIERS ON THE OUTER FLESHY COVERING OF THE SEED 


delicate inner pellicle of the aril will be found attached to it, which has been torn away 
by the raphe. After detailing these facts, I proceeded to show that the nature of 
the different coats of the seed may be always determined, with certainty, by the relation 
of one with the other, and from the position they bear in regard to the raphe and the 
chalaza of the inner integument, and that wherever the raphe is found outside the testa, 
any tunic exterior to the latter must be of a growth posterior to the development of the 
original coats of the ovule. From the position of the raphe in the seeds of Magnolia and 
Talawma, I consequently drew the infallible deduction, as it appears to me, that the more 
external fleshy envelope must be arilliform, the thick osseous nut must be the testa, and 
the inner integument with its thickened chalazå must be the true tegmen. 

Since the reading of the above memoir, the first volume of that truly excellent work, 
the ‘Flora Indica’ of Dr. Hooker and Dr. Thomson has appeared, in which they detail 
the nature of the outer tunics of the seed of Magnolia (in p. 73), entirely in accordance 
with the conclusion of Dr. Asa Gray, but the reasons on which they have adopted this 
conclusion appear to me to involve some points which materially affect the legitimaey of 
the inferences there deduced. I cannot agree in their opinion of the perfect accuracy of 
the account of the structure of the seed of Magnolia, as given by Gærtner, for though 
nothing contrary to truth is there stated, yet the most important point which bears on 
the present discussion, the existence of a raphe, is altogether omitted; and I feel con- 
vinced, that if that eminent carpologist had been aware of its existence, he would not 
have concluded that the outermost coating forms one of the true integuments of the seed, 
meaning by this term, those which are developments of the primine, secundine, &e. The 
authors of the “Flora Indica’ and the distinguished American professor do not notice the 
peculiar perforation in the summit of the crustaceous envelope through which the raphe 
passes, and they call this extremity the chalaza, a term which, in accordance with Gært- 
ner, I think ought always to be restricted to that peculiar thickening of the inner integu- 
ment around the point where the raphe becomes lost in its substance*. I have minutely 
described this process of the testa in the seed of Talawma as a distinct perforation which 
I have called the diapyle; it is of frequent occurrence in the extremity of the testa of the 
seeds of different plants, and is destined solely to the purpose just mentioned ; in fact 
it is the corresponding point in the original base of the primine through which the spiral 
vessels, represented in the figure of Dr. A. Gray, pass to communicate their nourishing 
influence to the secundine, and to the body of-the ovule, prior to the commencement of 
its inversion, and is the only point in the primine in which there exists any passage for 
these vessels, either before or after the anatropal action, and therefore the only point in 
the testa (or tunic, resulting purely from the growth of the primine), that could be 
traversed by the raphe. The presence of a diapyle in one of the tunics of a seed, is as 
certain an indication that this is the real testa, as the existence of a chalaza affords the 
surest proof of the nature of the inner integument. The actuality of the true chalaza, 


* « Chalaza nempe nobis dicitur parva areola saturatè colorata, aut tuberculum parvum spongiosum, aut callosum, 
quod | din vasorum umbilicalium internorum finibus, vel et ex chorii exsuccis reliquiis originem hit, et in 
'Perheie exteriore membranæ seminis interna conspicitur.”—Gærtn. de Fruct., Introd. p. 135. 


IN THE CLUSIACEÆ, MAGNOLIACEÆ, ETC. 83 


and its connexion with the raphe, in the relative position in which the latter is found, 
have evidently escaped the observation of these eminent botanists, as they had previously 
done that of Gærtner; it is, however, important to attend to these distinctions, as they 
form essential and convincing elements in this inquiry. 

Still more recently a paper from Dr. Asa Gray has appeared in * Hooker's Kew Journal 
of Botany’ (vol. vii. p. 243), “On the Development and Structure of the Integument of 
of the Seed of Magnolia.” This was written in reply to my observations on the arilliform 
nature of the outer tunic, to which I have above referred. In that note, this accomplished 
botanist warmly defends his former opinion, which he maintains with great ingenuity and 
candour, but I find nothing there stated that disproves the conclusion of most botanists, 
regarding the nature of the several seminal tunics in question, which I have here 
endeavoured to confirm. If I had not been convinced that the issue admits of the most 
simple and demonstrative proof, I should not have presumed to contest the united 
authority of the distinguished American professor, and of the highly talented authors of 
the * Flora Indica,' whose opinions are entitled to especial consideration. 

. The argument that the scarlet external envelope cannot be an aril, because the latter 

* must needs have an opening at the top," stands upon very feeble ground. This was the 
definition of St. Hilaire, who classed the different forms of arilliform coatings under two 
denominations: 1. the true aril where the envelope has an opening in the top; 2. the 
false aril, where the coating that surrounds the testa is entire. Dr. Planchon, however, 
who ably defended and extended the views of his friend, in regard to the nature and 
origin of the aril,has shown that this definition does not hold good, and among other 
cases, he cites a species of Tetracera from Java, where the perfectly true aril forms an 
entire tunic. I need hardly refer to Gærtner, who also divides the aril into two kinds, 
the complete and the incomplete; notwithstanding that in some of the instances of the 
first kind, he has mistaken the endocarp for an aril; others, which he enumerates, possess 
a complete arilliform covering, among them Nephelium, which in this respect offers a 
good analogy with Magnolia. Another instance is still more remarkable, because it is 
recorded by Dr. A. Gray himself, in the same admirable work, and within a few pages of 
his description of Magnolia: it occurs among the Anonacee, figured in the genus Asimina, 
(plate 27) and described (p. 57) as entirely covering the testa, without the mention of any 
aperture in the apex. There is indeed no reason why an aril, which ". development 
from the placenta, may not form an entire coating, as well as the true integuments of 
the seed, originally cup-shaped processes. The argument of the inadmissibility of an aril, 
because it is not open at the top, therefore completely fails. 


In regard to the remaining arguments of Dr. A. Gray, I can only 
been demonstrated, that the nourishing vessels from the placenta, as he has figured them, 


only penetrate the primine of the ovule of Magnolia at the gangylode* ; they € = 
do not enter at the opposite extremity; but if we examine the ripe seed, we e 


repeat what has just 


* For the sake of demonstration I have here given the name of gangylode to that common point of A of a 
primine, secundine and tercine in the original base, now the summit of the inverted ovule, which afterwards form the 


diapyle of the testa, and the chalaza of the inner integument of the seed. = 


84 MR. MIERS ON THE OUTER FLESHY COVERING OF THE SEED 


vessels of the raphe penetrating the outer coating at the hilum, that is to say, at the 
extremity diametrically opposite to that of the original gangylode, and after running 
along its inner surface, they find their way straight to the diapyle of the enclosed nut, 
through which they pass to lose themselves in the chalaza of the inner integument, both 
which points in the ripe seed (the diapyle and chalaza) correspond with the gangylode of 
the ovule. - 

I have endeavoured, by the accompanying figures, to render this question manifest. 
Plate XIX. figs. 56 & 57 represent an ovule of Magnolia after Dr. A. Gray, showing the - 
broad placentary sheath which encloses the spiral vessels of the raphe. Fig. 58 is a longi- 
tudinal section of the same, showing that the spiral vessels originate at the placenta and 
hilum, and that they terminate at the opposite extremity, at the point of the gangylode, 
which is the only point of the primine where these vessels traverse, in order to convey 
nutrition to the secundine and nucleus: it is here evident that the sheath or extension 
of the placenta containing these spiral vessels, must always remain exterior to the primine, 
as well as to the tunic resulting from its subsequent growth. Fig. 59 shows a section of 
the ripe seed of Magnolia, where the placentary sheath (seen in fig. 58) during the incre- 
ment of the ovule, has in the progress of its own growth, extended itself all over the 
primine, and has become enlarged into an arilliform scarlet coating, and we have proof 
that this coating is the result of such growth, by the fact that the raphe, retaining the 
same position and direction, still remains enclosed in its substance. The primine, during 
its increment, by the deposition of osseous cells within it, in the manner recorded by 
Dr. Gray, becomes converted into the bony testa, shown in figs. 60 & 61, where the diapyle 
or aperture is seen in its apex for the passage of the raphe, corresponding with the same 
point of the gangylode in the ovule, through which these same vessels there penetrated : 
we have evidence that this testa is the product of the primine, not only from this circum- 
stance, but because it is exterior to another integument, the tegmen, which covers the 
albumen, and which exhibits a chalaza in its apex, as seen in fig. 62. The above gives 
us an example of the development of the simple raphe. 

As an instance of a branching raphe, I will select the seed of Licania, shown in fig. 63; 
this has à basal hilum, and is the growth of an anatropal ovule. Referring to figs. 56 & 58, 
it is evident that the only point where the spiral vessels can enter the primine, is at the 
gangylode, the point which they passed through prior to the inversion of the ovule. If, 
a ss the hypothesis of Dr. Gray, the raphe insinuated itself into the 
en —— " must, in the case Of Licania, begin to send out its ramifi- 
nn a å «Suk it entered, that is to say, from the gangylode; but on the 
en nn à - + e vessels start from the opposite point of the hilum : the main — 
lar aig course (as in the case of the simple raphe) to the gangylode or 

DE » Mrowing out its redundant vessels in many lateral branches, while other 


the tunic resulting from its growth, that they must have 
th and have pierced their way through the outer pellicle 
hemselves into its substance at a point near the micropyle, 


IN THE CLUSIACEÆ, MAGNOLIACEÆ, ETC. 85 


a supposition that cannot be entertained for a moment. As therefore the spiral vessels, 
at the period of the inversion of the ovule, starting from the hilar point, must have been 
contained within a simple sheath, an extension of the placenta, and as these vessels during 
the subsequent growth of the ovule, are found to ramify from this point over its whole 
surface, it is clear that the sheath which contains them must have become here extended 
with them, in the same manner, as has been ascertained by actual observation, that it 
really does grow over the primine in other cases. 

This may be rendered still more palpable by reference to a simple model. This mode 
of demonstration was first ingeniously suggested by St. Hilaire in his *Legons de Bo- 
tanique ' (p. 541), and I will here repeat it with some modifications. Let us suppose a 
simple flower with a tubular calyx, closed before æstivation, enclosing a tubular corolla, 
which again invests a superior ovary: we have here a good illustration of an erect ovule, 
the calyx representing the primine, the corolla the secundine, and the pistil the main body 
of the ovule or nucleus. Let us farther conceive the same floral model supported on a 
pedicel of equal length, and that the flower be suddenly bent down upon its pedicel, 
becoming glued to the calyx: we have thus an excellent representation of an anatropal 
ovule, where the former base of the flower is now become its summit, and vice versé ; the 
calyx, corolla, and pistil still remain the analogues of the primine, secundine, and body 
of the ovule: the foot of the calyx or torus, through which the nourishing vessels pass to 
promote the growth of the several parts, corresponds to the spot I have called the gangy- 
lode, and the adherent pedicel will represent the nourishing vessels enclosed in form of a 
sheath, or extension of the placenta, and the origin of the future raphe. It is evident, 
that during the subsequent growth of these tunics, this raphe must always remain 
exterior to the primine, as we see it in this model. Now if, as it has been contended by 
Dr. A. Gray, the arilliform coat of the seed of Magnolia be only the primine of the ovule 
enlarged in growth, and if, as he admits, the raphe be found within this coating, it is 
evident, referring to our model, that the pedicel must have become detached from the 
calyx, and made to penetrate not through the original point of its attachment to the 
torus (corresponding with the gangylode), but in some unaccountable manner, and for no 
purpose, must have pierced its way through the calyx near its summit (ata spot corre- 
sponding to the hilum in the ovule), and thus have insinuated itself inside the > 
traversing its whole length in order to form a new line of pommi dation of > pss 4 
proceeding from the base of the pedicel to the torus, within, instead of without, the Er À 
this is so manifest an improbability, as to carry conviction in the simple statement of . 


the fact. 
Doctor Asa Gray now candidly confesses that 


inner integument, and therefore the true chalaza, 
this Se would naturally lead him, he has, in great measure, renounced his peel 
argument of considering the fleshy covering and the crustaceous nut, the one , erac E 
of the primine, the other of the secundine, and has now most nn "p ER 
entirely new view of the subject, suggesting that these two seminal envelopes 


i ; i the primine, forming a 
in fact one coating, both proceeding from the simple increment of ‘ids på om in 


drupaceous testa; his words are, “the external coat of the ovule 


å 


he had overlooked the existence of the 
and to avoid the intricacies into which 


LI 


86 MR. MIERS ON THE OUTER FLESHY COVERING OF THE SEED 


the seed, its outer portion forming the pulpy, the inner the crustaceous seed-coat.” This 
ingenious reasoning cannot however be maintained in view of the real circumstances of 
the case, for it does not in any degree remove the insuperable obstacle which I have urged 
against his doctrine, in the position of the raphe with regard to the several coatings of the 
seed and to the true chalaza. But supposing, for argument’s sake, we waive that objection, 
the reasoning is not tenable upon other grounds, because if such were the origin of these 
two envelopes, there ought to exist an intimate union of the fleshy exterior with the bony 
nut, having its several osseous and fleshy deposits enclosed within the two original epi- 
dermal pellicles of the primine. We find, on the contrary, that the osseous substance of 
the nut is furnished internally with a smooth skin, and externally with a distinct hardened - 
epidermis, which in Talauma I found to be black and polished in the living state; the 
fleshy coating is also furnished on both surfaces with a distinct reticulated membrane, the 
inner pellicle being clearly seen with a lens. The arilliform coating, in every case exa- 
mined, I have found provided with a double (that is an endodermal as well as an epi- 
dermal) membrane, showing it to be a distinct formation from the testa. Although 
perfectly free from the latter in Passiflora, &e., it is generally more or less adherent to it, 
especially in those cases where the raphe is spread over its whole area in branching rami- 
fications: by its close adhesion to the testa, it then forms a compound tunic, and when 
the external coating is fleshy and the raphe is simple, as in Magnolia, &c., it can often be 
easily separated from it in an entire state: this is what Gærtner calls, a baccate or fleshy 
testa*. 

My view of the nature of this development is simply the following: that in the act 
of inversion of the ovule, the spiral vessels destined for its nourishment and always 
retaining their original attachment toit at the gangylode, are drawn out, together with 
an enveloping portion of the placenta, so that by means of these spiral vessels and this 
placentary sheath, the same communication between the placenta and the gangylode of 
the ovule is maintained that had existed prior to the act of its inversion. This placentary 
sheath with its enclosed spiral vessels, appears like à prominent broad external band, as 
shown and figured by Dr. Asa Gray, both in relief and in section, in his analysis of Mag- 
nolia, to which I have referred : up to this point we are both in accord. This band 
afterwards becomes expanded by almost imperceptible degrees over the primine, until it 
finally envelopes it in the manner I shall presently demonstrate: it then becomes thick- 
ened by internal deposits, and assumes the form of a distinct scarlet fleshy covering over 
the testa, being quite arilliform in its structure and appearance. The testa is a distinct 
development, formed by the secretion of transverse crystalline cells, closely compacted 
within the substance of the primine, the deposition of which cells has been noticed and 
lyden Grayt. The only circumstance that bears any weight in the opposite 


* « å * " 
ùn WAR el ici aie seminibus baceatis, et respectu situs sui nonnunquam exceptionem a regulå facit, 
habet affinitatem, atque a een teneat locum; ut in Bizd atque Magnolid. Hinc proximam cum wilt 
sui superficie Es : ut in nt Å OE mie quod carnosa testa semper arctissimo nexu cum totå seminis 
: ctis, nec non in Gloriosd, &c., in quibus nullum inter carnem atque semen ipsum 


intercedit spatium liberum, sicut in seminibus arillati 1 i 
T Hooker's Kew Journ. Bot. vol, viii. ote viae Seytaliæ (Nephelii) et aliarum.”—Gærtn. Introd. p. 133. 


IN THE CLUSIACEÆ, MAGNOLIACEÆ, ETC. 87 


view of the question is one which certainly ought to have its due importance, and is one 
which I have never doubted in the smallest degree, viz. that Dr. Gray has watched the 
progress of the growth of the seed from the ovule and could never detect any subsequent 
‚or extraneous production over the primine. In the fullest credence of this assertion I 
still think we have convincing proof that such extraneous growth, though he failed in 
distinguishing it, must have taken place imperceptibly, in the same way that other 
observers have witnessed a similar gradual extension over the primine. 

The foregoing discussion has induced me to offer a few additional observations on the 
nature of the raphe. I have been led into this inquiry by the phænomena that present 
themselves in the Clusiaceæ, where I found it difficult to assign a reason, why the raphe 
should in one case consist of à single cord of vessels, almost in a free state, within the 
inner pellicle of the external arilliform coating, and wholly free from the testa, as inva- 
riably occurs in the C/usieæ, while in the Tovomiteæ, the raphe has no connexion with 
the enveloping aril, but appears imbedded in the slender chartaceous testa, not as in the 
former case like a simple cord, but spread in the form of several branching nervures con- 
tinually subdividing themselves, and distributed in new ramifications throughout the 
whole area of the testa. There appears here a difference of structure somewhat inexpli- 
cable, but after a little consideration, we may trace to one uniform rule the varying 
appearances of the raphe under these different forms and position. We owe to the pro- 
found investigations of our great countryman Mr. Brown, most of the knowledge we 
possess regarding the growth and development of the ovule and its original tunies, the 
changes they undergo during and after the anatropal metamorphosis, the gradual trans- 
formation of these tunics into the different coatings of the seed, and the share they afford 
in the nourishment and development of the growing embryo: but the phenomena 
attendant on the formation of the raphe, the modifications which these nourishing vessels 
undergo, and the different modes of their distribution, appear to have excited little notice, 
for we find few observations on the subject in the works of physiological botanista, oe 
this account I will venture to offer some remarks that have occurred to me in my pend- 


ing investigation of the Clusiacee. 
Let us now examine some of the circumstances attendant on 


Prior to this action, the body of the ovule or nucleus is generally ee ir aii 
an as d secundine are given : these cups gra 
are Am m co and conceal the nucleus, also called 


Size, and grow into complete tunies which envelope : . 

the ing: during iis ende stage, the vessels destined to nourish po Er hagen me 
diately from the placenta into the gangylode or common po 2 cues, nl 
With the secundine and primine then in contact with the ucl n iei 
gress of inversion, the gangylode, as before — zum "ue i e side of the 
‘and a portion of the placenta is drawn away with it and extended ove een 
primine, at first in the form of an elongated sheath or not d hide in the 
Carrying with it the spiral vessels proceeding from the — A quise cuiu fe its 
now remote gangylode: this placentary extension, whatever TTT at it may 
subsequent growth, must always remain outside the primine no ee de ia 

me agglutinated to it. Now these nourishing vessels may 


the inversion of the ovule. 


88 MR. MIERS ON THE OUTER FLESHY COVERING OF THE SEED 


i i e branched and extended with the 
vors RUM sin ke of the ovule, and thus spread into 
ehe seters but however they may become thus distributed, they must in 
pst eiselaned within their placentary-envelope, and, as above shown, be 
necessarily exterior to the primine. This placentary sheath, first seen as an adnate 
longitudinal band, and afterwards extended in the form of a complete investiture over the 
iue will be found to assume different phases of development : it may, during this 
itid ix expansion, be reduced to a degree of attenuation not thicker than a mere 
skin, and by desiccation of the matters secreted between it and the primine; both may 
become intimately fastened together, as we know occurs frequently with the primine and 
secundine, in which case the ramified bundles of wessels would thus become enclosed 
within this apparently simple, but really compound tunic, pose Whois of _— 
mosing nervures, such as I have figured in the testa of Tovomita and Qommiuhea , and 
by a careful dissection of this sort of tunie, we find the confirmation of this structure. 
In other cases, where the nourishing vessels remain compacted in one simple bundle or 
cord, we may conceive that the placentary sheath, which originally enclosed them, has 
extended itself over the primine in the manner deseribed, and has become developed in 
the form of such an arilliform tunic, as we find in the Clusiee and Magnoliaceæ, the 
raphe in such case necessarily remaining quite free from the testa. Inconsistent in result 
as the two opposite cases just referred to, may at first sight have appeared, it is evident 
from this explication, that both are in perfect harmony with one simple and uniform 
action, varied in effect according to the peculiar circumstances under which the secretory 
productions modify the nature of the developments. 
This extension of the placentary sheath may be either complete as I have described 
it, or only partial: such a partial expansion is known to occur in Turneracee, and the 
circumstances under which it is there developed, offer still further confirmatory proofs of 
the placentary origin of the arilliform expansion in the manner above detailed. St. Hilaire 
in his ‘ Flora Bras. Merid.’ pl. 120, figs. 4 and 5, exhibits the seed of Turnera herman- 
nioides, where the raphe’ proceeds from the hilum, one-third way long its ventral face, 
like a cord: beyond this to the summit, and half way down the dorsal face, it spreads a 
the form of a broad fleshy plate or incomplete tunic. The same development is shown in 
plate 121, fig. 5, where, in the seed of Turnera genistoides, the raphe is seen to extend 
above half way from the hilum, as a cord, whence it expands as far as the summit, in the 
form of two broad auricular plates, nearly the length of the seed, one lobe being seen 
upon each of the lateral faces of the testa, forming, as in the preceding species, a partial 
fleshy envelope. Another instance of the enlargement of the placentary sheath.oceurs In 
Asarum, called. by Gærtner an epiphysumt: Dr. Planchon also describes the seed of 
Asarum Canadenset, as being greatly swollen upon its ventral face, along the line of the 
raphe, by a large glandular mass extending from the base to the apex, and filled with oily 
vesicles, as in the fleshy coating of Magnolia; this he denominates a strophiole, and 


* Linn. Trans. xxi. tab. 26. figs. 22 & 31. os + Geertn. de Fruct. 48. tab. 14. 
1 Mémoire sur les développements des vrais et des faux arilles, p, 34. pl, 2. figs. 10 & 11. 


* 


IN THE CLUSIACEÆ, MAGNOLIACEÆ, ETC. 89 


Endlicher calls it a fleshy raphe, but this origin is evidently due to the same source as 
that of the outer tunic of Magnolia. 

It is here necessary to point out the distinction between the different kinds of arilli- 
form coatings of the seed. We have strong evidence to show that the scarlet covering in 
the Clusiee, Magnoliacee, &c., is derived from a growth of the placentary sheath, which 
is a production of the funicular cord, whose origin is coæval with the anatropal inversion 
of the ovule: other cases however occur, where another still more exterior development 
is generated at a subsequent period: this is well known to proceed from the main 
placenta, or more often from the short funicular cord, which is the foot-stalk of the 
placentary sheath by which the ovule is attached, sometimes forming a thick cup as in 
the Sapindaceæ, &c., or at other times extending itself like a fleshy, coloured plane sheet 
completely enveloping the seed, as in the Tovomiteæ, or which form branching fleshy 
segments embracing the seed, as in Myristica, Samydaceæ, &c. These several varieties 
derive their origin from a growth of the placenta and are not developed from the 
original coats of the ovule; they are therefore arilliform in their nature, but as they are 
produced under different cireumstances, I propose to confine the term aril (arillus) to the 
kinds of coating last described, always void of spiral vessels, and to denominate that sort 
found in the Clusiee, Magnoliaceæ, &c., the arilline (arillinus), in which the vessels of 
the raphe are always imbedded. Both may and often do exist at the same time, in the 
same seed, or they may make their appearance independently of each other. Å 

There is said to be still another kind of extraneous fleshy coating, produced over the 
testa in the manner first pointed out by St. Hilaire, to which he gave the name of false 
aril, and which was afterwards ably investigated by Dr. Planchon, who substituted for it 
the name of arillode*. This tunic is described as originating in the expansion or rever- 
sion (dedoublement) of the mouth of the exostome or foramen of the primine, which 
gradually extends itself over its whole surface, and forms an extraneous envelope around 
the testa. The formation of the arillode from such a source, rests upon the authority of 
Dr. Planchon, to which Lattach its due weight; and although in support of his views, he 
described, with great minutenesst, the gradual formation of the arillode in dm | 
latifolius, it is certain that in drawing the conclusion that the arillode proceeds from the 
extension of the exostome, he expressed at the same time some misglving on the subject. 
Examining therefore the details as there represented, and COMPARE, these ke E zem 

. A 1 e $ 
observations on the seed of Euonymus, I think it may be inferred, with equal, if not wi 


d Idem, p. 7. 
* Mem. ante cit. p. 10. T : tire ai-j rifié 

t “En disant que ce dernier sac (le faux arille) procède uniquement de — er : ter aem des 
l'exactitude à la clarté. Comme l'ombilic, en effet, est très voisin du — Sp - solution de continuité. 
bords de ce dernier, devrait rencontrer dans le funicule un obstacle à son extension et 


une partie 
Mais c’est 14 au contraire que P expansion est la plus épaisse, et même elle adhère avec la base du raphe sur une p 


å G jerni ie. Il faut donc nécessairement, 
de sa longueur de manière à ce qu'elle semble sur ce point naître ae eni er ; = er funicule. Je crois 
pour expliquer une pareille disposition, admettre une soudure congeniale entre l'expansio EEE 

le mieropyle est fort dificile à voir, lorsqu 


dévoir ajouter pour prévenir tous les doutes, que chez les — erture, et la cache complètement : 

lovule est déjà très développé, parceque le faux arille est plissé — de son en des bords de P'exostome.” 

mais en détachant avec soin l'enveloppe accessoire, on peut facilement s'assurer que 

—Loc. cit. p. 9. N 
VOL. XXII. : 


90 MR. MIERS ON THE OUTER FLESHY COVERING OF THE SEED 


greater probability, that the extraneous coating is produced from the funicular cord, 
rather than from the foramen or micropyle of the primine, and in such case the arillode 
i the aril. 
ces TR of examining the ovule during its growth, but have lately 
observed ripe and living seeds of Ewonymus Europæus. Here the outer coating is entire 
fleshy and scarlet, with a smooth inner skin, and we find beneath it, another polished, 
thinner, though somewhat fleshy tunic, that closely adheres to the seed. If this tunic be 
removed carefully from the thin pergameneous testa, it will be found to consist of two deli- 
cate reticulated pellicles, having cellular and fleshy matter interposed between them, the 
raphe being completely immersed in its substance, in the form of a ne cord, which 
originating at the basal hilum, proceeds along its face to the apex, where 3t pierces the 
inner pellicle of this tunic, passes into a small opaque speck in the summit of the testa 
(the diapyle), and is lost in the chalaza of the inner integument which is adherent to the 
shell. Here we have demonstrative evidence of the nature of these several envelopes ; 
the outer coat is manifestly a true and entire aril, for we cannot suppose it to be a deve- 
lopment of the primine, that is to say, an extension of its exostome, as Dr. Planchon 
almost doubtfully concluded, because it is altogether free from, and exterior to a more 
internal tunie which encloses the raphe: it follows, therefore, as a necessary Consequence, 
from the position of the nourishing vessels, that it must be a production emanating from © 
the main placenta or a growth from the funicular support of the seed. The fleshy 
epidermoid tunic which encloses the raphe, and which immediately invests the pergame- 
neous shell, appears to be an arilline, resulting from the growth of the placentary sheath : 
the thin pergameneous shell is, of course, the true testa, marked at its base by a small 
prominent nipple, close to the hilum, which is no doubt the thickened border of the true 
. mieropyle figured by Dr. Planchon, and from which he inferred that the growth of the 
aril had emanated: the apical speck through which the vessels of the raphe penetrate, is 
the diapyle: it is hardly necessary to add, that the radicle of the embryo, enclosed in 
albumen, points to the micropyle, while the extremities of the cotyledons are directed 
towards the diapyle. I do not find the aril pervious in the apex, as stated by Gærtner, 
and as figured by Dr. Planchon in another species, although this, no doubt, sometimes 
occurs; but in the instance above mentioned, the inner skin of the tunie, though slightly 
crumpled, is entire, while its outer pellicle is deeply plicated in flattened folds, so that 
the aril appears cleft into numerous fissures externally. 

Among the many interesting facts detailed by Dr. Planchon in the work just quoted, 
we meet (Joc. cit. p. 25) with an account of the circumstances under which the seeds of 
Opuntia become covered by two distinct extraneous envelopes, both exterior to the testa : 
the first is a somewhat thin, hard, coriaceous tunic, aecording to his observations; the 
second is a soft, mucilaginous, pulpy coat by which the former is encircled. The growth 
of the former was traced by Dr. Planchon from the period of the anatropal inversion of 
the ovule, which was carried to an extent of a complete gyration, so that the placentary 


sheath I have before described, appeared at first like an annular band around the 
periphery of the ovule; from this ring, on both sides, 


membranaceous expansions were 
seen gradually to extend themselves over the interve 


ning spaces, until they met in the 


IN THE CLUSIACEA, MAGNOLIACEÆ, ETC. 91 


centres, and thus formed one complete tunic, which finally assumed the solidity and 
texture of a crustaceous shell, which he called a * false testa”: this again became 
enveloped by a second placentary extension, in the form of a transparent soft pulp. 
These facts had been previously authenticated by the careful observations of Gasparini*, 
who describes, with great apparent accuracy, the curious phænomena attendant on the 
growth of the ovule of Opuntia, from its earliest development to the state of its ripe 
seed. After its anatropal inversion, the ovule is seen suspended by its short funicular 
cord (podosperm) from the hilum, which cord, in form of a placentary sheath, or thick 
cylindrical filament, is seen to extend itself round it, until it encircles the ovule like an 
annular ring: from this ring, on each side, a distinct membrane, at first very thin, 
expands itself by slow degrees, until at length, on both sides, it becomes extended in a 
complete tunie, over the entire surface of the ovary. After the period of fecundation, he 
farther observes, if we watch every now and then the growth of the ovary, during its 
transformation to the state of seed, we see the tunic just mentioned, as well as its very 
short podosperm, become covered, little by little, with a pulp. In proof of the fact that 
the production of this more external tunic originates in the extension of the podosperm, 
he states that sometimes, in some ovules, from some unknown cause, the above-described 
annular prolongation of the placentary sheath is not formed, and in such case the seed is 
not covered with the usual pulpy envelope. Some explanation, however, is here requisite, 
which I am enabled to give from the examination of the large seed of a species of Opuntia, 
collected by me in Chili many years ago. This has convinced me of the correctness of 
the details given by Gasparini and Planchon, with this exception, that the thin pellicular 
membrane, which both actually witnessed in the act of its growth and extension over the 
primine, and which the latter imagined became converted into the thick osseous shell, 
is no other than the intermediate epidermoid tunic, which I found still covering the 
Shell. We may feel assured that the deposition seen of osseous cells, to form the 
crustaceous shell, took place in the substance of the primine, and not in its arillinar 
covering, as Dr. Planchon inferred, for Gasparini makes no mention of such aD-ooour 
rence. That suchis the case, is manifest from the position of the raphe, and it is not less 
| clear, from the phænomena observed, that the membrane, as they saw it in the progress of 
its growth, is a production of the placentary sheath, and is therefore of the nature E "- 
arilline. So, in like manner, the pulpy envelope emanates from the placentary sheat rn 
funicle ; and that such is really its origin is proved by the curious fact related by — 
that when, as it sometimes happens, the placentary sheath is unformed, or grue 
pulpy matter takes place over the seed. From the circumstances above peas = 2 
safely conclude that the hard crustaceous shell in the seed of Opuntia = sr sd å 
its annular ring is the raphe, and that the intermediate tunic coating e sip 
arilline; while the more exterior pulpy envelope, whether originating in the p Ty 
sheath or the funicle, is still an aril, because it is void of spiral vessels. = 
Another striking confirmation of the fact of the gradual increment of the fles A 
over the primine, is eited by Dr. Planchon, and is the more important, because i 


occurs in Clusia, and bears immediately on the question at issue. Je de before 


* Osservazioni intorno alla struttura dell’ arillo. Rendiconto dell’ Accad. delle — T ga ^ er 


92 MR. MIERS ON THE OUTER FLESHY COVERING OF THE SEED 


quoted (p. 31), as an instance of the progressive formation of the false aril, he describes 
and figures (in plate 2. figs. 7 & 8) the ovule of Clusia flava, which he examined after the 
fall of the corolla, at which period he observed that the primine adjoining the funicle 
became enveloped for a quarter of its length by a cup-shaped arilliform process, and there 
can be little doubt that this expands into the entire fleshy coating, which at a later 
period we know envelopes the testa in the manner I have described in the Clusiee. 

M. Dutrochet, a very able physiological botanist, in discussing the nature of the aril, 
denies that its growth proceeds immediately from the placenta, and declares his convic- 
tion, formed after many years of patient investigation of the subject, in the following 
words :—“ L'opinion qui me paraît aujourd'hui la plus conforme å l'observation, est que 
l'arille est une extension de la partie inférieure ou de l'enveloppe corticale du funicule*.” 

Mirbel has furnished us with evidence of great importance in the solution of this 
inquiry: he minutely describes the growth and development of the ovule of Cueumis 
Anguria, in which he observed, after the period of its fertilization, the production of two 
distinct layers of cellular tissue over the primine, originating, no doubt, from an emana- 
tion and extension of the placentary funicle. His words are, “deux couches de tissu 
cellulaire, qui n’appartient pas primitivement à l’ovule, mais qui s’applique à sa surface 
et finit par lui servir d’enveloppe comme ses téguments proprest," thus affording un- 
questionable evidence of the extraneous growth of the arilline, evidence since confirmed 
by the observations of Gasparini and Planchon. I shall be able to show that the crusta- 
ceous tunic of the seeds of the Queurbitaceæ, hitherto held to be the testa, as well as its 
immediate soft envelope, are both of arilliform origin, and that the true integuments 
resulting from the primine and secundine of the ovule are to be found in the membrane 
that immediately invests the embryo, and which has always been considered the tegmen : 
this fact is attended by some curious phænomena which will be detailed in another place, 
where I will adduce many instances of the anomalous development of the raphe. I will, 
however, here allude to the peculiar structure of the outer tunics in many of the Cucur- 
bitacee : if the seeds of Citrullus, for example, be macerated in water, the outer shell 
will be found to eonsist of three distinct parts, which may be termed the epiderm, the 
ert ni — The epiderm is pellicular, transparent, and under the 
band ct are =- ze: reticulstions; the mesoderm is always fleshy, and 
ee mes SER = en mixed with pleurenchymatous deposits; the 
ylitders : cime: as m e eus 1s crustaceous, and consiste of transverse hollow 
iron RS = == = utinated together. The epiderm is a continuation, 
naine ee Dr sheath of the umbilical cord; the mesoderm is 
protect the more delicate s lint y agite cosi essa pres i ies in 
dietiust-depodt: sithi im "m die the raphe; but the crustaceous endoderm is a 
sac, enclosing the seed, and de ie er Forming å compressed and søger 
space filled with loose cellular tiene pus dts mouth (the hilum), within which is a 
sitrfned of the-eridoderin v iden which is also generally extended over the internal 
the’ fertilization of the pe — deposits must be formed at a period subsequent to 

» as will be evident from the copious details, illustrated by 


* Mém. Mus. se 
~ + Mém. Acad. Paris, ix. 622. tab. 1. fig. 10 e; fig. 11 4. 


IN THE CLUSIACEÆ, MAGNOLIACEÆ, ETC. 95 


admirable figures, of the growth of the ovule recorded in the valuable Memoirs of 
Brongniart and Mirbel, and, as will be seen, proved by the observations of the latter, 
cited in the foregoing page. It is not more unreasonable to conceive that the osseous 
deposit, forming the crustaceous covering, may in some cases be secreted in the arilliform 
coating, as well as in the primine of the ovule; in both instances, the secreted matters 
must pass through the same channel, and be supplied by the same vessels of the funicle, 
and such depositions at one point, instead of another, are probably regulated by the 
nature of the pre-existing tissues. In Zanonia and Fewillæa, the arilliform nature of the 
outer coating is better shown by the membranaceous state of the tunic, which is extended 
like a winged covering over the seed. 

From these circumstances we may infer, that the arilline need not necessarily be always 
fleshy in its nature, as in Magnolia; but that it may be either membranaceous, gela- 
tinous, coriaceous, or even osseous in its structure. Thus I have found from the position 
of the raphe, that the hard highly-polished tunic of the seeds of Drimys and Ilicium, 
usually regarded as testa, should be held to be a true aril: thus also the coriaceous 
coatings of many seeds will in like manner be found to be arilliform in their origin. Of 
that kind of seminal coating where the arilline is intimately combined with the testa, and 
where the raphe, greatly branched, lies imbedded between them, forming a compound 
tunic analogous to the structure already described in the Tovomiteæ, a very remarkable 
instance occurs in the Oleace@, where the raphe, instead of being spread into numerous 
branching nervures, exhibits itself by infinitely minute ramifications, as a dense network 
of most delicate spiral vessels, crowded together into a cottony web, like that of a spider’s 
cocoon, and fills up the entire space between the testa and arilline ; these tunics, aided 
by this interposition, are closely agglutinated into an apparently simple coating, but by 
maceration they may be separated from each other, and the interposed network may be 
drawn out into innumerable elegant spiral threads. This structure I have found in 
Tessarandra and Olea, and it probably exists in other genera of the family. Nearly the 
same development occurs in Caswarina, where a thick web of spiral fibres is found inter- 
posed between the crustaceous testa and the outer membrane, which is extended over it 
in the form of a wing. This structure, noticed many years ago by. Mr. Brown, and at 
that period described as a singular occurrence (Gen. Rem. p. 40), has since been confirmed 
by Schleiden, and figured in Schnitzlein’s ‘ Ieonographia " (Gen. 86). The Bu 
coating of the seed, here extended in the me wing, m covers the excessive deve- 
lopment of the raphe, will probably be found to be an arıllıne. = ÅT 
We have a = illastration of the arilliform nature of the external coating på p 
under somewhat variable forms, in the Passifloracee. In Tacsonia puma — hh 
the seed invested by a mucilaginous pulpy envelope, ee se pe 


i i i siderable vacant space between it and the osseous 
vesicle, quite detached, leaving å con der in gii: this 


testa, in every part save at a small point at the and the ar 
pellicle Te = vessels of any kind, except in its longitudinal raphe, Mee ^ P 
in its substance, appearing as a prominent white nerve, running ig e p 
the summit, where it finds a passage through a caruncular spot (the ; m i p7 
of the testa, beneath which it becomes lost in the chalaza of the inner integument. 


$ 


94 MR. MIERS ON THE OUTER FLESHY COVERING OF THE SEED 


Ryania (Patrisia), as shown in Delessert’s ‘Icones; iii. tab. 14, the seed is covered by a 
similar arilline, and along the whole length of a similar longitudinal raphe, à lateral 
cupular fleshy expansion is developed, manifesting the coexistence of aril and arilline, 
emanating from the same origin. Both these developments also occur in Paropsia, 
where the seeds are suspended by a long funicle, which, at the hilum, is expanded into 
a fleshy cup, that envelopes the lower half of the seed. In Acharia the longitudinal 
raphe becomes distended on each side, forming a lateral gibbous process (Ann. Nat. Hist. 
iii. pl. 9. fig. 15) analogous to that before referred to in Asarum. In Modecca palmata 
an entire and very thick fleshy tunic invests the osseous testa, marked by à prominent 
longitudinal keel, enclosing the raphe (Wight, * Icon.” tab. 201. fig. 12, 13, 14, &c.), and 
in Modecca Wightiana (id. tab. 179. fig. 3) we see precisely the same development, with 
the addition of another crenated hemispherical fleshy cup, covering the base of the seed. 
This is also seen in Modecca australis (Endl. * Icon.’ tab. 115). In all these cases, the 
raphe is perfectly free from the testa, and always forms part of a more external tunie 
more or less adherent to it, the arilliform nature of which has never been doubted; the 
structure is quite analogous to that found in Magnolia, &c., and distinct from the still 
more external development, the true aril. 

The mass of evidence here adduced, strengthened by the observations of botanists of 
high repute, showing the nature of the several metamorphoses which accompany the pro- 
duction of the raphe under its different forms, indicate the real placentary origin, whether 
mediate or immediate, and therefore the arilliform character of the several extraneous 
tunics, which assume such various textures and conditions around the testa,—restricting 
this latter term within the limit usually assigned to it—a simple development and growth 
of the primine of the ovule. The question appears to me so simple and manifest, that I 
should have considered it unnecessary to enter into such-full details in its support, if this 
point of structure had not been so positively denied by the high authorities to which 
I have referred: a desire for the solution of the truth has alone induced me to extend 
these observations to a greater length than otherwise would have been requisite. Many 
other interesting topics of physiological inquiry are connected with the farther considera- 
tion of this subject, and I have prepared another paper, in which are discussed many of 
the phenomena attendant on the peculiar direction of the raphe, especially in reference to 
the anomalies before alluded to, in Stemonurus, Anona, the Cucurbitaceæ, and other 
instances, with a view of tracing the causes of such unusual deviations from the ordinary 


. course of structure. 


IN THE CLUSIACEÆ, MAGNOLIACEÆ, ETC. 95 


EXPLANATION OF THE FIGURES. 
Tas. XIX, 


Fig. 56. Represents an anatropal ovule of Magnolia, after Dr. Asa Gray, seen on its ventral face, 
Fig. 57. The same viewed sideways. 
Fig. 58. À vertical section of the same. In all these the letter a shows the placentary sheath, enclosing 


b. the spiral vessels of the raphe; c. is the primine; d.the secundine; e. the tercine or nucleus ; 
J. the gangylode; g. the hilum. 


Fig. 59. Is a longitudinal section of the ripe seed of Magnolia. 

Fig. 60. Is the crustaceous testa, cleared of its fleshy covering, viewed sideways. 

Fig. 61. The same seen on its ventral face, showing the groove formed by the pressure of the raphe. 

Fig. 62. The inner integument, covering the albumen. In all, the same letters refer to a. the scarlet 
arilliform outer coating, being an expansion of the placentary sheath seen in figs. 56, 57 & 58, 
which has grown over and covered the primine, and now completely envelopes the testa; 
b. the raphe; c. the testa or development of the primine; d. the tegmen, or inner integument, 
resulting from the secundine; e. the albumen; f. the diapyle or scar of the gangylode, through 
which the raphe passes to reach the chalaza; y.the hilum; A. the funicular cord; i. the 
chalaza upon the extremity of the inner integument, where the raphe is lost; k. the embryo 
imbedded in albumen :—all much magnified. 

Fig. 63. Is a seed of Licania removed from its pericarpial covering :—nat. size. a. the tunic formed of 
three adherent membranes: the outermost is the arilline, the intermediate one is the testa, the 
innermost is the tegmen; 5. is the hilum; c. the vessels of the raphe, interposed between the 
arilline and testa, and dividing into continual ramifications, which spread over the whole area of 
the seed: they all branch from the hilum, which is situated at the extremity contrary to that 
of the chalaza, or point corresponding to the original base of the ovule, showing that these 
vessels must be exterior to the real testa, and enclosed within another distinct integument ; for 
if they were included within the substance of the testa, they would branch from the contrary 
extremity at the point corresponding with the original base of the ovule, the only point in 
which these vessels could have penetrated the primine, and therefore the testa. 

Fig. 64. Is the exalbuminous embryo, also nat. size; d. the cotyledons ; e. the radicle, placed near the 
hilum. 


Trans LiunnSve Vol XXII bab Pp 994 17 


åå 


SSA | 


X 
hund men 


y ? 


#JarmanSe 


Le 


IV. On several instances of the Anomalous Development of the Raphe in Seeds, and the 
probable causes of such deviations From the usual course of structure, especially in 
reference to Stemonurus (Urañdra of Thwaites), with some Prefatory Remarks on 
that Genus. By Joun Miers, Esq., F.R.S., F.L.S. $c. 


Read April 15th, 1856. 


IN a recent Number of Hooker’s Journal of Botany (vol. vii. p. 211), Mr. Thwaites has 
recorded a new genus, of which he gives the characters under the name of Urandra. 
This he refers to * Olacacee, tribe Icacine@.” I have read over his description with the 
utmost care, and am obliged to say that I perceive no difference whatever in the cha- 
racters of Urandra, and those I have detailed of Stemonurus, to which genus he confesses 
it is closely allied, differing only in all its flowers being fertile, in its small, not pulvinate, 
stigma, and in the structure of its fruit. The character founded on the constancy of the 
hermaphrodite flowers in the plant which he describes, cannot be considered of the 
smallest generic value, because this circumstance has been shown to exist not only in 
some species of Stemonurus, but in the contiguous genus Platea, in which, although 
some of the plants are unisexual, others are frequently hermaphrodite. I have also 
shown that the stigma in Stemonurus (Gomphandra, Wall.) is small, and not large and 
pulvinate as it had been described, but that it becomes subsequently immersed in the 
epigynous gland which crowns the ovary, a circumstance evidently not observed by 
Mr. Thwaites; and that it is this gland which assumes a pulvinate form on the summit of 
the fruit, and not the stigma, which may always be seen hidden in a small central depres- 
‚sion of the cushion. There remains therefore to be considered only the structure of the 
fruit; and Mr. Thwaites's details of the ovary, fruit and seed in Urandra, closely agree 
with what I have observed in Stemonurus. In this genus, as in all others of the 
Icacinacee, where the ovule is usually unilocular, the cell is always excentrically placed 
on one side of the pistil, the point of suspension of the ovules not being from the summit 
of the cell, but constantly inelined against the side on the line of the displaced axis of the 
ovary: the lobes of the stigma are always two or four, thus showing primé facie that 
the ovary is normally bilocular, and that the two ovules observed in the cell are really 
attached, near its summit, to the dissepiment, which, owing to the abortion of the other 
cell, appears to form the wall of the ovary. I was fortunate enough to meet with the proof 
of this conclusion in a ripe fruit of the closely allied genus Pennantia, where été 
is usually unilocular as in Stemonurus; but in the instance alluded to, the cm 
regularly two-celled, and only one seed was perfected in each cell, the sore c på 
ovule being still visible on the dissepiment at the point of attachment of each seed ; 
iti i ishing vessels proceeding from the base formed 
partition was of thin texture, and the nourishing vessels P bt 
a longitudinal nervure in the line of its axis, extending thence to the point of a 


o 
VOL. XXII. 


98 MR. MIERS ON SEVERAL INSTANCES OF ANOMALOUS DEVELOPMENT 


of the seeds near the summit of the cells. Here we have the most positive proof of the 
normal structure of the whole family, and this fact ought to be held in view in those 
cases which ordinarily occur, as in Stemonurus, where one of the cells has become oblite- 
ial abortion. 
vette who has given us the details of Urandra, has had the advantage of 
examining the fruit in a living state, and describes it to be a somewhat fleshy drupe with 
a subligneous putamen, its solitary seed consisting of an embryo formed of two small coty- 
ledons and a superior terete radicle loosely seated in the upper part of a longitudinal cleft 
or cell in a large albumen, which is thus nearly divided into two almost equal plates, by the 
intervention of two cellular strata that line this cavity, and whose margins nearly reach the 
testa on every side. This exactly corresponds with what I have observed in Stemonurus, and 
the same features are confirmed by Dr. Wight in his ‘Icones, under Stemonurus ( Gomphan- 
dra) polymorphus, where they are well delineated in plate 954. figs. 11 & 12, and where the 
chink above mentioned is shown distinctly in fig. 13. Mr. Thwaites examined the texture 
of the nucleus of Urandra under the microscope, and he describes the albumen to consist _ 
of cells, radiating from the more central tissue to the periphery of the seed; he states 
that the two strata lining its cavity are formed of cells similar to those of the albumen, 
but differing in containing no amylaceous granules: he considers them to be organically 
connected with the albumen, but I succeeded most certainly in separating them as easily 
as the integuments, and still preserve them in this state. At the time I published my 
Monograph of the Icacinacee, I had been able to examine only a single drupe of 
Stemonurus ; but I was fortunate enough afterwards to obtain another seed, through the 
kindness of the late Major Champion, which I analysed, and found that a portion of the 
embryo, in the first instance, had been partially eaten by a small insect, then observed 
within the cavity, so that the real cotyledons were gone, and I found a portion of the 
radicle, situated in the manner I had described, in the upper part of the axis of the 
albumen, and placed at one extremity of the two thin membranaceous strata described by 
Mr. Thwaites. I naturally concluded, by analogy, that these were the cotyledons; they 
occupied the same position, and were about the size of the foliaceous cotyledons I have 
delineated in Mappia fætida, and as these are figured by Dr. Wight (Icon. 955. figs. 10 - 
& 11) under Stemonurus Fetidus. In my second analysis, made soon after the publi- 
cation of my Memoir on Stemonurus, I discovered the embryo entire, with small coty- 
ledons, just as Mr. Thwaites has shown them; my attention was now, therefore, drawn to - 
the consideration of the nature of the two membranaceous strata, well described by that 
botanist. The result of this examination, and my correction of the form of the embryo, 
would consequently have been given in the forthcoming volume of my ‘ Contributions,’ 
where the details of Stemonurus and all other genera of the Icacinacee are figured. I 
sm, however, glad of this opportunity of rectifying my previous description, and of adding 
à drawing of my analysis of the seed. 
Another cireumstance mentioned by Mr. Thwaites calls for observation. In Stemonurus 
the albumen 9 f the seed is covered by the ordinary delicate membranaceous inner integu- 
ment, and this again is invested by a very thin testa, which is intimately agglutinated to 
it, and which is marked by fine hexagonoid reticulations. The raphe, prominent upon 


OF THE RAPHE IN SEEDS. 99 


the testa, and which by pressure leaves a corresponding groove on the surface of the 
albumen, is singular in its form, on account of its retroversion in the manner I have 
detailed, both here and in my Monograph of the genus. Its nature does not appear to 
have occurred to Mr. Thwaites, for he describes it merely as * a whitish raised line, which 
is very conspicuous on the outside of the seed, passing quite round it lengthwise, and 
consisting of à fillet of spiral vessels lying between the two thin coats or layers of the 
testa.” The nature of this fillet is soon ascertained by tracing it to its origin, and I have 
again referred to the parts of my two analyses which I have preserved, and these confirm 
my previous inference. As its form and position involve other considerations of some im- 
portance, it is desirable to describe it more in detail. On examining the putamen, which 
is the lignified endocarp of the fruit, we find it of an oval form, slightly flattened on one 
side, along which is observed a longitudinal groove, in which is imbedded a thick cord or 
bundle of fibres, which is easily raised from the channel, and is found to penetrate an 
aperture in the upper extremity of the groove: some fibres proceeding from the stigma 
here join it, and the cord now reduced to a thread, after its passage through that aper- 
ture into the cell, becomes attached to the testa of the seed, on a small protuberance 
near its apex; it then assumes the features of a prominent line upon it, and descends 
along one angle of the dorsal face of the seed, to near its base, then curves round this, 
and ascends along the other angle of the same face, to a hollow in the summit, beneath 
the apical protuberance just mentioned, where it disappears, describing in its course a 
horse-shoe, ‘or rather an oval curve: and this is the “ whitish raised line” described by 
Mr. Thwaites. The nourishing vessels are thus distinctly apparent throughout their 
entire course, from the external base of the putamen to the vanishing point of the thread 
just described, in one continuous line: the external portion of this cord is evidently the 
placentary axis of the dissepiment of the normal bilocular ovarium, for in the 2-celled 
and 2-seeded fruit which I have mentioned, where the seeds were separated by a distinct 
partition, this same cord was seen imbedded in the line of its axis, and became furcated 
on arriving at the summit, each branch passing through a double aperture into the two 
cells, where it attached itself to the apex of each seed, and continued in its course as the 
“whitish raised line” of Mr. Thwaites: this thread therefore bears the character of a 
raphe ; but as its form is very peculiar, it is well to compare it with the same organ in 
other genera of this family. € : à 
The development of the seed in Stemonurus is precisely similar to that in Pounontia, as 
I have elsewhere shown, agreeing even in the same relative size, shape, and position of 
the embryo, differing only in the singular extension of the raphe, and the presence of * 
large vacuity in the centre of the albumen, lined with thin pellicular plates. In P asset 
as in other genera of the Zcacinaceæ, the raphe originates as in Stemonurus ; am 
course is along the middle of the dorsal face, and it loses itself at the opposite = = 
ledonary extremity of the testa, according to the law of anatropal ia a vas 
latter genus, contrary to general rule, the descent of the raphe is along one P" sita x 
seed, and, after crossing the lower part, it ascends up the other Ke VO Meee eis poni 


from which it started, and is lost in the integument, close to the extremity of the radicle : 
which is always inclined a little towards the dorsal face, leaves 


the raphe in this course, 02 


100 MR. MIERS ON SEVERAL INSTANCES OF ANOMALOUS DEVELOPMENT 


by its pressure a corresponding groove close to the periphery of the albumen: the 
margins of the large cavity in the albumen, in which the much smaller embryo is 
suspended, also correspond with this line of circuit of the raphe, so that the albumen 
along this line is reduced to a very thin substance, and consequently is nearly separated 
into two parallel slabs. In Stemonurus I could not perceive the indication of any 
chalaza at either extremity of the inner integument, and there exists a more intimate 
union of this integument with the testa, which are both of delicate texture. In Pennantia, 
on the contrary, a distinct chalaza is visible at the cotyledonary extremity of the inner 
integument at the vanishing point of the dorsal raphe: the same exists in Mappia, and 
probably in all other genera of the Icacinacee. Mr. Bennett (Plant. Jav. rar. 131) 
quotes the observations of Mr. Brown and of M. Brongniart to show that the eversion of 
the raphe upon the dorsal face of the seed is of frequent occurrence in Celastraceæ, 
Rhamnacee, Aquifoliaceæ, and Caprifoliaceæ, and is fully confirmed by his own experience 
in the seeds of Euonymus and Rhamnus, where it is frequently everted, though often only 
laterally displaced ; and he entirely adopts the opinion of Mr. Brown, that this eversion of - 
the raphe in no way militates against the universal law of anatropal development ; for on 
examining these occurrences at an early stage of their growth, he invariably found the 
ovule with its raphe on the ventral side, or that next the placenta, and he observed that 
its subsequent appearance on the opposite or dorsal face, is due to an evident torsion of 
the upper extremity of the raphe, or short funiculus, during the increment of the seed, 
and not to any original development. The eversion of the raphe in the Jcacinacee, 
together with a Similarity in other essential points of structure, strongly prove the close 
affinity of this order to the Celastraceæ, Aquifoliacee, and other families which I proposed 
to img : buen s deron un (the Dryales), nearly allied to the Rhamnacee. 
pe : oned structure in Stemonurus, a curious deviation from the usual 
evelopmen of anatropous seeds occurs. In the organization of most of its parts there 
exists, as before mentioned, a perfect analogy with that found in Pennantia : the seed is 
albuminous, and the radicle of the embryo is superior and pointed to the hilum ; but while 
in Pennantia we trace the raphe in a straight line to the opposite or cotyledonary extremity, 
R^ red anatropous seeds, we find in Stemonurus, owing to the singular retroversion 
er > AS m Wee. beyond the point where it ought to have 
rn q escent. 9 the summit—to the point of its disappearance 
— de that this unusual eircumstance is not attended by a corresponding 
ag e es eer rine mr 
inversion of the embryo and its amet eum i Aper s ER 
We not see, from the course of the rod ih esee ius, imm d ee Eu 
er Saeed 9, an indication that a complete 
ae n op action of one or both of the integuments has occurred, while 
SE es . Ri ~ deg effected only a half-gyration, or simple anatropal move- 
E eura therwise account for the peculiar form of the raphe in this case? 
s: again to this point when I come to speak of other developments which appear 
ofer much analogy with this phænomenon. å : 


In ord: des ` 
er to prepare the way for this investigation, it is of some interest to determine 


OF THE RAPHE IN SEEDS. 101 


the nature of the two thin membranaceous plates, or cellular strata, that line the cavity 
of the albumen, as before described, and I think these may be referred, with little doubt, 
as we shall see in other well-authenticated cases, to the existence of a large persistent 
embryo-sac which has not been absorbed, as it usually is in most seeds. In the case in 
question, at the upper extremity of this cavity, the embryo only one quarter of its length 
is found, and here the radicle, pointing upwards, terminates in a long mucronate point, 
which is a kind of suspensor, by which it is pendent in the mouth of the cavity, and where 
it is enveloped in a small quantity of glutinous matter, which lines the internal surface of 
the sac. Instances of the existence of this cavity in the axis of albuminous seeds are not 
unfrequent, but the discovery that it originated in the persistence of the embryo-sac is 
due to Mr. Brown (App. King’s Voy. p. 21). 

The first consideration that arises is the question whether the existence of this persistent 
embryo-sac has any necessary connexion with, or has had any share in producing the phæ- 
nomenon of the retroversion of the raphe, and the inversion of the embryo in regard to it, 
in the manner before stated; whether the two latter unusual circumstances are independent 
of each other, or whether they are necessarily connected. The only method of forming 
any conjecture on this head, is to search for facts that have any collateral bearing on the 
question. In Diospyros we meet with a parallel case of a large cavity in the middle of 
an albumen, which is open at the hilum, and is in like manner lined with an adherent 
embryo-sac; the embryo is found in the outer end of the much larger cavity, the end of 
the radicle almost protruding out of its mouth: the albumen is deeply divided into 
numerous lamelliform folds, as in Amona, and is covered by two thin integuments, of 
which the inner one is plicated within the interstices of its lamellar clefts, while a distinct 
raphe, adhering to the outer integument, proceeds along a groove formed in one of its 
marginal sides, from the basal hilum to the summit, or opposite cotyledonary extremity, 
where it disappears. We have here therefore a parallel instance of the existence of an 
open cavity in the albumen, lined with the embryo-sac, but this circumstance is accom- 
panied by the presence of an ordinary raphe, while both the seminal tunics and the 
embryo have gone through the usual process of anatropal inversion. 

In Anona we meet with a complete peripherical raphe, as in Stemonurus, but the 
embryo-sac has disappeared; there is therefore no cavity, as the embryo lies tightly 
imbedded in the substance of the albumen, and the radicular end of the embryo, with a 


very short mucronate point, or suspensor, is directed to the spot where the raphe, after 


completing an entire circuit round the periphery of the seed, disappears close to the hilum. 
as indicated by 


In this instance we find that a complete gyration of the original tunics, 
the raphe, and only a semi-gyration of the embryo, have taken place ; pam m a 
perfect analogy with the structure of Stemonurus, excepting the presence en em - 
sac and its accompanying cavity. These two instances seem to prove t : e presen 
of this albuminous cavity and the complete gyration of the raphe do no — aá 
accompany each other, and they leave us as much in the dark as ever, regarding the ca 


of the phænomena observed in the seed of Stemonurus. 
In Opuntia we have another example of a completely annular raphe, på — 
imbedded in a small quantity of albumen, is quite campulitropal : the raphe 


102 MR. MIERS ON SEVERAL INSTANCES OF ANOMALOUS DEVELOPMENT 


pleting its circuit penetrates the testa through the diapyle, at a point where the two 
extremities of the recurved embryo are brought into close contiguity, although separated 
by a peculiar inflexion of the inner integument, close to the chalaza, so that this integu- 
ment is here bisaccate, one recess receiving the end of the radicle, the other the extre- 
mities of the cotyledons. In this case there appears no vestige of any embryo-sac, and 
although the raphe is completely cyclotropal*, only the cotyledonary or chalazal extremity 
of the embryo has accompanied it in effecting an entire revolution, while its radicular end 
has remained anatropal, having experienced only a half-gyration. This example therefore, 
although analogous in some respects, affords no assistance towards an explanation of the 
phænomena of Stemonurus. : 
In the Plumbaginacee, we have a remarkable instance of the complete gyration of the 
ovule under very different circumstances : in an early stage it is simply inverted according 
to the ordinary course of anatropal development, that is to say, a semi-gyration of the 
entire ovule on its centre, accompanied by the usual extension and adhesion of the placen- 
tary sheath or future raphe on the ventral side of the primine: subsequently a farther 
growth of the placentary sheath, or rather of the funicular attachment of the ovule, takes 
place, which becomes prolonged into a free cord, which, by its growth, again pushes the 
ovule onward, so as to turn it farther round on its centre, another half-revolution: the 
result is, that the funicular cord, arising in a free state from the base of the ovary, attains 
the summit of its cell, and from it the anatropal ovule is suspended; its cotyledonary 
or chalazal extremity thus returns to its normal position in regard to the carpel, after 
having performed a complete revolution on its centre, at the same time that it has only 
effected a half-revolution in respect to the point by which it was originally attached to 
the placenta. The embryo during all these changes continues perfectly straight; the 
placentary sheath or first expansion of the funicular cord remains adherent to the primine 
in the usual manner, while its subsequent extension is permanent as a free cord, so that 
the ovule, though strictly and simply anatropal in respect to the raphe, is in fact 
cyclotropal in.regard to the carpel. This development is again different to that 
observed in Stemonurus, inasmuch as the embryo has remained stationary, or what is 
almost tantamount, has made a complete gyration on its centre. One parallel circum- 
stance however exists in the form of the albumen, which, as in that genus, consists of two 
pun el plates united on their periphery by a thin annular zone corresponding to the 
commissure of the cotyledons. 7 
In Diospyros, to which I have before alluded, the embryo, which remains in its original 
sac after the expulsion of the redundant amniotic secretions, appears to have preserved its 
relative position in regard to the integuments and raphe, and to be truly anatropous. It 
would however be desirable to ascertain, by the examination of fresh seeds, whether it 


offers any indication of a suspensor. I conjecture from the appearance of the dried seed, 
that a portion of the sac will be found to protrude from the mouth of the albuminous 
cavity which it lines, and from its great tenuity to have withered away, for I have always 

* By this term, I Propose to distinguish an ovule, where its chalazal point, and with it the raphe, effects a complete 


. Cyclical gyration: it is an extensi 


OF THE RAPHE IN SEEDS. 103 


found the end of the radicle extending a little beyond the mouth of the cavity : this con- 
jecture is confirmed by the drawings of Gærtner (De Fruct. tab. 208). A similar per- 
sistence of the embryo-sac, and its forming the lining of a large cavity in the albumen, 
which is thus nearly separated into two parallel plates, and the occasional protrusion of 
this sac beyond the limit of the albumen, occur also in Strychnos and Ignatia, as may be 
seen from the figures given by Gærtner (De Fruct. tab. 189). The protrusion of a portion 
of the embryo-sac, and with it of the radicle, in the manner above conjectured to exist in 
Diospyros, is known to occur in Myzodendron, where it is well figured by Dr. Hooker in 
his * Flora Antarctica,’ plate 104. fig. 18 & 19; and in plate 105. fig. 17, 18 & 19*. A 
singular example of the persistence of the embryo-sac is afforded in Marantacee : this 
occurs in the genus Thalia, where two large vacuities are seen in its copious albumen, on 
each side of the embryo: these are the persistent sacs in which two abortive embryos had 
existed: this curious fact was first recorded by Mr. Brown nearly fifty years ago (Prodr. 
p- 807): it is one of the few instances that occur, where several embryo-sacs are generated 
in the same ovule. 

In regard to the origin of the albumen in seeds, it was shown by Gærtner (loc. cit. 
Introd. 188), and Mr. Brown confirms it as a well-established fact (Linn. Trans. x. 36), 
that it is merely that condensed portion of the amniotic fluid which remains unabsorbed 
by the embryo at the termination of its growth. This excessive portion of the fluid is 
generally expelled from the embryo-sac (quintine, Mirbel), and is deposited and consoli- 
dated in the cellular tissue of the quartine, which lines the tercine (nucleus, Brown) when 
- it constitutes the albumen, and in ordinary cases the sac becomes absorbed without 
leaving any trace of its existence. Besides the instances of its permanence just given, its 
existence under other peculiar circumstances was first shown by Mr. Brown, in the 
Nympheacee, where a portion of the redundant amniotic fluid forms a separate and 
second albuminous deposit, immediately surrounding the embryo, which he has aptly 
named the vitellus, and which in the ripe seed is found lodged in an exserted portion of 
the embryo-sac : it is therefore always seen at one extremity, outside of the great mass of 
the ordinary albumen: when this occurs, it is accompanied by the appearance of sr 
tubular vacant space in the axis of the albumen which is the opposite “ere > 
persistent embryo-sac, of which the other portion containing the VAN À eonun 
tion. This has been shown to exist among Dicotyledones, in the Nymphæaceæ, Le: 
bacee, Saururacee, Piperaceæ, &e., and in several cases among Monocoty — ( pe sj 
dix, King’s Voy. p. 21). A very excellent illustration of this 7 e n 
the admirable work of Dr. Asa Gray (Gen. Pl. Un. St.), where evidence of its preser 

3 ia, in pl. 38, 39, 43 & 44. 
shown in the seeds of Nymphæa, Nuphar, Cabomba, and Brasenta, MP? be formed 
Mr. Brown observes (loc. cit. p. 22), * that the albumen, Pops ly so called, ng (embryo- 
by a deposition or secretion of granular matter in the ufin af the vues pri 
Sac) “or in those of the nucleus itself” (tercine or quartine); or lastly, 


3 in Ceratophyllum the principal 
* Schleiden (Linnæa, xi. 526. tab. 11. figs. 12, 13) has shown the curious fact, that in Ceratop 


eee : le. 
i hi ine seed remains investing the plumule, 
Portion of the embryo is developed outside of the embryo-sac, which in the T. the embryo-sac around the plumule, 


and enclosed within the cotyledons; a small quantity of albumen is deposited in imple thin integumentary covering. 
but none is formed outside of the large cotyledons, which are invested only 2e 


104 MR. MIERS ON SEVERAL INSTANCES OF ANOMALOUS DEVELOPMENT 


substances” (amnios and nucleus) “having these distinct origins, and very different 
textures, may coexist in the ripe seed, as is probably the case in Scitaminee.” In that 
family the embryo is enclosed in a fleshy vitellus, and this again is surrounded by 
albumen: owing to the difference of texture and consistence of the vitellus, its albu- 
minous nature has been denied by some botanists. I have however met with an instance 
of a double albumen in Boldoa fragrans, a Chilean tree belonging to the Monimiacee : 
here the principal oval mass of the kernel of the seed consists of fleshy albumen appa- 
rently formed by two distinct deposits: the rather small embryo is seen with its cotyle- 
dons spread out quite flat by pressure, and as if they were seated astride upon the summit 
of the inner mass of the albumen, with its short radicle standing upwards; and the whole 
is enclosed within another concentric portion, much softer in substance, and formed of oily 
granules of much larger diameter than the close cells of the main inner portion, which is 
whiter, more dense, opaque, and fleshy, and from which it is easily separable in every 
part. Dr. Lindley (Veg. Kingd. p. 298) gives a good sectional analysis of this seed, but 
he has evidently not noticed the difference of the two kinds of albuminous deposit: the 
whole is covered by the proper inner integument, well distinguished at the extremity 
contrary to that where the embryo is seated, by a large dark chalaza, somewhat adherent 
to the outer tunic, which is recognized as being the testa, by its linear raphe, which is 
partly free near its origin. He justly claims for the Monimiaceæ à near affinity to the 
Myristicacee, for the form and position of the embryo quite correspond with those of 
Pyrrhosa, where, had its internal cavity been filled by a second deposit of albuminous 
matter, and were its outer coat not split into lamellar plates, we should have quite the 
seed of Boldoa : the whole plant of this latter genus, and especially its seed, bears a 
strong aromatic smell and taste, similar to that of the Nutmegs. The fact of the 
existence of a double albumen here fully realizes the prediction above quoted of 
Mr. Brown. This structure is of a generic, not of an ordinal character, for it does not 
exist in Mollinedia, å genus numerous in species, nor generally in Citrosma, although I 
have found it occur in one species of the latter genus. Brongniart also has demonstrated 
that albumen is generally formed in the quartine or cellular lining of the tercine, in which 
case he calls it the *perisperm, and at other times, though more rarely, within the 
embryo-sac, and then he gives it the name of « endosperm*.’ Boldoa hence possesses 
both a perisperm and endosperm, one concentric to the other, although the embryo-sac | 
has entirely disappeared ; but if the latter had remained persistent, if the endosperm had 
been absorbed, and the embryo and the perisperm had retained their present form and 
position, the seed of Boldoa would have been like that of Stemonurus, with the exception 
of the peculiar form and termination of its raphe. 

In this stage of the investigation, it is desirable to recall to our memory the nature and 
origin of the embryo-sac. This vesicle, called by Gærtner the sac of the amnios, was 
ges Malpighi and Grew, who showed that it originated from that point of 

es ch I have called the gangylode, or the point of vascular connexion of the 
TAN ia tenis with the main body of the ovule, and with the placentary sheath, 

rwards becomes the chalaza of the inner integument, the diapyle in the 


* Ann. Se, Nat. xii. 265. 


OF THE RAPHE IN SEEDS. 105 


testa, and where the raphe terminates. Gærtner farther affirms that the amniotic fluid 
is secreted at this point of the gangylode, which is also the opinion of Mr. Brown (Linn. 
Trans. x. 37). This point is sometimes distinguished by a peculiar protuberance, within 
the cavity of the main body, which Mirbel calls the chalazal appendage (appendice chala- 
zienne), and he says that at this point the embryo-sac is first developed. We are indebted 
to Mr. Brown (Append. King's Voyage) for nearly all the information we now possess 
respecting the nature and origin of the several parts of the ovule, and the changes they 
undergo during the growth and perfection of the embryo, and it is impossible to estimate 
too highly the importance and merit of these admirable observations, the truth of which 
is unquestionable. A large share of credit is due to M. Brongniart, who about the same 
time, and to M. Mirbel, who three years afterwards, contributed many important facts on 
this subject, all confirming the previous observations of Mr. Brown. The latter able phy- 
siologist brought together all this information in his two celebrated memoirs “On the 
Development of the Ovule,” &c. (Mém. Acad. Sci. Paris, ix. p. 609 & 629): he there 
gives a regular nomenclature to the several parts of these developments, and reduces all the 
evidence into a beautiful system, which has been adopted by all, and which has remained 
unquestioned now for nearly thirty years; but true as it may be in a general point of 
view, the evidence I have now brought together will show that the laws considered so 
universal, fail in their application in many cases, perhaps in far more numerous instances 
than are here offered, or will at first be credited. The existence of the two outer tunics, 
the primine and secundine, had long been known; but their remarkably peculiar cupuliform 
shape, seen alone in a very young state, was first shown by Mirbel, and by him also the 
metamorphoses of the main body of the ovule, called ‘chorion’ by Malpighi, and ‘ nucleus’ 
by Mr. Brown, who first explained its functions, were now more fully demonstrated. 
Mirbel, who called this ‘nucleus’ the *tercine, showed that it is lined with a peculiar 
tissue, which sometimes becomes transformed into another tunic, the * quartine,’ and which 
in Statice he describes as forming a completely closed vesicle. Another body, which he 
denominates the ‘ quintine,’ is still farther developed in the interior of the quartine : this is 
the * sacculus colliquamenti’ of Malpighi, the ‘amniotic sac’ of Mr. Brown, the * embryo- 
sac” of M. Brongniart, the point of origin of which has been noticed above; when this 
vesicle is wanting, which perhaps more frequently happens, it is then the quartine that 
becomes resolved into the embryo-sac. According to Mirbel, one end of this sac is 
attached to the point of the gangylode, while the other is fixed to the summit of the 
tercine or quartine: it gradually swells from top to bottom, thrusting away the tissue 
q gr y : lei 
that surrounds it. Brongniart first noticed the fact of the descent of the boyaux of gm T 
grains, through the stigmatic channel, but failed in tracing their farther parga T 
s 4 . : e - » 
mode of their action in effecting the fertilization of th ovule, a discovery ges Fu - 
According to the observations of more recent physiologists, it pd Re å 
every case of fertilization of an ovule, the boyau of a pouce ve 8 


stigmatic channel, and is conveyed through the apical orifices of the primine and secun- 


i i i he tercine, thus reaches the embryo-sac, in 
. dine (the micropyle), and passing through t A ee : pm 
the na of which is a small globular body, called by Meyen ‘the primary utricle, 


which body, consequent on its osculation with the boyau, subsequently swells J becomes 
VOL, XXII. 


106 MR. MIERS ON SEVERAL INSTANCES OF ANOMALOUS DEVELOPMENT 


the nascent embryo. We will not here touch upon the still disputed point, whether the 
process of fertilization is the result of mere impact, or whether the boyau enters into the 
primary utricle; but will pass on to the fact admitted by all, that a new globular deve- 
lopment subsequent to this action (the future embryo) is generated within the primary 
utriele, which is suspended within the sac by a delicate thread, often extending with 
the growth of the embryo in its young state: this is called the suspensor, which in some _ 
few instances becomes highly developed, though more frequently it is of no great length, 
sometimes remaining as a short distinct thread that terminates the extremity of the 
radicle, and which I have pointed out as existing in the embryo of Stemonurus. After 
the fecundation of the embryo, in the manner just mentioned, the embryo-sac as before 
stated, either by absorption, or by amalgamation with the surrounding tissues, generally 
disappears; but in some few cases, as we have seen, it remains persistent, and Stemonurus 
affords an instance of this occurrence. . | 
I have entered into these details upon the nature and function of the embryo-sac, with 
the view of considering, whether a different amount of circumversion of this sac, respect- 
ively to that of the tunics of the ovule, may not have taken place in Stemonurus, which 
would thus account for the phzenomena under consideration. May we not conclude, with 
some degree of confidence, that the embryo-sac has remained to form a cavity in the 
albumen, which has been copiously moulded around it, by amylaceous granules flowing 
from the redundant amniotic fluid deposited among the cellular tissue of the surrounding 
envelope? Has the embryo-sac in this case moved a half-revolution on its centre more 
than the coats of the ovule, or vice versd? I cannot venture to affirm the fact, but 
the evidence is certainly presumptively in favour of such a conclusion. This indeed 
appears to me the only key to the solution of the paradoxical difficulty in question. It 
is, however, a point that can only be settled by observation on the growth and develop- 
ment of the ovule of Stemonurus, and as it involves a topic of great importance in a phy- 
siological and structural point of view, I would earnestly impress it on the attention of 
every botanist who has an opportunity of examining the ovule in a living state. In the 


mean time I call attention to the following evidence, which appears to favour this view of 
the question. 


perfectly with the other. 

t I may also affirm without hesitation that the genus 
Peruviana, I have examined several species and made 
future time to give the full characters of Mollinedia, Ci 


Tetratome of Pöppig is identical with Mollinedia of the * Flora 
drawings and analyses of the living plants. I purpose at some 
trosma, and Boldoa, for which I possess ample materials. 


OF THE RAPHE IN SEEDS. 107 


hilum, near the bottom of the ventral margin of the nut, and at last arriving at the basal 
point, it terminates abruptly by penetrating the integuments: between this point and the 
hilum a very short space intervenes, which is obsoletely angular, and immediately at this 
angle, close to the margin, the small embryo is found imbedded in a large albumen, the 
radicle pointing outwards, and the two short fleshy cotyledons, greatly divaricated, being 
directed towards the vertex of the seed. Here then, as far as regards its direction with 
the hilum, the embryo is anatropal, having become inverted half a turn on its centre from 
its normal position, while the integuments have performed an entire gyration, by which 
the embryo is deprived of its anatropal characteristic. All the species of Mollinedia 
which I have examined have regular anatropal seeds, with one exception, where the 
raphe is cyclotropal, as in Citrosma. We have thus in Citrosma a verification of the 
fact observed in Stemonurus, the raphe terminating at a point exactly contiguous and 
corresponding to the radicle of the embryo, a fact quite incompatible with the laws of 
structure so beautifully expounded by Mirbel: it is a development that has hitherto been 
considered impossible; but I will proceed to show that these are not the only instances 
of the same occurrence we find in the structure of seeds. 

In the development of the seed of the Queurbitaceæ, we meet with a structure that 
offers much resemblance to the anomalies observed in Stemonurus; we find also other 
features, not only difficult of explanation, but contrary to all our ideas of the nature and 
function of the raphe. To aid us in the solution of these phænomena, we will first recur 
to the detailed account of. the development and growth of the ovule of Queumis Anguria, 
L., from the observations of Mirbel (Mém. Acad. Sci. Paris, ix. 621. tab. 1). The 
growth of the anatropal ovule is here shown in all its earlier stages, and we see that by 
the time of the fall of the flower, the micropyle of the primine and secundine becomes 
quite closed over the tereine; soon after which, a fine thread is observed to grow out of 
the micropyle, which was first noticed and described by Brongniart, in the ovule of Pepo 
macrocarpus and Momordica Elaterium. As the growth of the ovule proceeds, this 
tubular thread becomes covered by an envelope formed of cellular tissue: according to 
the drawing of Mirbel (Joc. cit. tab. 1. fig. 10), it is a prolongation of the mouth of the 
embryo-sac; but Griffith says (Posth. Notes, 153), that although he had observed the 
same fact in several instances in Cucurbitacee and Asclepiadaceæ, he was never able to 
trace any connexion with it and the embryo-sac, and was rather disposed to consider this 
thread as formed by the boyaux or pollen-tubes, an opinion confirmed by the observations 
of Schleiden, and which appears the more probable conclusion, as no remainder of this 
thread is visible in the ripe seed. At the period above mentioned, the bottom of the 
embryo-sac breaks away from the gangylode, becoming thus suspended by its — and 
the embryo is formed in its summit: there is no appearance 1n the raphe at this time to 
indicate any other change than the ordinary anatropal development. The same course of 

TE i i ir generally among the Cucurbitaceæ. We will 
metamorphosis is said by Mirbel to occur ge y ng ppt 
now compare these changes with the development found in the ripe "m d z = 
analysis of Citrullus Colocynthis as a fair example of the structure ot the ur 
family. 


Here the seed is obovate, much compressed, its hilum being an open slit along its 


P 


108 MR. MIERS ON SEVERAL INSTANCES OF ANOMALOUS DEVELOPMENT 


narrow basal extremity, which is enclosed in a short fleshy sheath, part of the funicle, from 
which it has broken away, and by which it is connected with the placenta, and suspended 
in the pulp of the fruit: a groove runs parallel with the margin on each face close to the 
periphery, forming in this manner a prominent zone all round the edge of the seed, which 
is broader towards the base, and which thus forms a short auricular expansion on each 
side of the hilum. On a former occasion (p. 92) I described the outer crustaceous 
tunic, hitherto considered as the testa, but which I shall prove to be of the nature of an 
aril, formed subsequently to the fertilization of the ovule, around its original integu- 
ments : it was there shown to be formed of three series of deposits, the epiderm, meso- 
derm and endoderm: the epiderm is a thin delicate membrane that covers the whole of 
the seed, and is extended beyond the hilum in the manner above-mentioned, as an exten- 
sion of the sheath of the umbilical cord; the fleshy mesoderm has been before deseribed, 
its numerous branching fibres being emanations from the bundle of simple vessels that 
fill the sheath of the funicular cord, and that surround the thread of spiral vessels pro- 
ceeding from the placenta, that terminates in the raphe of the seed; the principal portion 
- of the tunic is the endoderm, which is, in fact, a crustaceous compressed sac, enclosed in 
the above-mentioned vesicular terminal enlargement of the sheath of the umbilical cord ; 
it forms a casing closed all round the seed, except at the hilum, where it has a long open 
slit, within the mouth of which is a considerable open space filled with pithy loose cel- 
lular tissue, which fills up this interval between the pointed extremity of the enclosed 
nucleus and the mouth of the crustaceous covering: there is no connexion whatever 
between any part of this crustaceous coating and the enclosed seed, which lies quite free 
within it, and fills up its cavity. The raphe, consisting of a number of spiral vessels 
enclosed in a delicate tube, is first seen to issue from the umbilical cord; it then finds its 
way through one of the basal auricular lobes of the endoderm, in which a channel is seen _ 
for its passage, after which it enters into the clear space between this crustaceous aril 
and the integuments of the contained seed, and under the form of a perfectly free delicate 
white thread, it runs all round the margin of the seed, forming a complete circle, until it 
again reaches the space within the mouth of the hilum, and when in the midst of the 
deposition of tissue before mentioned, it changes its course suddenly inwards, and termi- 
nates in the conical neck of the integument of the seed, by which the latter becomes thus 
suspended : this conical neck is of a dark brown colour, and is evidently the true chalaza 
of the integument. The covering of the enclosed seed is thin and membranaceous, gene- 
rally of a greenish hue, is quite smooth and evidently composed of two, if not of three pelli- 
cular membranes, agglutinated into one thin integument. I have sometimes found this 
structure more clearly demonstrated where the nucleus has become withered from its 
fall ‚dimensions ; it then appears as a large, perfectly transparent cyst, enclosing the 
oR aang = by transmitted light displays the deeper colouring of the chalaza 
irent si this re which “ppears surrounded by another pellicle more trans- 
sn — à ect aig of the spiral vessels in the suspending thread, up to the 
Bits. covering-of the = Pa ere perfectly distinct. At the other end of the integu- 
digo tack edm es » the one contrary to that of its suspension, is seen another 

6, over the cotyledonary extremity of the embryo, and therefore 


OF THE RAPHE IN SEEDS. 109 


at the point where, according to the laws of structure, the chalaza ought to be found in 
all cases where a raphe exists: it is precisely in the situation ascribed to the chalaza by 
Mirbel, in his figure before quoted of the ovule, after the formation of the embryo. It is 
however only a false chalaza, and is most probably a thickening of the mouth of the 
micropyle (exostome of Mirbel) of the ovular integuments, a circumstance of frequent 
occurrence in certain seeds; or it may possibly be ascribed to a scar of the embryo-sac, at 
the point where it has broken away from the gangylode, as figured by Mirbel (loc. cit. 
fig. 10. letter f). That it cannot possibly be the true chalaza, is most evident, because 
it has had no apparent connexion with the outer crustaceous tunie, which exhibits no 
corresponding scar or trace of any former connexion at this point; and still more obvi- 
ously, because the raphe passes straight across it, without holding the slightest commu- 
nication with it. The embryo, which has no albumen, fills the whole cavity of the inte- 
guments in the ripe seed, the radicle being directed towards the hilum and to the 
terminating point of the raphe, and its extremity being covered by the true chalaza; its 
cotyledonary extremity is in the opposite direction, terminating at the areolar micropyle 
or false chalaza. The fact of the existence of the free cord of spiral vessels as above- 
mentioned (the raphe), running between the outer crustaceous tunic and the inner 
integuments of the seed, was evidently known by Mr. Griffith, who notices it (Posth. 
Notes, p. 182); but he omitted to observe the passage of this thread over the areole, 
which has always been considered as the chalaza, and also its course round the entire 
periphery, as well as its termination at the radicular extremity of the membranaceous 
integuments : the presence of the true chalaza at this point also escaped his observation. 
I do not find any mention of this curious structure by any other botanist. 

We thus observe the same phænomena existing in all the genera of the Queurbitacee 
that I have described as being found in Sfemonurus, Anona, and Qitrosma, that is to say, 
where the cord of the raphe makes a complete circuit of the seed, terminating at the same 
time at the radicular point of the integuments, instead of the cotyledonary extremity, as 
it ought infallibly to do, according to the acknowledged laws of structural development. 
What has been the nature of the metamorphoses within the ovule, that have produced so 
manifest a deviation from the ordinary course of structure? We have it demonstrated 
on the high authority of Mirbel, that up to the period of the impregnation of the ovule, 
the raphe, the chalaza, and the primine and secundine (then agglutinated into one coat), 
had only performed a simple anatropal inversion from their normal position; but we find 
that in the interval between this period and the ripening of the seed, all these parts have 
experienced a farther circumversion, so as to complete an entire circle, while the embryo- 
sac, or at least the embryo, has retained the same position which it held at the period of the 
simple anatropal inversion of the ovule. Mirbel notices, at this last-mentioned period, the 
first indication of the growth of the arillæform coating over the BEE Seton) at 
this epoch as two layers of cellular tissue coating that integument, and as pit i 
(loc. cit. fig. 10. letter 4): these layers of tissue are unquestionably the ru ; p d 
secretions that subsequently form the arillus, under the crustaceous cip a k , 
for which it has always hitherto been mistaken : it is doubtlessly owing a t a Lars am 
of this coating that the subsequent movement of the coats of the ovule has been hi 


110 MK. MIERS ON SEVERAL INSTANCES OF ANOMALOUS DEVELOPMENT 


from view, and has thus remained unnoticed. The assurance that such å movement 
must actually have taken place, is proved by the form of the raphe, and the very different 
position of the integuments in the ripe seed to that which they ought to have, according 
to the usual structural development. It is more than probable that the embryo-sac still 
remains immediately investing the embryo, in the Cueurbitaceæ, but this is not easily 
discernible, as there is no albumen, and as the primine and secundine, and probably also 
the tercine, become resolved into delicate membranes. Mirbel showed that about the 
period of fecundation, the embryo-sac had broken away from the gangylode ; and we must 
assume, in order to account for the changes that subsequently occurred, that. it also 
separated itself afterwards at the summit from the micropyle; but if the integuments 
received a subsequent amount of torsion, why did not the embryo-sac, or at least the 
embryo, follow the same movement? What could have retained the embryo in the same 
position it previously held, during the second inversion of the primine and secundine? 
The whole of this æconomy is so extraordinary, as to call for the especial investigation of 
some able botanist, aceustomed to accurate and delicate examination. I have simply 
announced the facts as I have found them, leaving it to be determined how far the hints 
above given, respecting the nature of these changes, may prove well-founded, or how far | 
the whole matter may be modified by farther evidence. 

The facts detailed in the preceding part of this paper clearly show, that the genus 
Urandra of Mr. Thwaites bears no relation whatever to the family of the Olacacee, 
and we have irresistible evidence that it differs in no respect from Stemonurus, 
with which genus it must remain, like Gomphandra, another of its synonyms; it 
agrees with it in its cupuliform, 5-toothed, persistent calyx, its five acuminated linear 
smooth valvate petals, its five alternate stamens, with thick filaments clothed at their 
summit with clavate hairs, oblong anthers somewhat divergent below, a conical ovary 
encircled at its base by a small annular gland, and unilocular, with two suspended ovules, 
a short style, and a small subcapitate stigma. The drupe and seed, as described by 
Mr. Thwaites, agree in every respect in their extraordinary development with what I 
have shown to exist in Stemonurus, and there is not one single feature, among those 
described, that I can discover, at variance with that genus. The plant of Mr. Thwaites, 
which appears to differ specifically from others on record, must therefore bear the name 
of Stemonurus apicalis. 

The remarks of Mr. Thwaites contained in his paper above cited, relative to the Ceylon 
species of Stemonurus described by me, will form the subject of a distinct notice. 


OF THE RAPHE IN SEEDS. 111 


EXPLANATION OF THE FIGURES. 


TAS. XIX. 


Fig. 1. Is a seed of Stemonurus polymorphus :—natural size. 
Fig. 2 & 3. The coriaceous endocarp deprived of its pulpy covering, seen on its dorsal face and on its 


Fig. 4. 


Fig. 
Fig. 


Fig. 
Fig. 


5. 


6. 


is 
8. 


side. 


The same on its ventral face, showing the channel in which is imbedded the cord formed of the 
axile placentary nourishing vessels. 

Is a section of fig. 3. magnified :—a. is a foramen in the summit of the channel through which the 
raphe passes into the cell of the putamen; à, the longitudinal channel in which the placentary 
cord is imbedded, and which passes into it through a tubular passage leading from the base. 
The seed removed from fig. 5. seen on its edge :—a. is the free portion of the raphe by which the 
seed is suspended ; it is a continuation of the axile placentary cord c; 5. is the raphe imbedded 
in the integument. l 

The same viewed upon its dorsal face, showing the singular course of the raphe. 

The same observed from above: the same letters refer to the same points in the three last figures. 


Fig. 9. The integument removed and spread out to show the form of the raphe more distinctly, 


Fig. 10. The albumen. 
Fig. 11. A transverse section of the same, showing the cavity formed by the persistent embryo-sac. 
Fig. 12. A longitudinal section of the same. 
Fig. 13. The persistent embryo-sac removed from the albumen with the embryo in its natural position. 
Fig. 14. The embryo removed. 
Fig. 15. Same still further magnified, 
Fig. 16. Is a seed of Anona sylvatica :—natural size. . 
Fig. 17. The same seen sideways. . 
Fig. 18. The same magnified, showing the marginal expansion of the crustaceous covering and its open. 
cavity at the base. | 
Fig. 19. One side of the cell removed, to show the enclosed seed and the hollow chamber at the base. 
Fig. 20. A transverse section of fig. 18, showing the raphe lying in a groove of the albumen along each 
side. : : : iling of 
Fig. 21. The seed removed, showing the annular form of the raphe and its connexion sy "e på 
the basal chamber of the shell: from the centre of the latter a mucronate point, as long as t 
hollow chamber, projects, and at each angle two threads are observed in the same space. 
Fig: 22, The same viewed edgeways, showing the peripherical form of the raphe. ter 
> 23. The raphe separated and remaining attached to = ui ris Mont hamide: 
ig. 24. A longitudinal section of the albumen with the enelosed embryo. à 
Fig. 25. The din portion of the same more highly magnified, showing the embryo, and the hollow in the 
base of the albumen caused by the impression of the basal chamber. 
Fig. 26 & 27. The embryo seen in front and sideways. 


Fig. 28. A fruit of Citrosma Aposyce, Mart. 
Fig. 29. A seed capped by a glandular enlargement of the style. 


F ig. 30. The same deprived of its cap. 
Fig. 31. The 


Fig. 32 & 33. The same magnified, seen in front and sideways, showing 
Fig. 34. A longitudinal section of the same, showing its hollow hilum an 


echinate nut. 
the flat expansion of its border. 


d the enclosed seed. 


same with its fleshy epiderm removed, showing an 


119 MR. MIERS ON ANOMALOUS DEVELOPMENTS OF THE RAPHE IN SEEDS. 


Fig. 35. The seed detached, showing the annular raphe :—a. commencement of the raphe at the hilum ; 
b. termination of the raphe. 

Fig. 36. The same seen edgeways. e 

Fig. 37. The raphe detached :—a. its commencement at the hilum; 5. its termination at the chalaza. 

Fig. 38. Section of albumen, showing a minute embryo near its base :—c. relative position of embryo 
with the commencement a, and the termination 5, of the raphe. 

Fig. 39. Base of albumen with embryo greatly magnified, the same letters referring to the same parts. 

Fig. 40. An anatropal ovule of Cucumis Anguria near the period of its fertilization ; after Mirbel. 

Fig. 41. The same seen just after impregnation; also after Mirbel :—a. the primine; 5. secundine; 
c. tercine or nucleus; d. quartine or embryo-sac suspended at the micropyle or mouth of the 
primine and secundine; e. base of embryo-sac broken away from its attachment to the chalazal 
point of the tercine ; f. the nascent embryo hung in the mouth of the micropyle by its suspen- . 
sor; 9. the gangylode or point of union of primine, secundine and tercine, and the future cha- 
laza; h. two layers of solid cellular tissue forming a distinct coating secreted and deposited over 
the primine; i. the raphe originating in the funicular cord, and terminating at g, the gangylode 
or chalazal point; Å. a pollen-tube inserted within the micropyle. 

Fig. 42 & 43. A seed of a species of Colocynth from the Pampas of Buenos Ayrés, seen in front and 
sideways. 

Fig. 44 & 45. The same magnified, showing its hollow hilum at f. 

Fig. 46. The same seen from below, showing the peripherical zone and the basal hilum. 

Fig. 47. A longitudinal section of the same :—a. the crustaceous tunic; 5. a nearly annular and perfectly 
free raphe lying between the crustaceous covering and the true integuments of the seed: it 
first passes through a tubular channel c on one side of the base of the crustaceous covering, and 
takes its course all round the enclosed seed, and again arrives at the base, disappearing in the 
loose cellular pith that fills the hollow space within the hilum ; the spiral threads appear to 
separate here and again unite to form a kind of suspensor to the seed; d. is a reddish-coloured 
areole on the summit of the integument, at the spot where the chalaza should exist according 
to ordinary rule, but the raphe certainly does not enter or terminate here, but passes freely 
over it, like a bisecting line; e. is another dark-coloured areole around the neck of the suspensor 
at the opposite extremity, which appears to unite with the raphe, for it contains spiral vessels ; 

J. is the hollow chamber within the hilum filled with loose pithy cells. | 

Fig. 48. The same viewed from the summit, showing d the false chalaza, and the manner in which the 
raphe crosses it. 

Fig. 49. The raphe detached, apparently connected by very minute threads with the neck of the 
suspensor, 

Fig. 50. The spiral vessels of the raphe highly magnified. 

Fig. 51. The seed invested by the testa and tegmen, both very thin membranes: d. the false chalaza; 


Fig. 53 & 54. The embryo seen in front and side 


å ways, 
8: 55. The same with one of the cotyledons 


removed, to show its nervation. 


+. 


THE 


TRANSACTIONS 
Ca 


OF 


THE LINNEAN SOCIETY 


LONDON. 


P 


VOLUME XXII. 
PART THE SECOND. 


MISSOURI 
BOTANICAL 
GARDEN. 


LONDON: 


PRINTED BY TAYLOR AND FRANCIS, RED LION COURT, FLEET STREET: 


SOLD AT THE SOCIETY’S APARTMENTS, BURLINGTON HOUSE; 


AND BY LONGMAN, BROWN, GREEN, LONGMANS AND ROBERTS, 


M.DCCC.LVII. 


PATERNOSTER- ROW. 


CONTENTS. 


# 


PART II.—1857. 


V. On a new form of Corynoid Polypes. By Pamir HENRY Gosse, Esq., F.R.S., 


Ad ae . page 113 

VI. Description of a new Species of Euplectella (Euplectella Cucumer, 0.). | By Professor 
OWEN, F.R.S., V.P.L.S. . re We eu 117 

VII. Om Brachynema and Phoxanthus, two new Genera of Brazilian Plants. By 
nonas BRSTRAA I ee 125 

VIII. On some new Fungi. By the Rev. M. J. BERKELEY, MA; ELS. . . . 129 


IX. On the Growth and Composition of the Ovarium of Siphonodon celastrineus, Griffith, 
especially with reference to the subject of its Placentation. By Josuru DALTON 
BOOKED, a AD RES US ae ABB 


X. Further Remarks on the Organs found on the bases of the Halteres and Wings of 
| dnseots. By d: B. Hicks, dog, M.D. Lond., FLS, .:: nce +. . . 141 


XI. On a new Structure in the Antenne of Insects. By J. B. Hicks, Esq., M.D. TP 
| | 14 


(^ xis] 


V. On a new form of Corynoid Polypes. By Purzre Henry Gosse, Esq., F.R.S., A.L.S. 
Read December 2nd, 1856. 


ALTHOUGH every discovery of a new species of animal ought to be recorded with as 
much precision as can be commanded, it is not every such discovery that is worthy of 
being made the subject of a memoir presented to a learned body; but the form and 
manners of a creature which has lately fallen under my notice appear so anomalous, 
that I am induced to bring it under the notice of the Linnean Society. 

In the summer of 1855 I happened to have, in one of my marine aquaria, several 
specimens of a Sabella, which I believe to have been identical with the Amphitrite vesi- 
culosa of Montagu (Linnean Transactions, vol. xi. p. 19). The largest of these had been 
some time in my possession, and, probably owing to.the habitual stillness of the water 
in the vessel, not holding in suspension the particles of mud, that ordinarily enter into 
the composition of tlie tube, the latest-formed portion was composed of pure transparent 
chitine, without any perceptible earthy element. This clear terminal portion of the tube 
I perceived to be occupied by a curious parasite. About twenty bodies having a most 
ludicrously-close resemblance to the human figure, and as closely imitating certain 
human motions, were standing erect around the mouth of the tube, when the Sabella 
had retired into the interior; and were incessantly bowing and tossing about their arms 
in the most energetic manner. : 

As soon as I had a little recovered from my surprise at this strange display, I began to 
examine the performers more in detail. A slender creeping thread, irregularly crossing 
and anastomosing, so as to form a loose network of about three meshes in width, sur- 
rounded the margin of the Sabella's tube, adhering firmly to its exterior surface, in the 
chitinous substance of which it seemed imbedded (see Pl. XX. fig. a). Here and there 
free buds were given off, especially from the lower edge; while from the upper threads 
sprang the strange forms that attracted my notice. "These were fusiform bodies, about 
45th of an inch in height, whose lower extremities were of no greater thickness than the 
thread from which they sprang, with a head-like lobe at the summit, separated from the 
body by a constriction, immediately below which two lengthened arms projected in a 
direction towards the axis of the tube. : i 

Such was the external form of these animals; and their movements were still more 
extraordinary. The head-lobe moved to and fro freely on the neck; the body swayed from 
side to side, but still more vigorously backward and forward, frequently bending into an 
arch in either direction; while the long arms were widely expanded, tossed wildly upward, 
and then uro downward, as if to mimic the actions of the most tumultuous human 
påssion. pes i 

Whenever the Sabella protruded from its tube, these guardian forms were pushed out, 


VOL XXIL ' 7 iy 


114 MR. GOSSE ON A NEW FORM OF CORYNOID POLYPES. 


and remained nearly in contact with the Annelid's body, moving but slightly; but no 
sooner did it retire, than they began instantly to bow forward, and gesticulate as before. 
These movements were continued, so far as I observed, all the time that the Sabella was: 
retracted ; and were not in any degree dependent on currents in the surrounding water, 
whether produced by the action of the Annelid, or by other causes. They were not 
rhythmical; each individual appeared to be animated by a distinct volition. 

Applying a higher magnifying power than I had yet used, to the animals, I found that 
the head-lobe enclosed a central cavity ; that the arms were also hollow, with thick walls, 
marked with transverse lines (internal septa ?), and muricated on the exterior; and that 
the body contained an undefined sub-opake nucleus (see fig. b), doubtless a stomachal 
cavity. l ) dig 

I cut out with fine scissors a segment of the tube, including two of the parasites, with 
that portion of the network of threads that carried them. They were immediately 
paralysed by the division of the threads; but those that remained on the tube were 
unaffected by the violence. The hiatus in the continuity of the circle was healed in a 
day or two; not by the approach of the divided edges of the tube, but by the shooting of - 
the threads across the chasm. One of the animals thus cut out is represented (at d), as 
it appeared immediately after the excision, magnified 240 diameters. When subjected to 
the action of the compressorium, with a power of 560 diameters, the arms were seen to be 
formed of globose cells, made slightly polyhedral by mutual pressure, set in single series 
(fig. e). The interior of these organs was divided by septa, placed at intervals of about 
the diameter. Some, at least, of the cells contained a small bright excentric nucleus 
(fig. f). 

When the tissues were quite crushed down by the pressure of the compressorium, a 
quivering motion was visible among the disjointed granules; but it was very slight. No 
trace of cilia, nor any appearance of ciliary motion, was perceptible during life. 

This larger Sabella-tube was not the only one infested with the parasites. I observed 
them on two, at least, of the smaller specimens, in the same situation, and with precisely 
the same movements. The extremity of one of these smaller tubes I cut wholly off, and 
placed in the live-box of the microscope. Two of the parasites only were on it, which 

were active at first, but in about an hour—probably from the exhaustion of the oxygen 
in the small quantity of water enclosed—they decomposed, or rather disintegrated; the 
outline dissolving, and the external cells becoming loose and ragged, and the whole 
animal losing its definite form. | 

One of these specimens, however, while yet alive and active, afforded me an observation 
of value, I had already associated the form with the Hydroid Polypes, and was inclined 
to place it in the family Corynide, considering the arms to be tentacles, and the head- 
lobe to be homologous with them in character, but abnormal in form. It appeared to be 
a three-tentacled Coryne, with the tentacles simple instead of capitate. But while I was 
observing the individual in question, I saw it suddenly open the head-lobe, and unfold it 
A een - a broad shovel-shaped expanded disk; not however flat, but with the 
pd ^m quts ng towards each other, like two leaves of a half-opened book (fig. 9). 
i; y reminded me of the great sucking-disk, which I had seen evolved from 


MR. GOSSE ON A NEW FORM OF CORYNOID POLYPES. 115 


the obtuse summit of Coryne Cerberus, as I have elsewhere recorded*; and confirmed 
my suggestion of the natural affinities of the form. 

The principal colony remained for many weeks under my observation, without any 
noticeable change. The terminal portion of the tube, which at first had been so dia- : 
phanous, gradually became more opake, and disfigured by the growth on it of confervoid 
threads, and by the entanglement of a multitude of minute Diatomaceæ in its surface. 
At length the Sabella spontaneously quitted its tenement: the parasites appeared at first 
unchanged and unaffected; but, strange to say, before the lapse of two days, they all 
gradually died away; as if their existence depended on the presence of their patron. 
They seemed to become feeble, attenuated, and almost motionless, before they dis- 
appeared. 

While they lived they afforded me much entertainment, as also to those scientific 
friends to whom I had opportunities of exhibiting them. When I used to see them 
surrounding the mansion of the Sabella, gazing, as it were, after him, as he retreated 
into his castle, flinging their wild arms over its entrance, and keeping watch with untiring 
vigilance until he reappeared, it seemed to require no very vivid faney to imagine them 
so many guardian demons; and the Lares of the old Roman mythology suggested for 
them a name. pr 

Bearing in mind the extraordinary eycle of phænomena that have been proved to occur 
in the reproduction of the Hydroid Polypes in general, and of the Corynidæ in particular ; 
it is highly probable that the animal, whose appearance and manners I have been 
describing, would, in the natural prolongation of its existence, have budded off some 
Medusoid forms, endowed with proper sexual functions. I detected, indeed, no trace of 
(so-called) ovarian capsules, nor any evidence of increase, except that of the gemination 
of the individual zooids from the common root-thread. This, however, by no means 
disproves the possibility, nor even the probability, of such developments, at å more 
mature stage of the polypoid condition. Had I discovered such, and were I able to follow 
out the life-history of the animal, and to show that it agrees in its reproductive phæno- 
mena with other Coryniform Polypes, it would still be an open question, —to Which 
condition—whether the Coryniform or the Medusiform—specific identity is to be assigned ; 
or, in other words, which ought to bear the nomina generica et trivialia, and to ag 
place in the System of Nature. Is the Medusa the animal, of which the hs ii " 
larva? Oris the Coryne the animal, of which the Medusa is but the detached and loco- 


motive sexual organ ? 7 
I incline to affirm the former of these two hypotheses; but yet, in accordance with 


. La h 
precedent, by which the Corynide and Campanulariade are reckoned Be de y 
of names, I may venture, provisionally at least, to register, under systematic appellations, 

the form before me, and wait for new light as to its future history. 

Genus Lar (Gosse). 
Zoophytum è familia Corynidarum, nudum, associatum sed sejunctum, erectum, 
retiformi saliens, tentaculis simplicibus filiformibus instructum. 
+ Devonshire Coast, p. 223, plate xiv. fig. 4. 


e filo radicali repente 


> 


Q 2 


116 — MR. GOSSE ON A NEW FORM OF CORYNOID POLYPES. 


Species, LAR SABELLARUM (Gosse). 


Species unica. L. tentaculis tribus, filo radicali circa Sabellarum tubos parasiticé repente. 


EXPLANATION OF THE PLATE. 
o. TAB. XX. 


Fig. a. The extremity of the tube of Sabella vesiculosa, occupied by a colony of Lar Sabellarum (mag- 
nified 70 diam.). 

Fig. 5. An individual Lar, displaying the hollow head-lobe and tentacles, "es the TT digestive 
canal (magnified 120 diam.). 

Fig. c. A portion of one of the tentacles, showing the T surface. 

Fig. d. A single Lar, with the thread, attached to an excised aon of the tube, paralysed and some- 
what contraéted (magnified 240 diam.). 

Fig. e. A portion of one of the tentacles, flattened by pressure ; emis the parietal cells, and the edges 
of septa (magnified 560 diam.). 

Fig. f. Nucleated cells from a crushed tentacle. 


Brans. Linn. Soo. Vol XXII. tab RO. p.116. 


RSS 


E Jarman 50. 


[PH Cose Hag, det, d 


[: 417.3 


VI. Description of a new Species of Euplectella (Euplectella Cucumer, 0.). 
By Professor Owen, F.R.S., V.P.L.S. 


Read February 17th, 1857. 


IN 1841 I communicated to the Zoological Society of London a description of a new 
generic form of reticulate Aleyonoid Sponge, represented by one of the most singular and 
beautiful, as well as the rarest, of the marine productions with which the researches of 
Mr. Hugh Cuming in the Philippine Islands had enabled him to enrich his famous 
Natural-History collection*. For this genus the name Zuplectellat was proposed, indi- 
cative of the exquisite regularity and complexity of the interweaving of its component 
threads. 

The characters of the genus are:—a cylindroid hollow form of body, closed at the 
wider end by an irregular network, and at the narrower end by the terminal tuft of finer 
filaments into which the parietal fibres are there resolved. 

The parietal fibres, or those that constitute the wall of the cylinder, are regularly dis- 
posed, and intersect. each other at definite and nearly equal distances throughout its 
extent. They consist of longitudinal (Pl. XXI. fig. 1, c, d, e), transverse (¢), and oblique 
fibres, the latter being of two kinds (o, 0), winding spirally round the cylinder, but in 
opposite directions: (see magnified view of part of the parietes, Pl. XXI. fig. 4). The 
longitudinal and transverse fibres are the thickest: they are arranged at intervals of from 
one to two lines, averaging one line and a half apart, and divide the cylinder-wall into 
square spaces (a) of about the latter diameter. The longitudinal fibres (fig. 4, b) are 
external to the transverse ones (£), to which they are bound by the oblique or spiral 
fibres; these are, some external, some internal, to the others, and they close by their 
decusastion alternate quadrate intervals (+) between the longitudinal and transverse fibres. 
The angles of the alternate open squares are intersected by finer and less regular oblique 
fibres, which reduce their area more or less to a circular form (fig. 4, a). : Lib the 

It appeared, in the first-described species, that the nne pilky. filaments into d^ : m 

“parietal fibres were resolved at the small end of the eylindroid, had been M pi d iva 
by violence from some other body. The subject of the present description, which has been 
; by my friend Dr. Arthur Farre, F.R.S., has 
rai confided EE NE hich it was attached by the 

been fortunately preserved, along with the foreign body to w SEPA Jes 
terminal filaments: such mode of attachment may now, therefore, be v pene 


characters of Euplectella as above defined. i mas ded on a specimen eight inches j 


The first-described species of this rare genus was : 
in length, of a slightly conical form, two inches across the base, and gradually and pro- 


gressively decreasing in diameter to the truncated apex, de is one inch and a quarter 
in longest diameteri. _ ring 


* Transactions of the Zoological Society of London, 
t Trans. Zool. Soc. vol. iii. pl. 13. a 


i iii. p. 205. + Gr. eù, well, rAéxw, I weave. 


118 | PROF. OWEN ON A NEW SPECIES OF EUPLECTELLA 


The present species (Pl. XXI. fig. 1) is six inches in length, two inches across the base, 
whence the cylindroid body gradually expands to near the middle of its length, where it 
presents a diameter of two inches seven lines, and then decreases to the truncated apex, 
which is about one inch and a quarter in diameter; but part of this appears to have been 
torn away. Thus the form of the body is ventricose, not regularly conical as in -Euplectella 
Aspergillum ; it more resembles a cucumber than the shell after which the first species 
was named, whence the present species may be named Fuplectella Cucumer. 

The next difference which strikes the eye is the absence of the oblique and wavy crests 
or ridges which project from the network of the cone, and especially the absence of that 

marginal plate which divides the reticulate terminal cap or lid from the wall of the 
cylinder, standing out like a ruff or frill in Zuplectella Aspergillum. - 

The convex reticulate lid or cap in Æuplectella Cucumer (Pl. XXI. fig. 3) is bounded 
simply by the marginal ridge (figs. 1 & 3,7), which represents the last or lowest* of the 
transverse fibres, but which is thickened by an aceession of the constituent fibrils, especially | 
from the oblique series, so as to project slightly like a rim or ‘bead’ in carpentry. Some 
slightly projecting fibrils from the track of the oblique series of fibres, chiefly multiradiate,. 
and of the kind figured in Pl. XXI. fig. 5, and which scarcely catch the eye, except when 
in relief at the border of the cylindroid, as at p, P, fig. 1, alone feebly represent the parietal 
and reticulate crests which so peculiarly distinguish the Zuplectella Aspergillum. In 
this species the gradual diminution of the cylinder is produced by the convergence and 
confluence of two contiguous longitudinal fibres at certain parts of the circumference. . 
The like convergence and final interblending of contiguous longitudinal fibres, as they 
pass from the free towards the fixed end of the body, is also manifested, though in a 
minor degree, in Euplectella Cucumer, as at c and 6, fig.1. But the gradual expansion of 
the cylindroid is made compatible with this diminution in the number of the longitudinal 
fibres by the divergence of many of the longitudinal fibres as they proceed from the 
marginal ridge, as at d, fig. 1, towards the widest part of the cylindroid, where their 
intervals are greater than in the corresponding part of Buplectella Aspergillum. Very 
few instances of confluence of longitudinal fibres take place before they reach the widest 
part of Euplectella Cucumer : the majority occur beyond it, as at e, e, fig. 1; and besides 
the diminution in number of the longitudinal fibres, they all converge as they approach 
the smaller and attached end of the cylindroid. Here the resolution of the several series 
of fibres into their constituent fibrils seems to take place, at least on one side, viz. that 
A E PL XXI. fig. 1, more abruptly than in Euplectella Aspergillum: on the opposite 

‚as In fig. 2, the fibres in Euplectella Cucumer begin to be resolved into the fibrils or 
"am iiie a ne la Aspergillum. The considerably greater length of 
E sd Å så offer — marked distinction between the two species: but as 

prectella Aspergillum was torn by violence and brought up by the fish- 


ds of attachment, no safe inference can be drawn as to their original 
ength in specimen. In the present example of rof 
Separated silky filaments penetrate, ple of Euplectella Cucumer the delicate 


and, as it were, permeate, the substance of a mass made, 


* i. : 
On the supposition that the Euplectella hangs dependent from its EE 2 


PROF. OWEN ON A NEW SPECIES OF EUPLECTELLA. 119 


up chiefly of portions of a coarse irregular siliceous sponge, which appears to be foreign to 
the proper body of the Buplectella, and includes some shells and other marine calcareous 
bodies. Some of the fine long fibrils, emerging from the mass, converge, as they are reflected 
back (fig. 2), and, after a few graceful bendings, again diverge into separate wavy locks of 
the most delicate hairs, having a silken or silvery lustre (h, h, figs. 1 & 2). The whole of 
this beautiful elongated filamentous medium of attachment of the Euplectella may be 
compared to a lock or tuft of the hair which Poets feign to have adorned the head of 
the Syren or Mermaid. 

The number of the longitudinal fibres at the base of the cone in Euplectella Asper- 
gillum is sixty, in Euplectella Cucumer it is fifty-three: their number at the apex in 
Euplectella Aspergillum is thirty, in Euplectella Cucumer it is thirty-five. I would not, 
however, be understood as confiding in these particular numbers being constant and cha- 
racteristic of the species. The fibres of the reticulate cap consist of converging and of 
connecting or transverse kinds : many of the former are continued from the confluence 
of two of the longitudinal fibres of the cylindroid, where they become bent at nearly right 
angles after leaving the connecting marginal band to form the cap: such fibres show a 
thickness proportionate to the additional material entering into their composition. The 
degree of irregularity in the converging and connecting fibres forming the reticulate cap 
is like that in Euplectella Aspergillum*. The superficial fine oblique series of fibres, with 
the superadded multiradiate spicula, terminate abruptly at the marginal rim in Euplee- 
tella Cucumer, just as the ridges of the cylinder terminate in Fuplectella Aspergillum. 

Dr. A. Farre notes, as the result of his study of the structure of the Huplectella Cucumer, 
that *the oblique lines are not formed out of one continuous line of fibre for each side, 
wound round and round, which is the idea of a spiral, but they consist of a double series of 
ellipses, placed at definite distances, which intersect each other at right angles, or nearly so. 
These form perfect ellipses only towards the centre of the specimen, for at either end they 
are necessarily interrupted. In these oblique fibres there occur the same confluence of two 
contiguous lines, in some places, and divergence or bifurcation of simple ones in others, 
as happen in the longitudinal series, and evidently with the same object of adapting their 
arrangement to the increasing or diminishing diameter of the cylinder. With regard to 
the relative positions of the longitudinal, transverse and oblique fibres, I find that these 
lie in several alternate series. First a bundle of about half-a-dozen longitudinal close- 
lying fibres. These run straight from end to end of the cylinder, " : 


' except where they bifurcate or combine. Then the looser bundles of oblique fibres 
decussate with the longitudinal ones, the fibres separating to pass over, under, and 
between them, and at the same time intersecting the fibres of the opposite oblique series 


in a similar way, 


* Compare PI. 13, Trans. Zool. Soc. vol. iii. fig. 2, with fig. 2 in PI. XXI. 


* 


120 PROF. OWEN ON A NEW SPECIES OF EUPLECTELLA. 


not with strict regularity, but in a sufficiently marked manner, and forming a rough 
pattern of those wicker-baskets in which the cross fibres are interwoven among the lon- 
gitudinal ribs. Lastly, the transverse fibres lie below these, and are, on that account, 
the least easily distinguished. "Then the same series of longitudinal, oblique and trans- 
verse lines is repeated, but more irregularly, until the walls of the cylinder acquire a 
depth of 2" at their thickest part, which occurs at about $ths of the entire length of 
the specimen from its operculated end."—(A. F.)  . 

The average diameter of the longitudinal fibres of the cylindroid in Æuplectella 
Cucumer is one-thirtieth of an inch: that of the transverse fibres is about one-fortieth 
of an inch: these oblique fibres present much smaller and more varying diameters. All 
these fibres consist of much finer fibrils, and these are composed of a delicate siliceous 
sheath enclosing still more minute fibrillules. The component fibrils of Zuplectella 
Cucumer, as in Euplectella Aspergillum, are of two kinds; one smooth, the other barbed 
at pretty regular distances like the hair of certain caterpillars (Pl. XXI. fig. 6): and 
some fibrils show both characters, and the gradual transition of the barbed to the smooth - 
part (fig. 7). In some of the barbed fibrils, which most abound in the resolved tuft of 
attachment, Dr. A. Farre discovered a terminal convex disk, with a border divided into 
four or five retroverted spines, simulating a small anchor: one of these is represented in 
fig. 6. The same experienced microscopist found that he could frequently detach, with a 
fine pair of forceps, from the exterior of the point of decussation of the oblique fibres, in 
the body of the cylindroid, minute multiradiate aciculi, like the one represented at fig. 5,— 
one ray representing the axis, from which four other rays would diverge at right angles 
and equal distances on the same plane. These **multiradiate spicula invariably consist 
of six rays, viz. a perpendicular spine, which projects above the surface of the cylinder, a 
small spieulum opposite to it which lies buried in the mass, for the purpose apparently of 


_ fixing the upper spiculum, and four basal rays (one very generally longer than the rest), 


which take a direction always exactly corresponding with the lines of intersection of the 


oblique fibres, with which they become blended."—(A. F.) These detached bodies may 


have been the commencement of the new-forming oblique fibres in the gelatinous sub- 
stance of the living Euplectella. ; 

On applying the test of fire, by subjecting the fibrils of the Zuplectella Cucumer to the 
flame of a candle, they generally splintered, and minute iridescent portions flew off in all 
directions. Some of these particles, being caught on slides of glass, showed under the 


microscope that they were parts of a delicate sheath of silex: their iridescent hue seemed 


to be due to the partially disintegrated constituent layers of siliceous matter entering . 
into the constitution of the sheath. Portions of fibrils, submitted in a test-tube to the 
flame of a candle, were observed to splinter, sometimes with a slight bend; and, on being 
submitted to a half-inch objective in the compound microscope, showed the siliceous. 


sheath variously cracked or splintered, enclosing a bundle of very minute fibrillules, with 


here and there a trace of charred gluten or organic matter. Submitted to the action of 
dilute muriatic, or nitric, acid, the fibrils underwent no other change than that of becoming 
rather more clear or less opake; and this without the extrication of bubbles, as from car- 
bonate of lime, and with not more evidence of any liberated gas than might be accounted 


PROF. OWEN ON A NEW SPECIES OF EUPLECTELLA. 121 


for by the solution of the gluten, to which act, perhaps, the greater transparency of the 
soaked fibril might be due. No diminution of diameter or other change of the fibril 
could be detected whilst it was under the action of the mineral acid. 

The fibrils of the Huplectella Aspergillum contain a greater proportion of organic 
matter, but have, as described in my original memoir, the same siliceous basis as in the 
present species. In consequence, however, of the different proportion of the glutinous 
and siliceous principles, they do not behave exactly like the fibrils of Euplectella Cucumer 
under the tests of heat and acid. In general, when subject to fire, they do not splinter, 
but merely bend; and when afterwards microscopically examined, show more trace of 
charring, and sometimes present increased opacity with the appearance of dead or frosted 
silver. Yet some show clearly their constitution of a siliceous sheath, including fine 
iridescent filaments or fibrillules. : 

It sometimes happened, in the case of fibrils of the Æwplectella Cucwmer submitted to 
the acid test, that a small amorphous mass would be adherent to part of a fibril; and 
such mass was quickly dissolved with the extrication of abundant gas-bubbles: the 
inference from this was, that some fragment of a foreign body of carbonate of lime had 
become accidentally entangled in the meshwork of the Buplectella. 

In the amorphous portions of sponge to which the Huplectella Cucwmer was connected 
by its long filaments, two modifications of reticulate structure were discovered by Dr. 
A. Farre. One, represented in Pl. XXI. fig. 8, is an irregular network, more or less bent, 
with subquadrate meshes, sometimes crossed by oblique threads. A second and more 
beautiful structure is represented in fig. 9. The meshes of this network are on nearly 
' the same plane, and of a more regular square form, with a short pointed spiculum pro- 
jecting from one side of each decussation of the threads, like the teeth of a harrow. Both 

figs. 8 & 9 give magnified views of the above structures, with the part magnified of the 
natural size. Fig. 9a gives a more magnified view of one of the squares, with its spines, 
of the * harrow’ structure. 

To the question put by almost every one to whom the Buplectella is shown, as to how 
the threads could have been so regularly yet intricately interwoven, I have sometimes 
replied, that there has been no such thing as interweaving in the case; that no thread, 
as such, was ever laid across another in the construction of the ÆEuplectella; that the 
analogy of human textile fabrics does not apply to this beautiful natural object. In 
artificial lacework the several stages of a complex result must be taken in the succession 
indicated by painful and exact calculation : in organic lacework different stages are done 
at once. Thus it is that the Divine works surpass those of man's utmost ingenuity. 
The threads of the Zuplectella were not first spun and then interwoven, but were formed 
as interwoven, the two processes going on simultaneously, or ‘pari passu? Just as in 
the cancellous texture of bone, the plates of bone are not first formed and then fitted to 
one another, as in building a house of cards; but the forming and the fitting go on 
together in the course of molecular growth. I presume also, that in the beautiful object 
which we call the Zuplectella, we have but its skeleton; and that, in the living state, 

d the exquisite structure of the flinty framework may be veiled by the delicate gelatinous 


enveloping organic tissue. 
VOL. XXII. 


R 


122 PROF. OWEN ON A NEW SPECIES OF EUPLECTELLA. 


The specimen of the Euplectella Cucumer here described, Dr. Farre writes, ‘was given, 
together with other presents, by the King of the Seychelle Islands to Captain Etheridge, 
R.N., in acknowledgment of some friendly services, and with an intimation that this 
was one of the rarest products of those regions. From Captain Etheridge it passed into 
the hands of Dr. Richard Payne Cotton, of Clarges Street, by whom it was recently 
presented to me.”—(A. F.) _ 

With regard to the first-described species, Zuplectella Aspergillum, Mr. Hugh Cuming 
writes to me :—* The Euplectella brought home by me from the Philippines was taken 
by a fisherman, in ten fathoms, rocky ground, off the island of Bohol, one of the southern 
islands of the Philippine group. The fisherman was employed in catching a species of 
cod which abounds in those islands, and finding, after some time, the fish did not take his 
bait, he drew it up, when to his surprise he found the above specimen attached to his 
hook, near the orifice; and fearing to injure it by disentangling the hook from such a 
fragile substance, he cut out that portion to which the hook was attached. On his arrival 
on shore at St. Nicholas di Zebu, he made a present of it to the governor of the town. 
On my arrival a few days after, I was introduced to the governor, who, upon knowing 
the object of my visit to the island, presented me with it as the greatest curiosity he had 
to offer me, as he had never seen the like before. On my showing it to the bishop of 
that city, and the principal inhabitants, they confirmed the opinion of its rarity expressed 
by the governor."—(H. C.) 

In conclusion, I beg to state, that although this paper is communicated to the Society 
as from one author, it is properly the work of two: many of the observations were 
originally made by Dr. Arthur Farre, which afterwards were confirmed by my own eyes; - 
and every recorded observation of my own has been repeated by, and received the sanction 
of my friend. In returning these acknowledgments for the considerate call which he 
pressed upon the describer of the original species to bring the second before the notice of 
naturalists, and for his generous relegation of this pleasing task to me, I desire, at the _ 
same time, to offer my best thanks to Mr. Cuming, who, with his own hands, transported 
his frail and unique specimen of Euplectella Aspergillum to the residence of Dr. A. Farre, 
in order to afford every facility of comparing the two species: and Ihave again to express, 
as at the close of my former memoir, my obligations to Mr. G. B. Sowerby, for his 


zealous devotion of much time and peculiar artistic skill to a faithful representation of 
this second species of the rare genus Huplectella. 


Fig. 


Fig. 
Fig. 
Fig. 4. 
Fig. 
Fig. 
Fig. 
. Fig. 


Fig. 


t2 


PROF. OWEN ON A NEW SPECIES OF EUPLECTELLA. 123 


DESCRIPTION OF THE PLATE. 
Tin XXL. 


- Euplectella Cucumer, nat. size; with the foreign sponge and other bodies to which it is attached :— 


A. wall of the cylindroid, or parietal portion; B. terminal grating or lid; a. parietal orifices ; 
c. confluence of longitudinal fibres ; d. ibid. prior to forming the network of the lid; e. ibid. as 
the cylindroid contracts towards its attached end; A. resolved filaments, reflected from the 
attached body; o, o'. oblique fibres; p. prominent lines from the oblique fibres, chiefly due to 
the multiradiate accessory spicula; r. rim or thickened transverse line at the attached border 
of the lid; £. transverse fibres. 


- The opposite side of the attached end of the Euplectella ERE with the substance to which it 


adheres :—h. the free ends of the resolved filaments: nat. size. 


. The reticulate grating or soldered lid of the Euplectella Cucumer : nat. size. 


A closed and an open interspace of the meshwork, magnified four diameters:—a. the open 
space; 5. longitudinal fibres; ¢. transverse fibres; Å. superficial oblique fibres closing an inter- 
space ; 0. deep-seated fibres; o'. intermediate fibres. 


. Å multiradiate spiculum, from the place of intersection of the superficial oblique fibres, magnified 


eight diameters. 


. Portion of a barbed filament with its terminal anchor, magnified twenty diameters. 
. Transition of the barbed to the smooth structure in one of the filaments, similarly magnified. 
. Å portion of a sponge from the mass to which the Euplectella adheres ; natural size, and mag- 


nified ten diameters. 


. Another portion of sponge from the same mass; natural size, and magnified ten diameters :— 


9a. one of the squares of the harrow-like structure, magnified twenty diameters. 


I PLU p I 


7 
y 


C 


ans Linn 


Ir 


Sn man _ nn. 


på u mo 


[ 125 ] 


VIL. Om Brachynema and Phoxanthus, tho new Genera of Brazilian Plants. 
By GEORGE Bentham, Esq., F.L.S. 


Read May 5th, 1857. 


AMONG the North Brazilian plants collected by Mr. Spruce, are two, which have 
appeared to me to present some interesting anomalies, preventing their being referred 
absolutely to any of our Natural Orders as at present defined. I therefore beg to lay 
_ before the Linnean Society the following descriptions, accompanied by drawings carefully 
. executed by Mr. Fitch, in the hope that their publication may be the means of further 
elucidating the immediate affinities of these plants. 


1. BRACHYNEMA RAMIFLORUM. (Tap. XXII.) 


This pit to which I have given the name of Brachynema, in allusion to the short 
filaments of the stamens, has a foliage which reminds one of the simple-leaved Galipeas 
or Ticoreas, whilst the structure of the flowers is nearer that of Æbenaceæ. I should 

indeed have had little hesitation in considering it as a true Ebenacea, slightly anomalous 
in inflorescence, in the shape of the corolla, and in the hermaphrodite flowers, were it not 
for the leaves. These have the general form and appearance, the unequal petioles, and 
the evident tendeney to articulation at the summit of the petioles, of several unifoliolate 
Rutaceæ. The surface of the leaf is not however covered with the minute resinous dots 
of most Diosmeæ, but the principal nerves when they approach the margin often emit a 
small lateral branch terminating in a gland on the margin itself, as is the case in some 
Diosmeæ. The same circumstance may however be observed in some Euphorbiaceae, in a 
few of the Malvoid families, and in some other Orders. The inflorescence of Brachynema, 
the shape of the corolla, the almost valvate slightly contorted æstivation of its thick 
lobes, have nothing in them incompatible with the gamopetalous Diosmeæ, but here all 
affinity ceases. The insertion of the stamens, the structure of the ovary, the fruit with 
its enlarged calyx, are nearly those of Ebenacee. The only seed which I could examine 
was probably not quite ripe. Within a striated closely adherent coating, it only pre- 
sented a uniform cartilaginous mass with å slight discoloration in the interior, in which 
however I failed in detecting the embryo. Outside, the longitudinal furrows of the seed 
show an approach to the ruminated albumen of some Zbenacee ; although, on the other 
hand, there is nothing to preclude the supposition that the whole may be an exalbuminous 
embryo. The coarse fibres more or less cohering to the external furrows, appear to be 
detached from the endocarp of the fruit. ; 

For the present, therefore, Brachynema must be considered as a doubtful Ebenacea, 

» anomalous in its foliage, in its hermaphrodite flowers, in its long-tubed corolla, and 
in the number of stamens equal to and alternate with the lobes of the corolla. 
I should add, that the flowers opp only on san older branches of the tree, the 


126 MR. BENTHAM ON TWO NEW GENERA OF BRAZILIAN PLANTS. : 


portions of branch bearing them are in my specimens detached from those which have the 
leaves. But besides that Mr. Spruce is far too careful a collector for me to suspect any 
accidental mismatching, the bark and wood of the flowering portions perfectly correspond 
with those of the leafy ones. 

BRACHYNEMA. 


Char. GEN.—Calyz cupuliformis, subinteger, post anthesin accrescens. Corolla infundibularis, tubo 
elongato, laciniis 5 crassis lanceolatis patentibus, æstivatione valvato-subcontorta. Stamina 5, brevia, 
ima basi corollæ v. cum corolla toro inserta, laciniis corollæ alternantia; filamento brevi dilatato ; 
anthera extrorsa, biloculari, connectivo apiculata, loculis longitudinaliter dehiscentibus. ^ Ovarium 
sessile, 5-loculare. Stigma subsessile, 5-lobulatum. Ovula in loculis solitaria, pendula. Fructus 
(drupaceus ?), pericarpio tenui, endocarpio crustaceo intus fibroso, unilocularis, monospermus. Semen 
lata basi affixum, sulcato-striatum, testa tenui arcte adnata. Albumen cartilagineum. Embryo 
ignotus (v. embryo crasso-carnosus exalbuminosus indivisus ?); ; 

Species unica B. RAMIFLORUM. Arbor, teste R. Spruce, gracilis, 30-pedalis. Rami subteretes, ligno duro, 
cortice scabriusculo. Folia sparsa, exstipulata, inzequalia, ovato-oblonga, anguste acuminata, integra 
v. obsolete sinuata, basi cuneata, membranacea, glabra, utrinque viridia, penninervia et transverse 
venosa, costa nervisque subtus prominentibus. Petioli majorum 4-6-pollicares, apice leviter inflexi 
et quasi articulati, minorum vix pollicares. Lamina majorum fere pedalis, 5 poll. lata, minorum vix 
4-pollicaris. Nervi primarii a costa utrinque 6-8, versus marginem arcuate et irregulariter con- 
fluentes, hinc inde juxta marginem ramulum emittentes in margine ipso glandula parva terminantem ; 
duo infimi oppositi, margini paralleli. Flores in ramos annotinos v. vetustiores ex axillis foliorum 
delapsorum nascentes, fasciculis subcymosis, 5-15-floris, sessilibus, ebracteatis. Pedicelli cras- 
siusculi, vix lineam longi. Calyx per anthesin cupuliformis, 11 lin. latus. Corolle tubus 7-8 lin. 

. longus, extus zonis transversis intensius coloratis variegatus ; limbi laciniæ crasso-triquetræ, patentes 
v. recurve, intus linea longitudinali pilorum intra tubum plus minus decurrente notatae. Staminum 
filamenta antheris breviora; pollen e granulis minutis globosis compositum. Ovarium crassum, 
subglobosum, vertice depresso-umbilicatum. Stigma pulviniforme, lobulis brevissimis subemar- 
ginatis. Fructus in specimine depresso-globosus, 6-8 lin. diametro. i 

Hab. In Brasilia boreali ad cataractas fluminis Aripecurù, in fl. Amazonum juxta Obidos fluentis 
(R. Spruce). ; : 

2. PHOXANTHUS HETEROPHYLLUS. (Tas. XXIL., XXIV.) 


Of the other genus, for which I propose the name of Phoxanthus, in allusion to the 
long pointed petals giving a tapering conical form to the bud, Mr. Spruce’s specimens 
afford ample materials for a full illustration, In the first hasty determination for the 
purposes of distribution, struck by the linear petals with stamens opposite to their dilated 
basc, and some ‚other points of resemblance: with Aublet’s figure of Mayepea, I had 
thought it probably a second species of that genus (which had already been referred to 
Linociera), and I accordingly labelled it Linociera (Mayepea) heterophylla. But a closer 
examination showed that I was in error. In the first place, Aublet's figure is incorrect. 
His specimen in the British Museum has opposite, not alternate, leaves, and from Dr. 
Solander's notes it appears that there are two, not four, stamens, and that it is altogether 
à true Limociera. If therefore Aublet’s dissections, especially fig. 4 of his Plate (t. 31), 
represent what he really saw, he must have had some flowers of Phoxanthus mixed up 
with those of his Mayepea, and have confounded the two plants. Further, I find that 
Mr. asi: plant has not generally four petals, as was accidentally the case in a flower I 


MR. BENTHAM ON TWO NEW GENERA OF BRAZILIAN PLANTS. 127 


first examined, but five, although very unequal in size. There are really five stamens, of 
which three are reduced to mere barren scales; all are opposite to the petals and inserted 
with them under å small five-lobed disk, and our plant must be sought for among the 
disk-bearing polypetalous families with compound leaves. The form and structure of 
the ovary remind at once of some Simarubaceæ, and if that family be extended as pro- 
posed by Planchon, it may well include Phoxanthus also. The stamens opposite the 
linear petals occur in Picrammia (whose place among Simarubacee is however doubted 
by Tulasne), and in the still more anomalous Alvaradoa ; the contortuplicate cotyledons, 
so remarkable in Phoxanthus, are also described in Picrodendron. The stamens, however, 
are very different in form from those which prevail in Simarubaceæ, and I have been 
unable to detect in our plant any of the bitter principle so universal in that family. The 
absence however cannot be absolutely ascertained from a dried specimen. The succulent 
pericarp is evidently very oily. 

Mr. Spruce describes Phoxanthus as a slender woody plant with a stem as thick as the 
arm, threading among trees to a great height, or, when standing alone, forming a small 
tortuous tree. The branches are distant, nearly simple, the lower ones near the ground 
alone flowering, and these have mostly simple leaves, whilst the upper barren branches 
have large pinnate leaves with three to nine leaflets often near a foot long. The flowers 
are small and numerous in large red panicles, the petals themselves being of a pale flesh 
colour. He gathered it in October 1852, in the forests of the Rio Uaupès, especially 
near the Falls of Panuré. 

PHOXANTHUS. 


Cnan. Gen.—Calyz brevis 5-fidus ; laciniis ovatis æstivatione imbricatis, 2 exterioribus vulgo minoribus, 
una interdum deficiente. Petala linearia, acutissima, basi dilatata, hypogyna. Stamina petalis 
opposita et cum iis sub disco hypogyno 5-lobo inserta ; 2 fertilia filamento brevissimo crasso, anthera 
biloculari, loculis connectivo crasso disjunctis rima transversali dehiscentibus ; 3 sterilia squamæformia. 
Ovarium subsessile, apice brevissime bilobum, lobis vertice stigmatosis, intus biloculare. Ovula in 
loculis gemina, versus apicem anguli interioris superposita pendula. Fructus drupaceus, abortu 
unilocularis, monospermus. Semen unicum, hilo lato juxta basin fructus affixum ; testa membranacea. 
Embryo exalbuminosus ; radicula crasso-carnosa incurva ad hilum spectans ; cotyledones insigniter 
contortuplicatæ. 

Species unica PHOXANTHUS HETEROPHYLLUS. Frutex alte scandens vel a 
subsimplices; inferiores floriferi juxta terram breves foliis plerisque simplicibus (unifoliolatis petiolo 
brevissimo); superiores steriles foliis imparipinnatis 3-9-foliolatis, petiolo 1-2-pedali. Stipulas non 
vidi. Foliola elliptico-oblonga, abrupte et anguste acuminata, basi rotundata v. acutiuscula, 6-12 — 
poll. longa, 3-4 poll. lata, coriacea, glabra, penninervia, impunctata. Paniculæ axillares, amplæ, 
laxæ, ramis divaricatis in vivo rubris, in sicco pube minuta rufescentibus. Flores parvi in ramulos 
ultimos thyrsoidei. Pedicelli vix semilineam longi, bractea minima suffulti. Calyx pedicello brevior, 

. campanulatus, laciniis membranaceis margine denticulatis. Petala valde inæqualia, majora 14-2 lin. 
longa. Æstivatio subvalvata videtur. Stamina fertilia petalis minoribus, sterilia majoribus opposita ; 
filamenta fertilium obovoidea ; anthera biglobosa, per anthesin introrsum incumbens, post pollinem 
ejectum ascendenti-erecta. Drupa 9-12 lin. diametro compressa subglobosa v. leviter incurva, ad 
insertionem sæpe depressa. Pericarpium tenue, carnoso-oleosum. Putamen lignosum. Semen peri- 


arbor parva tortuosa. Rami 


' carpio subconforme. 
Hab. In sylvis ad Rio Uaupès juxta fines Brasiliæ borealis (R. Spruce). 


i 


128 MR. BENTHAM ON TWO NEW GENERA OF BRAZILIAN PLANTS. s 


4 


DESCRIPTION OF THE PLATES. 
Tas. XXII. 


Brachynema ramiflorum, a leafy and a flowering specimen: natural size. 
3 


Fig. 1. Flower. 

Fig. 2. Corolla, cut open. 

Fig. 3. Stamen. 

Fig. 4. Calyx and ovary. 

Fig. 5. The same, vertical section. 

Fig. 6. Ovary, transverse section :—all magnified. 
Fig. 7. Enlarged calyx and fruit: natural size. 
Fig. 8. Fruit. 

Fig. 9. The same, transverse section. 

Fig. 10. Seed. 


. Tas. XXIII. 


Phozanthus heterophyllus, a flowering specimen : natural size. 
Fig. 1. Flower. 
Fig. 2. Petal. 
Fig. 3. Stamens and ovary, showing the largest staminodium or abortive stamen. 
Fig. 4. The same, seen from the other side, showing the two smaller staminodia and one of the lobes of 
the disk seen between them. 
Fig. 5. A single stamen. 
Fig. 6. Ovary and hypogynous disk. 
Fig. 7. Ovary, vertical section. 
Fig. 8. Diagram of the flower :—all magnified. 


Tas. XXIV. 


Phozanthus heterophyllus, a fruiting specimen, and a pinnate leaf from a sterile branch : both natural size. 
Fig. 1. Fruit: natural size. 
Fig. 2. The same, vertical section: somewhat magnified, as well as the three erg 


Fig. 3. Embryo, end view, showing the large ascending radicle, 
Fig. 4. The same, side view. 


å Fig. 5. The same, vertical section. 


Frans Lunn. Soo. Vol EXIT tab. 22. p.128. 


Frans Lunn. Soc. Vol. XXII. rad. 23. RE. 


6.Jarman sc 


Frans. Linn.Soc. Vol XXII. tab. 24. n. 128. 


6. Jarman sc. 


VIII. On some new Fungi. By the Rev. M. J. BERKELEY, M.A., F.L.S. 


Read May 5th, 1857. 


Å VERY valuable collection of Fungi E been lately made by Fendler in Venezuela, 
which are now in the hands of the Rev. M. A. Curtis of South Carolina, and will shortly 
be offered for sale. The collection abounds in new species, of which I have selected three 
as peculiarly interesting, with the intention of bringing one or two more under the notice 
of the Society at some future meeting. To these I have added a description of a new 
species of Mitremyces from South Carolina, and of a very curious genus of Hypogæous 
Fungi lately received from Tasmania, with many other good things, through the kindness 
of Mr. Archer. The Tasmanian collection will shortly be described in Dr. Hooker’s Tas- 
manian Flora, but this genus is so curious that it deserves previously a more especial 
notice. 


1. HyDNUM BRUNNEOLEUCUM, Berk. & Curt. Pileo flabelliformi : ne tenui luteo- 
brunneo glabro; hymenio pallido setulis validis consperso. * 
On dead wood. Venezuela (Fendler). : 


Pileus thin, flabelliform, vaulted, 13 inch long, nearly as much broad, yellowish-brown, 
smooth, slightly streaked behind. Hymenium whitish, sprinkled with many scattered 
strong bristles. 

This species is closely allied to Sy dd flavum, Berk. (Peziza flava, Swartz), a figure 
of which was published in the Annals of Nat. Hist. vol. x., from an original specimen in 
the British Museum. It occurs in Venezuela as well as Jamaica, and is no. 129 of - 
Fendler's collection. It differs from the present species not only in colour, but in the 
bristles being divided above. Both belong to Fries’ genus Kneiffia, but its characters 
` scarcely separate it with sufficient precision from Hydnum. H. brunneoleucum is exactly 
a Kneiffia, but then Hydnum luteum, which is clearly congeneric, approaches so closely 
to Hydnum, that I know not how to separate it. The two species have a Tag Peziza- 
like habit which at once distinguishes them from all others. - 


2. ORATERELLUS PAPYRACEUS, Berk. & Curt. Pileo centrali tellisin umbilicato cocci- 
neo glabro; stipite gracili karer fusco ; ed lævi ochraceo. 
Venezuela (Fendler). : 


Pileus 3$ inches across, extremely thin, umbilicate or very broadly Bifundibuliform, 
smooth, slightly fissured at the margin which is arched, bright scarlet, with a few vein- 
like peneilings in the umbilicus; stem 4 inches high, not a line thick, smooth, reddish- 
brown, equal except at the very base, where it is slightly dilated as it springs from the 
‘matted mycelium. Hymenium quite smooth, pee 

VOL. XXI. : s 


130 REV. M. J. BERKELEY ON SOME NEW FUNGI. 


This very beautiful Fungus, which is remarkable in the genus Craterellus for its 
elegant habit and extremely thin pileus, is conspicuous from the beautiful red of the 
pileus, resembling that of fading leaves of Ampelopsis quinquefolia. It is altogether one 
of the most charming species with which I am acquainted. 


3. SKEPPERIA, nov. gen. (TAB. XXV. A.) 
Stipes brevis, lateralis, in pileum tenuem utrinque arcte convolutum clavulamque obtusam referentem 
abrupte transiens; pileus externe cellulosus intus filamentosus.— Genus Stereo affine, Edmundo 
Skeppero plantarum Cryptogamicarum solerti indagatori dicatum. ` 


S. CONVOLUTA, Berk. & Curt. 
On dead sticks, Venezuela (Fendler, no. 286). ; 

About 1 line high. Stem rising from a little orbicular disk } of a line high, but con- 
fluent for some distance with the pileus and rather paler; pileus 2 of a line high, of a 
deep vinous brown, sprinkled with little heaps of granules arising from the breaking up 
of the outer coat, strongly convolute on either side, so as to form a little compact clavate 
body with-a deep groove on one side like that of a grain of wheat, composed externally of 
cells, internally of filaments which give rise to the hymenium. Hymenium pale, studded 
with cystidia. 

This singular Fungus looks at first sight like a little Mitrula, with which genus it 
may be confounded if the microscopical structure is neglected. The outer surface, how- 
ever, is covered with little heaps of granules consisting of a number of deeply coloured 
cells, beneath which smaller cells occur, which pass into branched threads, and these into 
an even hymenium consisting of closely packed sporophores with projecting cystidia. 

Few genera are more singular in their characters. Notwithstanding the resemblance | 
of Pistillaria, there is no close affinity. Skepperia will come next to Stereum, without 
however passing into it by any intermediate forms, 


4. MITREMYCES RAVENELIT, Berk. (Tas. XXV. B.) Pusillus; peridio exteriori in ver- 
rucas parvas liberas vel cohærentes dehiscente; interiori glabro; intimo conformi, 
nec spatio magno vacuo circumdato ; sporis ellipticis; ostiolo miniato. 

On the ground. Czsar’s Head, Mountains of South Carolina (H. W. Ravenel, Esg.). 7 

Rather more than 4 an inch high, including the stem. Stem short, consisting of intricate | 
mucedinous threads. Outer peridium cracking yp into warts of various sizes, which are 
sometimes scattered, but sometimes adhere together so as to form.a little lid. Inner 
peridium smooth, pale, terminated with a few triangular vermilion-coloured teeth; inmost | 
delicate, entirely filling the cavity of the second peridium ; flocci abundant, white, ter- 
minated with little racemes of obtuse processes; spores ovate, 3000 inch long, not 
granulated. ; 

Very distinct from all the other species in the larger size of the inmost peridium. The 

nearest ally is Mitremyces Junghuhnii, figured in the ‘Botanische Zeitung” for 1844, 

which has the teeth of the ostiolum pale, and the spores globose. 


REV. M. J. BERKELEY ON SOME NEW FUNGI. 131 


5. MESOPHELLIA, nov. gen. (Tas. XXV. C.) 
Peridium crassum, coriaceum, substratosum ; capillitium fasciculato-anastomosans, ad columellam cen- 
tralem suberosam liberam radians ; flocci subflexuosi; sporæ breviter fusiformes utrinque obtusius- 
culæ.—Genus Cyclodermati, Klotzsch, affine ; species hypogææ. 


M. ARENARIA, Berk, 

From 3-1 inch across, elliptic, somewhat depressed, subterraneous, clothed externally 
with white flocci which attach themselves to little grains of sand. After the external 
down has become exolete, dark branched veins are seen to run over the peridium, without 
however giving off free threads, as in Hysteromyces. Peridium single, coriaceous, appa- 
rently consisting of several compacted strata like wasp pasteboard; flocci pinkish-grey, 
radiating in little fascicles from the peridium to the large central corky columella, which 
is either entirely free, or attached by one or more tendinous threads; spores fusiform, 
short, slightly obtuse at either end, 5545 of an inch long, of the same colour as the flocci. 

This genus approaches close to Cycloderma, Klotzsch, which appears to have been seen 
by no other botanist ; but there is no inner peridium, the columella is not a prolongation 
of a stem, and the spores are not globose. It is a most interesting addition to the Hypo- 
gæous Fungi, and, like Cycloderma, connects Trichogastres with Myxogastres through 
Lycogale. The early condition of the plant is, however, quite unknown. The colour of 
the spores reminds one of Lycogale, and the veins of Hysteromyces. A Fungus was 
found by.Mr. Drummond on the Swan River, resembling a large cocoon, which is 
undoubtedly congeneric with this, if not identical. The specimens were, however, very 
imperfect. 


‘EXPLANATION OF THE PLATE. 


Tas. XXV. A. 


Fig. 1. Skepperia convoluta, nat. size. 

Fig. 2. Single plant, magnified. — 

Fig. 3. Section of pileus before the stem completely vanishes. 
Fig. 4. Section of ditto more highly magnified. . 

Fig. 5. Hymenium. ; 


‘Tas. XXY. B. 


Fig. 1. Mitremyces Ravenelii, nat. size. 
Fig. 2. Section slightly magnified. 
Fig. 3. Flocci from young peridium and spores. 
Fig. 4. Flocci from old peridium. 
s2 


25. A192. 


tab. 


Vol. 


Soc 


Linn. 


Jens. 


prise ©] 


IX. On the Growth and Composition of the Ovarium of Siphonodon celastrineus, Griffith, 
especially/with reference to the subject of its Placentation. By Josera DALTON 
Hooker, Esq., M.D., F.R.S. & L.S. (With a Plate, Tas. XXVI) 


Read June 16th, 1857. 


IN a learned memoir upon some remarkable plants in the Hon. E.I.C.’s Botanic Gardens, 
Calcutta*, Mr. Griffith established the genus Siphonodon upon a very curious Malacca 
tree of doubtful affinity and singular structure, and accompanied his description with 
many observations of the highest interest and importance to the student of structural 
and morphological botany. Amongst these observations is one, which, though published 
for now nearly fifteen years, has never attracted the attention of botanists, owing to the 
limited circulation of the Calcutta Journal, but, which, from its bearing on the subject of 
placentation, has a peculiar interest to myself; for it appears to me to be, if correct, the 
strongest proof hitherto adduced in favour of the theory which regards the placenta as 
terminating the axis, and not as being referable to the carpellary leaf. To this theory my 
own experience is opposed, and as I believed I had proofs of the invalidity of what ap- 
peared to me to be the most cogent arguments previously adduced in favour of it, I more 
particularly wished for an opportunity of testing the accuracy of Griffith’s statement 
with regard to Siphonodon. Mr. Griffith’s conclusion is, * I beg to propose this plant to 
botanists, as an instance in which the placenta is the termination of the axis, bearing 
around its base a verticillus of ovula, and produced upwards into a stigma, a single organ, 
surrounded for the most part by a style with which it has no connexion.” (Joc. cit. p. 255.) 
The foundation for this proposition is Griffith’s conviction that the ovules, of which 
there are many in one horizontal series, are developed on an axis in the centre of the 
flower, independently. of the carpellary leaves, and not enclosed by them or by any other 
organs except the perianth; and that the subsequently developed carpellary leaves form. 
a verticillus externally to the ovules, and rising upwards and inwards, finally enclose 
them. These conclusions appear to me to be founded on erroneous observations, and the 
chief object of the present communication is to lay before the Society my reasons for 
supposing them to be so. 
` I am indebted to my friend Dr. Thomson, who was well-aware of the interest attached 
to this plant, for specimens of Siphonodon flowers preserved in spirits, gathered from the 
same tree, I believe, in the Hon. E.I.C.’s Botanic Gardens, which was described by Griffith. 
These flowers are ih all stages of growth, from buds scarcely perceptible to the naked 
eye, to fully expanded flowers, which measure about 4inch in diameter: I find, how- 
ever (as is usual in flower-buds), that there is little relation between the size of the bud 
and the development of its reproductive organs. In all, I invariably find both whorls of 
the perianth to be fully formed in two closely imbricated series, before any traces of the 


* Calcutta Journal of Natural History, vol. iv. no. xiv. July 1843, p. 231, t. 14. 


134 DR. J. D. HOOKER ON THE GROWTH AND COMPOSITION 


ovules or carpellary organs are differentiated, and I shall therefore commence with the 
bud at this stage, as the first in which the structure and composition of the ovary can 
be studied. 

1. In the first stage (PL XXVI. fig. 2) the bud is a minute turbinate body, about 4 line in 
- diameter, with little external distinction between the flower and peduncle. On a vertical 
section (fig. 3), the double perianth is found to be wholly superior, the sepals placed highest, 
then the petals, then the stamens, whose anthers (s) form minute clavate bodies, sessile 
at the very base of the petals. All these organs are placed considerably above the plane 
of the centre of the flower, which presents a depressed area with a very low broad central 
mamilla, whose vertex scarcely rises to the level of the plane of the base of the stamina. 
At this period the ovary is manifestly plunged into the apex of the peduncle, and can 
by no means be regarded as invested by the true calyx, the bases of the sepals being 
obviously inserted in the margin of the broad obconic apex of the peduncle. In some 
buds at this period, the central mamilla of the axis (the future style of Griffith) is sur- 
rounded by a low broad ridge, or annulus, correctly described by Griffith as belonging to 
the carpellary leaves. 

This is Griffith's earliest stage; he, however, regards the central mamilla as the pistil- 
lum itself, a view which is irreconcileable with the fact that the ovarian cells are in the 
next stage developed below the plane passing through its base, and removed from the 
circumference of its base. 

2. The next marked stage (fig. 4) is that in which the anthers are fully formed and their 
polliniferous cavities defined, and hence well distinguishable. At this period the central 


mamilla has risen, and presents the appearance of a style consisting of a short column — : 


with a depressed conical hemispherical base; it is more or less invested by the annulus (a), 
which was obscurely seen in the first stage; this annulus has five notches at its mouth, 
is very thick, and points upwards and inwards towards the apex of the column, being 
usually applied to its base. The relative position of the stamens and perianth to the plane 
of the ovarium is the same as in the first stage; vascular bundles are developed in the 
petals, and are traceable continuously downwards into the peduncle, and branches from 
these bundles supply the stamens also. The stamens arch over the annulus, and the con- 
niving apices of the anthers almost reach the central column ; they are opposite the five 
notches in the annulus, and five slight ridges on the annulus correspond with its teeth 
and with the interstices between the anthers. 

At this period the ovules are first developed. A whorl of about twenty minute cavities 
are traceable in a plane below the point of junction of the annulus and conico-hemi- 
spherical base of the column ; the ovarian cavities (0) point downwards and outwards. The 
ovules appear as minute papillæ, occupying the base of each cavity towards the axis; 
they present no distinction of parts whatsoever. No vascular bundles are discernible in 
the ovarium. | | 

In this stage I have included the second and third stages of growth described by 
Griffith, because, after examining an extensive series of buds, I find very great irregu- 
larities in the time of appearance of the ovarian cavities, relatively to the development 
of the annulus. In some buds I find the ovules formed before the annulus has reached 


‘ 


OF THE OVARIUM OF SIPHONODON. 135 . 


the rounded shoulders of the column, in others not till after it has enveloped that organ. 
It is at this period, however, that the important difference between my observations and 
Mr. Griffith’s occurs, and as it is upon this that the whole question of the placentation 
turns, I have taken every means of endeavouring to arrive at the truth. Griffith says, 
that at the period when the base of the central column becomes concealed by the growth 
of'the annulus, very young ovula may be detected, apparently attached around the whole 
base of the column, and not presenting any manifest relations with the carpellary leaves. 
He adds, that at this time a double long section had not shown him any solution of 
continuity resulting from foreibly pulling back the carpellary leaves (annulus) and thus 
exposing the ovula. On the other hand, I, at no period, find the ovules to be developed 
freely at the base of the column; I have sought in vain between the annulus and column 
for any such appearance, removing the annulus piecemeal with the utmost caution; and 
in every double vertical section that I have made, I have found the ovarian cavities to be 
developed even at a much earlier period than that indicated by Griffith, below the junction 
of the annulus and column, manifestly within the substance of the ovary. The cavities 
containing the ovules cannot at any period be exposed without tearing the annulus, which, 
however, owing to the delicacy of the tissues and the excessive mfnuteness of the organs, 
it is sometimes difficult to avoid doing. 
3. In the succeeding stage (fig. 5), the anthers are fully formed, ånd, arching inwards, 
conceal the whole vertex of the ovary, leaving the apex of the column alone exposed 
between their apices (fig. 8). The eylindrical portion of the column has lengthened, and is 
terminated by a capitate stigma-like head; its conical base is proportionally smaller and 
more depressed. The annulus is very much thickened and grown in all dimensions; it 
has advanced up the column, and presents on a double vertical section two broad shoulders 
embracing the column, which it thus sheaths in a tube; its upper surface is moulded by 
the pressure of the anthers which lie between the five radiating ridges accurately described 
by Griffith. The five teeth (figs. 8? & 64) terminating the ridges are now more promi- 
nent, turn upwards, and are very cellular and fimbriated; and a distinct groove is formed 
on the surface of the annulus, close to its orifice and concentric with it. The ovarian 
cavities are fully formed, linear-oblong, and all point outwards, but some upwards and 
some downwards. The ovules are horizontal with an obscure lateral raphe, and each is 
conformable to its cell. The column has an evident cuticle, as has also the contiguous wall 
of the annulus. At this period vascular bundles appear in the ovary, and are traceable 
downwards from each ovular insertion, and transparent lines are continued from each 
ovule upwards into the annulus, to whose inner face these ascending lines are parallel and 
contiguous. The tissues of the sheath of the annulus are pale and translucent. Mr. 
Griffith says, that at this period the stigmatic surfaces will be found at the conical base of 
the central column, at the point of its junction with the base of the annulus; he says, 

«the lines of communication, by which each isolated ovulum is placed in the usual con- 
_ ditions to receive the male influence, will be found to terminate on the circumference of 
the conical base of the central column.” On the contrary, I find the cuticle to be uni- 
formly extended over the points indicated; but its tissue is there rather looser, the cells 
which form it being elongated inwards, giving when torn the appearance of stigmatie 


136 DR. J. D. HOOKER ON THE GROWTH AND COMPOSITION 


tissue, and causing that fragility of that part of the ovarium which led to Griffith’s sup- 
posing that the ovarian cells were formed subsequently to the freely developed ovules. The 
true stigmatic canals I find terminating in the papillose fimbriated teeth of the annulus. 

4. Towards the period of expansion of the flower (fig. 6), a considerable change is found — 
to have taken place in the relative proportions and positions of the ovary and perianth. 
Owing to the upward direction of the development of the ovary, it is now half-superior, 
in the usual acceptation of that term. The stamens (fig. 9) form a closely imbricating 
whorl, appressed to the surface of the annulus, the stigma-like head of the central column 
appearing in the space between their apices, and the five cellular fringed teeth of the 
annulus also projecting. 

On removing the stamina, the protuberant surface of the annulus is more strongly 
5-ridged, and the groove round its orifice is deeper. A double vertical section shows 
that the column is much elongated, and its surfade covered with a more evident euticle, 
as are its capitate stigma-like apex and the adjacent walls of the annulus. The external 
surface of the annulus is also covered with a similar cuticle, except at the groove sur- 
rounding its orifice, and on the fimbriated teeth, which are manifestly the true stigmata, 
and are formed of loos papillose tissue (fig. 13). The ovarian cavities are at this period 
found to be considerably displaced, some being carried a little upwards and others down- - 
wards, so that two or three are often exposed at one vertical cut. Unrollable spiral 
vessels are now found in the vascular cord that passes downwards from the ovules, but 
. none in the transparent line that passes upwards from the ovules parallel to the inner 
walls of the annulus, these lines being formed of loose conducting tissue, terminating in 
the stigmatic teeth at the orifice of the annulus. A transverse section of the annulus 
(fig. 10) now shows five pale spots of loose cellular tissue; these are the stigmatic canals, 
placed very close to the inner walls of the annulus. 

5. At the period of expansion (fig. 7) the stamens are found to have risen from the 
surface of the annulus, and the stigmatic teeth are sphacelated and covered with pollen- 
grains, which adhere to its surface and to the surface of the mouth of the tube; but I 
have not succeeded in tracing the pollen-tubes into the stigmatic canals, probably owing 
to the excessive minuteness of the pollen in this plant. 

The differences between my own and Mr. Griffith’s observations are, first, in the ovules 
being developed in cavities of the ovarium, and not free at the base of the conical central 
body, which he has mistaken for a placental development ; secondly, in the presence of a 
cuticle over the whole surface of the said body and on the contiguous walls of the 
annulus; thirdly, in the absence of any vascular bundles passing from the ovules upwards 
into the annulus; and fourthly, in the absence of any stigmatic puncta at the base of the 
central column, and of stigmatic lines passing from these inwards to the ovules. The 
- groove round the mouth of the annulus, the absence of cuticle over it, and the structure 
and consequently the stigmatic nature of the teeth of the annulus, seem further to have 
escaped Mr. Griffith’s notice. 

If my observations are correct, the composition of this very remarkable ovarium is — . 
reducible to the known modifications of carpellary structure, and may be regarded UN 
consisting of a whorl of five connate carpels adnate to a central fleshy axis or torus; 


OF THE OVARIUM OF SIPHONODON. 137 


this torus being produced, as in Nymphea, Victoria, &c., into a styliform and stigma-like 
body. Each carpel bears two ovules on each of the ten marginal placentæ, making twenty 
ovules in all. The annulus is the free upper portion of the five confluent carpellary leaves ; 
the five ridges are the lines of junction of these; the five stigmata are each double, 
formed by the terminations of the confluent placental margins of the adjacent carpels, as 
in Papaveraceæ and many other Orders. 

There is one point, however, to which further allusion is necessary, as possibly in part 
explaining Mr. Griffith’s views; and this is, the above-mentioned line of loose cellular 
tissue that extends from the base of the central column to the ovarian cavities, and which 
is met at the ovarian cavity by the true stigmatic tissue. It is very possible that this line 
indicates the existence of a stage in the early state of the ovarium in which the carpellary 
leaves were not completely closed; for though I feel satisfied that the ovules of this plant 
are at no period independent of the carpellary leaves, and are indeed formed in cavities of 
those leaves and from their margins, it does not follow that there may not have been a free 
. opening to these cavities, or one closed only by a very lax tissue. It is indeed held by some 
. botanists, that all carpellary leaves are congenitally open, and close more or less com- 
pletely afterwards; an opinion which is not as yet absolutely proven, though I cannot 
but think that the open ovary of Conifere* and its allies is a strong argument in its 
favour. This however, if true, by no means reconciles Mr. Griffith's observations, or his 
theoretical view of the structure of the ovary of Siphonodon, with my own. 

The structure I have described in Siphonodon suggests a different view of the affinities 
of this obscure genus than those which have been doubtfully adopted by Griffith, though 
in the absence of ripe seeds it will be difficult to establish these, and I shall not therefore 
attempt to do so now. 

In submitting this very singular plant, then, as a strong proof of the validity of those 
laws of carpellary placentation which it has been supposed to have subverted, I cannot 
refrain from expressing my admiration of the learning which Mr. Griffith has displayed 
in his discussion of the view he somewhat hastily adopted ;—of the guarded manner in 
which he expresses his opinion ;—of the full weight he gives to every structural point that 
seems to him to militate against it, and of the candour with which he states every adverse 
argument that suggests itself to him. Though I believe his observations and conclusions 
to be erroneous, it must be recollected that the plant is a very anomalous one, its parts 
exceedingly small, and that my experience assures me that specimens preserved in spirits, 
such as I examined, are in many respects much better for determining structural points 
from than living ones are. Mr. Griffith’s paper further abounds in acute observations on 
many other points in the structure of Siphonodon, to which I have not alluded; and it 
contains, in a note, a short abstract of the only accurate account hitherto published, so 
far as I know, of the development and true nature of the ascidia of Nepenthest. 

* It appears more consonant with the known laws of vegetable morphology to regard the Coniferous ovary as an 
arrest of the usual tendeney of ovaries to close, than to suppose the ovaries of most Phænogams to be congenitally 
closed, and that of Conifers open. 

+ Some years ago I prepared drawings of the development of the pitchers of Nepenthes, from plants in the Royal 


Garden at Kew. These confirm Griffiths observations in every particular, and prove the pitchers to be modifications of 
excurrent midribs. Each pitcher commences as a gland at the anterior apex of the conical mamilla, which represents 


VOL. XXII. T 


138 : DR. J. D. HOOKER ON THE GROWTH AND COMPOSITION 


— Before dismissing the subject of carpellary placentation, I would state that the other 
‘arguments in favour of free axile placentation, to which I have alluded at the beginning 
of this communication, are, the free central placenta of Primula, and the position of the 
ovule of Tagus. I am not aware of any deviation from the free central placentation 
having been observed in Primulacee except by Prof. Henslow, who kindly placed in my 
hands for examination a monstrous flower of Primula vulgaris, in which two ovaria pre- 
sented ovules developed on the confluent margins of some of the carpellary leaves, far 
removed from the base of the ovary, and wholly unconnected with the central placenta, 
which still existed, but in a reduced form, and bore ovules. | 

The other case is that of Taxus, adduced I believe originally by M. Schleiden, who 
considers the ovule as terminating the bracteate peduncle or rachis which bears it. That 
this view is not tenable appears to me to be proved by the fact that I have found two and 
even three ovules at the apparent apex of the peduncle. I further cannot but consider 
the bracteate peduncle of the female flower of Taxus* as a genuine cone, the majority of 
the bracteæ of which bear no ovarian seale and ovule. When more than two ovules are 
developed in this cone, these almost invariably wither and disappear at a very early 
period; but I have examined partially diseased cones of Taxus, in which the second ovule 
had enlarged and persisted in a scale below the terminal one. 

With such a preponderating mass of evidence in favour of carpellary placentation, 
derived from the normal condition of both simple and compound pistilla, in which the 
relation of the parts cannot be misinterpreted, and from monstrous states of ovaries whose _ 
composition is more obscure, it does seem to me unphilosophical to call for the operation — 
of a second law to explain the comparatively few exceptional examples of placentation 
that are not at once referable to the carpellary. Of these exceptional cases, Primulaceæ, 
Caryophylleæ, and their immediate allies have always been supposed to be the most 
difficult to explain away, but Caryophyllec, both in their normal state and as monsters, 
present many cases of carpellary placentation, and Professor Henslow’s Primula demands — 
a double law for one flower, if the free axile placentation be not considered a modification | 4 
of the carpellary. j D 

If it be admitted that the floral whorls are formed from leaves, even when appearances - 


a leaf in the earliest stage of its development. A contraction below the gland separates it from the lamina of the leaf, 
which is independently developed with a convolute vernation. 

While upon this subject, I may further mention, that the bilobed lamina of Dionæa is unquestionably the true — 
- lamina of the leaf of that plant ; and that it is inflexed in early vernation, as in Drosera, to which it is so closely 
allied, and to which it is further united by means of Aldrovandra. 

Of Aldrovandra I may remark, that this very rare and singular plant has been rediscovered by Dr. Thomson in the 
immediate vicinity of Galcutta. It was originally found there by Dr. Roxburgh, but unsuccessfully sought for by 
Dr. Wallich, Griffith, Voigt, and indeed every Calcutta botanist. Though unknown in any intermediate spot between 
Europe and Calcutta, I can find no difference whatsoever amongst the specimens from these widely-sundered localities. — 

* The fleshy cup of Taxus, which is usually called a disk in botanical works, but which is regarded as an arillus by 
Schleiden, is undoubtedly the outer coat (primine) of the ovule; it is developed at an early stage as an annulus in 
immediate contiguity with the second coat (secundine) : this second coat in Taxus, as in many other Conifers, becomes 
the outermost covering of the seed, or tegmen. I have ventured to allude to this here, as I am not aware that the 


' age of the integuments of the seed and ovule of Conifere has been correctly stated in any general systematic 
work. 


OF THE OVARIUM OF SIPHONODON. 139 


are against this view, and when it may be impossible to demonstrate it; and that 
this law allows of no exception, —it seems natural to suppose that the law of the formation 
of ovules in one of these whorls should be regarded as no less absolute, and that any 
explanation that does no violence to nature should be accepted to account for apparent 
- exceptions, in preference to abandoning the general law and adopting a particular one for 
those exceptions. Under this point of view I should prefer agreeing with those botanists 
who consider that all placentation is axial, and that carpellary insertion is to be ex- 
plained by supposing axes to be produced upon the carpellary leaves, rather than conclude 
that nature resorts to two totally distinet and opposite principles in effecting the same 
purpose with the same organs. 'The instances, however, of the ovules being normally 
developed on the margin of the carpels, are so extremely numerous and convincing, and 
the cases of insertion which are not at once reducible to this relation of parts are so 
few and so scattered through different Natural Orders, that I cannot but regard the 
latter as insufficient to invalidate the law in question, and to demand the operation 
of another. 


EXPLANATION OF THE PLATE. 


Tas. XXVI. 


Fig. 1. Flowering branch of Siphonodon celastrineus :—of the natural size, from Mr. Griffith’s drawing. 
Fig. 2. Bud at the earliest period of the appearance of the parts described :—of the natural size. 

Fig. 3. Vertical section of ditto; s. stamens. 

Fig. 4. Vertical section of upper portion of more advanced bud, the ovular cavities having just appeared. 
Fig. 5. Another bud further advanced, the stigmata formed, and vascular cords to the ovules. 

Fig. 6. Flower about to expand, cut vertically. The ovules are displaced, and the annulus swollen. 
Fig. 7. Flower after impregnation, with the stamens still attached. 

Fig. 8. Annulus from fig. 5, seen from above; t. the stigmatic teeth. 

Fig. 9. Stamens seen from above, concealing the annulus. 

Fig. 10. Transverse section of annulus, showing the five stigmatic cords or passages. 

Fig. 11. Portion of ditto much more highly magnified. 

Fig. 12. Portion of annulus, with stigma, conducting tissue, ovules, and vascular cords. 

Fig. 13. Portion of another annulus with stigmata, an ovule and its vascular cord. 

` All but figs. 1 & 2 more or less highly magnified. 


$ 


T2 


Brans. Linn. Spo. Vol XXIT. tab. 26. p.140 


JD Booker, MD. del. G Jarman, sc. 


[ 141 ] 


 X. Further Remarks on the Organs Jound on the bases of the Halteres and Wi ings of 
Insects. By J. B. HICKS, Esq., UD. Lond., F.L.S. 


Read June 2nd, 1857. 


IN à former paper, published in the Journal of the Linnean Society *, I endeavoured to 
describe a curious and beautiful structure to be observed at the base of the halteres of 
‚the Diptera, and a similar but less elaborate condition on the subcostal nervures of the 
wings of both Diptera and other insects, and stated that generally the organs in question 
on the posterior wings of the 4-winged tribes were the more numerous: I also proceeded 
` to show, that into the halteres, as also into each of the wings of insects in general, could 
be traced a nerve of considerable size, varying in the different tribes and genera in that 
particular, and that the nerve proceeding to the posterior wings was generally the larger, 
and also that the one supplying the halteres of the Blow- and Drone-fly was the largest 
nerve in. the insect, except the optic. I may here repeat, that each of these structures 
consists of very thin and transparent, hemispherical, or more nearly spherical projections 
from the cuticular surface, beneath which the wall of the nervure is deficient, so as to 
allow a free communication with its interior ; these organs are arranged in rows on the 
halteres, and in variously shaped groups in the wings. I concluded that the nerve pro- 
ceeded to these structures, inasmuch as there was no muscle in the wing or halteres, and 
that consequently the nerve must be one of sensation ; and looking at the elaborate and 
regular disposition of the structures composing it, I was induced to think the organ was 
one of special sensation, and that, situated as it was near the great thoracic spiracle, as 
well as exposed to constant and rapid currents of air, it seemed not improbable that it 
was connected with the olfactory sense. 
At that time I had not been able to prove the actual connexion between the nerve and 
these vesicles, but I have now the pleasure of exhibiting a drawing (Tab. XXVII. fig. 1) 
— made from a specimen which I have preserved, showing most distinctly the nerve in its 
whole course, giving off in its passage by the groups of vesicles, numerous branches which 
can bé traced into their bases. This may be seen by any one taking due care in the 
preparation of the specimen. The drawing is from the second wing of a Longicorn beetle 
(Strangalia elongata); though perhaps every wing will not show it well, on account of 
the position of the nervures. I may here mention the method employed, which is one, 
I believe, frequently in use :—After cutting off the wing and washing it well in water or 
spirits of wine, and draining off the major part by blotting-paper, I immerse it in spirits 
of turpentine for a week or two, after which it is placed in Canada balsam between glass 
in the usual way, taking care not to heat it, as that renders the nerve too transparent. 
. In those parts which are too dark for observation, I have been enabled to render them 
colourless by chlorine: a little chlorate of potash, and a few drops of hydrochloric acid, 


. * Vol. i. p. 136, Nov. 1, 1856. 


142 DR. J. B. HICKS, FURTHER REMARKS ON 


mixed in a small bottle, with the addition of a little water when the effervescence is brisk, | 
will in a short time remove sufficiently the colour of most insects. Some parts which are - ' 
quite black, ås the elytra of beetles, wings of bees, &c., require a day or two, and some 
even a week. The hydrochloric acid has also a valuable property in rendering the nerve- _ 
tubes more perceptible. This plan of bleaching will be found invaluable in examining 
the structure of antennæ, which are often quite black, as I shall hereafter show. 4 
I have also shown the distribution of the nerve in the wing of Tenthredo viridis: the — 
bundle of nerves going to the vesicles was well marked in the specimen from which the 
drawing was taken. i "o 
These examples will, I think, serve to point out the intimate connexion between the 
— vesicles and the nerves. Whether the whole nerve be distributed, in all instances, to these 
organs, I am inclined to doubt, since a small branch seems to pass beyond the point | 
where they cease. j i 
I shall now endeavour to trace these organs through the different tribes, so far as I have 
been able. i 
In the Hymenoptera I find that they exist far more extensively than I supposed when 
my former paper was read ; their detection has been mainly owing to the use of chlorine, 
‘so few of this Order having light-coloured nervures in the wings. The figures here given - 
represent these organs in Ophion lmteus (Tab. XXVIII. fig. 2) and Tenthredo viridis 
(Tab. XX VIII. fig. 3), two species which are good examples of the tribe, and in which the 
nervures are the most transparent. These organs in the Wasp (Vespa vulgaris), Honey- ' 
bee (Apis mellifica), Andrena fulvicans, and Tenthredo lucorum, having a great resem- - 
blance to the above, I have not thought it worth while to figure them. 
They consist of two groups on the upper, and one scattered group on the under side of _ 
the subcostal nervure, amounting in Ophion to 200-300 above, and perhaps 100 beneath, 
with a smaller group at the end of the nervure. 
With regard to the Diptera I have nothing to add to my former remarks, excepting | 
that I have found the vesicles in Hippobosca equina distinctly marked both in the — 
halteres and wings. . The diameter of each vesicle is 3300 inch. 
In Tipula oleracea they are also well shown on the wings. 
In the Hemiptera they are very scanty; I have observed and drawn those on the — 
second wing of the Tree Bug, a species of Pentatoma. HM 
In Notonecta glauca, or the Water-boatman, they are even more simple. `° 
In Corysus I have been unable to find them. I have carefully examined the elytra of © 
the common Bed-Bug (Cimex lectularius), but cannot find anything definite; some papilla - 
occur on them, but they do not seem to be arranged in any distinctive manner. i 
In the Coleopterous group we find them highly developed, as may be observed in the — 
- drawing of these organs in Strangalia elongata in Plate XXVII. fig. 1. 
They occur in numerous groups on the subcostal nervure, mostly at the widest part, 
but are also scattered along it to the joint of the wing, where we find about ten or twelve — 
large vesicles in a group, after which they cease. | 
a the Carrion Beetles (Necrophorus) they are very well developed, as also in Silpha, 
where they are rerharkably large, considering the size of the insect. 


ORGANS FOUND ON THE HALTERES AND WINGS OF INSECTS. 143 


In Lucanus Cervus, Melolontha vulgaris, and Cetonia aurata they are not so large, but 
still are numerous. i 

In Carabus granulatus they are found in the rudimentary second wing, more diffused 
over it than in some other genera, inasmuch as other nervures than the subcostal are 
furnished with them, though not to the same extent. 

In an apterous Beetle which I examined, I did not find any similar organs on the part 
of the side corresponding to the position of the wings. 

With regard to the distribution of the nerve in the elytron of the Coleoptera, there are 
some curious facts, which, I think, have not been hitherto noticed. If the elytron be 
treated in the way above mentioned, and first bleached if it be dark-coloured, the 
branching tracheæ will at once be recognized, accompanied by the nerve, which gives 

out branches as it proceeds. Now, in all the elytra of beetles I have examined, there is a 
number of distinct projections from the whole dorsal surface, in many hemispherical, in 
some like small papillæ just raised above the cuticular surface; these eminences are 
arranged in a tolerably regular manner, and are quite distinct from other irregularities of 
surface. The hemispherical forms are very transparent, the wall of the elytron being 
deficient beneath them, as in those before described. There are two or three papillæ at 
the highest point of the vesicle. Some, when viewed by polarized light, exhibit a dark 
cross, something after the manner of starch-cells; those of Melolontha show it well. 

In many beetles the vesicle or papilla is accompanied by a hair, which always, I 
believe, when so situated, overarches it, as if for the purpose of affording protection. 
Now, into each of these vesicles and papillæ a branch of the nerve can be distinctly 
traced, as I have shown in the two drawings taken from Melolontha (Tab. XXVII. fig. 2) 
and Carabus (Tab. XXVIII. fig. 1). In the case of Melolontha the hair may be seen 
overarching the vesicle. ; 

With regard to the Lepidoptera I have found nothing new, except that the organs 
are well developed in the female Vapourer Moth (Orgyia) on the rudimentary wing. 

In none of the Orthopterous specimens hitherto examined have I been able to find 
any structure like that described in the other groups. Å nerve, small in some, but large 
in Mantis religiosa, is to be found entering the wings, yet I have been unable to make 
out anything satisfactory, except a few scattered vesicles on one or two of the nervures. 
~ In the Neuroptera these organs can be distinctly found, though not quite so fully 
developed as in most of the preceding groups. In Libellula depressa they are not, rela- 
tively to the size of the insect, by any means so numerous as in the Coleoptera. I have 
drawn those of Agrion, where they are but few ; and those of Phryganea, which are rather 
more simple. In the Flea (Pulew irritans), I have been unable to detect anything 
connected with this subject. i 

I have not been able to distinguish any difference between the sexes in reference to 
these structures, their relative size corresponding to that of the individual, which is 
generally larger in the female. The species of a given genus, especially of the Dipterous 

- group, exhibit a marked similarity in the appearance of these organs ; so that it is possible, 

by examining them, to tell to what genus any Fly, whose,halteres are presented to you, 


belongs. ' 


144 DR. J. B. HICKS, FURTHER REMARKS ON 


In regard to their relative development in the various Orders, so far as I have already 1 
gone, I am enabled to say, that they are most perfect in the Diptera, next so in the — 
Coleoptera, rather less so in the Lepidoptera, but slightly developed in the Newroptera, 1 
scarcely at all in the Orthoptera (though this assertion may be hereafter modified), and 
that only a trace of them exists in the Hemiptera. 

I have found in each of those divisions the most numerous and largest organs in 
those insects which are notorious for their acute scent, ez. gr. in the Carrion Beetles, 
Blow-flies, and Bombycidæ, and the smallest and fewest in those that are either very | 
sluggish in habit, as the Bugs; or in those that hunt their prey by sight, as the Libellule; — 
or those having a large development of the eye, as T'abanus, &c. 


Note.—Since the above paper was read, I have been favoured, through the kindness of 
Mr. J. O. Westwood, with the following extracts from Keller’s * Geschichte der gemeinen 
Stubenfliege’ (pp. 22, 23), taken from copies of the two versions in the Hopeian Library 
at Oxford. The original version in German, dated 1764, is Mr. Westwood's copy, and 
the French, dated 1766, is Mr. Hope's. 

“Les baguettes sont placées derrière les stigmates. La couleur jaune de l'une de ces 
baguettes (très grossie, pl. 4. fig. 3) est bien de beaucoup plus vive que celle des vésicules 
résonnantes [alulets], cependant elle paroit pourtant être faite de la méme matière, du 
moins la téte de la baguette, g, et ne consiste qu'en une simple membrane, bordée d'un. 
chassi, d’une matière un peu plus solide, qui les divise dans la même proportion en deux . 
hémisphères. L’on voit encore mieux ces parties dans les Cousins qui de méme que tous — 
les insectes à deux ailes sont pourvüs de ces baguettes et qui ont aussi deux vésicules | 
résonnantes trés petites, que dans les mouches, puisque les tiges des vessies presque trans» - 
parentes sont plus longues. . Aussitôt que nous examinons la vessie antérieure au travers | 
d'une loupe nous ne pouvons faire aucune autre conclusion, si non, qu'elle soit creuse, 
ou pour mieux dire que ce soit une vessie enflée. Nous pouvons conjecturer la méme 
chose ici par comparaison. Mais cette conjecture devient d'autant plus certaine, si nous 
examinons la baguette d'une mouche morte, où nous trouvons sa tête affaissée et con: 
tractée comme une vessie vuide d’air (pl. 3. fig. 27 €). 
 * Nous verrons dans Ja suite que cette cavité des baguettes est faite dans des desseins 
trés importants. La verge [peduncle] garnie de poils très courts, ou la tige a le plus 

d'épaisseur près du corps; elle est inflexible, et presque garroté par en haut de plusieurs 
nerfs [oben mit vielen nerven als wie mit Schnüren umwickelt; mit einem worte; $ 

beschaffen, dass man ihm gleichsam seine Stürke ansicht]; en un mot, elle est faite de 
manière que l'on peut juger de sa force par les dehors." KC 

The plates in both versions of the work are from the same coppers. In the figures of | 
the halteres a rough view is given of the groups of vesicular projections at the base of. 

those organs. The individual vésicles, however, are not represented as distinct from 
each other. By the * nerves," described as being outside, and as giving an idea í 
strength, I presume the author means * nervures,” which he considers the rows 9 
vesicles to be, and not nerves,derived from the ganglionic centres.—J. B. H. 


ORGANS FOUND ON THE HALTERES AND WINGS OF INSECTS. 145 


EXPLANATION OF THE PLATES. 
(All the figures are magnified.) 


Tap. XXVII. 


. Preparation from the wing of a Longicorn Beetle (Strangalia elongata), showing the subcostal 


nervure giving off branches passing to the vesicles: å, trachea; n, subcostal nervure. 
Elytron of Melolontha vulgaris: t, trachea; n, nervure ; à, portion more highly magnified, showing 
the termination of the nerves. 


4 | Tas. XXVIII. 


. Portion of the elytron of a species of Carabus: a a, trachea; b b, nerves proceeding to papillæ ; 


c c, markings in elytron, probably solid between upper and under wall. 
Portion of anterior wing of Ophion luteus: a, base of subcostal nervure, upper side; 5, end of 
nervure; €, under side of nervure. 


. Tenthredo viridis. A, portion of anterior wing; a, upper, b, under side. B, portion of posterior 


wing; a, upper, b, under side. C, section of wall of nervure through the organs; diameter at 
su rface zyngth of an inch. 


VOL. XXII. 4 U 


Trans. Lina. Soo. Vol XXII. tab. Z7. MILE. 


à nn ei... a, 


IB. Hicks, MD. dzl. €. Jarman. sc. 


[ 147 ] 


XI. On a new Structure in the Antenne of Insects. 
By J. B. Hicks, Esg., M.D. Lond., F.L.S. 


' Read June 2nd, 1857. 


THE object of this paper is to call attention to some peculiar structures to be found in 
the antennæ of insects, which have not been hitherto noticed. If we submit the antennæ 
to the bleaching process which I have before described (see pp. 141-2), these structures 
will be well brought into view ; they vary somewhat in form in the different genera, but 
all, I believe, possess the same elements, and are formed on the same principle. 
I shall first describe those of the Diptera. 
The antenna of the common Blow-fly (Musca vomitoria) is usually described as consisting 
of three joints, the terminal one being dilated and elongate, having a plumose seta or 
bristle on one side. There are many genera of Diptera which possess a similar antenna, 
and it is on this third dilated joint that the structures about to be described are found. 
When the antenna is not so constructed, although many of the joints present similar 
appearances, yet I believe they will not be found on the first and second joints, but almost 
invariably, if not always, to commence on the third, as may be seen in the accompanying 
figures. 
First, of the Blow-fly (Musca vomitoria) (Tab. XXIX. fig. 1. Over the whole surface 
- of the elongated third joint may be noticed at first sight, a multitude of transparent dots, 
apparently vesicles, but on closer examination and in profile, they will be found to be level 
_ with, if not rather below the general surface (fig. 1,0 1). These dots after a careful inves- 
tigation prove to be perforations of the inner coat of the wall of the antennå; closed in 
externally by a very thin membrane, which can be seen, when the antenna is crushed, 
on the lines of fracture. 
ind this closed perforation is a sac, rather larger than the perforation itself, which, 
| en the antenna is broken up, may be found floating about in the balsam (fig. Adi. 
i The diameter of tl se perforations of the inner wall of the antenna is about syy inch. 
They are placed very close together. The area on each half of the antenna covered by . 
these structures, has a surface-measurement of 4; inch by 35 inch, and as each square of 
go inch has within it 3025 (that is, a square having 55 on each of its sides), the total 
number embraced within the space occupied by them will be about 9075; and the total 
for both sides will therefore be about 18,150. But it will be proper to deduct from 
this number about 1000 for the space occupied by the apertures (presently to be described), 
‚ and some possible irregularities in the disposition of these organs; we may therefore 
assign 17,000 for the whole surface of each antenna. 
But besides this multitude of peculiar structures, there are numerous apertures leading 
into cavities, which may be either simple or composed of one to five chambers, all commu- 
v2 


148 DR. J. B. HICKS ON A NEW STRUCTURE 


nicating with the common aperture. These apertures are protected slightly by hairs which 
arise from the edges, and also a little way within them (fig. 1,62,0,d 2). There are about 
eighty of these cavities on one side of the antenna, and probably about the same number on 
the other; their diameter varies from 3555 to 1250 inch. The largest sacculated chamber 
is about gl; inch. By focusing down to the floor of this cavity, papillæ may be seen, each 
in the centre of an hexagonal area. On viewing these structures in profile, as at fig. 1, d, a 
there may be seen beyond, and adherent on all sides to their walls, a firm granulated 
mass of considerable thickness, and which appears to be tubulated nearest the walls of 
the sac. This latter investigation is attended with some difficulty, and requires much 
care, but I think I am not far wrong in stating that the hexagonal areas correspond 
to the tubes which extend inwards from the walls of the chambers. This is shown at 
fig. 2, d 2. | 

In Helophilus pendulus (Tab. XXIX. fig. 2), the smaller organs first described are not 
so numerous as in Musca vomitoria, while the cavities are more frequent, though less 
chambered. . Hairs also arise from the interior of the cavity, as is shown at fig. 2, d2. — 

In many antennæ of this shape, there are one or two openings very much larger than 
the rest, in the lower part opposite the origin of the seta; these are well shown in Mesem- 
bryum meridianum. They possess the same structure as the cavities before described. 
These cavities, and their chambers or sacculi, will, I think, be found to consist of the same 
elements as the smaller organs; the chambers being compounded of groups of these, and 
seeming to be formed by the infolding of the external surface, as is shown by the presence 
of hairs inside them. > 

The above structures I have found in all antennæ of this form, varying in relative 
numbers in the different genera. i 

Before I take leave of this form of antenna, I would throw out a suggestion with 
regard to its true structure. The seta or bristle (Tab. XXIX. figs. 1 & 2) is 3-jointed, 
the last one plumose or not, as the case may be. Now it seems to me that these are the 
last three joints of a 6-jointed antenna, the third joint of which is more or less dilated on 
one side, thereby throwing aside the last three. This I think will be seen to be the true 
explanation of the seta, when it is considered what peculiar and important structures are 
developed on the third joint, which is probably dilated for their reception. | 
. In Tabanus (Tab. XXIX. fig. 3) the third joint is much dilated, but unequally, so that 
the four other joints, although continued in the general direction of the antenna, are on 
one side of the axis. On all the five joints from the third to the apex, organs are found 
exactly like the smaller closed perforations in the Blow-fly, their diameter varying from 
. 060 t0 zooo inch. Between each is a tooth-like projection of the cuticle, like a hair. 
The largest are scanty, and chiefly found on the lower part of the third joint. 

In Bombylius the antenna is 5-jointed, the third joint only possessing these closed 
perforations, and being much elongated and slightly dilated; the diameter of the pores — 
varies from 3757 to 3357 inch. : 

In Hippobosca equina, the antenna is 5-jointed; the third, fourth and fifth joints being 
devoted to the reception of the same structures, They are depressed beneath the general 
surface, in the form of a saucer, at the bottom of which is seen the thin membrane across 


IN THE ANTENNÆ OF INSECTS. 149 


the perforation of the inner wall. The last joint is plumose, exactly resembling the third ` 
joint of the seta in Musca, Mesembryum, Eristalis, Volucella, and other similar forms. 

` In Tipula (Tab. XXIX. fig. 4) these organs (precisely like those in Hippobosca, Bom- 
bylius, &e.) are to be found on the third joint of the antenna, which is more elongated 
and dilated than the rest, as is seen in fig. 4, a. Their diameter is 3257 inch. 

In Ctenophora the third joint is much more dilated, as at fig. 4, 6, c. In Ctenophora 
bimaculata (fig. 4, c) this dilatation is well marked on one side, and on this part these 
organs are readily observed, and the nerve distinctly seen to proceed to them, as in fig. 4, el. 

In Bibio Marci the antenna is composed of nine joints; the first two are free from any 
of the above appearances ; but the last seven all possess them on one side, and a nerve 
can be traced distinctly to them. . But the most remarkable condition of this antenna is, 
that from the third to the eighth inclusive, on the outer aspect of each joint is a cavity 
of considerable size, similar to those described in Mesembryum, the appearance of the 
floor being the same. The aperture leading to this cavity is about ybg inch diameter, 
and around its wall is a layer, apparently of tubuli, as around the chambered cavity of 
the Blow-fly, and to each of these a nerve can be very plainly seen to be given off from 
the antennal nerve as it passes by them. 

In this insect the advantages of the use of chlorine are strongly shown, the whole of 
it being of a jet-black, which in a day or two becomes quite colourless. 

The antenna of Asilus cristatus is very like that of Bombylius in shape and arrange- 
ment; the organs are, however, larger and more scanty. 

The club-shaped antenna of the Butterfly approaches that of the Blow-fly very closely, 
in possessing both the small transparent dots as well as the chambered cavities. I have 
figured the antenna of Argynnis paphia (Tab. XXIX. fig. 5). Both the above structures 
- extend nearly to the base of the antenna, but more sparingly towards the lower end. The 
small organs are surrounded by various cuticular markings, and vary from 3555 to 5565 
inch (fig. 5,51). The openings into the chambers are in diameter from 5355 to 1355 inch 
(fig. 5, 6 2, 2). The chambers of the cavity vary from two to six; in the centre of their 
- base is a projection or papilla, and the floor viewed from above presents the same ap- 
pearances as in the Blow-fly. 

I may remark that the small organs can be in no way mistaken for the roots of the 
scales, their form being very dissimilar. 

I have not carried out the examination in the antennz of Moths, but have found these 
small organs in the dilated extremity of the divisions of the antenna ‘of the Drinker Moth. 

In the Hymenopterous group we find these organs very extensively spread over the 
antenna, the external form varying somewhat in aspect; that of the Hive-bee (Apis 
mellifica) being a beautiful illustration, showing as it does the structure on a larger scale 
than any I have as yet described; and as the antenna is very free from cuticular markings, 
the nerve is very distinctly seen to give off a branch to the groups, which in this insect 
only occupy one side of the antenna (Tab. XXX. fig. 6). 

Each structure consists (viewed from above) of round transparent spots, about 1787 
inch diameter, but on a side view they are seen to be depressions of the surface, the internal 
“wall being perforated, with a very thin membrane closing in the perforation, which is 


150 DR. J. B. HICKS ON A NEW STRUCTURE 


probably the external layer of the antennal wall continued over it; and this is Pee 
the condition in every variety of these structures (fig. 6, a 2). 

In fig. 7 is represented the antenna of Eucera longicornis ; the whole surface is occupied 
by a number of saucer-like depressions, which are arranged so close that they touch each 
other on all sides. In the centre of these is clearly seen the tubular perforation of the 
internal wall, closed in by the very thin membrane. The diameter of the perforation 
is about 7753 inch; the whole saucer-like depression is 4559 inch. If a fracture be made 
in the antenna, the above thin membrane will become detached and float about in the 
balsam. Beneath the perforation will be found, by very careful examination, a membra- 
nous sac (fig. 7, c). These sacs touch each other all round, the whole forming a thick 
layer on the internal surface of the wall of the antenna. The number of these structures 
in this insect is about 2000 for each of the ten joints furnished with them ; the total will 
therefore be about 20,000. 

In fig. 8 is shown the antenna of another Hymenopterous insect (Andrena fulva). 
It has the same construction as the last-mentioned, except that there are hairs between 
the saucer-like depressions; and each hair as well as the depressions being surrounded 
by a ring of very minute tubercles, the whole has a pretty appearance. i 

In the Ichneumons which I have examined, I find the appearance different, thon the 
real structure is not so. The perforation remains the same, but the thin transparent 
membrane closing it in, is elevated above the general surface, and extends lengthwise 
beyond the perforation, while it is generally not so wide. The shape of this membrane 
is not unlike that of an inverted canoe, and a keel-like ridge extending longitudinally 
through its centre adds to the similitude. The perforation beneath it is oval. I have 
endeavoured to show this at fig. 9, a, 5, c. ; 

> extreme length of the external arched membrane I have found in one specimen to 
be gig inch; the longest diameter of the subjacent perforation is 44555, the shortest 
3700 inch. 

The antenna of Vespa vulgaris and of V. crabro is arranged on the same principle, but 
the organs are more numerous and not quite so regularly em They are well 
shown in the pupa state. 

In Nomada the appearance is much the same as in Apis mellifica. i , 

But the most beautiful of all the structures I have as yet met with is that presented i in à 
the antenna of Libellula depressa (Tab. XXX. fig. 10). This antenna has six joints. The 
third, fourth and fifth have on one side the following organs. 1st, a simple, nearly round 
sac, formed in the internal membrane and projecting inwards; the external opening 
being closed in by the delicate thin membrane, as noticed in the other insects (fig. 10, å 
& 6 2). 2nd, a sac as above, but opening inwardly into another convoluted membranous 
sac, which is shown at fig. 10, 41. These are more simple towards the upper joints; and 
in some of the simpler sacs there seems to be å small cavity at the back. Whether there 
be any membrane separating the outer and inner sac, it is difficult to say ; but I faney there 
is no separation of the kind. 

The nerve may be distinctly seen to pass to these structures, and as the cuticle has 
scarcely any marking, the whole is plainly visible, and forms a very interesting object. 


IN THE ANTENNÆ OF INSECTS. | 151 


The diameter of the simple sacs is about 3357 inch, that of the largest from sopọ to 
1300 inch; the length of the longest sac 445 inch. 

In the Grasshoppers (Tetrix) (Tab. XXX. fig. 11) I find a structure very similar to 
that of the simple sacs of Libellula, and formed in the same manner ; but there is a distinct 
papilla (of which there is some trace in Libellula) in the centre of the floor of the sac 
(probably where the nerve is connected with it), and from this papilla a small chain of 
minute bodies extends to the centre of the membrane which closes the aperture. This 
connexion is well seen ‘in profile (fig. 11, e 1), and, when viewed from above, the point of 
contact with the external membrane is seen in the centre, as is indicated in fig. 11, B. 

The diameter of those represented in fig. 15, is about 5555 inch. There are about 
twenty-two joints in this antenna, the lower of which possess about twelve sacs; the 
number inereases towards the apex, till about from the fourteenth to the last there 
are 100 on each, so that we may estimate the total number on each antenna to be at 
least 1000. 

In Gryllus, Gryllotalpa, and Blatta, I have found these structures in each joint of 
the filiform antenna, generally from two to four in number, and commonly occupying 
the same position and aspect of the joint. I have drawn those of Gryllus domesticus at 
Tab. XXX. fig. 12. 

In the Coleoptera, so far as I have examined, similar organs are to be met with. 

In the attenuated antenna of the Longicorns, they are placed near the upper end of 
each joint, where they form groups of eight to ten, one group on each side. 

In Silpha clavicornis they are scattered mostly on one side of the thickened joints, the 
membrane by which the aperture is closed projecting hemispherically. 

In Melolonthå vulgaris the whole surface of the lamella is thickly covered by organs. 
which are apparently of similar structure; these are figured in Tab. XXX. fig. 12. 
` They consist of an irregularly-round cup-like depression, the centre of which rises up 
nearly to the external surface of the antenna. From this projection a tube runs inwards 
quite through the internal layer of the wall of the antenna, while the external end 
- which forms the central projection is closed by a delicate membrane. s i 
. . The diameter of the cup is about 5545, while that of the central projection is 3757 inch. 

The external layer of the cuticle is marked by hexagonal lines so disposed that each of 
the cups occupies one of the areas considerably enlarged. Tan 

The nerve entering the antenna is large, and at the base of each lamella sends into it a 
branch, which dividing into four or five smaller branches, spreads out over the whole 


internal surface. 
The number of these organs on each antenna 1s as > y Å 
each lamella is about 2 inch by 7 inch; each square of 4, inch has on each side 45 o 


these structures; the square would therefore hold 2025. As there are four of these 
areas in the entire side of the lamella, the number would be 8180. The sum for both 
sides will be 16,360, and as there are seven lamellæ on each antenna, the total number 
of these organs on the whole antenna will be 114,520. A deduction of 2000 or 3000 
may be made for irregularities, difference of size, &c. ; still, making the most liberal 
. deduction, we may say there are at least 100,000 on each antenna. 


follows. - The area of one side of 


152 DR. J. B. HICKS ON A NEW STRUCTURE 


As in some of the Diptera and Lepidoptera, so in this species, the antenna has cavities 
in which we can detect from three to ten organs similar to those on the outside. 
With regard to the physiological relations of these organs, I think we may perceive in 
them a structure analogous to that of the reputed auditory organs at the base of the 
antennules in certain Crustacea; but whether the resemblance be sufficient to allow us to 
conclude that they have a similar function, it is perhaps premature to assume. Their 
essential parts seem to be, a cavity in the antenna-wall, filled with fluid, closed in from 
the outer air by a very delicate membrane, and to the back of which a nerve certainly 
proceeds. . There seems to be some correspondence between the chain of minute bodies 
passing through the cavity in the Grasshoppers, and the otolithes in the Crustacea; but at 
present it is only in this tribe I have found such an arrangement, though subsequent 
observations may extend the number in which it exists. If these organs are connected 
with the auditory sense, the opinion, held by Carus, Straus-Dürckheim, Oken, Bur- 
meister, and Rennie and Kirby, that the antennæ are organs of hearing, will not be - 
without foundation. In considering the function of these structures, we must bear in 
mind that all the auditory organs hitherto discovered in the Invertebrata have been in 
aquatic animals, and consequently, when the elements of an auditory organ are found in 
an air-breathing animal, it will be necessary to consider the difference between the force 
with which sound is transmitted from water to water through a membrane, and through 
the same from air to water. If we assign an olfactory function to these organs, one 
difficulty presents itself, viz. that for the odorous particles to affect the nerve; they must 
reach it through a membrane and a stratum of fluid. 
The careful anatomical examination of the antennæ in all the genera of Insects would 
be a work demanding much time and many labourers, but productive of a rich harvest. 
If the method: of proceeding I have recommended be adopted, viz. the bleaching of the 
parts before they are placed in Canada balsam, I have no doubt many interesting varieties 
of these organs would be met with. The comparison of the habits of the insect with the 
structure of the antennal organs would afford a further confirmation of their nature. At 
the same time a comparison should be instituted between these organs and those I have 
before described in the base of the halteres and wings, especially with reference to their 
relation to the sense of smell. | 


I may add, that the magnifying powers required to show the structures in question 
best, are the 4th and 4th-inch objectives. - ! 


Note.—Since the reading of the above paper, I have seen (through the kindness of — 
Mr. J. O. Westwood) a copy of the rare work of Erichson, * Dissertatio de Fabrica et Usu 
Antennarum in Insectis," 1847, in which he mentions the existence of “ pores," which 
are shut by a thin membrane from the interior part of the antenna, and that their dispo- 
sition varied in different insects; but their peculiar form in Ichneumons, Wasps, &c.,also . 
the sacs, cavities, and nerves proceeding to them, he has failed to notice.—J. B. H. . 


IN THE ANTENNÆ OF INSECTS. 153 


. EXPLANATION OF THE PLATES. 


(All the figures are magnified.) 


TAR. XXIX. 


Fig. 1. Antenna of the Blowfly, Musca vomitoria :—a, antenna, showing both the smaller organs and the 
chambers; à 1, the smaller organs, having a diameter of zopp of an inch; 5 2, the apertures to 
the chambers, having a diameter of 355 to 1757 of an inch; c, aperture of chambers; c’, ditto 
as seen focused lower down; d, section of wall of antenna, showing 1, the smaller organs, and 
2, the chambers and their sacculi, the diameter of the largest being +45 of an inch. 

Fig. 2. Antenna of Helophilus pendulus :—a, antenna; 6 1, the smaller organs, having a diameter of 7757 
of an inch; c 1, do. do.; c 2, apertures to chambers, diameter +55 of an inch; c 28, do. do. 
focused lower down; c 2b, focused still lower; d, section showing the smaller organs at 1, 
and the chambers with their sacculi at 2. 

Fig. 3. Antenna of Tabanus bovinus :—a, antenna, with the third to the eighth joints furnished with the 
organs; à, enlarged view of the organs of various sizes. 

Fig. 4. a, Antenna of Tipula oleracea; b, antenna of Ctenophora ——?; c, antenna of Ctenophora bima- 
culata; d, section of organs; d, organs as seen from above, diameter ziyy of an inch. 

Fig. 5. Antenna of Argynnis paphia :—a, antenna showing the position of the organs; 41, small closed 
perforations of various sizes, the smallest 4,5, the largest 5753 of an inch; 5 2, openings to 
cavities, from 3357 to 1357 of an inch; c, last joint of the antenna, showing the openings to the 
cavities ; d, section of chambered cavities, with a projection or papilla in the centre of each, 


Tas. XXX. 


Fig. 6. a, Part of antenna of Apis mellifica; a 1, groups of organs; a 2, nerve giving off branches; b, 
view of organs, more highly magnified. 

Fig. 7. a, One joint of antenna of Eucera longicornis; b, saucer-shaped depressions, diameter from edge to 
edge 1757 of an inch; c, section of wall of antenna, showing sacs or cells beneath; c1, membrane 
closing in the openings in internal membrane, diameter 775; of an inch. 

Fig. 8. a, One joint of antenna of Andrena fulva; b 1, saucer-like depressions, with membrane in centre 
closing in the perforations; à 2, hairs; 1 and 2 are both surrounded by a ring of very minute 
tubercles ; c, section of wall of antenna. 

Fig 9. a, One joint of antenna of a species of Ichneumon; b, section of wall of antenna, showing the 
transparent membrane overarching and extending beyond the perforation of the internal wall; 
c, enlarged view of organs; membrane as seen from above, very penus, stretching. over 
like an inverted canoe; the opening beneath is the perforation of m internal membrane. 

Fig. 10. a, Antenna of Libellula depressa, showing the 3rd to the 5th joints with organs placed on one 
side of the nerve; 2, portion of wall of the 3rd joint; å 1, large sacs with plicated walls, closed 
externally by a delicate membrane ; 5 2, small simple sacs, also closed by membrane ; b 3, nerve 
proceeding to sacs; c, sacs of 5th joint. : 


- VOL. XXII. ^ 


154 DR. J. B. HICKS ON A NEW STRUCTURE IN THE ANTENNÆ OF INSECTS. 


Fig. 11. a, Antenna of Tetrix ——?, showing about 100 sacs to each joint ; 6, enlarged view of two organs 
hoa above; 4 1, central spot where the connexion with papilla takes place; c, profile view of _ 
sacs, closed from the external air by a membrane, the centre of which is —— 
papilla by means of the delicate line c 1. 


Fig. 12. Antenna of Melolontha vulgaris :—a, enlarged view of surface from above; 4, profile view ail 
: organs; c, a lamella showing the nerves. Diameter of cups about riye, of centre projections 
sy cf am inch. 


4 
i E 
å r* å i 
i k 
x 
e p? 46 ^ 
! "T^ * » N 
ae HS Ot è SE s 
w Án aic: MA 
r ta 
M t 
sekt E * å 
* 
- å 
+ w 
+ 
n 
* * 
x d. ® 
> fu. 
xa 
4 
* 
m 
^ v À 
wes e 
å LA 
+ + 
eis ^: 
E s ” ^ 
t på 4 $ 
) Gå £4 Ge y + * 3 * a 
* x s 1 
-+ » E * * å 
FE ) 
H 


` 


Trans. Tinm. Soo Vol. XXII. tab. 30. s. 153. 


(979.0. gar 
9 0 hi 


1 


Fig.12. 


6. Jarman, so. 


THE 


TRANSACTIONS 


OF 


THE LINNEAN SOCIETY 


OF 


LONDON. 


VOLUME XXII. 
PART THE THIRD. 


á 


LONDON: 


PRINTED BY TAYLOR AND FRANCIS, RED LION COURT, FLEET STREET: 


SOLD AT THE SOCIETY’S APARTMENTS, BURLINGTON HOUSE; 
AND BY LONGMAN, BROWN, GREEN, LONGMANS AND ROBERTS, PATERNOSTER-ROW. 


M.DCCC.LVIIL — 


* 


E 


CON TEATS. 


PART III.—1858. 


XII. Observations on Entozoa, with notices of several new species, including an account 
of two experiments in regard to the breeding of Teenia serrata and T. cucumerina. 


By T. SPENCER Coppoip, MD EUS Lu å win a ee op page. 166 
XIII. On the Arrangement of the Cutaneous Muscles of the Larva of Pygæra bucephala. 
By Joun Lussock, Eg, FRA, FES PAGES e NST 


XIV. On the Agamic Reproduction and Morphology of Aphis —Part I. By Tuomas 
H. Hvuxssy, F.R.S., Professor of Natural History, Government School of Mines. 
Communicated by G. Busk, FRA El rv 


XV. On the Agamic Reproduction and Morphology of Aphis.—Part II. By Tuomas 
H. Huxzey, F.R.S., Professor of Natural History, Government School of Mines. 
Communicated by G. Busk, E-R.S, FDS. . 2 . . . . . . . . . 221 


XVI. On the external Anatomy and Natural History of the Genus of Annelida named 
Palolo by the Samoans and Tonguese, and Mbalolo by the Fijians. By JON 
Denis MACDONALD, Assistant-Surgeon of H.M.S. * Herald,’ employed on Survey- 
ing Service in the South-western Pacific, under the command of Captain H. M. 
DENHAM, R.N., F.R.S. Communicated by G. Busk, F.R.S., F.L.S. . . . 287 


XVII. On the probable Metamorphosis of Pedicularia and other forms; affording pre- 
sumptive evidence that the Pelagic Gasteropoda, so called, are not adult Jorms, 
but, as it were, the Larvæ of well-known genera, and perhaps confined to species 
living in deep water. By Joux Dents MACDONALD, Assistant-Surgeon of H.M.S. 
* Herald,’ employed on Surveying Service in the South-western Pacific, under the 
command of Captain H. M. Dunnam, R.N., F.R.S. Communicated by G. Busk, 
NG NN 


XVIII. On the Anatomy of Eurybia Gaudichaudi, as bearing upon its Position amongst 
the Pteropoda. By Jomn DENIS MACDONALD, Assistant-Surgeon of H.M.S. 
* Herald,’ employed on Surveying Service in the South-western Pacific, under the 
command of Captain H. M. DENHAM, R.N., F.R.S. Communicated by G. Busk, 
ERS FRS... : | Mn ces ae D 


XIX. Notes on Phoronis hippocrepia. By F. D. DYSTER, Esq., ELA... 251 


XX. Synopsis of the Fructification of the Compound Sphæriæ of the Hookerian Herba- 
rum. By FREDERICK Currey, Zsq., M.A., PRS, ELS > + r 257 


[ 155 ] 


XII. Observations on Entozoa, with notices of several new species, including an account 


of two experiments in regard to the breeding of Tænia serrata and T. cucumerina. 
By T. SPENCER CoBBoOLD, M.D., F.L.S. 


Read December 3rd, 1857. 


IN the month of February 1856, I commenced some experiments with the view of con- 
firming the statements of continental helminthologists in regard to the development and 
habits of the cestoid Entozoa. Unforeseen circumstances prevented my carrying out these 
investigations to any great extent; nevertheless, as I have, during my past connexion 
with the Anatomical Museum of the University of Edinburgh, collected and examined 
numerous forms of Entozoa, I desire to lay before the Society the result of these casual 
searchings, at the same time offering an apology for the necessarily incomplete and frag- 
mentary character of my remarks. 

-Excluding Man, the number of vertebrate species examined by me, with especial refer- 
ence to the presence or absence of Entozoa, amounts to seventy, in forty-five of which 
worms were detected. This proportion, however, must not be regarded as a fair criterion 
of the frequency with which these higher orders of animals are infested; for although a 
great number of individuals—from twenty to forty—of the common species were examined, 
only one or sometimes two of the species more difficult to procure came under notice. 
In a statistical point of view, certain particulars in regard to the age of the animal, the 
time that had elapsed after death before it was examined, and other conditions materially 
lessening the chances of the detection of Entozoa, would have to be taken into considera- 
tion. The forty-five species found to contain worms may be arranged as follows :— 


Common Stickleback, Gasterosteus aculeatus. — Toad, Bufo vulgaris. 
Fifteen-spined Stickleback, G. spinachia. Frog, Rana temporaria. 

Minnow, Leuciscus phoæinus. Lesser Newt, Lissotriton punctatus. 
Rockling, Motella quinquecirrata. Serpent, Bothrops ? 


Viviparous Blenny, Zoarcus viviparus. 
Smooth Blenny, Blennius pholis. 


Lucky Proach, Cottus bubalis. Kite, Falco m 
Haddock, Morrhua eglefinus. Kestrel, F. tinnunculus. 
Coal-fish, Merlangus carbonarius. Peregrine, F. peregrinus. — 
Cod, Morrhua vulgaris. Sparrow-hawk, Accipiter nisus. 
Ling, Lota molva. Honey m ce e 
Holibut, Hippoglossus vulgaris. beas - = 0 ut 
Skate, Raia batis. Reds å 0 — 
Pipe-fish, Syngnathus acus. Curlew, Numenius i 

Grey Gull, Larus glaucus. 


Angler, Lophius piscatorius. 4 i 
: Kittiwake, L. tridactylus. 
. Orthagoriscus mola. Å 
Pun-fah, OM it Guillemot, Uria troile. 


Sturgeon, Acipenser sturio. Y 
VOL, XXII. j 


J 


156 : DR. T. S. COBBOLD ON ENTOZOA. 


Auk, Alca torda. Rabbit, Lepus cuniculus. 
Red-throated Diver, Colymbus septentrionalis. Dog, Canis familiaris. 
Golden Pheasant, PAasianus pictus. Lion, Felis leo. 
Capercailzie, Tetrao urogallus. Ox, Bos taurus. 

Sheep, Ovis aries. 
Mouse, Mus musculus. Giraffe, Camelopardalis giraffa. 
Cat, Felis catus. Porpoise, Delphinus phocæna. 


In this list, notwithstanding the unfavourable circumstances above alluded to, I have 
found no less than sixty forms of Entozoa infesting forty-five vertebrate species; and there 
cannot be the slightest doubt, as subsequent observations will show, that this figure affords 
but a very low estimate of the liability of invasion to which these creatures are exposed. 
It is also worthy of remark, that out of those species in which Entozoa were not found, 
only one belonged to the fish tribe; whereas a very careful examination failed to detect 
any worms in several kinds of birds and mammals. Among those species not harbouring 
Entozoa, but in which their presence might have been anticipated, were the following :— 
Wild Duck, Swan, Woodpecker, Starling, Pheasant, Tawny Owl, large Water Newt, Water 
Rat, Squirrel, Stoat, Otter, Badger, Fox, and common Brown Bear. 

Six or eight species of Entozoa have been described as inhabiting Gasterosteus aculeatus, 
but I have only met with three; namely the Tenia filicollis (Pl. X XXI. fig. 1); an imperfect 
specimen of cestoid worm corresponding in some respects with the Schistocephalus dimorphus 
of Creplin, and a small trematode presumed to be the Monostoma caryophyllinum in a very 
young state (figs. 2,3). The head of T. filicollis, according to the descriptions of Dujardin 
and others, is simple, globular, unarmed with the usual proboscidiform retractile tubercle, 
and unprovided with lateral suckers. In those which have come under my notice, a very 
distinct proboscis was to be seen while the specimens remained fresh, but there was no 
indication of hooks; these may have fallen off. Four well-developed suckers surrounded 
the head, and the water-vascular canals were unusually conspicuous throughout the 
entire segmental series. In an adult individual, these tubes, four in number, terminated 
superiorly so close to the cup-shaped bothria, that it was difficult to divest oneself of the 
notion that they communicated with the latter. 

After a fruitless search for Entozoa in many specimens of Gasterosteus spinachia, one 
example at last yielded an extremely minute trematode (figs. 4, 5), enclosed in a cyst of 
cellular tissue and attached to the subperitoneal surface of one of the ovaries. While 
under microscopic examination it was accidentally swept off the slide, but an outline of - 
its structure has been preserved. A feature in this specimen consists in the presence of 
an unusually large ventral appendage, the free extremity of which is bifid and each 
division apparently perforated. The hermaphroditic signification of this organ is evident; 
but the circumstance of the ends of the male and female reproductive tubes being thus 
greatly extended, may in some measure be regarded as accidental. Traces of a bifurcate 
stomach were plainly discernible, the canals being slightly sinuous. As I am unable to 
refer this worm to any known species, I recognize it provisionally under the signification 
of Monostoma dubium. 


In Motella quinquecirrata—the Gadus mustela of Linnæus—Rudolphi mentions the : 


DR. T. S. COBBOLD ON ENTOZOA.. 157 


occurrence of a cestoid entozoon. This species of Bothriocephalus I have not seen; but 
in a specimen of the above-named fish its place was supplied, as it were, by two other 
worms—the Distoma fulvum (figs. 6, 7, 8), hitherto observed in the Ling and Gadus medi- 
terrameus, and the common Filaria piscium, which is especially abundant in the Cod-tribe. 
So far as I am aware, the Distoma fulvum has not yet been figured under that name, but 
after a careful investigation I am satisfied that it is identical with the D. simplex of 
Rudolphi and Fasciola Bramæ of Müller*. Dujardin, after giving the specific characters 
of D. simplex, appends an account of two unnamed trematodes which he found in a 
specimen of Gadus quinquecirratus from the coast of Brittany; these also appear to me 
to be one and the same species, and I think he would have been justified in considering 
them as merely altered forms of D. simplex or D. fulvum. Dujardin moreover makes 
D. simplex synonymous with the Fasciola ZEglefini of Müller and the D. Wachnie, first 
described by Tilesiust. After consulting the figures and descriptions of these authors, I 
consider D. Wachnie to be the original representative both of D. simplex and D. fulvum, 
but I believe F. Zglefini may be properly regarded as a distinct species. The living speci- 
mens of D. fulvum examined by me alternately elongated and contracted themselves very 
vigorously, thus varying the length of the body between the twelfth and fortieth part 
of an inch. In this way the neck sometimes became extremely attenuated, contrasting 
strongly with the large and prominent ventral sucker. Under a half-inch lens, the largest 
specimen, as now preserved, exhibits the intromittent organ everted and the convoluted 
uterine tube crowded with ova. 

The only entozoon I have observed in the viviparous Blenny is a single specimen of 
Ascaris aucta, R. It was coiled in a cyst beneath the peritoneal surface of the liver, the 
gland being otherwise diseased. In an example of Blennius pholis, four specimens of 
Echinorhynchus tereticollis, R., were obtained from the intestine, in different stages of 
development. The largest exhibited a bulging of the cuticle near the middle of the body, 
probably the result of injury. 

Cottus bubalis.—Several were examined and only one found infested—that by a single 
tape-worm, which occupied the intestine immediately below the pancreatic cæca. This 
entozoon, long known to inhabit Cottus scorpius, and by Miller termed Tenia Scorpii, 
has been more fully described by Leuckart, Eschricht, and Van Beneden, under the better 
title of Bothriocephalus punctatus. The extreme transparency of this worm, when alive, 
produced, during its active movements, very puzzling appearances, and had I not m- 
viously entertained the persuasion that all tape-worms were destitute of a true digestive 
tube and buccal cavity, I should have felt entitled to affirm, that this species at least was 
provided with a continuous alimentary canal. The anterior cephalie segment, while 
- extended, greatly exceeded in length each of the ten or twelve succeeding segments, and 
when contracted appeared rather broader. Müller’s figure gives an idea of considerable 
disparity in this respect, but in other particulars the specific resemblances ‘were at once 
recognized. Toward the lower part of the so-called neck, the joints exhibited at the lateral 
margins indications of division, which became gradually more defined towards the tail. 

* Zoologia Danica, tom. i. p. 33. pl. 30. fig. 6. 


+ Mém. de l’Acad. des Sciences de St. Pétersb. tom. ii. p. 363. tab. 19. figs. 8-10. ^ 
: > 


158 DR. T. S. COBBOLD ON ENTOZOA. 


In the Haddock and others of the Cod-family we meet with many kinds of Entozoa 
and the worms found in one species seem liable to occur in all members of the group. 
Most of the Trematoda infesting fishes are exceedingly minute, and it is absolutely neces- 
sary for their identification that specific descriptions should be accompanied with accurate 
figures. I am led to this remark from the difficulty of making out the identity of å 
Distoma found associated with Echinorhynchus acus, R., and Bothriocephalus rugosus, R,, 
in the intestine of a Haddock. The trematode in question agrees in some respects with 
D. scabrum and D. appendiculatum, especially the latter, but differs in a few essentials ; 
moreover, D. appendiculatum is not mentioned as infesting the Gadidæ. Provisionally 
therefore I propose to regard it as a distinct species, and shall briefly describe it under 
the title of D. rachion (paxiaioc), as follows (figs. 9, 10) :—Body of a faint pink colour; 
length 3 lines; flat, covered with minute spines; obtuse at either extremity ; posterior 
half of the body rather broader than the front; oral sucker large, not quite terminal; 
ventral sucker comparatively small, near the centre of the body; cesophagus narrow, 
twisted, bifurcating immediately above the ventral sucker ; gastric canals very capacious; 
genital pore large and conspicuous; internal seminal vesicle, uterine tube, ovary, testes, 
and contractile vesicle particularly distinct. Some other details in regard to the struc- 
ture of this trematode will be found indicated in the explanation of the accompanying 
figures. In regard to Bothriocephalus rugosus I have only to observe, that in conse- 
quence of the head being found in all instances firmly rooted within one of the pancreatic 
cæca, it was impossible to ascertain the precise form of the cephalic segment. Dujardin 
remarks the same peculiarity in specimens obtained by him from Gadus pollachius and 
G. merlucius. The single Echinorhynchus, obtained from the same Haddock, exhibited 
when alive an organ lying near the proboscideal sheath—the so-called lemniscus, con- 
cerning the use of which we are still ignorant. In common with others, I have observed, 
in another acanthocephalous species, two of these organs, one on each side of the sheath. 
They have no connexion with the retractor muscles; but the latter, in the example under - 
consideration, could not be recognized through the semi-opake integument. A Haddock _ 
examined by me on the 8th of May, 1854, contained, in addition to two specimens of the 
above-mentioned Bothriocephalus, several small cysts enclosing Filariæ, and a larger 
solitary cyst occupied by a Tetrarhynchus (figs. 11 to 19 inclusive). These require a 
passing notice. The minute sacs were imbedded in the submucous tissue of the aliment- 
ary canal, and scarcely exceeded half a line in breadth. The single large cyst, situated 
beneath the peritoneum outside the intestine, measured about a quarter of an inch long, 
and one-eighth of an inch broad. It was oval, semitransparent throughout, and on divi- 
sion of the peritoneal investment, easily removed. When detached, the fibro-cellular in- 
vesting capsule or ectocyst offered a pyriform outline, and from its narrow extremity there - 
proceeded a stalk-like prolongation. The ectocyst on being torn open disclosed the tetra- 
rhynchoid Scolez, the head and body of which were invaginated, but became readily evolved 
by gentle pressure applied to the bulky caudal vesicle or endocyst. Amplified forty dia- 
meters, the internal structures were well seen, the proboscidiform tentacula and * sclerous 
particles” being especially conspicuous. I have here, with due consideration, applied the 
term “sclerous” to the subcutaneous corpuscles so abundant in the caudal vesicle, in 


DR. T. S. COBBOLD ON ENTOZOA. 159 © 


order to signify my adhesion to the view of Siebold and others as to their earthy nature. 
Helminthologists express various opinions in regard to their structure and function. All the 
older writers speak of them as **ova ;" whilst other, more recent observers, following Esch- 
richt, ascribe to them a nutritive function. The late Prof. William Smith of Cork apparently 
adhered to this latter view, for in his interesting paper on * Measled Pork," he designates 
these corpuscles “ assimilating cellules*," whilst the distinguished Belgian entozoologist, 
Prof. Van Beneden, attributes to them a still higher organization, calling them cutaneous 
glandst. The ordinary cellules of the parenchyma were found to be comparatively small, 
requiring the aid of a quarter-inch lens for their satisfactory definition. The epidermic cells 
showed in profile very fine markings, indicating a columnar epithelium; and immediately 
beneath the corium there were outlines of anastomosing tubes, which appeared to represent 
a water-vascular system. The armature of the retractile proboscides is rather complicated ; 
each proboscis being made up of numerous segments, and every joint supporting qua- 
druple rows of four hooks, sixteen in all; the superior set being disproportionately large. 
Having thus cursorily alluded to five different forms of Entozoa infesting the Haddock, 
I pass to the consideration of worms found in the Coal-fish, common Cod, and Ling. 
Merlangus carbonarius. Morrhua vulgaris —Numerous examples of Ascaris clavata 
(R.) occupied the intestinal canal of one individual. No other species of Entozoa were 
detected in the limited number of specimens examined. In a Cod examined on the 15th 
of March, 1855, two specimens of Bothriocephalus rugosus had severally attained a length 
of nearly fifteen inches, and their anterior segments, for an inch or more downwards, were 
so firmly impacted within the pancreatic cæca, that it was found impossible to dislodge 
them without injuring the filamentary head and neck. As if to make the anchorage 
doubly secure, the cartilaginoid thickening of the invaded pancreatic caecum had dege- 
nerated into a caleareous and contracted cylinder, twisted upon itself in various ways. 
Lota molva.—On the 31st of March, 1855, two kinds of nematoid worms, differing in 
relative size, were obtained from the intestinal tube. The smaller species, about half 
an inch in length, may be referred to an entozoon described by authors under a variety 
of names. It is the Cucullanus marinus of Zeder and Müller, the C. foveolatus of 
Rudolphi, and the Dachnitis globosa of Dujardin (figs. 20-23). The vaginal orifice of 
the female is situated near the middle of the body, being protected by two prominent folds 
of integument. The spicules of the male, two in number, are sabre-shaped, and there is 
a spacious sucker-like depression in front of the anal opening. The larger round-worms 


* Microscopical Journal, vol. v., description of pl. 2. 
+ Recherches sur les Vers cestoides, p. 23 :—“ ils consistent 


un mucus destiné å lubrifier la surface du corps.” agi des i 
Since the above was written, a paper by M. E. Claparède has appeared in Siebold and supine MN f. 
Wissensch. Zoologie, vol. ix. p. 99, on the subject of the calcareous corpuseles of the Trematoda, in which it 1s shown 


that these bodies are lodged in the dilated extremities of branches of the so-termed » er il wi 
system of vessels. This interesting fact seems to have been demonstrated to the satisfaction of Joh. er, Virchow, 


Lachmann, and others, and to have been previously known to, though not published by, ppn — en. 
respect to it, it may be remarked that so far back as 1830, Laurer pointed out that jo i sig 
system of vessels terminated in minute vesicular dilatations in Amphistomum conicum, ough a s 


could not be detected by Nordmann in Diplostomum volvens. 


dans de courts cæcums légèrement ramifiés, et sécrétent 


160 DR. T. S. COBBOLD ON ENTOZOA. 


above mentioned cannot, so far as I know, be satisfactorily referred to any species of 
Ascaris or other nematode at present described; and although unwilling to multiply 
species in a group of Entozoa already extended numerically far beyond the proper limits, 
there is in the present instance no alternative. Ascaris acanthocaudata (mihi) may be 
identified as follows :—Body 10 to 15 lines long, half a line broad; diameter slightly 
increased toward the head, suddenly narrowed at the tail; mouth trilobular; caudal 
extremity armed with several minute spines (figs. 24-26). A preserved specimen in my 
collection exhibits the transverse and longitudinal muscular layers with unusual clearness; 
but the most interesting structure, which can be seen with a half-inch lens, and its minute 
tissues with a quarter-inch objective, consists of a broad double band of partially disin- 
tegrated cellules, extending from head to tail on the under side of the body. The centre 
of this band is occupied by a well-defined canal apparently containing a delicate white 
thread. It does not give off any branches, but terminates superiorly in an abrupt manner 
immediately below the trilobular mouth. From this point there is a faint appearance 
of radiating lines towards the margin of the lobes, but I cannot positively say that they 
are nervous filaments. Notwithstanding this doubt, I am confident that the long-enter- 
tained opinions of Otto and Siebold, in regard to the existence of a nervous cord in the 
Nematoidea, are correct. 

Hippoglossus vulgaris.—A. specimen dissected on the 4th of May, 1854, yielded two 
forms of thread-worm—Filaria piscium and Ascaris collaris*. Examples of the former 
were coiled within the mesenteric folds, while the latter were chiefly enclosed in cysts, in 
and upon the mucous membrane of the intestine. A fully-developed Ascaris collaris . 
(Pl. XXXII. fig. 27) presented the following characters—here given on account of certain 
difficulties in the way of identification :—Body an inch in length, comparatively thick; 
head obtuse, trilobular valves of the mouth very small; cesophagus narrow, communicating 
with a broad intestine, bounded on either side by two slender cæcal appendages of un- 
equal length and vesicular at the free endst; caudal extremity rounded, rather narrower 

than the head; anal orifice in front, not far distant. In addition to these nematodes, 
the intestine contained numerous specimens of Scolex polymorphus, and there was also a 
minute trematode enclosed in a capsule beneath the peritoneal membrane. The anatomy 
of various Scolex-forms has been beautifully illustrated by Prof. Van Beneden and 
Wagener? ; nevertheless I remark upon one or two particulars in passing. In this Scolex 
(figs. 28, 29) the simple structure of the sclerous particles formerly alluded to is most 
satisfactorily seen, and it is difficult to understand why such discrepancy of opinion should 
prevail in regard to their nature. Siebold long ago recognized their dermo-skeletal cha- 
racter. The four presumed water-vascular canals take their origin by two single trunks, 
one on each side of the lower part of the proboscidiform sucker; there is no appearance 

of intercommunication, however, between them and the cavity of the cup. The ency: 
trematode corresponds in many respects with the Fasciola Platesse of Miller, answering 
more closely, however, to the Distoma atomon found by Rudolphi in Pleuronectes flesus — 
* According to Miescher, Steenstrup, Wagener, &c., the Filariæ are only young Nematoidea. 


T In Cheiracanthus robustus, Diesing remarks the presence of four similar lemnisciform bodies. 
1 Die Entwickelung der Cestoden, &c. 1854. 


DR. T. S. COBBOLD ON ENTOZOA. 161 


(figs. 30-32). It is not unlikely that these are one and the same species, seeing that the 
principal difference, according to Dujardin, consists in the colour of the body and in the 
relative size of the ventral sucker. A living Distoma varies much in form during con- 
traction, and the same worm will present appearances when preserved very unlike those 
seen when it was fresh. I have found this to be the case especially in the entozoon 
under consideration, the ventral sucker presenting, during life, a most unusual breadth. 
The oral sucker was comparatively small; the simple gastric cæca, uterine tubes and con- 
tractile vesicle being severally large and conspicuous. 

Raia batis. Syngnathus acus.—Of Entozoa infesting the former, I have only to notice 
the extreme abundance of Bothriocephalus coronatus (R.) occupying the chambers of the 
spiral intestine. Under an amplification of 400 diameters, the advanced ova exhibit a 
well-developed Scolex, provided with rudimentary hooklets and sclerous particles. In 
the common Pipe-fish I have found several specimens of Filaria piscium. 

Lophius piscatorius.—The voracious habits of the Angler guarantee the presence of a 
variety of worms, and, with the exception of Orthagoriscus, no fish is perhaps more 
copiously infested. A specimen, dissected May 12, 1854, yielded three species of Entozoa, 
namely Ascaris rigida, Scolex polymorphus, and Distoma gracilescens. Most of the 
nematode individuals were imbedded in folds of the peritoneum and mesentery, the other 
kinds occupying the intestinal canal. Published deseriptions of Distoma gracilescens 
(figs. 33-37 inclusive) being few and imperfect, I offer the following notice of its more 
obvious characters :—Body of a pale brown colour, semitransparent, one-sixth of an inch 
long, flat, linear, beset all over with minute tubercular spines, those about the head being 
more cogently developed ; anterior half somewhat narrower than the posterior; oral sucker 
oval, not quite terminal; ventral sucker circular; sheath of the penis large, and placed 
immediately below the ventral cup; uterine tube broad and tortuous, occupying the 
inferior half of the body; vitelline organs consisting of two elongated, botryoidal masses, 
commencing a little below the oral sucker and passing down on either side of the neck; 
testes bulky, transparent, placed toward the lower and back part; in front of these, two 
smaller vesicular bodies, corresponding to the ovary and seminal vesicle; contractile 
vesicle very large, with thick muscular walls. à 

Orthagoriscus mola.—1 have dissected, in its entirety, but one specimen of this remark- . 
able fish. It was a very young individual, and was captured off Anstruther, on the coast 


of Fife, September 6, 1856*. There were no Entozoa in the intestinal canal, but the 
liver and retractor muscles of the anal fin contained several examples of the Gymnorhyn- 


chus reptans of Rudolphi (figs. 38-46 inclusive). Professor Goodsir has given an au 
description of this cestode in the Edinburgh New Philosophical J ournal for 1841, under 


the title of G. horridus, regarding it as a new species. If I may be allowed to differ, I 
itted to notice the two lowermost 


do not think the circumstance of Bremser's having om! 2 me 
bservance of the jointed condition 


rows of exaggerated hooks on the proboscis, or his non-o : 
i . reptans, 
of the body, as sufficient evidence that these cha rs were not present in his G. rep: 
x i , 26 inches ; 
* Its dimensions were as follows :— Length from hend to tail, 18 inches ; between tips TS im^ WC 
greatest depth of the body, 12 inches ; length of pectoral fins, 2 inches and a ga 0 pervure, 
Two or three other individuals were taken a few weeks previously in the Firth of Forth. 


162 DR. T. $. COBBOLD ON ENTOZOA. 


It requires considerable enlargement and a good lens to render the comparatively large 

size of the hooks obvious, and the articulations of the body are but very faintly indicated 

toward the anterior part of the animal. Notwithstanding the fish in question had been 
dead several days and cast aside as refuse by the salesman of whom it was purchased, the 
Entozoa coiled within the muscles were alive, and on being removed to a tumbler of sea- 
water, they continued to live until the third day following, when they were placed in 
spirit. During the active movement of contraction and elongation, the proboseidiform 
tentacula were freely protruded and withdrawn. It was difficult to ascertain the exact 
number of hooklets upon these processes. I think every circular row carries sixteen, and 
there are about one hundred of these rings of hooklets on each proboscis; if this estimate 
be correct, the total number of hooklets is 6400. The form of the proboscis is clavate, 
and its free extremity more or less rounded; in relative size the hooklets are tolerably 
uniform, those towards the tip being rather larger; the two lowermost rows of the series 
are very highly developed. 

Acipenser sturio.—On the 25th of April, 1855, I obtained a great many examples of 
Dachnitis sphærocephala of Dujardin (Pl. XXXIII. fig. 51) and Distoma hispidum (Abild- 
gaard) from the spiral intestine. Contrary to the statements of Creplin, I find the hooks 
surrounding the head of the latter remarkably conspicuous and disposed in two rows, 
fourteen in each. I think it would be preferable to place this species in the subgenus 
Echinostoma. (Pl. XXXII. figs. 47, 48; Pl. XXXIII. figs. 49, 50.) Dujardin retained 

it among the true Distomata with hesitation. The integumentary spines in front are 
prominent, and directed outwards with a slight inclination backwards; those below the . 
ventral sucker become gradually smaller, and finally degenerate into mere tubercles at 
the posterior extremity. 

Rana temporaria.—From different individuals I have obtained Filaria rubella(?), Ascaris 
nigro-venosa, A. acuminata, Oxyuris ornata, Distoma cylindraceum, D. clavigerum (figs. 
52, 53), and Polystoma integerrimum, the latter being fully half an inch in length*. 

Bufo vulgaris.—With reference to parasites, I have only examined one or two examples 
of the common Toad, and in the intestine of an individual dissected on the 20th of 
April, 1855, found a single specimen of Ascaris acuminata associated with numerous 
small flukes. These trematodes are fully as large as the Distoma cygnoides, frequently 
infesting the urinary bladder of the Frog; nevertheless they may probably be referred to 
a smaller species—the D. clavigerum of Rudolphi, their length varying from one to two 
lines and a half. The vitelline ducts, the uterine canal with its crowded ova, and the long 
intromittent organ, are readily distinguishable in preserved specimens. 

Lissotriton punctatus.—I here wish to call attention to a striking illustration of the 
predilection for a particular species which certain Entozoa exhibit in selecting their habi- 
tation. A careful dissection of some thirty or forty Water-newts—consisting of our two 
most common species, the large black and the lesser speckled Salamanders in nearly 
equal proportions—has shown, that while in almost every instance worms were present . 


* The anatomy and development of these and other forms of Frog’s flukes have recently been illustrated by — 
Dr. H. Å. Pagenstecher in his attractive monograph, entitled * Trematodenlarven und Trematoden,” Heidelberg, 1857- 
Also in Dr. Wagener's “ Entwickelungsgeschichte von Distoma cygnoides.” 


DR. T. S. COBBOLD ON ENTOZOA. 163 


in Lissotriton punctatus, not a single entozoon of any kind could be obtained from 
Triton cristatus; a fact rendered more significant inasmuch as the specimens were all 
obtained from the same locality, namely from ponds at the summit of the Braid Hills 
near Edinburgh. In the stomach of one of the large species there was a small trematode- 
like body, but on microscopie examination it proved to be a young horse-leech. From 
specimens of the lesser Newt, procured at intervals during the months òf April, May and 
June, 1855, there were obtained in the first place numerous examples of Ascaris acumi- 
nata, the presence of which was by no means invariable; secondly, an abundance of 
Echinorhynchus anthuris; thirdly, multitudes of Opalina intestinalis, a ciliated infusorial 
animalcule believed by Agassiz to be in reality only a larval planarian; and fourthly, a 


' considerable number of Trichodine. The first three groups were found in the stomach 


and duodenal portion of the intestinal canal; the fourth occupying the cloaca and ducts 
in its immediate neighbourhood. The only remark I have to offer respecting Ascaris 
acuminata has reference to its size, which is variously estimated by different authors ; 
my specimens average about 8 lines in length, which accords with Mr. Bellingham’s 
statement in his Catalogue of Irish Entozoa. This leads me further to observe, concern- 
ing the definition of species, that much unnecessary stress is laid upon the relative size of 
individuals, bulk alone, it would appear, being frequently regarded as a criterion of spe- 
cific distinction. In very many instances at any rate it cannot be denied that this is the 
case. I believe the error is so prevalent in the older lists, that for every new species 
now added, two old ones might with propriety be expunged. In the present communica- 
tion, it is not my intention to show how the truth of this assertion may be sustained by 
an appeal to facts; nevertheless, if it were desired, sufficient illustration might be afforded 
from the materials in hand. The second group, represented by Echinorhynchus anthuris 
(figs. 54-62 inclusive), deserves attention, forming as that species does, on account of its small 
size and extreme transparency, an excellent subject for microscopic investigation; but a 
minute description is not needed, in consequence of the admirable manner in which it was 
originally described by its discoverer, M. Dujardin. General details being given in the de- 
scription of the figures, attention in this place is only invited to a brief notice of the peculiar 
ova. These characteristic bodies in their early state are perfectly round, appearing simply 
as nucleated cellules in the interior of the so-called ovaries, which are also spherical, and 
float loose in the cavity of the body; the nucleus may be taken to represent the germinal 
vesicle, but it cannot be recognized in the fully developed ovum. In this latter condition 
each ovum encloses a perfectly transparent cell, lobed at both ends after the fashion 
of Trichina, and an irregular but very distinctly granular yelk-like mass; these are 
placed side by side; and from the granular body proceeds at either end a coiled 
thread or chalaza advancing to the extremity of the external envelope and vegeta 
blended with it; the wall of the transparent cell exhibits in profile very delicate E 
dulations. In regard to the infusorial Trichodinæ, which may yet m out to be cedi 
forms of some higher group, I have only to observe that some were in the encysted con- 
dition. - 
 Bothrops.—For an opportunity of dissecting å species of this genus Iam — to 
Mr. Edwards, Demonstrator of Anatomy in the University of Edinburgh. € - 


VOL. XXII. 


164 DR. T. S. COBBOLD ON ENTOZOA. 


amples of Pentastoma proboscideum occupied the mesenteric folds, but with reference to - 
their structure I am unable to add any new or otherwise interesting facts. : 

Passing to the consideration of Entozoa in birds and mammals, my remarks under the : 
former head will be very short, as they are designed to indicate little more than a record 
of worms found in partieular species. ; 

Falco milvus.—In the duodenum of a Kite opened on the 16th April, 1855, there were 
present five or six specimens of Ascaris depressa, several examples of Trichosoma Falco- 
mm, and multitudes of Hemistomum spathula (Diesing). The latter—better known as 
the Amphistoma macrocephalum of Rudolphi—presented a bright grass-green colour, 
owing to the quantity of bile in the intestine. Outside the gut there was a minute botry- 
oidal fatty mass, consisting of four unequal lobules united together and attached by two 
filamentary stalks; each of these lobes contained an encysted nematode. 

Falco tinnunculus; F. peregrinus; Accipiter misus; Pernis apivorus ; Strix otus.— 
Specimens of Ascaris depressa were obtained from a Kestrel on the 21st of January, 1856, 
and being very numerous they completely choked that part of the intestine in which 
they were lodged; a solitary individual was also procured from the duodenum of a Honey 
Buzzard on the 30th of May of the previous year. From the cellular aponeurosis at the 
back of the abdominal cavity of a Peregrine, I also obtained in April of the same year 
a single specimen of Filaria attenuata, measuring nearly 104 inches; and from the 
stomach of a Sparrow Hawk dissected in J anuary 1856, an example of Spiroptera lep- 
toptera. This last entozoon I have also found associated with Hemistomum spathula in 
the small intestine of the Long-eared Owl in the month of J anuary. 

Totanus calidris; Numenius arcuata.—T have taken Tenia variabilis from the small 
intestine of the former of these allied species in J anuary, and also 7. spherophora (figs. 
63-67 inclusive) from the Curlew at the same period in great abundance, the latter 
entozoon being situated midway between the gizzard and cloaca. As this cestode is only — 
imperfeetly known, some additional facts in regard to it may prove acceptable. The 
head is correctly described by Diesing as obcordate, but no mention is made of the arma- 
ture of hooks surrounding the rostellum; this is not to be wondered at, considering the — 
facility with which they drop off after death, or during life, on even the most gentle 
handling. One cannot judge how many it carries from the number found in the prosco- — 
leces, where there appear to be six hooks arranged as usual in three pairs on the proboscis; — Å 
and in no case have I seen the adult cestode with its full complement. The anterior seg- 
mente are extremely narrow, but well defined immediately below the head, the sucker- 
bearing and proboseideal divisions appearing to represent very distinctly two rings of — 
the segmental series, or in other words, the first two modified individuals of the colony, 
if we can suppose with Van Beneden each proglottis to represent an independent animal. 
The middle and succeeding segments become gradually broader and deeper toward the | 
caudal extremity, the lateral margins showing a bilobular outline; the intromittent — 
organs are placed consecutively on one side only, their bulk being comparatively large; — 
and the external wall of the sheaths is closely beset with minute spines directed back- 
wards when the organs are protruded. The segments near the tail seemed ready to 


burst from the volume of the contained ova, most of which latter, when withdrawn and 


DR. T. S. COBBOLD ON ENTOZOA. 165 


examined separately, displayed internally that advanced condition of the embryo termed 
proscolex by Van Beneden. 

Larus glaucus ; L. tridactylus; Uria troile; Alca torda; Colymbus septentrionalis.— 
In the months of December and January, 1854-55, I obtained from the common Grey Gull 
abundance of Tetrabothrium cylindraceum (the Bothriocephalus macrocephalus of Dujardin) 
and Echinostomum spinulosum. The latter (figs. 68-72) is usually described as a Distoma, 
but ought certainly to be generically separated. I find the disc surrounding the head 
capable of being elevated so as to form a kind of hood; and the hooks being connected 
together by an extension of the integument, the coronet resembles the fin of a fish. The 
vitelline organs have à peculiar zigzag conformation, the cæca alternating in parallel 
rows. In the small intestine of the Kittiwake I have also constantly found Tetrabothrium- 
cylindraceum, and from the same situation in the Red-throated Diver quantities of 7. 
macrocephalum, which Diesing—correctly, I think—regards as distinct from the former. 
From the œsophagus and proventriculus of the Auk and common Guillemot I have pro- 
cured, during spring, numerous examples of Ascaris spiculigera; one dissection in the 
case of Uria troile exposing two of these nematodes lodged in the auricle of the heart. 
The intestines of another specimen of this bird contained several individuals of a cestode, 
which has been vaguely indicated by Abildgaard under the title of Tænia Colymbi Troiles ; 
I have not been able to satisfy myself, however, that this species is distinct from Tetra- 
bothrium macrocephalum. 

Phasianus pictus; Tetrao urogallus.—Both ceca of a Gold Pheasant, dissected on the 
7th of January, 1856, were found enlarged to three or four times their natural width, owing 
to the presence of a multitude of dark-coloured tubercles, about the size of peas, the inte- 
rior of each of which contained a coiled nematode, the Ascaris vesicularis of Frölich. 
From the intestine of a Capercailzie, examined in the spring of the previous year, were 
procured numerous specimens of Trichosoma longicolle ; and also from the subcutaneous 
areolar tissue overlying the great pectoral muscle, a solitary entozoon resembling Ligula 
reptans. 

Mus musculus ; Felis catus.—Fresh experiments are not required to determine the now 
well-established fact in regard to the Cysticercus fasciolaris of the Mouse being the imper- 
fectly developed Tænia crassicollis of the Cat; indeed, long before their actual identity 
was demonstrated, the frequent occurrence of a cestoid condition of the Scolex within the 
liver suggested an hypothesis which has since proved correct. In my collection there is a 
specimen of this helminth in the tænioid condition taken from the liver of a White Mouse, 
which is nearly as long as that figured by Bremser. Such examples are by no means 


uncommon. In addition to Ascaris mystax and Dochmius tubæformis, the er 
ave 


worm I have seen in the domestic Cat is the Tænia elliptica, but, like Dujardin, 


been unable to obtain its head. ve 
Lepus cuniculus; Canis familiaris.—The experimental researches of Von Siebold, 


Küchenmeister, Leuckart, and other continental helminthologists, have sufficiently esta- 


blished a mutual relation between Cysticereus pisiformis and Tenia serrata, hereby 
law of alternate generation. 


affording an additional instance of the truth of Steenstrup’s 


I am not aware if any entozoologist in this country has attempted to repeat 23 experi- 


166 DR. T. S. COBBOLD ON ENTOZOA. 


ments, and I therefore regret that cireumstances have prevented my carrying out more 
fully an intention of verifying some of those particulars which have recently attracted our 
notice. Two of a few experiments, however, having been completed and attended with 
partially satisfactory results, I take this opportunity of placing them on record :— 

On the 5th of February, 1856, I administered to a Dog three minute Cysticerci, obtained 
from the fresh livers of two wild Rabbits. These eysts were perfectly round and measured 
only the twelfth of an inch in diameter, and were evidently not to be referred to the 

common C. pisiformis. A fortnight after (19th) six similar cysts were obtained from 
the liver of another wild Rabbit and given to the same Dog. Ten days subsequently the 
canine animal was destroyed by chloroform, when, on laying open the small intestines, six 
specimens of Tenia cucumerina were detected. Three of these individuals were severally 
about 12 inches in length, the others measuring only 3 inches. Thus far, therefore, 
but for the circumstance of three of the nine cysts administered being undeveloped or lost, 
no doubt whatever could be entertained as to the complete success of this experiment. 
As it is, we bave the interesting result of three nearly completely developed tape-worms, 
the growth and number of which exactly correspond with the circumstances that ap- 
parently led to their presence in the viscera of the Dog; and in addition we have three 
other individuals, only one-fourth of the bulk of the former, which may legitimately be 
regarded as the partially developed representatives of three of the six Cysticerci that were 
subsequently administered. It would appear, in consequence of the C. pisiformis being 
frequently present in the liver of the Rabbit, as well as in the mesentery, that our minute 
Cysticercus has not hitherto been recognized as distinct; at least, I find no notice of it in 
foreign works, and shall therefore in future speak of it as C. cueumerinus. In order to im- 
part additional value to the above experiment, a young tame Rabbit was next procured, 
and the three larger Tænias (whose caudal proglottides contained multitudes of imper- 
fectly formed ova) were given to it, the living worms being greedily devoured with portions 
of a cabbage-leaf. Eleven days after, the Rabbit was destroyed, and the liver found to 
contain numerous minute cysts corresponding to those of C. cucumerinus, the great omen- 
tum likewise containing four specimens of C. pisiformis. No inference of any value can 
be deduced from the presence of both kinds of Cysticerci in this case, for they are almost 
always present in tame Rabbits, young or old, whether they have eaten tape-worms or not, 
On the same day, 11th March, portions of the liver were given to a young Dog, and on the — 
17th many examples of C. pisiformis from the omentum of another tame Rabbit were 
given to the same animal. Two or three days before the second worm-feeding, this dog 
had commenced ejecting its ordinary food, and I have no doubt that by far the greater 
part of the more recently introduced worm-feedings shared the same fate. This throwing — 
up of the stomachal contents was followed by total abstinence; and as the dog appeared 
weak, it was accordingly destroyed on the 20th, thus allowing only three days for the 
development of any pisiform Cysticerci that might chance to be retained, and but nine — 
days for the small liver-cysts. The result was as follows :—Seven tape-worms were pre- —— 


sent in the intestine; of these, four were specimens of Tenia cucumerina, varying from a 


to 10 inches in length, and the remaining three were examples of T. serrata, the segmen- — | 
tation of which was scarcely manifested, the longest individual measuring less than an 


DR. T. S. COBBOLD ON ENTOZOA. 167 


inch. On the whole, therefore, it is reasonable to conclude that the latter experiment 
confirms the statements of Leuckart and others as to C. pisiformis being the young of 
T. serrata, whilst both experiments satisfactorily demonstrate the breeding of 7. eueume- 
rina from a minute Cysticercus, probably hitherto unnoticed (figs. 73-79) *. 

The other species of Entozoa casually observed by me in Lepus cuniculus and Canis 
familiaris, ave Fasciola hepatica and Oxyuris ambigua in the former, and Ascaris margi- 
nata in the latter. 

Felis leo; Bos taurus; Ovis aries; Camelopardalis giraffa.—From a partial exami- 
nation of the viscera of two Lions, the only entozoon detected was Ascaris leptoptera. 
I have carefully dissected specimens of Fasciola hepatica and Cenurus cerebralis, but 
have no new facts to offer in respect of their organization or development. From a Giraffe 
I have obtained two forms of Cercariæ and numerous specimens of a very large fluke 
(Fasciola gigantica, mihi), details of which with eoloured figures have already been 
published t. ; 

Delphinus phocena.—1 have dissected, either in whole or in part, several individuals 
without detecting Entozoa; but from a specimen shot in the Firth of Forth, and kindly 
forwarded to me by J. Jardine Murray, Esq., in the month of April 1855, several 
interesting forms were obtained. The pulmonary vessels, both arteries and veins, and 
likewise the smaller bronchial ramifications on the left side of the thorax more especially, 
were extensively occupied by two species of Strongylus, or in other words, by the Pros- 
thecosacter inflexus and convolutus of Diesing; two examples of the former being likewise 
found in the ventricles of the heart. The small intestine of this Porpoise was com- 
pletely choked for the space of 8 or 9 feet by five very large tape-worms so closely 
impacted together that the gut presented all the appearance and firmness of a solid 
cylinder. Four of the worms measured severally between 7 and 10 feet in length, and 
the fifth about 18 inches. This worm constitutes a fresh addition to our sterelminthoid 
fauna. Accepting Prof. Van Beneden’s classification of the cestoid Entozoa, the name 
now proposed will place this new genus between his diphylloid Echinobothrium and the 
pseudophylloid Bothriocephalus. Diphyllobothrium stemmacephalum (mihi) may be briefly 
characterized as follows:—Length upwards of 100 inches, greatest breadth #ths of an 
inch; head arched, supported by a narrow neck, the latter rapidly increasing m breadth ; 
bothria two in number, compressed, shallow, subsessile, together forming a semicircular 
festooned crown; segments „45th to jth of an inch broad from above downwards, marked 
by 10 or 12 longitudinal furrows, the lower porder of each slightly overlapping the suc- 
ceeding segment ; reproductive orifices conspicuous, widely separated, both placed in the 
mesial line (figs. 79-83 inclusive). 
real species of Tape-worm be indicated, otherwise 


und Blasenwürmer, p. 98 et seq.) actually denies 
if proved correct, would almost 


* It is highly important that the cysticercal condition of every 
the value of our breeding experiments is lost. Von Siebold (Band- ev 
the hitherto recognized specific distinetions of five well-marked cestodes. His view, 
sanction a revival of the transmutation theory. 

T Description of a new Trematode infesting the Giraffe, &c. 
of the British Association for 1856. The fluke discovered by Professor Busk in 
Lankester) exceeds in size not only the Distoma gigas of Nardo, but this species 
it is generically distinct. 


Edin. New Phil. Journ. for 1855. See also Reports 
the duodenum of a Lascar (D. Buskii, 
also in a slight degree, from which 


168 DR. T. S. COBBOLD ON ENTOZOA. 


Another still more interesting entozoon, and one which was only discovered after a very 
careful dissection, is a small fluke, from the gall-ducts of the liver. This minute trema- 
tode is furnished with intestinal cæca of a zigzag form, and it is of especial interest as 
establishing the clear-sightedness of those Entozoologists who have insisted upon a generic 
distinction between the organization of the common fluke (Fasciola) and of the Distomata . 
properly so called; we have here, in short, an intermediate condition between the complex 
alimentary cæca of the one genus and the simple bifariously divided tube of the other. For 
the generic and specific recognition of this species I offer the following nomenelature and 
characters :— Campula oblonga. Length $thto$thofaninch; breadth th; oral and ven- 
tral suckers conspicuous ; reproductive pores immediately in front of the latter ; integument 
everywhere clothed with minute spines; vitelline organs largely developed; gastric cæca 
of a zigzag form, somewhat irregular (figs. 84, 85). This entozoon occupies the peripheral 

- branches of the hepatic cæca, which are very much. enlarged and thickened at the infested 
points, sometimes enclosing from fifteen to twenty individuals at one spot. In the stomach 
of the Porpoise there were also specimens of the crustacean Lerneonema and a Filaria, 
evidently belonging however to the partially digested fish-remains within that viscus. 

To facilitate reference, the following is a list of the worms noticed in this paper :— 


LarvarL ENTozoa. 


Coenurus cerebralis. 
Cysticercus cucumerinus. 
—— Giraffe. 

—— fasciolaris. 
pisiformis. 

Scolex polymorphus. 
tetrarhynchus. 


NEMATOIDEA. 
Filaria attenuata, R. 
piscium, R. 

——— rubella ?, R. 
Trichosomum falconum, R. 
longicolle, R. 
Spiroptera leptoptera, R. 


Prosthecosacter convolutus, Dies. 


inflexus, Dies. 
Oxyuris ambigua, R. 
ornata, Dyj. 

Ascaris acanthocaudata, 7. S. C. 
acuminata, Schrank. 
aucta, R. 

clavata, R. 

collaris, R. 

— depressa, R. 
leptoptera, R. 

——— marginata, R. 

— mystax, R. 

——— nigrovenosa, R. 


Ascaris rigida, R. 
spiculigera, R. 
vesicularis, Fröl. 
Dachnitis globosa, Dy. 
sphærocephala, Duj. 
Dochmius tubæformis, Du. 


ACANTHOCEPHALA. 


Echinorhynchus acus, R. 
anthuris, Duj. 

; tereticollis, R. 

Pentastoma proboscideum, Brem. 


CESTOIDEA. 


Bothriocephalus coronatus, R. 

punctatus, R. 

—— rugosus, À. 

Ligula reptans?, Dies. 

Gymnorhynchus reptans, R. 3 
Diphyllobothrium stemmacephalum, T. S. C. . 
Tetrabothrium cylindraceum, R. 
macrocephalum, R. 

Schistocephalus dimorphus ?, Crep. 

Tænia crassicollis, R. 

elliptica, BatscA. 

—— filicollis, R. 

serrata, Goeze. 

sphærophora, R, 

variabilis, R. 


DR. T. $. COBBOLD ON ENTOZOA. 169 


TREMATODA. 
Fasciola gigantica, 7. S. C. Distoma clavigerum, R. 
hepatica, Linn. ——- cylindraceum, Zed. 
Campula oblonga, 7. S. C. —— fulvum, R. 
Hemistomum spathula, Dies. —— gracilescens, R. 
Echinostomum spinulosum, R. |o —— hispidum, Abildg. 
Monostoma caryophyllinum ?, Zed. —— rachion, T. S. C. 
dubium, T. S. C. Polystoma integerrimum, R. 


Distoma atomon, A. 


The names here abbreviated are those of Rudolphi, Bremser, Dujardin, Diesing, Linneus, Zeder, 
Creplin, Frölich, and Abildgaard. 


NR WO tO 


c9 
. 10. One of the above, enlarged 40 diameters. It exhibits the regularly arranged dermal spines 


ll. 
> 12. 
«13, 


. 14. 
. 15. 


. 16. 
17; 


DESCRIPTION OF THE PLATES. 
Tap. XXXI. 


. Tænia filicollis. Natural size. 

. Monostoma caryophyllinum, imperfectly developed. Natural size. , 

. The same, showing an oral sucker and cells in the interior. Magnified 300 diameters. 

. Monostoma dubium (mihi). Natural size. 

. The same, displaying the elongated ventral appendage and internal gastric cæca. Enlarged 200 


diameters. 


. Four examples of Distoma fulvum. Natural size. 
. One of the above, showing the position of the suckers and the extremely attenuated neck during 


elongation. Enlarged 20 diameters. 


. The same, during contraction ; it also exhibits the intromittent organ and ova within the uterine 


tube. Magnified 60 diameters. 
Three specimens of Distoma rachion (mihi). Natural size. 


investing the whole of the body; an oral and ventral sucker, the lining membrane of the former 

consisting of polygonal cells; a narrow æsophagus communicating with a broad, elongated, 
horse-shoe-shaped digestive cavity—in front of the latter a wide uterine tube symmetrically 
disposed in coils on either side and filled with ova; and an internal seminal vesicle. The ovary 
is placed immediately above the caudal vesicle, and is surmounted by the testes. 
Transparent cyst from the subperitoneal surface of the intestine of a Haddock. Natural size. 
The same, ruptured. Magnified 4 diameters. 
Scolex of Tetrarhynchus, escaped from the cyst. 
large caudal vesicle, the surface of which is slightly segmented an 
sclerous particles. Magnified 4 diameters. 


The same with the body and proboscides evolved. Natural size. ; hylloid 
Part of the Scolex, enlarged 20 diameters. It exhibits the four narrow and pointed phylloi 


bothria, a portion of the transparent caudal vesicle with its sclerous particles, and more particu- 


larly the four proboscides, two of which are partially withdrawn from their sheaths. 


An oval sclerous particle, magnified 200 diameters. 
Section of a proboscis or tentacle, showing the pecul 
Magnified 200 diameters. 


It shows the head and body enveloped by the 
d dotted by the highly refracting 


jar arrangement of the hooks and hooklets. 


170 


Fig. 18. 
Fig. 19. 


Fig. 20. 
Fig. 21. 


Fig. 22. 
Fig. 23. 
Fig. 24. 
Fig. 25. 
Fig. 26. 


Fig. 27. 


Fig. 28. 
Fig. 29. 


Fig. 30. 
Fig. 51. 
Fig. 32. 


Fig. 33. 
Fig. 34. 


Fig. 35. 
Fig. 36. 
Fig. 37. 
Fig. 38. 


| Fig. 39. 
Fig. 40. 


Fig. 41. 
Fig. 42. 


DR. T. S. COBBOLD ON ENTOZOA. 


Lower part of one of the proboscides, showing the retractor muscle within the sheath. Magnified 
60 diameters. 

Anastomosing tubes seen beneath the skin, at the lower part of the body, in front of the pro- 
boscideal sheaths. Magnified 260 diameters. 

Two specimens of Dachnitis globosa; male and female. Natural size. 

Anterior extremity of the same (male), showing the position of the mouth and muscular æso- 
phagus. Magnified 260 diameters. 

Tail of the same, showing the sabre-shaped spicules and a depression immediately in front. 
Magnified 260 diameters. 

Section from near the centre of the female, showing more particularly the vaginal sheath and 
prominent folds of the vulva. Magnified 200 diameters. 


Ascaris acanthocaudata (mihi). Natural size. : 
Section of the same, magnified 260 diameters. It exhibits the longitudinal muscular fibres with 


their nuclei, and a broad band of partially disintegrated cells, in the centre of which is a canal 


occupied by an apparently nervous thread. 
Tail of the same, with numerous minute spines at the tip. Magnified 260 diameters. 


Tas. XXXII. 


Anterior part of Ascaris collaris, exhibiting the small trilobular mouth, muscular cesophagus - 
and broad intestine; also four cæcal appendages, two on either side of the digestive tube. 
Magnified 200 diameters. 

Several examples of Scolez polymorphus. Natural size. 

Head of Scolex polymorphus, showing the four suckers, the retracted proboscis, the so-called 
water-vascular canals, the sclerous particles, carmine pigment-cells, and columnar epithelium. 
Magnified 400 diameters. 

Cyst from the peritoneal cavity of a Halibut. Natural size. 

Distoma atomon removed from the cyst. Natural size. 

The same, showing the suckers, the simple gastric ceca, uterine tube, and contractile vesicle at 
the inferior extremity. Magnified 20 diameters. __ 

Four examples of Distoma gracilescens. Natural size. 

One of the same, magnified 60 diameters. It displays the spinose tubercles of the skin, the 
position of the suckers, the central penis-sheath, the botryoidal vitelline organs on each side of 
the neck, the two (or four ?) testes, the uterine tube, the ovary and seminal reservoir, and the 
large muscular contractile vesicle*. 

Bunch of vitelline organs. Magnified 100 diameters. 

Section of the uterine tube, with its double row of contained ova. Magnified 100 diameters. 
Ovum. Magnified 260 diameters. i 
Outline of a dissection of the Short Sun-fish, showing in particular several Gymnorhynehi 
enclosed in cysts within the retractor muscles of the anal fin. Reduced to 4th of the natural size. 
Head, neck, and subcervical enlargement of Gymnorhynchus reptans enclosed within a transpa- 
rent sheath. Enlarged ird of the natural size. 

Anterior fourth of the body of Gymnorhynchus reptans removed from its investing capsule. 
Natural size, 

A few articulations from the posterior part of the body. Natural size. Å 
Club-shaped proboscis exhibiting a multitude of small hooklets serially disposed in rows of ert 


* Von Siebold, in his ‘Lehrbuch der vergleich. Anat.,’ speaks of three or four testes in D. appendiculatum ad — 
D. cygnoides. ; : . : 


Fig. 43. 


. Proscolex removed from one of the caudal segments. 
. Another still more advanced, showing the inverted roste 


DR. T. S. COBBOLD ON ENTOZOA. 171 


teen to each circle; the hooks of the two lowermost rows being very large. Magnified 20 
diameters. å 


Diagram of one of the circlets showing the arrangement of the lesser hooks. Magnified 60 
diameters. 


. One of the large hooks, to show the blunt extremity directed diagonally outwards and down- 


wards. Magnified 260 diameters. 


. A small hook, showing its sharp and curved tip pointed vertically downwards. Magnified 260 


diameters. 
Head, viewed from above, to illustrate the form and aspect of the two lateral bipartite suckers 
and the position of the closed orifices of the proboscideal sheaths. Enlarged 8 diameters. 


. Echinostomum hispidum. Natural size. 
. The same, enlarged 15 diameters. It exhibits the uterine tube, testes, ventral sucker and inte- 


gumentary spines. 


Tas. XXXIII. 


. Another view of the head, showing the oral sucker. å 
. Cutaneous spines. The uppermost one is from the neck ; the central, from the first row sur- 


rounding the head ; the lowermost, from the second cephalic circle of spines. Magnified 200 
diameters. 


. Dachnitis sphærocephala. Natural size. 

. Distoma clavigerum. Natural size. 

. The same, enlarged 12 diameters, 

. Section of the intestinal tube of the speckled Salamander, laid open to exhibit the mode of attach- 


ment of Echinorhynchus anthuris to the mucous surface. Natural size. 


. One of the males, enlarged 8 diameters. : 
. Caudal extremity of a female, showing the artificially distended and transparent integument, the 


muscular sac enclosing the reproductive organs, the central oviduct, and certain peculiar acces- 
sory glands. Magnified 200 diameters. 


. One of the loose globular ovaries containing ova in their nascent condition. 

. Another ovary more advanced. | 

. Å series of ova isolated from the above to exhibit their different stages of development. 
. Fully developed ovum enclosed in a cyst. 
. Front view of the same. 

- Ovum detached from its enveloping cyst, 


showing the lateral position of the yelk and the pecu- 


liar Trichina-like form of the membrane enclosing the albumen. The last ‘six figures are severally 


magnified 350 diameters. 


. Tenia spherophora. Natural size. 
. Head of the same. Magnified 200 diameters. 
. Section from three of the segments showing the extern 


al reproductive organs and imperfectly 
developed ova. Magnified 200 diameters. 

llum with its three pairs of hooks. 
Magnified 500 diameters. 


. Echinostomum spinulosum. Natural size. 
. The same, enlarged 10 diameters. 

. Posterior view of the head, showing the fi 
. Diagram illustrating the arrangement of the vitelline cæca. 
- Ovum. Magnified 300 diameters. 

. XXI, 


n-like disposition of the cephalic spines. 


2A 


172 


Fig. 73. 
Fig. 74. 


Fig. 75. 
Fig. 76. 
Fig. 77. 


Fig. 78. 


Fig. 79. 
Fig. 80. 


- Fig. 81. 


Fig. 82. 
Fig. 83. 
Fig. 84. 
Fig. 85. 


valet 


DR. T. S. COBBOLD ON ENTOZOA. . 


Three examples of Tænia serrata of about seventy hours’ growth. Natural size. 

Head of one of the above. Magnified 10 diameters. 

Tenia cucumerina. Natural size. 

Head of the same, enlarged 25 diameters. 

Joints of the same, showing the male reproductive organs opening at both of the lateral margiris, 
also the ova and water-vascular canals. Magnified 25 diameters. 

Male organs, consisting of testes and vas deferens, and the penis retracted within its sheath. 
Magnified 60 diameters. : 

Head and anterior segments of Diphyllobothrium stemmacephalum. Natural size. 

Section from near the centre of the body. Natural size. 

Head and neck, enlarged 100 diameters. 

Lateral or marginal view of the same. 

Appearance of the bothria when viewed from above. . 

Campula oblonga. Natural size. 


The same enlarged 20 rast exhibiting more particularly the peculiar zigzag conformation 
of the digestive cæca. 


5 = 


“ibis 
si 


sc 


r 


6 Jarman 


Linn. Soc. Vol XXI tab 4] ^.J69. 


nans 


E 


IS Cobbold del. 


59 


Trans. linn. Soc Vol IL tab. 52, h.1( 


G Jarmanv sc. 


TS.Cobbold del. 


Trams Linn.Soo. Vol XIII tab. 88,n.169. 


G Jarman so. 


IS Cobbold del. 


[ 173 ] 


XIII. On the Arrangement of the Cutaneous Muscles of the Larva of Pygæra bucephala. 
By JOHN LUBBocK, Esq., F.R.S., F.L.S., F.G.S. $c. 


* i Read February 18th, 1858. 


LYONET'S celebrated memoir on the larva of Cossus ligniperda, and Straus-Dürckheim’s 
still more beautiful work on Melolontha vulgaris, are the most perfect, and indeed the 
only monographs we have at all complete, on the general anatomy of any insects. 

The very complicated arrangement of the cutaneous muscles in the caterpillar of Cossus 
inspired me with the wish to see how far the muscles of other larvæ would agree with, or 
differ from, this type. 

With this view I dissected with great care some larvæ of Tipula oleracea and of 
Ctenophora bimaculata; but on comparing two or three specimens together, I found to my 
astonishment very considerable variations. It-then occurred to me to compare several 
specimens together, in order to ascertain the nature and amount of these variations. The 
larvæ, however, of the different species of Tipulidæ so much resemble one another, that I 
could not feel certain that these supposed variations were not rather specific differences. 

In order to remove this element of doubt, I selected the larvæ of Pygera bucephala, 
which were obtainable in any numbers, and could not be confused with those of any 
other species. Moreover, as some of these caterpillars live on oak- and some on beech- 
leaves, I was curious to observe whether this difference in the food induced any alterations 
in the muscles. It did not, however, appear to have any such effect. > c 

The full-grown caterpillars are about two inches long, and of a brown colour with dark 

longitudinal bars. DE : 

I have divided my paper into three sections :— 

First, a description of the cutaneous muscles of the larva of Pygæra. 

. Secondly, a description of the variations observed in these muscles ; and : 

Thirdly, a comparison, as complete as possible, of these muscles in Pygera with the 
corresponding muscles of other larvæ. : : 

In examining closely Lyonet’s drawings, I have always found them inconveniently 
small: moreover he has represented alternately the two sides of the insect, so as to reverie 
the directions of many of the muscles; and as he has also figured the same muscle in dif- 

ferent plates, it is difficult to ascertain the exact number of muscles in each segment. 
` In my drawings I have always represented the left side of the insect, and hayo never 
introduced the same muscle into two plates. — — eh 

I propose to commence my description of the muscles at the third abdominal segment, 

and then, passing forward on the one hand and backward on the other, to point out the 


differences which exist in the muscles of the anterior and posterior "o à 
1 À rA 


174 MR. LUBBOCK ON THE CUTANEOUS MUSCLES 


No. 1 is a broad and strong muscle which is inserted into the skin near the front of the 
segment on one side of the dorsal vessel, and passes straight back to be inserted into the 
fore part of the following segment (fourth abdominal). This musele is identical with 
that marked A by Lyonet. 

No. 2 (B of Lyonet) is in fact treble, and consists of three muscles joined end to end; 
posteriorly it is inserted at the ventral side of the preceding, and passes forward parallel 
to it; but instead of being attached at the fore part of the third abdominal segment, it 
unites with the corresponding muscles of the two preceding muscles, and is inserted. Hr 
the fore part of the first abdominal segment. 

No. 3 is absent from this segment. 

No. 4 (D of Lyonet) lies under No. 1, and is T in front, immediately under and 
behind it; posteriorly, however, it does not pass quite so far back, but is inserted at the 
line of separation of the third and fourth abdominal segments. It is not wholly straight 
in its direction, but inclines a little downward in front. 

No. 5 (G of Lyonet) lies under the preceding, and is in some respects opposite to it; 
that is to say, the anterior insertion is on the dorsal side, and the posterior on the ventral, 
but on the same transverse lines. 

These first five muscles must tend to contract the skin of the back. 

No. 6 (E of Lyonet). This muscle lies at the ventral side of 2, and just at the dorsal 
side of the great lateral trachea. Its attachments are hidden by the transverse muscle 
35, which will presently be described, and are on the same transverse lines as those of 
No. 1. 

No. 7 (H of Lyonet) This muscle rises on the ventral side of No. 6 and on the same 
transverse line as the posterior insertions of Nos. 4 and 5, and passing forward and upward 
under Nos. 6 and 8, is attached on the dorsal side and near the anterior end of No. 6; this 
muscle and the next following bear the same relation to No. 6 that Nos. 4 and 5 do 
to No. 1. 

No. 8 (F of Lyonet) rises posteriorly between 5 and 6, and passing forward and down- 
ward between 6 and 7, is inserted at the ventral side of the anterior end of 6. 

No. 9 (I of Lyonet). This muscle is a little more than half as long as 5 and puede 
to it, lying on the dorsal side of the posterior half. 

Nos. 10 and 11 (L of Lyonet) are attached between the posterior insertions of 1 and 6, 
one on each side of the posterior end of 8, and pass straight forward for two-thirds of the 
length of the segment. 

Nos. 12 to 15 (Q of Lyonet) rise almost on the same transverse line and underneath 4, 
and pass forward and downward, so as to cross under 9 and 5, for one-third of the length 
of the segment. 

I have thus described all the longitudinal muscles in the upper half of the segment, and . 
shall now pass to those in the lower half. 

No. 16 (d of Lyonet) rises on the same transverse line as 4 and 5, and "—— on 
the line of division between the third and fourth abdominal segment, on the ventral side 


of the great lateral trachea, and passes straight forward to the front of - _ abdo- 
minal segment. 


OF THE LARVA OF PYGÆRA BUCEPHALA. 175 


Nos. 17 and 18 (c and of Lyonet) are parallel, similar to, and on the ventral side of 16. 
"Nos. 19 and 20 (ff and e of Lyonet) arise under the posterior ends of 16 and 17, and are 

inserted in front under the anterior ends of 17 and 18. They are therefore not quite 
parallel to the axis of the body, but pass a little downward in their course forward, and 
are moreover nearer together in front. They appear to correspond to the two muscles 
marked f and e by Lyonet; these two, however, are not parallel to and at the side of 
one another, but the latter overlaps the former in front. 

No. 21 (a of Lyonet) is parallel to 16, 17 and 18, but is altogether rather further back, 
so that its attachments in some respect alternate with those of 16, 17 and 18, though they 
are not nearly midway. 

No. 22 (i of Lyonet) lies under the preceding muscle, but inclines to the median line 
behind. It is generally bifid in front. 

No. 23 (f of Lyonet) rises on the dorsal side of 24 and runs parallel with it, to be in- 
serted under the fore end of 16. 

No. 24 (part of g of Lyonet) rises just in front of the ganglion and on the same trans- 
verse line as 21, and passing forward and upward joins 23, and is inserted with it under 
the fore end of 17. : 

No. 25 (part of g of Lyonet) rises near the middle line of the body, and passing dia- 
gonally forward and upward joins 24, and is inserted with it under the fore end of 17. 

No. 26 (4 of Lyonet) rises on the same transverse line as 56 and 17, close to the dorsal 
side of the posterior end of 23, and under 25, and passes forward and upward parallel to 
that muscle, to be inserted under its anterior attachment. 

No. 27 (part of k of Lyonet) is wanting in this segment. 

No. 28 (part of k of Lyonet) rises on the ventral side of the hind end of 19, and passing 
forward and downward towards the ganglion is attached to a ridge, which commences 
near the middle line of the posterior end of the segment and is continued forward, at the 
same time curving upward. It is broadest in front, where also it is often divided into 
several heads. ; 

Nos. 29 and 30 (p of Lyonet). These two muscles lie on the ventral side of, and parallel 
to, the preceding. These three muscles vary considerably in form and relative size. 

No. 31 (part of % of Lyonet) rises under the posterior end of 18, and passing forward 
and upward at about one-third of the segment at an angle of 50°, is very much hidden by 
the fold of skin forming the boundary of the two segments. Å 

No. 32 (part of £ of Lyonet) lies parallel to, and on the dorsal side of, the preceding. 

Nos. 33 and 34 (part of ¢ of Lyonet) lie parallel to, and on the dorsal side of, the 
preceding. 

I have now completed the description of the longitu 
to that of the transverse series. 

The first two of this series pass over or 
the muscle 6; the other transverse muscles lie 


the skin, 
rt i i 6 of this 
No, 35 ising i osite and on the dorsal side of 1 
(part of 8 of Lyonet), arising in part opp 5 DU Ue pecelíng 


segment, and partly opposite the hind part of the correspon 


dinal muscles and come therefore 


rather inside the great lateral trachea and 
between the longitudinal series and 


176 MR. LUBBOCK ON THE CUTANEOUS MUSCLES 


ihent, passes upward, with a slight inclination backward, dividing at the same time 
into two heads, which are inserted on the ventral side of 2, and nearly opposite the an- 


terior attachment of 1. 
No. 36 (part of 0 of Lyonet) is parallel to the preceding and has similar attachments, 


but lies rather further back. 
No. 37 (part of « of Lyonet) rises under the fore end of 16, and running forward and 


upward is inserted under the middle part of 35. 

No. 38 (part of « of Lyonet) bears the same relation to 36 that 37 does to 35. 

No. 39 (part of 7 of Lyonet) is attached under the fore end of 19, and passes baekward 
and upward to be inserted into the trachea close to the spiracle. 

Nos. 40 and 41 (m of Lyonet). These two muscles rise anteriorly under 20, and pass 
diagonally backward and upward for about one-third of the length of the segment. 

No. 42 (n of Lyonet) is parallel to and on the ventral side of the preceding. 

Nos. 43 and 44 rise on the dorsal side of 28 and pass backward and upward along the 
posterior margin of 42, opposite to about the middle of which they are inserted. 

No. 45 (part of Z of Lyonet) may perhaps be regarded as a continuation of 39. It is 
inserted into the posterior and lower side of the trachea, close to the spiracle, and passes 
backward and upward to be inserted close to the upper end of 49. 

Nos. 46, 47 and 48 (B of Lyonet) These three transverse muscles lie on the outer side 
of the following, and are therefore covered by it in the drawing: 46 and 47 are usually 
joined together at the upper end. With the two following they are concerned in the 
movements of the proleg. - 

No. 49 (y of Lyonet) is attached below the proleg, and passing upward to the inner 
side of the preceding, is inserted just above it. ; 

No. 50 also is attached at its lower extremity to the proleg, and, passing upward and 
backward, is inserted under the great lateral trachea. 
` No. 51 (8 of Lyonet) ought almost to be considered as two muscles, since, though it 
rises singly close to the anterior end of 34, it divides almost immediately. Both branches 
pass upward, inclining at the same time, and especially the posterior branch, backward, 
to be attached under 6. 

No. 52 (e of Lyonet) is å short transverse muscle which lies altogether on the outer side 
of the anterior branch of 51. 4 

No. 53 (r of Lyonet) rises at the lower end of 48, and passing straight down, at the same 
time expanding in width, enters the proleg. ; 

Under this muscle lie four other small ones, which also are attached to the proleg by 
their lower ends. ; er " 

No. 54 (perhaps part of # of Lyonet) is å transverse muscle which lies on the outside of 
31 and 32, and partly of 28 also. ‘This muscle and the following are often difficult to — 
find, from their being covered by the skin-fold. oe ow? 

No. 55 (perhaps part of ¢ of Lyonet) is in this segment generally completely double, and — 
lies parallel to and just behind the preceding. m 

No. 56 is a series of about eight transverse muscular fascicles which lie on the outside 
of 28, 29 and 30, and are much hidden in a fold which is bounded by the ridge already 


OF THE LARVA OF PYGÆRA BUCEPHALA. 177 


mentioned in connexion with those muscles. The fascicles of which this series is com- 
posed vary considerably in number and form in different specimens. 

No. 57 (perhaps part of z of Lyonet) rises near the middle of the upper side of 56, and 
passes straight back to the posterior end of the segment. 

No. 58 (perhaps part of v of Lyonet) rises close to 57, and passes backward and down- 
ward to the posterior end of the segment. 

Besides these muscles, there are two small ones (86 and 87) which lie under and parallel 
to 53, and another (88) which runs under and transverse to 46, 47, 48 and 53. 

Anterior Segments.—The first two abdominal segments are very similar to one another, 
and may therefore conveniently be considered together. The chief differences which exist 
between them and the third abdominal segment depend on the absence of prolegs. 

It has been already mentioned that the muscle 2, belonging to the second abdominal 
segment, is quite free from the skin, being attached by its ends to the corresponding 
muscle of the preceding and succeeding segments. The other superior longitudinal muscles 
very closely resemble those of the third abdominal segment. In the first abdominal 
segment there are the two following variations :—First, Nos. 10 and 11 are either united 
together in front, or at least closely approach one another, —a tendency which is more 
developed in the two posterior thoracic segments, where they have completely coalesced ; 
and secondly, No. 6 sends off a branch from the upper side of its anterior end. 

Nos. 10 and 11 reach to the fore end of the first abdominal segment, and converge in 
front or sometimes unite. 

In the ventral longitudinal series, No. 18 of the first abdominal segment is considerably 
swollen anteriorly, so as to overlap 21, which in this segment is inserted as nearly as 
possible on the same transverse line as 16, 17 and 18, and under the latter. 

No. 22 of the same segment is single in front. 

i No. 27 was absent in the third abdominal segment; in the first two it rises on the 
dorsal side of 28 and passes forward parallel to that muscle. i 

Nos. 29 and 30 are small in the second abdominal segment, having the same attach- 
ment posteriorly, but not passing so far forward. In the first abdominal segment it is 
difficult to distinguish them from 31 and 32. 

In the first abdominal segment, the four small muscles marked 29, 30, 57 and 58 appear 
to correspond with those so named in other segments; but it is difficult to determine 
Which is which. : : å 

No. 85 is completely simple in the first abdominal segment, where also 36 is wanting. 
In this segment also there are only three muscles to represent 40, 41, 42, 43 and 44, 
though it is difficult to say which of the five have disappeared. 

In both these segments 46 is inserted under or rather outside of 49 above, 
of it below; the dorsal end of 49 is inserted just above 46, and above 51 below; and 48 
is in general quite single. 

Nos. 47, 48, 50, 52, 53 and 56 are completely absent in both segments. i 

Third Thoracic Segment.—The muscles in this segment differ so much from seit 
that have been previously described, that in many cases I feel very doubtful whether 


have used the right numbers to represent them. A careful examination of the thorax in 


and in front 


178 MR. LUBBOCK ON THE CUTANEOUS MUSCLES 


different species would throw much light upon this question, but I have not at present 


the materials necessary for the investigation. 
Nos. 2 and 3 are entirely absent. 
No. 4 passes straight forward, but does not reach the front of the segment. 


No. 5 is completely double. One branch rises at the side of 1, and passes diagonally 
forward and upward to be inserted under it. The other rises under the lower side of 1, 
and passes forward parallel with the other branch, to be inserted under its upper part. 

No. 6 rises on the same transverse line and a little on the ventral side of 1, and passes 
back near the bind end of the segment, inclining at the same time downward. 

No. 7 lies under or sometimes at the ventral side of, and more or less parallel to, the 
preceding. 

No. 8 is double, and instead of passing straight forward as in the abdominal segments, 
inclines to the ventral side in front. Both branches rise at about the middle line of the 
front of the segment, and diverge gradually. The one is inserted under the hinder end 
of 1, and the other under and at the side of the lower branch of 5. 

No. 9 rises under the posterior ends of No. 5, and passes forward and upward, almost 
to the middle line of the back and for three-fourths of the length of the segment. 

Nos. 10 and 11 have coalesced in this segment. 

No. 16 is almost entirely covered by 17. 

No. 18, instead of being parallel to 16 and 17, as in the abdominal — rises under 
the posterior end of 17, and passing forward and downward, crosses the middle line of 
the segment between the two nervous chords, which are here at some distance from one 
another, and is attached to the fore margin of the segment on the ventral side of No. 21. 

Nos. 19 and 20 lie under 17 and 21. 

No. 21 rises on the fore margin of the segment and at the ventral side of 17, and 
passes back, under 18, to the same transverse line as 16 and 17. 

No. 22 rises on the posterior margin of the segment under 16, and passing forward and 
downward, is attached to the skin under the fore end of 21. 

No. 28 rises under the posterior end of 21, and passes forward and upward to the fore 
end of 17, where it is inserted. 

Nos. 24 and 25 are attached in front as usual, but instead of extending the whole | 
. length of the segment to which they belong, are attached to the middle line of the ventral 
side, and are not above half the usual length. : 

No. 26 rises close to the ventral chord not very far from the posterior end of the seg- — 
ment, and passes forward and upward to be attached under the fore end of 20. 

No. 27 is entirely absent. 

No. 28 rises on the dorsal side of 30 and under 20, and passes forward parallel to 30 and - 
under 26 to the middle ventral line. à 

No. 30 rises under 23, and passes downward and forward to the posterior end of 26. 

Nos. 29, 31, 32 and 33 are altogether absent. 

No, 34 is almost as in the first abdominal segment, but lies under instead of over 5 
and 55. 

No. 35 is as in the abdominal segment. 


OF THE LARVA OF PYGÆRA BUCEPHALA. 179 


No. 36 is absent, as in the first abdominal segment. 

Nos. 37 and 38 in the thoracic segments pass more forward, so that they are situated at 
the hinder end of a segment rather than at the front. 

No. 39 is absent, as there are no spiracles in the two posterior thoracic segments. 

No. 40 is double behind. No. 41 is parallel to 40, but not more than half as long. 

No. 42 rises at the lower side of the fore end of 41, and passes upward and backward 
about as far as 40. 

Nos. 48, 44 and 45 are absent. 

No. 46 rises under 17, just behind 38, and passes straight upward, and divides at the 
same time into two diverging branches, both of which are attached to the skin under 7. 

Nos. 47 and 48 are not present. 

No. 49 much resembles 46, in front of which it is situated. It is however more deeply 
forked and is shorter, its lower end not passing so far down. It also passes inside 67, 
while 46 lies outside that muscle. 

No. 50 rises under the posterior end of 7 by a double head, and passes forward and 
downward to be inserted under the middle of 16. 

No. 51. This muscle rises immediately behind the upper end of the hinder branch of 46 
and passes straight down, diverging however a little from 46, to be attached under the 
upper side of 16. : 

No. 52 is small ; it rises at the lower end of 50 and passes to the hinder side of the lower 
end of 51. 

No. 53 is absent. 

No. 54 rises just behind the lower end of 51 and passes upward, going at the same time 
under 51, and being attached under the middle of 46. 

No. 55 rises close to the lower end of the preceding, and passes upward and backward 
about as far as the posterior end of 7. | 
_ No. 56 differs very materially from the series of muscles so marked in the abdominal 
segments. It is here double. The two fascicles rise just in front of and rather above 57, 
and passing downward and forward, at the same time diverging from one another, are in- 
serted opposite the posterior end of the muscles 23, 24 and 25, belonging to the opposite 
side. 

Nos. 57 and 58 are represented by three small muscles which rise under 25 at a little 
distance apart and pass forward, converging at the same time, and are inserted into the 
legs. They frequently coalesce so as to form only one muscle. 

We have now completed the muscles in the third thoracic segment which appear to 
represent those of the abdomen. There remain yet fifteen to be described. 

Nos. 59 and 60 rise under the fore end of 1, and pass upward and backward re 
dorsal line, where they are attached close to the anterior end of 9. 

No. 61 rises at the lower side of 60 and passes back parallel to that muscle, på rd 
. ås far, and at the same time dividing into two, so that its lower branch is attached close 

to the fore end of 12. — d the 

No. 62 rises close to the posterior end of 60 and passes ne Ys 
fore end of 4. In some specimens T could not find this muscle. 


D 
VOL. XXII. "T 


180 . MR. LUBBOCK ON THE CUTANEOUS MUSCLES 


No. 68 rises partly under the fore end of 7 and partly between 7 and 1, and passes 
backward and downward to the upper end of 49. 

No. 64 rises under 7, and just on the lower side of 63, and runs parallel to that muscle, 
It does not however pass so far back. It is generally completely double. 

No. 65, again, rises just below 64, and passes back parallel to it, but is about twice as 
long, and reaches to the upper end of 42. 

No. 66 rises under the posterior end of 17, and passes forward and upward to the 
posterior end of 64. 

No. 67 rises on the lower side of 42 and passes upward to the fore end of 66. 

No. 68 rises under the lower end of 49 and passes forward and upward to the anterior 
end of 8. 

No. 69 rises at the fore end of 68 and passes to the lower end of 50. 

. No. 70 rises at the hinder end of 68 and passes to the hinder end of 40. 

No. 71 rises at the lower end of 70 and passes to the upper end of 54. 

No. 72 rises at the hinder end of 63 and passes to the lower end of 50. 

No. 73 rises under the middle of 17 and passes downward and forward to be inserted 
close to 24. 

No. 74 rises at the upper end of 73 and diverges from it toward the upper side of the 
lower end of 56. 

No. 75 is behind, and almost parallel to, the preceding. 

No. 76, rising under the fore end of 8, passes backward and downward to be inserted 
under the middle of 17. 

No. 77 rises at the posterior end of 76, under 17, and passes straight back to the hind 
end of the segment. Å 

No. 78 also rises at the hind end of 76 and passes downward and backward. It is a 
short and small muscle, consisting generally of four distinct fascicles. 

First and Second Thoracic Segments.—The muscles of the thoracic segments differ from 
one another, as might be expected, much more than do those of the abdomen. It is 
unnecessary here to mention the muscles which agree with those of the third thoracic 
segment, and I will therefore confine myself to pointing out the differences, using the 
third thoracic segment as the standard of comparison. 

Second "Thoracic Segment, however, does not differ very much from the third, and this 
agreement is even more striking in the small muscles than in the large ones. 

Nos. 4 and 7 are thinner and weaker. 

The single muscle representing 10 and 11, which are again distinct in the first thoracic 
segment, is here expanded in front, and usually double. 

Nos. 16, 19 and 20 are represented by only two muscles. 

No. 18 is double at the posterior end. 

No. 22 rises at the upper end of 16, and passes backward and upward to the posterior 
endof6. X à 

No. 52 runs from the lower ends of 50 and 72 to those of 54 and 55. 

Nos. 57 and 58 in most of my specimens had united together, and were smaller than 
in the third thoracic segment; but this is not, I presume, always the case. 


OF THE LARVA OF PYGÆRA BUCEPHALA. 181 


Nos. 59 and 60 have united together. 

No. 61 overlaps 60 at the fore end. 

No. 63 is generally, if not always, double. 

First Thoracic Segment.—In the posterior part of this segment several muscles are 
wanting, and in the anterior they differ widely from the usual type, being principally 
modified so as to effect rotations of the head. 1 

The most striking alteration is the presence of the muscle marked 82 (C+of Lyonet), 
which is attached in front to the posterior border of the head, and passing through the 
present segment is attached by four heads along the middle dorsal line of the second 
thoracic segment; nor did it vary in any of the specimens examined by me. There does 
not seem to be any muscle comparable to it in the other segments. There are yet three 
other muscles which seem to be peculiar to this segment. No. 80 rises close to the fore 
end of 82, and passing over 19, 20 and 26, is attached by two heads near the middle 
ventral line, close to 56. No. 81 rises with the preceding and follows nearly the same 
course, but passes under 19, 20 and 26. No. 83 rises from the fore side of the spiracle 
and passes forward to the upper end of 46. 

No. 1 is in this segment divided into two or three separate muscles, which are attached 
side by side behind, and fixed to the fore part of the segment, or rather into the mem- 
brane connecting the head with the thorax. 

No. 4 is attached in front, close to 1. 

No. 5 is double; in some cases however it is united in front. The one fascicle rises 
under or at the side of 1, the other rather further down, and both are inserted on the 
ventral side of 4. 

No. 7 lies at the lower side of 6 and parallel to it. ; 

No. 8 perhaps is wrongly identified, since it here lies under 10 and 11, rising close to 
the fore end of the latter and passing back to the hind end of the former. No. 9 is 
much broader than usual. : 

Nos. 10 and 11 are both present, and offer no peculiarity, except that 11 is expanded 
and sometimes double in front. ; 

Nos. 12, 13 and 14 are absent. 

No. 15 is a straight muscle, longer than, and lying at the side of, 9. - ; 

No. 16 is completely double; in front it lies at the upper side of 17, and under it behind. 

No. 18 rises under the hinder end of 21, and passes forward with a slight inclination 
downward to be inserted at the lower side of the fore end of the same muscle. 

Nos. 19, 20 and 26 are parallel to one another; they rise under 17 and 21, and 
pass forward and upward to be inserted at the anterior end of the segment, not far 
above 16. : 

No. 22 rises under the fore end of 16 and passes backward and upward to the hind 
end of 6. 

Nos. 23, 28, 29 and 34 seem to be absent. 

_ No. 31 is perhaps wrongly identified. It rises under 
In front of the spiracle. 
Nos. 37 and 38 appear to be absent. 


16 and passes upward and forward 


2522 


182 MR. LUBBOCK ON THE CUTANEOUS MUSCLES 


No. 40 is longer than in the second thoracic. In one specimen it was single behind, 
and inserted between 51 and 66. 

Nos. 46 and 49 are both quite single, and much larger than in the segments following. 

No. 50 is wanting. 

No. 51 rises at the posterior end of 40, and passes downward to be inserted under 17. 
This muscle may perhaps be rather the representative of 54. 

No. 52 rises close to the lower end of 49 and passes downward and backward. 

Nos. 54 and 55 are absent. 

Nos. 59 and 60, There are three or four separate fascicles which apparently represent 
these two muscles. They lie in front of 9, and are parallel to the direction of the 
segment, | 

No. 61 is almost or sometimes completely double. It rises at the fore end of 82, and 
the two branches diverge slightly, both being inserted under 59 and 60. 

No. 62 consists of three or four small oblique fascicles which rise along the front of the 
segment under the three preceding muscles and pass backward and upward. 

Nos. 63 and 64 are absent. | 

No. 65 rises close to the lower end of 49 and passes upward and forward. It is very 
doubtful whether this muscle is homologous with that marked 65 in the other segments. 

No. 66 is much less conspicuous than in the other thoracic segments. 

No. 67 rises under the upper end of 49 and passes to the hinder end of 41. 


Nos. 68 and 69 rise in front as usual, but pass back parallel with one another to the 
upper end of 51. 


Nos. 70 and 71 are parallel with 65. 

Nos. 72, 73, 74 and 75 are absent. 

No. 79 lies under 77 as usual, and in this segment is hidden by it. 

No. 80 rises close to the anterior end of 19 and passes downward, at the same time 


with a slight inclination backward, to the middle line of the segment, close to 56. At 
its lower end it is double. 


No. 81 resembles 80, but is smaller; it lies also rath 
instead of on the inner side of 19, 20 and 26. 

No. 82 has been already described. 

No. 83 is devoted to the spiracle, from which it passes forward and upward to the 
upper end of 46. å 

Nos. 84 and 85 rise at the lower end and a little in front of 19, and diverging from one 
another, pass backward and downward ; 85 is inserted near the posterior end of 57 and 
58, and 84 on the ventral side of them, near their fore end. 

Posterior Abdominal Segments.—Having thus described the thoracic segments, I now 
return to the abdomen. The fourth, fifth and sixth abdominal segments do not mate- 


Fund differ from the third. The muscles, however, become narrower and thus occupy 
ess space. 


No. 26 is double behind, thus indicating the more complete separation which is to 
take place in the following segment. 


No. 


er in front, and passes on the outer 


28 is absent. In the fifth and sixth segments there is also a muscle which rises 


OF THE LARVA OF PYGÆRA BUCEPHALA. 183 


just above the upper end of 46, and passing downward is inserted just below the lower 
end of 53. : 

No. 52 is absent. 

No. 55 is single in the sixth segment. 

Nos. 16 and 17 approach nearer to one another, and separate a little from 18. 

Nos. 47 and 48 are absent. 

No. 49 in the fifth and sixth segments is completely double, and stronger than in the 
fourth segment, which indicates that the prolegs of these segments act in a somewhat 
different manner. 

The gastric muscle ç rises, as in the larva of Cossus, on the upper side of 21. 

Seventh Abdominal Segment.—Owing no doubt principally to the absence of prolegs, 
this segment differs much from those that immediately precede it. Thus 47, 48 and 56 
are entirely absent. 49 and 50 also, though present and well-developed, are much shorter 
than usual. — 

On the other hand, in the specimens I examined there were two new muscles lying 
under 10 and 11, and inserted below them in front. 10 and 11 approach one another in 

front, and sometimes even unite together. 

No. 25 has completely united with 24, and 26 lies on the lower side of these two muscles. 

To represent 31, 32, 57 and 58, there are four small variable muscles which seem to be 
of little importance. 

No. 46 also is very thin and weak, so that it is probably of little functional utility, and 
only present to fulfil the general law of arrangement of the muscles. 

No. 51 is single and small. It rises at the lower end of 49 and passes to the front of 
34, as in the anterior abdominal segments. 

No. 52 is absent. 

No. 53 is parallel and similar to 54, though a little shorter; these two muscles are very 
unlike in front, but become gradually more and more alike. 3 

No. 56. As there are no prolegs to this segment, the series of muscles marked 56 is 
absent, 

Nos. 57 and 58 are represented by two small muscles parallel and similar to 31 and 32. 

Eighth Abdominal Segment.—In this segment the muscles have become much narrower 
and weaker, and 31, 32, 33, 34, 50, 53 and 56 are absent. 

. Nos. 12 and 13 have almost coalesced, so as to form one small muscle, and the same 
has taken place with 14 and 15. i ne 

No. 4 has entirely disappeared, while 3, on the other hand, is double; as also happens 
in the larva of Cossus. re 

Nos. 7, 8, 10 and 11 have become parallel, and seem to form ee: war 

In the Ninth Abdominal Segment the diminution of the muscles in size and num! en = 
proceeded still further; 3, however, is double, as in the preceding segment, gn ar 
its appearance again. A 

Indeed the DEG of missing muscles is much greater than of those which are pre- 
sent. These are, Nos. 1, 2, 3, 4, 5, 6, 7, 16, 17, 18, 21, 37, 38, 50 and Ble 

The first six of them resemble the same muscles in the preceding segment. 


184 MR. LUBBOCK ON THE CUTANEOUS MUSCLES 


No. 7 rises at the posterior end of 16, and passes forward and upward nearly to the 
upper end of 37. : 
Nos. 16 and 17 have coalesced. E 
Nos. 50 and 51 are even larger than usual, and pass quite back to the posterior end of - 
the segment. 51 is double in front and attached under 16. : 
Besides these muscles, there are attached to the upright wall of the end of the body, - 
and in a series along the lower middle line of the segment, a number of muscular fascicles 
which are distributed to the rectum and posterior parts of the alimentary canal; in the | 
same place also are situated the muscles which move the posterior prolegs ; I have not, 
however, examined these muscles with care enough to enable me to describe them satis- 
factorily. 3 
Comparison of the Muscles in different Specimens.—Any one who has carefully exa- 
mined Lyonet's drawings of the cutaneous muscles in the larva of Cossus ligniperda, or. | 
still better, has made himself acquainted with the organs themselves, must have been 
struck with amazement at their number and complicated arrangement. 1 | 
The muscles in most animals are penetrated by the arteries and capillaries, and are 
supplied by them with blood; but this not being the case in insects, it is perhaps neces- 
sary that their muscles should be divided into numerous fascicles, in order that the blood 
may have free access to them. "Whether for this or some other reason, they are in the 
shape of thin ribands, the thickness of which varies very much in different muscles and - 
in different parts of the same muscle, but is in the largest about 37th of an inch. 
The largest muscular fibres in man, which are not penetrated by blood-vessels, are, I : 
am: informed by Professor Huxley, goth of an inch in diameter. We can, however, - 
en argument from this comparison without knowing the permeability of the musde 
and the power of penetration possessed by the blood. It is also worthy of notice, that the 
pg of insects are separated into very small fascicles, as if, from the violent and - 
dre e muscles, their particles were more quickly deteriorated, and required - 
quently to be removed and replaeed by others. The muscles of 


insects, however, ar = : : TEC 
RR e not only very much divided, but are also very compliqués in their à 


e head and 
pourra qu’étonn 


de I 2 ^" yn 
ENER, e quil y en å méme qui le fixent à beaucoup moins." It might 


s in insect larvæ, as for instance 16, 17, 18, 19 and ~ 
of muscles rather than as whole muscles, and that i | 
very much diminished. | 

~h objection, we should expect to find these pu 
do so, R umber; and as, except in certain instances, they do = : 
he cui must, I think, admit that Lyonet was ini in his mode of estimating e 
. Some few, | 


however, of t d 
amount of thes. he smaller muscles do vary; and to determine the number and ; 


se variati ief object ; 
lons was my chief object in undertaking the present paper. 


OF THE LARVA OF PYGÆRA BUCEPHALA. 185 


No naturalist had previously undertaken this task, though Lyonet has mentioned here 
. and there certain variations observed by him. 

It would have taken a great deal of time, and seemed to me scarcely worth while, to 
make this comparison in all the segments of the body ; and I have therefore confined 
myself to the third, fourth, fifth and sixth, thus including a thoracie somite, two abdo- 
minal without prolegs, and one with these appendages. 

These four segments I compared carefully in nine different specimens, five of which 
were fed upon Oak and four upon Beech. But in order to determine a few minor 
points, I dissected perhaps as many more specimens in a cursory manner, which how- 
ever confirm me in the opinion that the larger muscles scarcely vary at all. Indeed 
the only mutation observed in them at all worthy of notice was that in one case the 
muscles of the third abdominal segment, 16, 17 and 18, were not attached to the skin 
in front, but were continuous with the corresponding muscles of the preceding segment. 

The differences which do exist are not exactly those which I expected to find; whilst 
some things which I rather thought might vary, remain on the contrary always the 
same. 

Let us take for example Nos. 4 and 5. In the first abdominal segment 4 passes on the 
inner side of 5; the contrary, however, might apparently have been the case, without 
alteration of the places of insertion, or of the mode of action of the two muscles; and I 
thought it probable, therefore, that 5 might sometimes lie on the inner side of 4. 
But although six segments in at least fifteen specimens were examined, making in all 
ninety instances, the arrangement was invariably the same in every instance. 

This remark is applicable to several other muscles, for instance 6 and 8, 7 and 8, and 
23 and 18 in the third abdominal segment; and, indeed, I never observed a case in which 
a muscle varied in this respect in its relation to another. 

The muscles 4 and 5, however, offer a different arrangement in the thorax to that 
which is so constant in the abdomen. Here, if they are rightly identified, which perhaps 
` may be doubted, though I believe that I am correct, 5 is completely double and lies on 
the inner side of 4. 

This altered arrangement appears to be necessitated by the altered position of the 
fore end of 4, and by the presence of 60 and 61. : 

Again, 8, which in the following segments lies on the inner side of 11, in the first 
thoracic segment has shifted its position and passes on the outer side. 

A careful examination of the Plates will show a few more instances of this fact, which, 
however, are not very frequent. : diei) 

The relative sizes of the different muscles appeared to vary very little in different 
specimens, except indeed in those muscles which might be termed the variable 
muscles. i : 

Such were especially 12, 13, 14 and 15; 28, 29 and ws 56; 59 and 60; anis m 
lesser degree, 31, 32, 33 and 34; 57 and 58; 11; 23 and 24 in the third thoracic segment, 
8 in the third thoracie segment, and 40, 41 and 42. 

If we suppose, as seems probable, that muscles composing 
usually act together, and that the same is the case with the oth 


the series 12, 13, 14 and 15 
er three series, 28, 29, 50 ; 


186 MR. LUBBOCK ON THE CUTANEOUS MUSCLES 


31, 32, 33, 34; and 40, 41 and 42; then the variations observed in these muscles will be 
of very little importance to the animal. 

There are, indeed, other muscles and series of muscles which might å priori have been 
considered equally unimportant, and have therefore been expected to show similar varia- 
tions. Our knowledge, however, of the mechanism of the muscles, and of the functions 
they subserve, is so small, that we must be very cautious in forming an opinion on the 
relative importance of different muscles. 

It may perhaps prove that the variability of particular muscles is rather an indication 
of a different arrangement hereafter to be discovered in certain neighbouring groups. 
We must, however, suspend our judgment upon these facts until we are better acquainted 
with the myology of other insect larvæ. 

I expected to find, in specimens in which the variable muscles were divided into more 
fascicles than usual, that this tendency to the division of muscles would pervade the 
whole animal, or at least the whole segment; this, however, did not appear to be at all 
the case. It seemed also probable that the variable muscle would either be symmetrical, 
or at least would tend to be so, in the two opposite sides of the same animal; but in the 
instances in which both sides were examined, this rule was not found to hold good. 

Differences in the Third Abdominal Segment.—In the fourth specimen, No. 1 sends 
off a small branch on the dorsal side at the front end. 

No. 5, in the sixth specimen, is partly overlapped by 4 in front. 

- The series 12 to 15 consists normally of four small muscles situated in the hinder part 
of the segment, and passing forward with a slight inclination downward. However, in 
the third specimen which I examined, the fore end of 15 was turned under 14. No. 12 
was double in front, and 13 was almost entirely double, being only joined together in 
front. In the fourth specimen 13 and 14 were completely double, and 12 and 15 were 
divided for the anterior portion. The two branches of No. 15 were unequal, the dorsal 
one being the longer, and inserted under the anterior end of 14. In the fifth specimen, 
12, 13, 14 and 15 were double in front. In the sixth and eighth specimens, on the con- - 
trary, 12 and 13 were united together; and in the latter are shorter than, in the former 
the same length as, 14 and 15. 

No. 7 is usually single, but in one specimen it was divided into two, for about the 
- posterior one-third. 

No. 20, in the sixth specimen, a little overlapped 19 in front. 

No. 22 is usually double for about its anterior half ; sometimes, however, it is more 
deeply cleft, and at others it is almost undivided. Sometimes again it is trifid, and in 
this case the three divisions may differ in size and in degree of division. 

No. 25 varies a good deal in width in different specimens. 

Nos. 28, 29 and 30 also vary a good deal. Sometimes all three are of about the same 
width. Sometimes 28 is much the largest, and is bifid, trifid, or even quadrifid. At 
others 28 is smaller than 29, and the latter is more or less bifid or trifid. Sometimes 
both these muscles are bifid, and in one instance 30 was completely double. 

AA isdem 84. Sometimes 31, sometimes 32, sometimes 34, is double; some- 
nd 34 are double. Sometimes, on the contrary, 33 and 34 have coalesced. 


OF THE LARVA OF PYGÆRA BUCEPHALA. | | 187 


No. 35 in one specimen was divided into three at the upper end. 

No. 37 is sometimes å good deal expanded at its upper end, and sometimes divided 
into two. 

No. 38 in the same way is more or less divided at the upper end. 

No. 51 is more or less divided, the two main branches of which it consists being some- 
times united for a large proportion of their course, and sometimes, on the other hand, 
almost entirely separate. 

No. 52, which is usually single, was, on the contrary, in one specimen double. 

No.56. This series of muscles varies extremely. There are usually from 7 to 11 fascicles, 
but they were not alike in any two specimens I examined. 

No. 57 was double in the second specimen, and 

No. 58 in the fifth. 

Second Abdominal Segment.—The cases of variation in this muscle are almost the same 
as those already deseribed. 

No. 1 in the fourth specimen was completely double. 

No. 10 in the second specimen was double for the anterior half. In the fourth specimen 
10 and 11 were very small and united in front. In the ninth specimen 11 was double in 
front. 

Nos. 12, 13, 14, 15, 27, 28, 29, 30, 31, 32, 33 and 34 offered various differences of the same 
nature as those already mentioned; besides which, in the fourth specimen, 14 was much 
longer than 13 or 15. 

No. 26 was completely double in the second specimen. 

No. 38 was completely double in the fifth specimen; in which also 

Nos. 40 and 41 were united by their lower ends. 

No. 54 in the eighth specimen was completely double. ai 

First Abdominal Segment.—Nos. 12, 18, 14, 15, 27, 28, 29, 30, 31, 32, 33 and 34 differ as 
much as usual, ; 

Nos. 10 and 11 are sometimes united in front, sometimes they only converge and do 
not join. 

No. 18 in some specimens does not cover the fore end of 21. 

No. 37 in the fifth specimen is double. 2 ; 

Third Thoracic Segment.—Nos. 12, 13, 14 and 15 differ as in the above-mentioned 
segments; 29, 30 and 31, on the contrary, are much more regular. | 

No. 8 in the fifth specimen was completely double; in the sixth it was 
greater length, but united in front. 

No. 20, in the ninth specimen, was double in front. 

; Nos. 23 and 24 differed considerably in different specimens. T 
1, however, perhaps the most usual. 
No. 62. This småll muscle was in two cases entirely absent. 


No. 68 was double in two speci 
pecimens. ; ; e 
Nos, 59 and 60 are sometimes united; in the fourth specimen they were gar = 
“one si de, and had joined on the other. In the ninth specimen the iran en " 
VOL. XXII. 


double for the 


he arrangement figured 


188 MR. LUBBOCK ON THE CUTANEOUS MUSCLES 


instead of lying at the side of 60, was inserted under, that is to say, at the outer side of 
it. In two specimens there was a small muscle lying at the dorsal side of 59. 
No. 61 is sometimes simple behind, sometimes divided. 


I now proceed to compare the muscles of Pygæra with those of Cossus ligniperda, as 
described by Lyonet, and of one or two other insects which I have more or less com- 
pletely dissected. 

The following Table shows the letters and numbers used for the corresponding muscles 
in the abdomen :— 


1=A 1624 sant 46 = 
2=.B 17.2536 Re 47 = »y 
5 = C Ig me uS ¿ 48 = 
4=D 9=f uc] 49 — 
5G 20 = e Bi " 50 = 8 
6=E 21 = a ur) 51 = 
7=H 24cm yo 02 m. 
8=F 28 =f 39 2. [^ | 53 = „Sin the first figures 
ger E 24 = | 39 — I (part) ex Ur 
Tesi: 25 =f | 40=m 54 = 
11 = 26 =} ee at? 
ms Hai | 42 T 56 absent 
= =p 43 = 57 = 
14= (Q 20. nue + un he 
15 = 30 | " 45 = 1 (part) 


Thus, with very few exceptions, every muscle of the third abdominal segment in Pygera 
can be referred to its homologue in Cossus. 

There are, however, some few differences: B in Cossus is only as long as the segment, 
_ as is also the case in the larva of Pontia rapi ; while 2, the corresponding muscle in Pygera, 

has united with the same muscle of the two preceding segments,—the three together 
forming only one muscle as long as the first three abdominal segments. 

: Again, G shows traces of subdivision, and is covered in part by F, while 5 is single, and 
lies on the dorsal side of 8. I also is treble, while 9 is single. L is treble, while 10 and 
11 only form two muscles; on the other hand, Q is only treble, while the series from 12 
to 15 is quadruple. 

€ crosses over ff, so as to cover it behind instead of lying at the side of it. % does not 
seem to be present in the third abdominal or any of the posterior segments of Pygera, 
and only makes its appearance in the second abdominal segment. 

There seem to be three fascicles, marked Z, to correspond to 33 and 34; and 35 and 36, 


which are marked 6, are represented by several fascicles. 
First and Second Abdominal Segments.—In these segments there are no further differ- 
ences of importance. 35 and 36, however, which in Pygera are reduced to a single 
muscle, continue in Cossus compound as before. 
Nos. 46, 47, 48, 49 
f and e resemble 1 


OF THE LARVA OF PYGÆRA BUCEPHALA. . 189 


to give up this task, and do not think that it can be effectually accomplished until we 
know the arrangement which exists in some of the intermediate genera. 

Some, however, of the more remarkable peculiarities of the thoracic segments in 
Pygæra are equally present in Cossus. Thus, the altered position of 1, 8, 8', 18, 22 and 
82 resemble those of D, C and E, c and e and C+. The remarkable muscles, however, 
marked 76 and 77, do not appear to have any close representative in Cossus. 

This comparison, however, is made far more difficult by the arrangement of Lyonet's 
figures, and by the fact that the same letters are certainly not always used for the corre- 
sponding muscles in different segments. 

In the first abdominal segment of the larva of Pontia rapi the larger muscles were 
arranged nearly as in Pygæra, except that 2 was attached to the skin at the posterior end 
of the segment, as in the larva of Cossus. In the larva of Disphragis ceruleocephala the 
muscles 1 and 2 on the one hand, and 16, 17 and 18 on the other, were represented by 
numerous separate fascicles, amounting to at least ten in each instance. 

This separation of the fibres composing a muscle into separate fascicles is carried to a 
much greater extent in the larvæ of Coleoptera, or at least in Dyticus and the wood-feed- 
ing Lamellicorns, which alone I have examined. In these two groups each of the larger 
muscles is represented by at least twenty separate fascicles, which makes it far more 
diffieult to distinguish the arrangement of the muscles. 

The reserves of fat in the larvæ of these Lamellicorns are stored up in large vesicles, as 
much as ‘01 of an inch in diameter, and which, being connected together into thin mem- 
branes, like a mesentery, have a beautiful bead-like appearance to the naked eye. 

The muscular system of the larva of Tipula oleracea and of Ctenophora bimaculata* offers 
us many interesting points of difference. Being unprovided with legs, these little creatures 
move by resting one part of their body against some solid object, and then pushing the 
anterior part forwards. To enable them to do this with facility, an immense number 
of muscles are attached to the inner side of the skin. The total number, indeed, falls 
a little, though very little, short of that in Cossus; yet the real complexity is greater, 
since the average number of muscles in each of the body-segments is rather over m enty 
in Tipula, while in Cossus it is about sixty. This surely shows greater complexity than 
the larger total, which is merely made up by the irrelative repetition of a lesser number 
of muscles in a greater number of segments. 

The joints of the back in the larva of Cossus, like those of Pygæra, allow | 
play than those of the ventral side, and we find the muscles entirely in accordance with 
this structure. On the contrary, in the larva of Tipula and of Ctenophora bimaculata, 
the back can be bent almost, if not quite, as much as the belly; and if this were not already 
known, it might be deduced from the arrangement of the muscles. 

Lyonet remarks (p. 154, 7. c.) that his “a,” which corresponds to my 2 adr) 
quable, en ce que, pendant que les autres muscles droits se terminent aux some pes 
anneaux, son extrémité postérieure passe au 4, 5, 6, 7, 8, et 9 anneaux cette L 
*t s'insère assez avant dans l'anneau qui suit; ce qui vraisemblablement a été ain 


much less 


1, “est remar- 


i d found with, those 
: Mr. Walker was kind enough to name for me specimens bred from larvæ resembling, an 
Which I dissected, = 
Cc 


190 , MR. LUBBOCK ON THE CUTANEOUS MUSCLES 


ménagé pour faciliter l'ondoyement que fait le corps: de la chenille quien elle marche, et 
qui en rend le mouvement progressif plus aisé que s’il étoit vermiculaire.” 

It is hardly necessary to say, that there is no similar muscle on the dorsal side in 
Cossus or in Pygæra. In Tipula, on the contrary, there is a dorsal muscle to which 
these remarks would well apply. . 

The great flexibility of the back shows itself, however, in many other ways. In Tipula 
there lie upon each side of the muscle 6, which is here represented by four distinct 
fascicles, five muscles, corresponding to 16, 17 and 18 on the ventral side; those on the 
other side being apparently a great development of 2, which becomes almost as important 
as 16, 17 and 18 together. Moreover, 1 and 21, instead of having their anterior termi- 
nations a little behind those of 16, 17 and 18, completely alternate with them; and, 
again, there are other muscles whose posterior insertions are close to those of 1 and 21, 
while they pass as far forwards as 2 on the one hand, and 16, 17 and 18 on the other. 
These arrangements must make the body much more supple. i 

Once more, there are strong oblique muscles both at the back and at the ventral side, 
which must give the animal great powers of twisting itself forward with a corkscrew-like 
motion. | 

I hope, however, at some future time to be able to lay before the Society a complete 
account of the muscles of these larvæ, which I have dissected and figured with great care, 
and shall content myself at present with mentioning one other remarkable peculiarity. 

The muscles of the first four body-segments are alike, and very different from those of 
the posterior somites; we must suppose, therefore, either that the thorax of insects has 
in reality four segments, or that the muscles of the first abdominal or last cephalic seg- 
ment have become developed like those of the thorax; in fact, that the first segment of 
the abdomen, or the last of the head, has detached itself from its usual position and 
joined itself to the thorax. 

The latter hypothesis seems to me more probable, and it is also supported by the position 


of the anterior ganglia, which are contained in the second segment (first body-segment) — 
and that of the labium, which also rises in this segment, although its upper surface has — 
partially coalesced with the under side of the head. I am reminded, however, by Prof. 
Huxley that neither of these conditions | 


is conclusive. The anterior abdominal ganglia 
not unfrequently move forwards into the 


not u thorax, so that the cephalic ganglia may perhaps 
in this case be moved back; and the labium, not being homologous with a pair of coalesced 
appendages, but simply a protrusion of the sternal surface, may perhaps in this case have 


SØR - with it a portion of the sternum of the first thoracic segment, in addition to 
Dr. mnd : € com øy Segment, of which it normally consists. 

thoraeie ER : = er the auiyression: that the last segment of the head put on à 

thrown td Tom arve of certain Hymenoptera. Mr. Westwood, however, bas _ 
wun great doubt upon this Supposition, if indeed he has not disproved it altogether*. . 


* See the Trans. i. 
8. Ent. Soc. vol. ii. p. 125, and Todd’s Cyclopædia, article Insecta, p. 871. 


OF THE LARVA OF PYGÆRA BUCEPHALA. 191 


DESCRIPTION OF THE PLATES. 
Tan. XXXIV. ann XXXV. 


In Tab. XX XIV. the insect is supposed to have been cut open down the side, the internal organs to 
have been removed, and then the skin to have been flattened. The muscles of one side only are 
represented. 

In the first figure of Tab. XXXV. the muscles figured in the former plate are supposed to have been 
cut away, 50 as to expose many others which would at first have been partly concealed. 

In the second figure of Tab. XX XV. the remainder of the cutaneous muscles are represented. 

Of course the muscles represented in the last two figures would have partly appeared between those 
figured in Tab. XXXIV., but I thought it would cause confusion if they were inserted more than once. 

All three figures refer to the three thoracic and first four abdominal segments, and are magnified 


about 8 times. 


a t " 


Frans. Linn. Soo. Vol. KHL tab. 34. p 191. 


rO, 


BEER REE VORES re 


PLL Ag. 


Middle 


[dø] 


J 


XIV. On the Agamic Reproduction and Morphology of Aphis. —Part I. By Tuomas 
 H. Huxzey, F.R.S., Professor of Natural History, Government School of Mines. 
: Communicated by G. Busk, F.R.S., F.L.S. 


Read November 5th, 1857. 


$ 1. Preliminary Remarks. 

$ 2. The Viviparous Female, and the Development of the Pseudova. 

$ 3. The Oviparous Female, her Reproductive Organs and Ova. 

$ 4. The Development of the Pseudovarium in the Viviparous Female. 
$ 5. Summary; and Comparison of Germs and Ova. E 
$ 6. Hypothetical Explanations of Agamogenesis. x 

$ 7. Classification of the Phenomena of Agamogenesis. 


$ 1. Preliminary Remarks. 


“J'AI souvent pensé qu’on pourrait, dans l'histoire des sciences, désigner les époques par 
les principales découvertes. Par exemple, 1665 seroit l’époque de la Gravitation; 1789, 
l'époque de la méthode naturelle en Botanique; et, si parva licet componere magnis, les 
années 1740 à 1750 seroient l’époque des Pucerons*.” e ot EF 

Without, perhaps, being disposed to go so far as the enthusiastic French investigator - 
of Plant-lice, no physiologist will deny that the experiments conceived and attempted by 
Réaumur, but first successfully carried out by Bonnet, established faets of the highest 
importance, and raised questions which still disturb the very foundation of his science. 

But what were these great facts, established by Bonnet and his successors or contem- 
| poraries, Trembley, Lyonet, Degeer, Kyber, and others? | | 

If the moderns paid due attention to the labours of their predecessors, an accurate 
answer to this question should be found in every accredited text-book on zoology. But 
it will be found, on the contrary, that important errors have crept into the current con- 
ceptions respecting the reproductive processes and mode of life of the Aphides, and that 
at.the present day the state of general information as to the natural history of these 
singular creatures is in many respects rather behind, than in advance of, that of the past 


generation. ie : 
Bonnet's wonderfully patient and laborious researchest proved, beyond all doubt, 
exual influence; 2ndly, that the 


Ist, that the viviparous Aphis may propagate without s i 
brood thus produced nray give rise to young in the same way ; that these may repeat this 
asexual prolification; and so on for as many as ten broods; 3rdly, that the viviparous 
Aphides and their brood may be either winged or apterous; 4thly, that, under snp 
conditions, winged or wingless males appear and copulate with ae females, which, 
in thé instances observed by Bonnet, were wingless. ME j b ü 
These are the statements put forth by Bonnet on the evidence of direct observation 


à i * ité d'Insectologie, 1745. 
* Duvau, Mém. du Muséum, xiii. 1825. $ Fre d = ogie, 


= 


194 PROF. HUXLEY ON THE AGAMIC REPRODUCTION 


and experiment, and they have been confirmed by every subsequent original observer 
whose works I have perused. Besides these matters of fact, Bonnet states, as his strong 
opinion, that there is no fixed limit to the process of agamic, viviparous reproduction, and 
that, under favourable conditions of warmth and nourishment, it might be continued for 
* thirty generations" (Z. c. p. 102), or, in other words, indefinitely. 

The accurate and pains-taking Degeer, who gives an elaborate account of some seven- 
teen species of Aphis, affirms as the result of his researches, “that the winged Aphides 
are never oviparous*." He describes at length the apterous males of certain species 
(P. lisse du Pin, P. du Pommier, P. du Genévrier), and shows that apterous, oviparous, 
and winged viviparous broods may coexist, as in Aphis Rose. 

Degeer considers that, as a general rule, the oviparous females and the males are pro- 
duced by alate viviparous females. 

The next important original memoir on the Aphides is that published in Germar's 
Magazin der Entomologie for 1815, by Kybert, evidently a most careful observer, but 
somewhat wanting in method and clearness as a writer. Kyber is in perfect accordance 
with Bonnet and Degeer; and more than this, he experimentally proved the justice of 
Bonnet's supposition, that the duration of the agamie reproductive power is practically 
indefinite, and is chiefly, if not wholly, dependent on conditions of temperature and 
- nutrition. He says (p. 34) :— 

*I never saw a male in copulation with a winged female in any species. It was 
always the apterous females which were attacked by the males; for in many species apte- | 
rous females remain among the families. Neither have I ever seen winged females lay 
eggs. This has, indeed, been already remarked by Degeer.” 
_ Ina note Kyber adds the caution, that he has not observed more than twenty species 

in copulation, and does not wish to extend his conclusions beyond these. 

The fourth note to this important paper contains the following remarkable observation : 
—......“ The winged females especially, in which, even after frost has set in, fully-formed 
young may always be found, when the apterous females of the same family have long 
been laying eggs. On the 21st November, 1812, I still had winged Aphides (Haberblatt- 
låuse) in my possession, although the apterous ones had copulated and laid their eggs in 
September, —a remarkable circumstance without doubt, and one whence important con- - 
clusions with regard to the mode of propagation of the Aphides are likely to flow. Pos- 
sibly, many winged females survive the winter together with their young.” (p. 10.) 

In other parts of his memoir (p. 2 et seq.), Kyber adduces strong evidence in favour of 
the hybernation of the viviparous forms of some species; which Degeer had already 
proved to be the case with respect to the remarkable * Puceron des Galles du Sapin.” 

5 the Aphis Dianthi, Kyber was never able to observe either copulation or oviposition ; 
a nn being any natural term to the number of asexual broods which. 
an ie = T that he raised viviparous broods of both this species and 

: ecutive years, without any intervention of males or oviparous 


* Degeer, Mém. sur les Insectes, 1774, vol. iii. p. 74. 
t v Erfahrungen und Bemerkungen über Blattliuse von J. F. Kyber, Diacon. in Eisenberg. 


AND MORPHOLOGY OF APHIS. 195 


females, and that the energy of the power of agamic reproduction was at the end of that 
period undiminished. The rapidity of the agamic prolification throughout the whole 
period was directly proportional to the amount of warmth and food supplied. 

Duvau, in his already cited “Nouvelles Recherches sur l'histoire naturelle des Pucerons,” 
read before the French Academy of Sciences in 1825, states that he had carried the series of 
successive agamic generations in the Aphis of the Bean (fève) to eleven, which was one 
more than Bonnet had obtained. The process lasted seven months, and the last young 
was born on the 27th December, but died on the 29th. Duvau, however, kept some alive 
until January, and naturally asks whether it is not probable that, under favourable cir- 
cumstances, the agamic process may be continued throughout the winter. The average 
length of life of his Aphides was thirty days, or a little more; but the representative of 
the ninth generation lived from September 29th to December 19th, or eighty-one days. 
Like those of preceding observers, Duvau's researches clearly show the influence of tem- 
perature on the fecundity of the viviparous Aphis. 

It is in Morren's in many respects valuable paper on the Aphis Persice, published 
in the ‘Annales des Sciences Naturelles’ for 1836, that the germs of the two most 
notable errors which have crept into the natural history of the Aphides may be found. 
At p. 76 the following passage occurs* :— 

“The influence of temperature on these animals is obvious; in other Aphides, and 
under ordinary circumstances, the female lays her eggs when she has wings and after 
copulation with the male, who is winged at the same epoch. Oviposition takes place in 
this manner at the seventh generation for some—at the ninth or even at the eleventh for 
_ others; before it, female larvæ alone are produced.” 

. Morren here supposes himself to be simply repeating what he has read. But so far 

as I am acquainted with the older literature of the Aphides, he is entirely mistaken. I 
can nowhere discover that either Réaumur, Bonnet, Degeert, Kyber, or Duvau have 
. Observed winged oviparous females in any species; nor do the statements of any of these 
observers justify the belief that the sexual forms always appear after a certain number of 
generations. All that Bonnet affirms is, that his particular experiments came to an end 
accidentally after the production of a certain number of agamic generations, which is, of 
course, quite another matter. ‘ 

- When Morren details his own observations, his results are in exact accordance with 
those of the older observers. “In the Aphis Persice,” says he, “ I have very frequently 
seen (and I have shown the phenomenon to my colleague, M. Burgraeve) that the winged 
and fertilizable female never contained ova and never laid any, but that she contained 
little living Aphides, which are born fully developed, and provided with legs, proboscis, and 


* « L'influence de la température sur ces animaux est manifeste ; chez les autres tage et — V. rer Y 
ordinaires, la femelle pond des œufs lorsqu’elle est ailée, et après un accouplement e le mále nde t ^tm à 
Cette ponte se fait ainsi à la septième génération pour les uns, å la neuvième ou même å la onzième pour les i 
avant elle, il y a seulement naissance de femelles naissant à l'état de larves."—Morren, l. å Gd T belies 

T Degeer's account of the gall-forming Puceron du Pin is an en — m - - ard a: daté m 
only an apparent one. Degeer expressly states that he never saw the winged form of this species in cop ER 

besides, it is not a true Aphis at all. , 4p 

VOL, XXIF 


196 PROF. HUXLEY ON THE AGAMIC REPRODUCTION 


antennæ. It was only in November that the apterous females presented eggs in their 
ovaries and oviducts, and for that effect, a considerable degree of cold was necessary*.” 

Morren describes the male, female, and agamic organs of reproduction, but less com- 
pletely than Von Siebold, who, in 18397, carefully investigated the Aphis Lonicere, and — 
first demonstrated the existence of the spermatheca and colleterial glands in the oviparous 
females. Von Siebold distinguishes three forms of this species, two winged and one 
apterous. The large winged Aphides were all viviparous; the smaller, males. The 
apterous forms were oviparous, and the progeny of the alate females. 

Steenstrup says of the Aphides (Alternation of Generations, p. 108), * The propagation 
of these creatures through a series of generations. has been already long known, In the 
spring, for instance, a generation is produced from the ova, which grows and is metamor- 
phosed, and without previous fertilization gives birth to a new generation, and this again 
to a third, and so on for ten or twelve weeks; so that in certain species even as many as 
nine such preliminary generations will have been observed; but at last there always 
occurs a generation consisting of males and females, the former of which after their meta- 
morphosis are usually winged; fertilization and the depositing of eggs take place, and 
the long series of generations recommences in the next year and in the same order.” 

In the first edition of Professor Owen’s ‘ Lectures on the Invertebrata,’ published in 
1843, however, Morren’s errors are adopted, extended, and enunciated as the law of pro- 
pagation of the Aphides, in the following terms :— 

* In the last generation, which is the seventh, the ninth, or the eleventh, according 
to the species of Aphis, the fertilizing influence would seem to have expired}, and deve- 
lopmental force exhausts itself in more frequent and numerous moultings, in the for- 
mation of wings, and in the modification of the female organs already described. Many 
males, which, like the females, acquire wings, form part of the produce of the last brood, 
which takes place in autumn. They rise in the air, frequently migrate in incalculable 
numbers, unite, and the females then produce eggs, which are glued to twigs and leaf- 
stalks, retain their vitality throughout the winter, are hatched in the spring, and give 
birth to the apterous and larviparous females, which continue to produce successive gene- 
rations of similar females until the close of summer.” (p. 235.) d 

It has not been my good fortune to discover, either in Prof. Owen's writings or those of 
his predecessors, any evidence in support of the singular statement contained in the last 
paragraph of this citation, which is incorrect in all important respects, and has, indeed, 
been omitted in the second edition of the * Lectures.’ 

Mr. Walker, in the first of his long and valuable series of papers on the Aphides 
(Annals, vol. i. 1848, p. 259), writes thus :— 


* “Or chez le puceron du pêcher j'ai vu un grand nombre de fois, et j'ai montré le phénomène à mon collègue, 
M. Burgraeve, que la femelle ailée et propre à la fécondation ne renfermait point des œufs et n'en pondait point, 
mais qu'elle renfermait des petits pucerons vivants qui naissent tout développés avec leurs pattes, leur trompe, et leurs 
antennes. Ce ne fut qu'en Novembre que les femelles sans ailes présentaient des ceufs dans les ovaries et les oviductes, 
et pour cela il fallait un froid déjà assez vif.”—Morren, l. c. p. 76. - 

$ Ueber die inneren Geschlechtswerkzeuge der viviparen und oviparen Blattlåuse. Froriep’s Neue Notizen, 1839. 

1 This phrase is little more than a translation of a passage in Morren which will be given below. 


AND MORPHOLOGY OF APHIS. 197 


* T am indebted to my friend Mr. Haliday for the following translation of an extract 
from Erichson’s Bericht, &c., 184.4, Ent. Zeitung, pp. 9, 81, 133, 410. Ratzeburg observed 
a species of Aphis on the Birch, which continued to produce a living progeny from 
August into winter without either male or female appearing. Bouché and Kaltenbach, 
in explanation, remark that the males in this family are not always winged. However, 
in the May following, Ratzeburg, continuing his observations, found the winged females, 
and afterwards (in October) winged males also, which paired with them. The species was 
then identified as 4. oblonga, Von Heyden. For the male to pair with a winged female 
(continues Mr. Walker) is a very unusual case among Aphides*.” In fact, I have hitherto 
found, in Mr. Walker’s long list of 101 species, no case of an oviparous winged female 
observed by himself. Mr. Walker states as a known fact, that Aphis Rose habitually 
lives through our mild winters. 

In his work on ‘ Parthenogenesis’ (1849), Prof. Owen modifies his previous statement 
so far as to say, in a note (p. 59), that the perfecting of the female generative organs in 
Aphis “is not attended by the acquisition of wings; or if they be developed in the ovipa- 
rous female, they soon fall. I have, however, retained them in the diagram for a better 
illustration of the analogy. Many of the virgin viviparous Aphides acquire wings, but 
never perfect the generative organs.” 

The diagram referred to exhibits two figures, (b) and (i), which, for anything that 
appears in the text, might be taken to be the author's representation of male and female 
Aphides. On comparing them with the illustrations of Morren's memoir, however, it is 
at once obvious that they are copies of his figures 1 and 2, of which fig. 2 does really 
represent a male; while fig. 1, on the other hand, is not an oviparous, but a viviparous 
female. In the explanation of his figures, Morren indeed merely says of fig. 1, “ Femelle 
vue en dessous ;”” but it requires no great amount of attention to his text to observe his 
distinet statement (already quoted), that the winged female is viviparous, and not ovipa- 
rous. I am obliged to be thus particular in explaining these unusual circumstances, as 
otherwise the existence of a typical figure of a winged oviparous female Aphis, in the 
work of an accredited author, might be brought forward as conclusive evidence of the 


ordinary occurrence of such femalest. 


* On turning to Ratzeburg's notice in the * Entomologische Zeitung, 1844, p. 410 (Fortgesetzte Beobachtungen 
über die Copula der Blattläuse), which is the last word of the correspondence between Kaltenbach, pons and him- 
self on this subject, I find his precise words to be these :—'* Wie gross war daher mein Erstaunen, als ich bei meiner 
ersten, nach der Rückkehr angestellten Excursion, am 22 October gleich auf den ersten Blick unter der Menge von 
ungeflügelten Individuen, welche die des vorigen Jahres bei weitem übertraf, auch geflügelte Puppen und geflügelte 
Männchen bemerkte, und wie gross war meine Freude, auch gleich darauf mehrere der letztern in der Begattung zu 
finden, also in einem Acte, den ich bei Blattläusen selbst noch nicht hätte beobachten können.” Subsequently, Ratze- 
burg states that he was able to observe the copulatory process early and late, at any time between the 22nd October 
and the 16th November. - å 

It will be observed that there is not a word here about such winged females as Ratzeburg, in å —— 
states he saw in May of the same year. The winged pupæ are apparently, from the context, the nn. Bi " em 
and the forms with which the winged males copulated were the wingless females. So that here, as = : å " 3 sd 
posed cases of winged, oviparous true Aphides I have looked into, the evidence, when closely examined, bre er i 

t Professor Owen, in the last edition of his * Lectures on the Invertebrata,’ p. 410, quotes Léon Dufour as having 


. » i a > , i A les 
witnessed the coitus of the male Aphis * with the winged female. The reference is to ger så in Anna 


198 PROF. HUXLEY ON THE AGAMIC REPRODUCTION 


When the natural history of the Aphides is freed from the mythical additions which 
have accumulated around, and obscured it, I believe the following propositions may be 
said to be established on good evidence :— 

1. Ova deposited by impregnated female Aphides in autumn are hatched in the spring. 

2. From these ova, viviparous, and in the great majority of cases apterous, forms 
proceed. 

3. The broods to which these give rise are either winged or apterous, or both. - 

4. The number of successive broods has no certain limit, but is, so far as we know at 
present, controlled only by temperature and the supply of food. 

5. On the setting in of cold weather, or in some cases on the failure of nourishment*, 
the weather being still warm, males and oviparous females are produced. 

6. The males may be either winged or apterous. 

7. So far as I am aware, there is no proof of the existence of any exception to the law 


that the oviparous female is apterous. 

8. Viviparous Aphides may hybernate, and may co-exist with oviparous females of the 
same species. 

So much by way of clearing the ground. I now proceed to the particular subject of 
this paper, which is primarily, to describe the nature of the process by which the agamic 
young arises within the body of its viviparous parent. But very few investigators have 
applied themselves to this question, and those who have are unfortunately in diametrieal 

contradiction to one another as to the most important points. 

Prof. Leydig published a notice on this subject in the ‘Isis’ for 1848, which I have 
not seen ; but subsequently his views, fully stated and accompanied by figures, were pro- 
mulgated in Siebold and Kölliker's Zeitschrift for 1850, vol. ii. Heft 1. He maintains 
“that the germ of the (viviparous) Aphis is developed out of cells, and its embryo is as 
much composed of cells as one which has proceeded from a fecundated ovum” (1. c. p. 65). 
And he particularly details the manner in which one of the large cells contained in the 
terminal chamber of the proliferous organ of the viviparous Aphis becomes detached, 
enlarges, and is converted into the embryo. Although Leydig does not absolutely say as 
much, his observations lead to the conclusion that there is no histological difference be- 
tween the agamic germ in its youngest state, and a true ovum at a corresponding period. 

Von Siebold implies, and Prof. Owen, Victor Carus, and the late Dr. Waldo Burnett 
assert, with different degrees of distinctness, on the contrary, that there is a clear histolo- 


d i i LE] y 
me ten ee I have carefully, and more than once, scrutinized this volume of the ‘Annales, 
vel of RER = to discover the passage referred to. Léon Dufour has, in fact, two memoirs in the first 

of the ‘Annales’ for 1844. The first is on the « Anatomie générale des Diptères ;” the second, “ Histoire 


des Métamorphoses et de I Anatomie du Pioph a ei xpected there 18 no 
il P t 
asionis, As might be € » 


r ær Royale des Sciences, 1703), however, and I find only this: 
m É d Hi * a em 
Bets vis es ke que les pucerons vivent une année entière, et que pendant l'hiver ils se retirent dans des trous, 
== nt au printems pour pondre leurs œufs, comme le font les mouches ordinaires.” 
Hausmann's “ Beiträge” in Illiger's Magazin, Bd. 2. 


AND MORPHOLOGY OF APHIS. 199 


gical difference between the primary germs of the viviparous Aphis and true ova,—Carus 
and Burnett reiterating their opinions even since the publication of Leydig’s views. 
Finally, Mr. Lubbock, in his late valuable memoir on Daphnia, (Phil. Trans. 1857) has 
expressed his inability to find any germinal vesicle in the germs of the viviparous Aphis, 
and, so far, may be ranked among Leydig’s opponents. 

I have recently resumed some investigations commenced two or three years ago on 
this interesting subject. My object was originally purely morphological,—the Aphis sug- 
gesting itself as a very convenient subject for working out the general development of 
Insecta ; but I have found myself unable to. refrain from wandering out of my direct 
course, and attempting to further the solution of the great problem of Agamogenesis, or 
asexual reproduction. 

My observations are in the main in accordance with those of Leydig. On many 
minor points, however, we are at variance; and besides this, there are matters of great 
interest, upon which Leydig does not touch, but on which I hope to be able to throw some 
light. For, besides yielding an answer to the question as to the existence or absence of 
any histological distinction between a bud and an ovum, the investigation of the vivi- 
parous and oviparous Aphides affords decisive evidence as to the soundness of certain 
explanations of the phenomena of Agamogenesis in general; and finally, the study of 
the general development of Aphis furnishes data of great importance in Articulate Mor- 
phology. 

I propose in the present memoir to follow out these lines of inquiry. I will in the first 
place describe the minute structure of the essential reproductive organs or * Pseudovaria” 
of the viviparous or agamic female; and the development of its germs or pseudova (as I 
propose to term them) will be considered. Secondly, the reproductive organs of the ovipa- 
rous female and the development of the ova will be described. Next, I shall speak of the 
manner in which the proliferous apparatus or pseudovarium of the viviparous female is 
developed within the germ; and I shall compare together the agamic and sexual repro- 
ductive processes. I shall then endeavour by means of these facts to refute a hypothesis 
' which has been offered in explanation of Agamogenesis ; and finally, I propose to consider 
the Morphology of the Articulata so far as it is elucidated by Development. 

The species of Aphis, the reproductive organs of whose viviparous form I am about to 
describe, appeared this autumn upon a plant of the Ivy-leafed Geranium which hangs 
in my study, and for the last two months has been regularly giving rise to broods, some- 
times winged and sometimes apterous, without any appearance of males or females. — 
With respect to the external characters of the reproductive organs, I have nothing of 
importance to add to Siebold’s or Morren’s description. 


§ 2. The Development of the Pseudovum. 


The terminal chamber of any of the cæca of the pseudovarium is a rounded or oval body 
| (P.XXXVI. fig. 1, A), united by a delicate ligament (a), proceeding from its free end, with 
the ligaments which pass from the other cæca of the same side, to form the common pseud- 
ovarian ligament. Thewall of the chamber is a delicate transparent membrane (b), in which, 
here and there, rounded endoplasts (or nuclei) are imbedded ; while others lie on its inner 


200 PROF. HUXLEY ON THE AGAMIC REPRODUCTION 


nti itheli continuous with the contents of the cham- 
e mere = pret se by a homogeneous pale periplastie 
peng å ed a doves clear spheroidal cavities (e) whose walls are 
eene e» a of the periplast. The cavities have on an average a dia- 
es ang an ies In the centre of each is a rounded opake body (/) like 
gas om aA of the wall of the dilatation, and, indeed, obviously of the same — 
i fluid I have examined this tíssue, it began after a time to alter. In the 
very weak syrup which I ordinarily employed, the change consisted partly in Wm 
increased definition of the walls of the clear cavity, but more particularly in the € 
up of the periplast into spheroidal masses, each of which contained a single ees e Y 
its endoplast*. The resemblance of such a body to an ovum with its germin vesi på 
and spot is complete; nor would it be possible for any one ignorant of the origin 
the body to say that it was other than an ovum. Water instantly alters the appa 
of the tissue, completely destroying its distinctive character. Dilute glycerine shrivels 
up the vesicles and alters the appearance of their central endoplast, probably by endos- 
mose. Acetic acid renders the periplast dark, and gives an exceedingly marked definition 
to the parietes of the vesicle. To see the appearances I have described as normal, the part 
must be examined perfectly fresh, and in a solution of sugar neither too dilute nor too 
concentrated. 
In certain specimens the contents of the lower part of the terminal chamber are dif- 
ferent from those of the upper. As much as a third of the whole chamber may be bests: 
pied by a mass of periplast containing only a single clear vesicle. Such 6 CORE is 
figured in fig. 1, Pl. XXXVI. Fig. 2 exhibits a further advance in the same direction; thie 
mass, which, from its close resemblance to a true ovum, I have called a pseudovum, having 
enlarged so much as nearly to equal the contents of the terminal chamber, from which it is 
distinguished by a slight constriction. In figs. 3 and 4, the constriction has become — 
marked, until at length à penultimate chamber is formed, connected only by a narrow neck 
with the terminal one, fig. 4. It is on an average about s5sth of an inch in diameter. The 
epithelial layer (c) of its wall is ordinarily well developed, and when water is added swells 
up, So as to separate the periplastic substance of the pseudovum from the wall. The pert 
plast itself exhibited no structure, and appeared unchanged except in size. The e 
vesicle was sometimes unchanged, sometimes enlarged, but otherwise unaltered. Of its 
endoplast I was sometimes unable to diseover any trace; on other occasions I found a 
few granules in its place (fig. 3); and, once, two particles, each rather more than half - 
diameter, appeared to lie side by side in the interior of the vesicle. : 
The marked contrast between the perfect distinctness of the endoplast in the vesicles 
contained in the ultimate pseudovarian chamber, and its apparent absence in the ye 
similar vesicle of the mass contained in the penultimate chamber, or in the lower part 0 


the last one, was the more striking, as the two could be readily compared under the same UE 
circumstances and in the same field of view. 


å li dig (1: e. p. 63) appears to regard this as the first state of the ovigerms, and he has overlooked ME 
epithelium, å 


AND MORPHOLOGY OF APHIS. 201 


Finally, the vesicle itself ceases to be visible (fig. 4), and the penultimate chamber 
contains only its epithelium and a mass of apparently structureless substance ;—I say 
apparently structureless, because the addition of water made the mass more clear, and at 
the same time rendered an irregular areolation and scattered granules visible in its sub- 
stance. Whether the areolæ are the outlines of delicate vesicles, and the granules their 
endoplasts, or not, are points which I could not satisfactorily determine; at any rate, I 
could never observe anything like the regular structure observable in the contents of this 
chamber when a little larger. 

Fig. 5 represents such a chamber, zrrth of an inch in length. The endoplasts of 
the wall are seen lying in or upon it, and oceupying its interior is a distinct oval mass of 
substance agreeing in appearance with the periplast of the pseudovum, but distinguished 
from it by containing a great number of elear spheroidal cavities not more than s3loth 
of an inch in diameter, each of which contains a central endoplast of not more than 
Tooooth of an inch. These cavities are closely packed, but not flattened against one 
another. The walls of the cavities react differently on the addition of acetic acid to the 
rest of the periplast, becoming darker and more sharply defined. In fact, each cavity 
is what is commonly termed a nucleated cell, while the intervening periplast is the so- 
called intercellular substance. 

I have here stated merely the histological faets which may be observed by any one 
who will take the trouble to examine with sufficient care the ultimate and penultimate 
pseudovarial chambers of a few viviparous.4phides. Of the existence of these states, 
and that the order in which I have detailed them fairly represents the order in which 
they succeed one another in nature, I have no doubt; and I therefore look upon it as an 
established fact, that the primary steps in the agamic development of Aphis are, first, the 
enlargement of the periplast around one of the pseudovarian vesicles, and its detachment 
ås å separate body, which, from its resemblance to an ovum, I will call a “ pseudovum;” 
secondly, the contemporaneous formation of a distinct chamber—the penultimate cham- 
ber of the pseudovarium ; thirdly, the disappearance of the vesicle of the pseudovum, and 
the conversion of the latter into a germ-mass composed of cells imbedded in intercellular 
Substance and containing minute endoplasts. 

I should be sorry, however, to express an opinion as to the exact nature of the process 
by which these changes are effected, with anything like the same degree of confidence. 
Three hypotheses present themselves :— € 

Ist. The pseudoval endoplast divides and subdivides, so as to give rise to the endoplasts 
of the germ ; or. 

2nd. The pseudoval endoplast is resolved, and the endoplasts of the germ are developed 
autogenously in its periplast ; or— : 

9rd. The pseudoval endoplast disappears, and the endoplasts of the germ are supplied 
from the epithelium of the walls of the pseudovarial chamber. : 

Of these three hypotheses, I strongly incline towards the first, as most in accordance 
With what we know of histological development in general. The whole progress of 
modern research, in fact, goes to show that cells and endoplasts hardly, if ever, arise 
autogenously, but are the result of the subdivision of pre-existing cells and endoplasts. If 


202 PROF. HUXLEY ON THE AGAMIC REPRODUCTION 


this be the case, however, the second hypothesis is excluded, and the third is improbable 
in itself, and is supported by no evidence. In the absence of such evidence, the marked 
contrast in size and appearance between the epithelial endoplasts of the penultimate 
chamber and those of the germ tends to show that the two have no direct relation to one 
another. ! 

'Those who have followed the details of the development of the pseudovum and its 
resulting germ, given above, will not fail to admire the clear insight of | Morren, when he 
affirmed that the agamic offspring of Aphis was developed by “the individualization of a 
previously organized tissue.” A more neat and expressive definition of the process could 
not be given : and as Morren nowhere entertains the absurd doctrine that an organized 
tissue must be as complex as *mucous membrane” or * muscular fibre," which has 
been attributed to him, the criticisms to which his views have been subjected on this 
ground are sufficiently baseless. No one will pretend to deny that the pseudovarium is 
“ organized," nor that the pseudovum is a portion of it which has become “individualized.” 
. But I subjoin Morren's words, that the reader may form his own judgment as to his 
merits :— 

“A dire vrai, je me refuse à émettre une opinion au milieu d'un tel dédale, et je tiens 
pour plus philosophique d'avouer son ignorance dans un phénoméne oå la nature nous 
refuse méme l'apparence d'une explication. S'il fallait une explication à toute force, 
j'admettrais que la génération se fait ici comme chez quelques entozoaires, par individuali- 
sation d'un tissu précédemment organisé. La génération n'est pas pour cela spontanée: 
une génération spontanée doit étre la production d'un étre organisé de toutes pièces, | 
lorsque les élémens inorganiques se réuniront pour produire un animal, une plante. 
Cette génération est impossible et n'a jamais lieu. Une génération équivoque est celle oà 
des tissus organisés préalablement par un étre déjà pourvu de vie, s'individualisent, c'est à 
dire, se séparent de la masse commune et participent encore, aprés cette séparation de 
l'état dynamique de la masse, c'est à dire, de sa vie, mais à son propre profit. C'est 
ainsi qu'un tissu produit un entozoaire*. C’est de la vie continuée. Mais supposez 
que la vie ait assez d'énergie pour imprimer au tissu que s'individualise la forme de 
l'espèce productrice, et vous avez la génération des pucerons. Cette énergie se perd 
au bout de quelques générations, et une nouvelle impulsion devient nécessaire, c'est celle 
du mále. ; 

* Voilà à tout hasard, une hypothèse que dans ma jeunesse j'aurais embrassée avec 
plaisir; mais aujourd'hui je préfère douter: les faits que j'ai exposés plus haut valent 


mieux qu'une théorie.”—Morren, sur le Puceron du Pécher, Annales des Sc. Nat. série 2. 
vi. 1836, p. 90. 


$3. Description of the Oviparous Female Aphis and of the Development of 
the Ovum. | 


, Throughout the two months during which the Ivy-leafed Geranium, on which gie 1 
viviparous Aphides are living, has been in my possession, neither males nor females have : 


* I need hardly remark, that no evid 


ne | 
: ence of the development of Entozoa, in the way supposed by Morren, 


AND MORPHOLOGY OF APHIS. « 203 


made their appearance. Therefore, being extremely desirous to compare the process of 
the development of the germ'with that of the ovum, before completing this paper, I began 
in the last days of October to seek for oviparous females of some other species, 

An Oak-tree in the Zoological Gardens at length supplied me with that which I sought. 
The small twigs and leaves afforded habitation to a number of minute wingless Aphides, 
all so nearly equal in size, that I did not doubt their non-viviparous, and hence in all 
probability their oviparous character. 

Microscopic examination fully confirmed my suspicions; for not only were the Aphides 
full of ova, but I found multitudes of similar ova adhering to the plant in the axils of the 
leaves, and more particularly between the outer bracts of the buds*. 

These Aphides were very different from my viviparous species. They were about th 
of an inch in length. The general hue of the body was pale green ; but it was diversified 
in the dorsal region by four longitudinal rows of blackish rounded spots, one spot in each 
row being seated on the tergum of most of the somites, from the prothorax backwards. 
Hence, there were nearly as many transverse rows of four spots each, as segments of the 
body. The two median spots in each row were larger, and situated close to the middle line. 
The external spots were more upon the sides of the body. The spots upon the mesothorax, 
and thence to the sixth abdominal somite inclusive, were the largest and most conspicuous. 
Each spot was constituted by a dark elevation of the integument, which supported a 
tuft of long setæ, knobbed at their extremities like the glandular hairs of certain plants. 
The hairs were not confined to these localities, however, but were scattered over the head 
and other parts of the body. The eyes were red, and produced into a small tubercle on 
their posterior margins. The distal portions of the antennæ, and the tarsi, were blackish. 
The antennæ were not more than equal to half the body in length ; they were seven-jointed, 
the penultimate joint being somewhat swollen at its extremity. Both this and the pre- 
ceding and following joints were so sculptured as to appear, at first, minutely annulated. 
The basal joint was the thickest of all, the second less thick, but stronger than the others. 
` The proximal half of the antennæ was sparsely setose. The promuscis was short, extend- 
ing, when deflexed, no further than the posterior edge of the prothoracie sternum. The 
abdomen tapered into a cone beyond its sixth somite, on whose dorso-lateral region the 
very short trumpet-mouthed siphons were situated. The abdomen was terminated by 
two subcylindrical rounded setose tubercles, of which the lower was the larger. They 
had the anus between them, and acted as anal valves. The posterior limbs, when fully 
extended, hardly reached beyond the end of the abdomen. 

The eggs when first laid are of a dark green hue and very soft; afterwards they ap- 
pear to become black. : 

The vulva of the oviparous Aphis (B) opens between the eighth and ninth abdominal 
sterna, the eighth (8) being a little prolouged, so as to form a sort of inferior lip to the 
vaginal aperture (Pl. XL. fig. 1). The vagina (C) is a thick-walled tube provided with a 


* I do not think that my Aphis of the Oak is identical either with that described by Réaumur, or that ee 
by Bonnet. None of my specimens attained the size of theirs, nor do either of those vem mention the peculiar 
dorsal markings: of my species: furthermore, the proboscis in both Réaumur's and — wem long ; dl. it 18 
very short, The proper specific names of both the Aphides alluded to in this memoir will be discussed in a note. 


YOD: XXII, dim 


a^ 


204 PROF. HUXLEY ON THE AGAMIC REPRODUCTION 
layer of external transverse, and internal longitudinal, striated muscles. After entering 
the sixth abdominal somite, it divides into two branches—the oviducts (DD), whose walls 
exhibit the same muscularity, but are less thick. Both vagina and oviducts are lined by 
a well-developed epithelium. 

The oviducts divide into four ovarian cæca, whose delicate structureless wall is un- 
provided with muscles, and lined by a columnar epithelium. Each cæcum is ordinarily 
divided by constrictions into six chambers. Of these I found the posterior (that nearest 
the vulva) (E) always empty, and of nearly the same length, though of a much smaller 
diameter than that which precedes it, or the fifth from the apex of the ovarium. This 
fifth chamber (F) always contained a fully formed ovum, provided with a chorion and 
an opake coarsely granular yelk. 

The fourth chamber (G) is smaller than the fifth ; it contains a coarsely granular vitelline 
mass in which no germinal vesicle can be perceived, and which ordinarily has no invest- 
ing membrane. 

The third chamber (H) is still smaller; and its contents are usually only slightly gra- 
nular, so that the germinal vesicle and spot of the ovum in this chamber are beautifully 
distinct (fig. 2). 

The second chamber (I) is the smallest of all; the germinal vesicle and spot of its rudi- 
mentary ovum can be easily seen; and but very few fine granules are deposited in the 
substance which will eventually form the yelk. 

A clear cord-like mass (q), commonly divided longitudinally, so as to appear double, 
traverses this chamber, and can be traced into the next. 

The apical chamber (K) is as large as the third, but is longer transversely than longitudi- 
nally, while the reverse is the case with the third chamber. Its outer wall is formed by 
a continuation of the same structureless membrane as that which constitutes the rest of 
the cecum. The epithelium (p), which is particularly thick in the upper part of the second 
chamber, especially at the neck or constriction between the first and second, is suddenly 
attenuated as it spreads on the inner face of the wall of this chamber, and becomes very 
thin from the flattening of its cells. From having the characters of a cylinder-, it takes 
those of a pavement-epithelium. 

It is at first extremely difficult to understand the nature of the contents of the apical 
chamber. All its anterior part appears to be filled with about a dozen closely appressed 
bodies (2), which, if examined without due attention, or under a low power only, may easily 
be confounded with ova. Each of these bodies has a sort of wedge shape, such as would 
result from the compression of rounded masses in a spherical envelope which they nearly 
fill. Its apex is turned inwards ; its base outwards. Each consists of a thick transpa- 
rent outer coat closely investing a denser and well-defined membranous sac. The latter 
contains a clear substance, in which many irregular granules are imbedded. The lines of 
separation between the appressed sides of these bodies are well seen, either in a sectional 
sr view. In the latter case, they appear as polygonal meshes; in the 
si tha gas BE epee aga bodies from one another, and bounding their curved d 
rhe dede rå epithelium. On tracing the lines of separation towards the cen 

e ends of these bodies, they become lost, and a mere clear, homogeneous 


AND MORPHOLOGY OF APHIS. 205 


substance seems to occupy the whole central part of the chamber ; but on carrying the 
eye backwards, this clear mass is seen to be continuous with the two cords which I have 
above described as entering the second chamber (PL XL. fig. 3). 

The histological constitution of these bodies is at once sufficient to convince the observer 
that they are not ova, and I regard them as glandular masses which secrete the matter 
of the clear cord-like bodies which descend into the second and third chamber. 

The ova themselves, or rather the rudiments of the future ova, are not always to be 
seen with ease; and if the epithelium of the lower part of the apical chamber has become 
much altered, they cannot be detected: for they are visible exclusively in this part of 
the chamber, of whose epithelial cells they are, as I believe, merely a modification. 
However this may be, germinal vesicles and spots of all sizes intermediate between that 
of the ovum of the second chamber and that of an ordinary epithelial cell are seen in 
close contact with the parietes of the chamber. I have detected as many as six in this 
position. When the chamber is subjected to compression they may be set free, and are 
then seen to be surrounded by a zone of clear substance, the rudimentary vitellus. 
Under similar circumstances, the “glandular bodies” may also be isolated; when they 
present themselves as vesicles surrounded by a clear homogeneous substance, which is 
frequently prolonged at their apical extremity. It is gradually dissipated, and the inner 
sac set free by the action of water. 

I have not seen any ovarian ligament in the oviparous Aphis. 

The structure which I have described was wholly unexpected and new to me; and I am 
not aware that anything similar has yet been noticed in the ovaria of Insects*. I am 
inclined to believe that the glandular bodies contribute directly to the formation of the 
vitellus, because I have more than once seen cases, like that figured in Pl. XL. fig. 3, 
where the clear cord-like body appeared to pass directly into the mass of the ovum. 
There was always a widely open communication between the first and second, and 
between the second and third chamber; but the passage between the third and fourth 
was closed by the meeting of the epithelial lining. Does each ovum, as it is given off 
from the ovary, and passes backwards, carry with it a gelatinous mass, the product of one 
half of the glandular bodies, and only cease to be connected with these glands when it 
has taken the third place? 

Three cæca open into the dorsal side of the lower part of the vagina; of these the 
anterior single one is the spermatheca, the posterior pair are the colleterial glands 
(PL XL. fig. 1, m, n). r : : 

The spermatheca (x) is a sac with a narrow neck, dilated at its extremity, which opens 
considerably in advance of the colleterial glands, while its enlarged end lies between 
them. The duct of the spermatheca has thick walls continuous with those of the vagina; 
but its dilated portion is thin, and has a yellowish colour. It contains a multitude of 
large filiform spermatozoa bent upon themselves, and is very tough and resisting. 

The colleterial glands (m) are subeylindrical, but are constricted inferiorly where they 


* Unless, as I am strongly inclined to suspect from Leydig’s description, and from a bar gm — ‘pad E 
part of the ovaria of Coccus, the corresponding chamber of that insect’s remarkable ovaria presents a similar ure, 


See, however, the note which concludes this paper.) 252 


206 — PROF. HUXLEY ON THE AGAMIC REPRODUCTION 


open close to the vulva. They consist of a delicate etmuerarciess coat lined by a thick 
layer of granular substance, whose cellular composition is very indistinct in the fresh 
state, but becomes obvious on the addition of acetic acid. 

‘The interior of the gland contains a clear, viscid, strongly-refracting substance, appa- 
rently separated from the epithelial lining by a membranous layer. I am in doubt, 
however, whether this apparent membrane be anything more than the folded and wrinkled 
outer layer of the viscid matter. When the Aphis is suddenly placed in glycerine or 
subjected to slight pressure, a drop of the colleterial secretion not unfrequently exudes 
and manifests its viscidity by leaving a long trail. 

The fully-formed ovum (Pl. XL. fig. 1, F) measures about th of an inch in length. It 
is oval, rather smaller anteriorly, and of a deep green hue, in consequence of the colour of 
the yelk. The chorion is a tough transparent membrane, about 5555th of an inch thick, 
and presents no external sculpturing or internal structure. Internal to the chorion is 
a delicate vitelline membrane which immediately invests the yelk. It is, however, con- 
nected with the chorion posteriorly. When the egg is heated with caustic potass, the 
yelk is driven away from the sides (eventually dissolving), and with it the vitelline 
membrane on the sides and at the anterior part of the ovum; posteriorly, however, I 
always found it adherent. The yelk itself is very coarsely granular ; so that there would 
be no chance of discovering the germinal vesicle, even if it existed. 

The recent observations of Leuckart and Meissner on the micropyle of the ovum in 
Insects naturally induced me to look for such a structure in the egg of Aphis. 

Leuckart, in his elaborate essay, clearly shows that the micropyle may be single or 
multiple, and may occur at either or both poles of the egg; but unfortunately he gives us 
less information respecting the ova of the Homopterous Hemiptera than regarding those 
of any other great group of Insecta. Cercopis, in fact, is the only genus of this division 
in which he has observed the micropyle with certainty, and here there are two, one on 
each side of the anterior pole. 

: The anterior extremity of the chorion in Aphis (Pl. XL. fig. 4, B) presents a small 
conical papilla, in which I have been unable to discover any aperture. Internally, how- 
ever, the corresponding surface of the chorion appears as it were rough and uneven ; and 
when caustic potass is added, it, like the rest of the inner surface of the chorion, exhibits 
a vay curious marking, as if so many circles or more irregular figures were impressed 

: upon it The thickness of the papilla is about Zvooth of an inch; and in young ova å 
delicate filiform appendage more than once appeared to be continuous with it: this, how- 
ever, was invariably absent in fully-formed ova. 

At the opposite pole (fig. 4, A), the ovum presents a curious appendage, about s$oth of. 
seg in length. When the ovum is in its natural position within the ovary, the 
= "e E eg which closes over it below, leaves a sort of chamber in which this 
VE så tp d I fees closely applied against the chorion, is received. lu 
3 er inh st appendage appears like a rope with loosened strands, 
M ibd assi xs and is seen to be coated with a clear gelatinous substance, 
hadde ER es rod- ike filaments of about zygth of an inch in length are ım- 
: . caustic potass, this clear substance and its imbedded particles are 


AND MORPHOLOGY OF APHIS. 207 


dissipated, and the central cord becomes less distinct ; but I have never yet seen it dissolved, 
and sometimes it seems altogether to resist the reagent. The rounded tubercle of the 
chorion to which it is attached, however, now clearly exhibits a central funnel-shaped 
body, continuous with the axis of the appendage, and appearing like a canal (fig. 4, C). 

Is this a micropyle, and what is the nature of the appendage? I regret that I have 
not the leisure to pursue the inquiry far enough to answer this question satisfactorily ; but 
I incline to think that the micropyle is really situated here*. 

The albuminous papilla surrounding the bundle of spermatozoa in the impregnated 
ova of Musca, Dexia, and Melophagus (Leuckart, l. c. pl. 7. figs. 1, 2, 4, 5), reminds one 
strongly of the envelope of the appendage in Aphis. 

The micropyles of Libellula, Dexia, and Musca, again, exhibit a sort of “mouthpiece ” 
formed by a prolongation of the chorion surrounding the micropylar aperture. 

The account which I have given of the reproductive organs of the oviparous Aphis is 
in general agreement with that of other observers. Morren describes the reproductive 
organ of the wingless oviparous female of 4. Persicæ thus :— 

“The ovigerous cæca well deserved their name; for no foetuses were any longer visible 
in them. Each was exactly composed of three chambers, of which the first or terminal 
was enlarged and spherical, and filled with twelve to twenty-four little, well-formed ova, 
yellow in the centre, and white peripherally. These ova descended into the second cham- 
ber, and then elongated and enlarged; but in general they acquired their hard covering 
only in the third or last chamber, which in all the females was occupied by a very large 
ovoid greenish ovum. These ova became covered at the same time with the sebific liquid ; 
for some were seen to be provided with a little appendage intended to fix them to the 
bodies in which the parent lays them. This appendage was mucous, and arose from a 
thickened viscous liquid.” (2.:c. p. 89.) ! n 

1 recognize in Morren's “twelve to twenty-four ova” the ovarian glands which I have 
described. His microscope was obviously inadequate to show him the true ova; but it 
seems difficult to suppose that in this species there is, as he maintains, neither colleterial 
glands nor spermatheca. His objection to Dutrochet’s statements appeårs to me to be well 
founded, for Dutrochet examined a viviparous female; but I strongly suspect that he has 
himself overlooked the “ sebific” apparatus in the oviparous forms. a 

Von Siebold states that the ovarian cæca of the oviparous Aphis Lonicere are divided 
into only two chambers :— ` : å 

“In the undeveloped state the whole tube forms only a simple pyriform appendage of 


* After deseribing the cup-like micropyle at the anterior pole of the ovum of the Louse, Leuckart (/. c.) goes on 
sn there is at the posterior pole of the ovum a structure 


“ Besides this micropylar apparatus at the anterior pole, 
å S s attached rather on one side of the centre of 


Which attracts attention. It may be described as a blunt cone, which i x 
the posterior pole, and has ahis a peculiar striated appearance by reason of its longitudinal folds, and band-like 
thickenings. The interior diameter of this structure measures gl" ; the upper is less, about vr 5 and ee z 
about the same. A hollow space is contained within this body, so that it might be compared tos pn; bu i er 
3s if from the roof, or cupola, as it might be termed, of this bell, a number of closely appressed elevations = DE n 
depended. With respect to the import of this remarkable apparatus, I will only throw v the remis TIAM 
an apparatus of attachment. Fora long time I thought I had discovered in it a second micropylar apparatus ; 


“nounced this view when I was unable to discover any aperture in it.” (p. 140.) 


208 PROF. HUXLEY ON THE AGAMIC REPRODUCTION 


the oviduet; but as development proceeds, the upper globular chamber becomes by 
degrees separated by a constriction, and at the same time a great difference makes its 
appearance between the upper and the lower chambers : for the lower chamber contains 
a finely granular mass which gradually becomes modelled into an oval egg; the upper 
chamber, on the other hand, is filled with vesicular bodies, in which smaller vesicles con- 
taining a nucleus are distinguishable. If these bodies are to be regarded as germs of 
ova (Wollte man diese blasenfórmigen Körper als Eier-keime betrachten), we may assume 
that these Aphides were capable of bringing forth more than eight ova.” 

Von Siebold then goes on to describe the colleterial glands, and the spermatheca, which 
had not before been seen. If the ovaries of Aphis Loniceræ are not constructed on a totally 
different plan from those of the species I have described, it is, I think, pretty clear that 
Von Siebold, like Morren, has mistaken the ovarian glands for the rudiments of the ova. 
. Indeed, his phraseology indicates that he himself had no great confidence in his interpre- 

tation of the parts. 
$ 4. The Development of the Pseudovarium. 

In the viviparous female, the germ increases in size, and gradually becomes separated 
from the terminal chamber by the successive development and separation by constriction 
of new pseudova. The number of chambers between the terminal one and that nearest 
the vagina, therefore, varies until it attains its maximum, which is necessarily regulated 
by the ratio between the time required for the perfection and birth of a larva, and the rate 
at which new pseudova are detached from the pseudovarium. In the species of Aphis 
which I examined, I found ordinarily four or five such chambers. Germs between zooth 
and 55th of an inch in length presented the following characters (Pl. XX XVII. fig. 1 :— 
They exhibit a central darkish matter, surrounded by a clear cortex. The latter is com- 
posed of a single layer of a substance similar in appearance to that composing the mass of 
the germ above described, while the central substance is obscured by a number of minute 
granules which hide its internal structure. N evertheless, I have occasionally detected 
what I believe to be endoplasts, scattered through its substance, as in Pl. XXXVII. fig. 1, 
which represents a germ in this stage treated with very dilute acetic acid; and as in a 
more advanced condition we shall find such bodies easily recognizable, I do not doubt 
that the central substance has the same fundamental composition as the peripheral layer. 
The central mass, it will be observed, completely simulates the vitellus- of an impregnated 
ovum; and I will therefore term it a « pseudovitellus.” The peripheral clear layer i$ 
on the other hand, in all essential respects comparable to a blastodermic vesicle ; and I see 
no reason why it should not be called a blastoderm, since the term is not necessarily con- 
fined to the product of impregnation. 

Ina more advanced condition (fig. 8), the blastoderm has become thicker in all parts, 80 
as to consist of at least two or three layers of “cells;” but the thickening shows itself 
ev ae ee side of the distal end of the germ (that turned towards the vagina), 
Scale in ina m. ri twice as thick as in other parts. A linear bui 
ET URN p x thickened layer (fig. 4); and at the same time indibi 
"ui en ie etween the distal extremity of the thickened portion and 

: | rm: it is as if the latter were giving way at this point. In some 


AND MORPHOLOGY OF APHIS. 209 


specimens the cell-cavities of the inner portion of the thickening were particularly well 
marked ; and the coarsely granular central substance exhibited a tendeney to break up into 
large globular masses, which became particularly distinet on the addition of water. 

It is in the largest of these germs that the resemblance of the pseudovum to an ovum 
is completed by the formation of a pseudovitelline membrane (fig. 8, a). This structureless 
homogeneous membrane is, doubtless, developed by a process of excretion, either from 
the pseudovum or from the walls of the chamber which contains it. It completely enve- 
lopes the pseudovum, and acquires greater thickness and strength as development proceeds. 

The embryo first becomes clearly fashioned in pseudova between „45th and 41th of an 
inch in length (Pl. XX XVII fig. 5). At the distal extremity, in the region of the thickening 
of the blastoderm, the latter appears separated into two portions, the outer of which forms 
a sort of hood over the inner. The hood eventually becomes the hinder part, if not the 
whole, of the abdomen of the larva. It is continuous, on the side answering to the dorsal 
side of the larva, with the rest of the blastoderm, which now, instead of enclosing the 
pseudovitellus, lies partly beneath and partly behind it. That portion of the blastoderm 
which lies behind the pseudovitellus, and parallel with the hood, is the rudiment of the 
sternal region of the thorax ; and I shall hereafter term it the thoracic segment of the blas- 
toderm. That part of the blastoderm which lies beneath the pseudovitellus will become 
the sternal region of the head; and I shall therefore call it the cephalic segment, while 
the hood itself is the abdominal segment of the blastoderm. 

The thoracic segment, it will be observed, is in this stage bent up at right angles to 
the axis, and reaches the dorsal region, which it bounds posteriorly. The cephalic seg- 
ment, on the other hand, hardly extends upwards at all, but lies in one plane; so that the 
anterior end of the embryo is almost wholly formed by the pseudovitellus. The latter is 
aggregated into a few large globular masses, which are in immediate contact with the 
pseudovitelline membrane on their dorsal surface. - Å 

The pseudovitellus is in immediate contact inferiorly with a layer of the blastoderm of 
å more pellucid aspect than the rest, and separated from it by a more or less distinct line 
of demarcation. This layer (g) could be detected only on the dorsal face of the thoracic 
and cephalic segments, and owed its superior transparency to the comparatively large 
` size of the clear cavities surrounding its endoplasts. 

That portion of the layer which covered the posterior portion of the thoracic segment 
Was particularly remarkable for the size and clearness of its cells and their endoplasts (7). 
In the progress of development, the central portion of the alimentary canal occupies a 
place nearly corresponding to the centre of the clear layer; while, if we trace out the site of 
the rest of the mass in larger and larger embryos (Pl. XXX VII. figs. 1, 3, 4, 5), we find 
it always retaining the same relative position to the reflected abdominal hood, but gradu- 
ally enlarging, and eventually becoming subdivided into five oval lobes upon each side, 
each of which surrounds itself with a membrane, and assumes the form of the terminal 
Chamber of one of the pseudovarial emca. It would be a great mistake to suppose that 
It is only one of these chambers, however ; it is in fact the rudiment of an entire cæcum ; 
and before the embryo leaves the parent, it becomes divided into three chambers by the 
Sradual development and metamorphosis of pseudova in the way described above. 


210 PROF. HUXLEY ON THE AGAMIC REPRODUCTION 


The granular pseudovitellus takes no part whatever in the formation of the reproductive 
organs. In embryos of gj;th of an inch in length, I could very plainly observe a clear 
space with an endoplast in the middle of each of its spheroidal masses (Pl. XXXVIII, 
fig. 3). Similar masses constitute a larger or smaller proportion of the corpus adi- 
posum of the larva and adult insect; and I believe that the latter proceeds from the 
former. 


§ 5. Summary and Comparison of Germs and Ova. 


I will now sum up the results of the observations which have been detailed in the pre- 
ceding pages. 

1. The pseudovarium consists of vagina, oviduets, and pseudovarian cæca. 

2. The vagina is unprovided with either spermatheca or colleterial glands. 

3. The pseudovarian cæca are each divided into many chambers by constrictions. 

4. The apical chamber contains bodies which are not distinguishable from the germinal 
vesicles and spots of the true ovaria. 

5. These bodies, surrounded by a mass of clear substance representing a yelk, are set 
free as pseudova, and are then undistinguishable from true ova. 

6, The pseudova are eventually converted into cellular germs, apparently by the same 
process as that by which an ovum is converted into an embryo. 

7. In these germs the central part becomes a granular pseudovitellus, the peripheral 
a blastoderm ; the rudiments of the different organs next appear, and the germ becomes 
surrounded by a pseudovitelline membrane. 

8. Eventually the pseudovitellus probably becomes the corpus adiposum. 

9. All the other organs are developed from the blastoderm, which becomes distinguished 
into two layers. From the outer of these the muscles, nerves, limbs, and tegument are 
developed, while the inner gives rise to a part of the alimentary canal (?) and to the 
reproductive organs or pseudovarium of the larva. | 

10. The pseudovarium contains no particle of unchanged tissue of the germ, but is a 
considerably differentiated and readily distinguishable mass. The mass divides into ten 
lobes anteriorly; and these lobes become the pseudovarian cæca. Before the larva is 
horn, each cæcum is divided into three chambers, the two posterior of which contain rudi- - 
mentary embryos. 

11. The genital apparatus of the oviparous female consists of a vagina, oviducts, and 
ovarian ceca. The latter are multilocular; and the vagina is provided with the sperma- 
theca, and the two colleterial glands first demonstrated by Von Siebold. 

12. The rudiments of the ova are undistinguishable from those of the pseudova. They 
are developed in the lower part of the apical ovarian chamber, the upper part of which is 
occupied by the bodies I have termed ovarian glands. The ova are not at first enveloped 
in a chorion. 


13. In the lowest chamber the ova are provided with a chorion, vitelline membrane, and 
what appears to be a micropyle. 


If these propositions are correct, I see no valid objection to the conclusion, that the 
agamic offspring of Aphis is developed from a body of precisely the same character as that 


AND MORPHOLOGY OF APHIS. 211 


which gives rise to the true egg. The pseudovum is detached from the pseudovarium in 
the same way as the ovum from the ovarium. In both cases, the act of separation is in 
every respect a process of gemmation. : 

From this point onwards, however, the fate of the pseudovum is different from that of 
the ovum. The former begins at once to be converted into the germ ; the latter accu- 
mulates yelk-substance, and changes but little. Both bodies acquire their membranous 
investment rather late; within it the pseudovum becomes a living larva, while the ovum 
is impregnated, laid, and remains in a state of rest for a longer or shorter period. 

Although, then, the pseudovum and the ovum of Aphis are exceedingly similar in 
structure for some time after they have passed out of the condition of indifferent tissue, 
it cannot be said that the sole difference between them is, that the one requires fecundation 

and the other not. When the ovum is of the size of a pseudovum which is about to 
develope into an embryo, and therefore long before fecundation, it manifests its inherent 
physiological distinctness by becoming, not an embryo, but an ovum. Up to this period 
the influence of fecundation has not been felt; and the production of ova instead of pseud- 
ova must depend upon a something impressed upon the constitution of the parent before 
it was brought forth by its viviparous progenetrix. 

In this respect, the ova of Aphis exhibit the same relation to the pseudova as the 
ephippial eggs of Daphnia (whose development has been so well described by Mr. Lub- 
bock) bear to the agamic eggs; for the histological change in the ovarium of Daphnia, 
which precedes the development of the ephippial eggs, is clearly shown by Mr. Lubbock 
to have no relation to fecundation. 

Let me remark on yet another interesting, though perhaps only partial, analogy. Von 
Siebold has shown that the ova of the Queen bee produce females or males, according as 
they are fecundated or not. The fecundated ovum produces a queen or a neuter accord- 
ing to the food of the larva and the other conditions to which it is subjected; the un- 
fecundated ovum produces a drone. Now, what have we seen in Aphis? The fecun- 
dated ege produces viviparous Aphides, which are the equivalents of the neuter bees ; 
and from them are eventually produced males and oviparous females. The oviparous 
females are fecundated and lay eggs which produce only viviparous or neuter Aphides. 

On the view which Dr. Carpenter and myself take of the zoological individual, the 
whole produce of a single fecundated ovum of the Aphis is as much the Aphis individual 
as it is the Bee individual. Consequently we have two equivalent and related series. 


å äi Ova requiring impregnation, and males. 


Bee, Impregnated ova : Neuters or females roducing4 Females which give rise to ova requiring 
Aphi producing 4 |... P à 
phi. Impregnated ova Viviparous neuters impregnation, and males. 


The fact that in the one case the males are developed from pseudova —€— fully- 
formed true ova, "and: in the other from pseudova resembling — " 
Makes no essential difference in the analogy, but only demonstrates pe CR 


impossibility of drawing any absolute line of demarcation histologically between ova 
and buds. 


VOL. XXII. T 


212 PROF. HUXLEY ON THE AGAMIC REPRODUCTION 


$ 6. Hypothetical Explanations of Agamogenesis. 


The majority of writers on the wonderful phenomena of Aphidian life, have been con- 
tent to state the facts more or less clearly ; but Morren, who has done this so clearly and 
philosophically, has in addition carelessly thrown out à hint of a mode of explaining 
them. The agamic Aphis, he says, is a portion of organized tissue which individualizes 
itself :— 

* Suppose that vitality is sufficiently energetic to impress, on the tissue which indivi- 
dualizes itself, the form of the producing species, and you have the generation of the 
Aphides. This energy becomes lost at the end of a certain number of generations, and 
a new impulse becomes necessary. It is that of the male. In my youth I might have 
adopted with pleasure such an hypothesis as this; but now I prefer to doubt: the facts 
which I have set forth are worth more than a theory.” 

The hypothesis is, however, to my mind, in no essential particular distinguishable 
from that hypothetical explanation which has been propounded by the author of the 
well-known work on * Parthenogenesis.” Substitute for “energy of the male,” in the 
foregoing passage, “spermatic force;" and the difference between the two hypotheses 
becomes evanescent. 

But this is a question of minor importance as compared with the value of the hypo- 
thesis in itself; and it is with regard to this latter point that I now propose to make a 
few remarks. 

Professor Owen’s views are, I believe, fairly stated in the following extracts from the 
work cited :— 

“ We find derivative germ-cells, and masses of nuclei like those resulting from the final 
subdivision of germ-cells, retained unchanged at the filamentary extremities of the - 
branched uterus forming the ovaria of the larval Aphides.”—1. c. pp. 7, 8. 

* According to my own observations, the germs are perceptible in the embryo Aphis, 
above the simple digestive sac, before any organs have been formed for their reception. 
And with regard to the nature of the organs when formed, I may remark that the con- 
tinuity of the ovarian tubes with the oviducts in all insects, is such as to render the 
negation of the term ‘ovary’ to those two bodies from which the slender extremities of 
the eight oviducal and uterine tubes proceed in the larval Aphis, to say the least, quite 
arbitrary. My examinations agree with those of Siebold, in determining scarcely any 
appreciable difference between the ovaria of the oviparous and those of the viviparous 
females. The contents of the ovarian tubes differ, inasmuch as they contain oval masses 
of granules or nuclei, comparable to the germ-mass in its state of minutest subdivision, 
in the virgin Aphides, and not ova with the germinal vesicle as in the oviparous females.” 
—Ibid. p. 38. 

“The completion of an embryonic or larval form by the development of an ovarian 
germ-cell, or germ-mass, as in the Aphis, without the immediate reception of fresh sper- 
matic force, has never been known to occur in any vertebrate animal. 
roa er which renders this seemingly strange and mysterious generation of an - 

precedent coitus possible, is the retention of a portion of the germ-mass 


AND MORPHOLOGY OF APHIS. 213 


unchanged. One sees such portion of the germ-mass taken into the semitransparent 
body of the embryo Aphis, like the remnant of the yelk in the chick. TI at first thought 
that it was about to be enclosed within the alimentary canal, but it is not so. As the 
embryo grows, it assumes the position of the ovarium, and becomes divided into oval 
masses and enclosed by the filamentary extremities of the eight oviducts. . . ." — Zbid. 
p. 69-70. 

* [t would be needless to multiply the illustrations of the essential condition of these 
phenomena. That condition is, the retention of certain of the progeny of the primary 
impregnated germ-cell, or in other words, of the germ-mass unchanged, in the body of 
the first individual developed from that germ-mass, with so much of the spermatic 
force inherited by the retained germ-cells from the parent cell or germ-vesiele as 
suffices to set on foot and maintain the same series of formative actions as those which 
constituted the individual containing them.” —Tbid. p. 72. 

“The physiologist congratulates himself with justice when he has been able to pass 
from cause to cause, until he arrives at the union of the spermatozoon with the germinal 
vesicle as the essential condition of development—a cause ready to operate when 
favourable circumstances concur, and without which cause these circumstances would 
have no effect. 

What I have endeavoured to do has been to point out the conditions which bring about 
the presence of the same essential cause in the cases of the development of an embryo 
from a parent that has not itself been impregnated. The cause is the same in kind, 
though not in degree, and every successive generation, or series of spontaneous fissions, 
of the primary impregnated germ-cell, must weaken the spermatie force transmitted to 
sueh successive generations of cells. 

“The force is exhausted in proportion to the complexity and living powers of the 
organism developed from the primary germ-cell and germ-mass.”—1bid. pp. 72, 73. 

These statements are repeated in the recently published second edition of Prof. Owen's 
Lectures on the Invertebrata. l 

The paragraphs I have cited contain two kinds of propositions—assertions with respect 
to matters of fact, and deductions from those assertions. The former are, according to my 
observations, incorrect; and, as I conċeive, the latter are unfounded. 

As regards the first citation, for instance, the contents of the apical chambers ef the 
pseudovaria are not by any means identical with those * resulting from the final subdivision 
of germ-cells retained unchanged," as the most cursory comparison of the two structures 
will show. ; 

In the second citation it is affirmed that the germs are perceptible in the embryo before 
any organs are formed for their reception. This, again, is an error if my observations ped 
correct. "The absence of figures, and the too vague and general character of the descrip- 
tions in Prof. Owen's work, render it very difficult to understand what lie really has seen; 
but I imagine that he has taken the substance which constitutes the rudiment of the whole 
pseudovarium, and which becomes differentiated partly into pseudova, partly into tur walls 


- of the organ, for a mass of germs. What is meant by “those two bodies €— the 
| F 


PROF. HUXLEY ON THE AGAMIC REPRODUCTION 


214 
slender extremities of the eight oviducal and uterine tubes proceed,” and which are supposed 
There are no such bodies, that I can discover. 


to be ovaries, I am at a loss to divine. ) l 
In the latter part of the same citation, the existence of a histological difference between 


the contents of the pseudovarium and those of the ovarium is asserted. But there is 
assuredly nothing in the former to which the description can apply; and I re-affirm the 
impossibility of drawing any histological line of demarcation between the pseudova and 
the young true ova. : 

How any one who carefully studies the development of Aphis can arrive at the con- 
clusion that a portion of the germ-mass is taken into the body of the embryo Aphis, * like 
the remnant of the yelk of the chick," I know not ; and, for the reasons mentioned above, I 
even doubt if I clearly apprehend what is meant. Dr. Burnett (l. c. p. 73) assumes that 
what is intended by * portion of the germ-mass" is what I have termed the pseudovitellus. 
In that case the statement is erroneous ; for the pseudovitellus takes no share in the for- 
mation of the pseudovarium. If, on the other hand, the true rudiment of the pseud- 
ovarium is indicated, the statement in question is equally incorrect; for this is never out 
of the body, and hence can hardly be taken into it, nor can that out of which the so- 
called * oviducts” are produced be properly said to become * connected with them,” or to 
“aid in forming their filamentary extremities.” 

When the basis of a hypothesis is shown to be incorrect, the hypothesis itself is com- 
monly considered to be disposed of; but possibly in the present case it may be urged that, 
although the contents of the pseudovarium are wholly dissimilar “ to the germ-mass in 
in its state of minutest subdivision,” they are nevertheless so little changed that my 
criticism of the phrase is trivial. To this I reply that, whether the alteration be small or 
great, it is as great as that which occurs in the terminal cæca of a gland, or in a true 
ovarium, and that the tissue of the apical pseudovarian chamber is far more differentiated 
than the indifferent tissue which constitutes the youngest portion of an ordinary epithelium 
or epidermis. 

Whatever conclusions are based upon the resemblance of the tissue of the pseudovarium 
to that of the embryo, must therefore apply in equal or greater force to the tissues which 
I have just named ; and, unless reason can be shown to the contrary, whatever powers are 
possessed by the one, in virtue of this similarity, must be possessed in equal or greater 
degree by the other. 

But in this case what becomes of the hypothetical explanation of the asexual reproduc- 
tion of Aphis, under discussion ? 

The condition of such reproduction is, according to the hypothesis, the retention of 
* certain of the progeny of the primary impregnated germ-cell unchanged," « with so 
much of the spermatie force, inherited by the retained germ-cells from the parent-cell 9r 
germ-vesicle, as suffices to set on foot and maintain the same series of formative actions 
Mei which constituted the individual containing them." ; 
= for the sake of argument, that the amount of histological change 1» 
GE mass is unimportant. I am ready to suppose even, in accordance W* 

ypothesis, that its cells retain sufficient “spermatic force” (whatever that may be) 


AND MORPHOLOGY OF APHIS. 215 


to commence an independent life. But I ask, how does this explain agamogenesis ? 
Why does not the epithelium of the ovarium (which is as little or less changed) give rise 
to young without impregnation ? Why are not the young cells of glands, which are as 
little changed, * parthenogenetic” ? Why, finally, does not the deep substance of our 
epidermis and epithelium, which absolutely more nearly resembles embryonic tissue than 
the structure of the pseudovarium does, give rise to young ? 

It may be replied, however, that the supposed ** spermatic force” is exhausted by the re- 
peated subdivisions of the germ-cell before it becomes a part of the deep epidermie tissue ; 
for it is one condition of the hypothesis, that every successive generation or series of spon- 
taneous fissions of the primary impregnated germ-cell must weaken the “ spermatic force” 
transmitted to such successive generation of cells. 

I presume, however, that the original “ spermatic force” is at least as strong in a Man as 
in an Aphis. The average size of the embryo-cells in Aphis is at least not greater than 
in Man, and the specific gravities of their essential tissues are not very different; so that 
we may fairly assume that as many embryo-cells go to form a given mass of Aphis as of 
Man. In that case the impregnated embryo-cell must subdivide as often ; and therefore 
the “ spermatic force ” must become as much exhausted in forming, say, a grain or a pound 
of Aphis, as in giving rise to the like quantity of human substance. 

In his Lectures, Prof. Owen adopts the calculations taken by Morren (as acknowledged 
by him) from Tougard, that a single impregnated ovum of Aphis may give rise, without 
fecundation, to a quintillion of Aphides*. I will assume that an Aphis weighs -sth of 
à grain, which is certainly vastly under the mark. A quintillion of Aphides will, on this 
estimate, weigh a quatrillion of grains. i 

He is a very stout man who weighs two million grains; consequently the tenth brood 
alone, if all its members survive the perils to which they are exposed, contains more 
substance than 500,000,000 stout men—to say the least, more than the whole population 
of China! And if the law cited above be correct, the “ spermatic force” in each cell of an 
Aphis of this brood must be diminished 500,000,000 times as much as that of a single 
human cell; nevertheless the * spermatic force” of the Aphis cell is enough to impel it 
to the production of young, while that of the human cell is not ! 

When to these considerations I add, that it has been shown that the agamic propaga- 
ion of the Aphis may, under proper conditions, be continued for four years without 
interruption, in which case the “ spermatic force" in the later broods must stand in an 
infinitely minute ratio even to that contained in the cells of the tenth generation, the 
reductio ad absurdum by simple arithmetic, of the so-called explanation, appears to me to 
be sufficiently obvious. 

For the sake of argument, however, I am willing still to suppose for ogni — 
‘gamogenesis does take place in consequence of the retention of a ** spermatic force. Bat 
I must ask, how does this phrase constitute an explanation of the phenomena? Nothing 
is more common than the misuse of the word * force” on the part of ma = a 
versed in the Phraseology, than trained in the severe methods, of physical science. The 
"patient inquirer every now and then calls in the aid of molecular force, or chemical 


* I have not thought it worth while to add, in the products of the generations preceding the tenth. 


216 PROF. HUXLEY ON THE AGAMIC REPRODUCTION 


force, or magnetic force, or od-force, to account for the existence of a mass of phenomena 
which will not arrange themselves under any of his established categories —forgetting that 
a * force,” the conditions of whose operation (that is, whose laws) are undetermined, is but 
a scientific idol, at once empty and mischievous,—empty, because it is but a phrase with- 
out real meaning; mischievous, because it acts as an intellectual opiate, confusedly satis- 
fying many minds and obstructing the progress of inquiry into the real laws of the 
phenomena. If we show that a fact is a case of a law, we explain that fact; but expla- 
nation by reference to an undefined * force,” of questionable existence, is simply ‘ignorance 
writ large.” 

Now, how does the hypothesis fulfil the indispensable conditions of a genuine explana- 
tion? In the first place, what proof is there of the existence of such a force as *sper- 
matic force.” All that we know is, that an ordinary ovum will not undergo those changes 
which constitute development without the contact of the spermatozoon. Hence it is 
concluded that some force contained in the spermatozoon is the efficient cause of all these 
changes. But what would be thought of the artillerist who should imagine he had ex- 
plained the propulsion of a bullet by saying it was ‘trigger force’? Or to take an 
illustration from phenomena of a like order to those under discussion: a seed will not 
grow unless it is exposed to a certain amount of warmth and moisture; but have I 
explained the growth by saying that it is the effect of ‘heat and moisture force’ which 
becomes diffused through the seed ? 

The very existence of this “spermatic force,” then, is a gratuitous assumption; and if 
we seek for its laws of action, we find but two stated: first, that it becomes weakened by 
the successive divisions of the germ-cell; second, that “the force is exhausted in pro- 
portion to the complexity and living powers of the organism developed from the primary 
germ-cell and germ-mass.”’ i Se 

I have shown to what singular consequences the first assumption leads us; it remains 
only to consider the second. If it be true, the occurrence of agamogenesis in the animal 
kingdom must bear an approximatively inverse ratio to the complexity of the organization 
of the different groups. Let us examine one or two subkingdoms in this point of view. 
Among the Annulosa, the Rotifera and Turbellaria possibly possess it to a small extent; 
the Nematoidea do not possess it at all. Many Trematoda possess it; others, such as 
Aspidogaster, have nothing of the kind. The Acanthocephala are not known to possess it; 
the Echinodermata are regarded by Prof. Owen as possessing it, but their different families 
show every gradation from simple metamorphosis to something like agamogenesis. A few 
Annelida possess the power in a marked degree; in many, nothing of the kind is known: 
en E: the Earth-worm and the Leech have it not. Of the Orustacea, Som 

x >ranehiopoda, exhibit it in the highest perfection ; but no trace of it an 
pr in. Copepoda, Cirripedia, Pæcilopoda, Edriophthalmia, or Podophthalmis: 
pos fe n N: and Arachnida the process is not known :- but we find it in the ROS 
ER = Soins this not, so far as we know at present, in Aptera or Ort E 
co FE So CURIE Hymenoptera and Lepidoptera ; and there is every reason 
Se y Nen in isolated, though perhaps in many, genera of these orders. 

| again: agamogenesis occurs in the Polyzoa and Ascidioida, not in the Bre 


AND MORPHOLOGY OF APHIS. 217 


chiopoda. It is not known to occur in any of the Lamellibranchiata; and among the 
higher Mollusca the nearest approach to it is presented by the animal (whatever it is) 
which gives rise to the * Synapta-schnecken” (high Gasteropods), and by the Hectocoty- 
ligenous Cephalopoda. 

After this simple statement of well-known facts, I need not remind even the tyro in 

zoology, that there is no evidence of an inverse relation between the occurrence of aga- 
mogenesis and complexity of organization. 

I have hitherto, in the course of this argument, confined myself in the main to the deve- 
lopment of Aphis; but it is only just to observe that the author of the hypothesis brings 
forward yet another original observation in support of his large generalization :— 

* In the freshwater polype, the progeny of the primary impregnated germ-cell retained 
unaltered in that body, may set up, under favourable stimuli of light, heat, and nutriment, 
the same actions as those to which they owed their own origin ; certain of the nucleated cells 
do set up such actions, those, e.g. in the Hydra fusca, which are aggregated near the adhe- 
ring pedicle or foot; and the result of their increase by assimilation and multiplication 
is, to push out the contiguous integument in the form of a bud, which becomes the seat 
of the subsequent processes of growth and development; a clear cavity or centre of assi- 
milation is first formed, which soon opens into the stomach of the parent; but the com- 
munication is afterwards closed, and the young hydra is ultimately cast off from the surface 
of the parent *.”— Lectures,’ 2nd ed. p. 124. 

I have had occasion carefully to watch the process of gemmation not only in Hydra, 
but in many species of all the other subdivisions of the Hydrozoa; and I venture to assert 
that no such process as that described by Prof. Owen takes place in any one of them. 

The bud is from the first in communication with the cavity of the body, of which it is 
å mere diverticulum, whose walls are a little thickened at the extremity. No special cell 
or group of cells can be discovered as the centres whence growth proceeds. No “inte- 
gument” is pushed out by any thing beneath it; but the outer layer of the body of 
the animal thickens and grows pari passu with the growth of the bud. No especial 
accumulation of derivative germ-cells can be seen in any part of the body of any Hydro- 
200» , and before gemmation commences there is no distinguishable difference of texture 
between the part in which gemmation commences and any other portion of the body. 
Furthermore if a complex Hydrozoon, such as a Physophora or Agalma, be examined, it 
will be found that there is no histological distinction whatsoever between that part of the 
body which is to give rise to a free swimming generative zooid, and that which produces 
merely a bract, a tentacle, or a stomach. : 

In this case then, as in that of the Aphis, the hypothesis receives no support from, but 
Is totally opposed by, facts; and I unreservedly adopt the conclusion (long na clearly and 
well expressed by Dr. Carpenter), that * spermatic force ” is but a name without definite 
Meaning, applied to that which is not proven to exist, and the assumption of whose 
existence, even, does not help us a single step towards the understanding of the wonderful 
Phenomena of agamogenesis. 


2 * 99 
* I have cited this passage from the ‘ Lectures’ rather than from the work on “ Parthenogenesis,” as they may 
Supposed to contain the expression of the author’s latest views, 


218 PROF. HUXLEY ON THE AGAMIC REPRODUCTION 


Truly we may say, with Degeer (J. c. p. 129), * Les Pucerons sont des insectes bien 
capables de déranger tout système formé de génération, et de mettre en déroute tous ceux 
qui s'efforcent d'expliquer ce mystère de la nature.” 

But the question may be asked : if the “spermatic force” be a myth, what és the cause 
of the phenomena? Considering that the groundwork of modern physiology is not a 
score of years old, I do not think the confession of our inability to answer that question 
at present is any opprobrium to science. 

When we know why, in a mass of tissue of identical structure throughout, one part 
becomes a brain, and another a heart, and a third a liver—when we can answer these 
every-day questions of the sphinx, we may attempt her more difficult riddles without 
running too great a risk of being devoured. 

At the present time it seems to me well nigh hopeless to look for an explanation of 
these phenomena. Some such classification of them, however, as will indicate their 
analogies with other vital manifestations, may fairly be attempted, and, when successfully 
carried out, will prove the first step towards an explanation. 


$ 7. Classification of the Phenomena of Agamogenesis. 


It does not seem to be very difficult to effect such a classification. In the course of the 
development of the total product of a single impregnated ovum (which, with Dr. Carpenter, 
I regard as the zoological individual), one of two things may occur: either all the living 
products may remain in connexion with one another, or they may become separated from 
one another. The former case I term Continuous, the latter Discontinuous Development. 

In continuous development, the size may increase, the form and texture remaining 
unchanged—constituting simple growth; or, the size remaining unchanged, the form 
and texture may alter—constituting simple metamorphosis; or the two processes may be 
combined, as in all those changes which we term gemmation, without separation from 
the parent. | 

Discontinuous development differs from continuous only in this, that the products of 
the growth and metamorphosis of the embryo become separated into two or more portions, 
which when they retain their vitality independently are termed * zooids.” | 

When the produced *zooid” is capable of development into an independent organism 
without the influence of an act of conjugation with another zooid, I term the process 
agamogenesis. The producing zooid may be devoid of sexual organs, as in the Salpe, 
many Hydrozoa, many Trematoda—in fact, in the great majority of cases of agamogenesis. 

I term the first producing zooid of the individual the protozooid ; the produced zooids, 
deuterozooids. In some cases the deuterozooids acquire sexual organs, and give rise to 
ova and spermatozoa ; but in others they produce new zooids : thus broods of tritozooids, 
&e., will be produced. When the producing or protozooid possesses no sexual organs, 
I think Prof. Owen’s term of *metagenesis” might well be applied to the kind of 
agamogenesis ; but where the protozooid possesses sexual organs, and its buds have all 


the histological characters of ova, then the process may fairly enough be termed parthe- 
nogenesis. 


AND MORPHOLOGY OF APHIS. 219 


Finally, the produced zooid may be incapable of development into an independent 
organism, unless it conjugate with another zooid. In this case we have sexual reproduc- 
tion, or gamogenesis. 

The natural character of this classification of the various modes of development is 
manifest when it is thrown into a tabular form :— 


Growth. 
Continuous 1 Metamorphosis. 
Gemmation without fission. 


Development. * . [Metagenesis. 
Agamogenesis. 
Parthenogenesis. 
Discontinuous $ 


(Gemmation with 
fission). | Gamogenesis. 


Whatever hypothesis we may entertain with respect to the nature of these processes, 
and however we may think fit to conceive the nature of the * individual" I think it 
must be admitted, that all the phenomena of development in the animal kingdom (and I 
would venture to add, in the vegetable kingdom also) fall under one or other of these 
heads. 

Furthermore, all these modes of development pass into one another. Growth and 
metamorphosis are combined in all animals. Gemmation, so long as the gemma continues 
attached, is but a peculiar kind of growth and metamorphosis. From the fixed bud to the 
separate one, we have all gradations; and fission is little more than a peculiar mode of 
budding. 

Free gemmation is * metagenesis” when the bud is not developed within the homologues 
of the sexual reproductive organs; it becomes “ parthenogenesis” when the bud is deve- 
loped within such organs ; finally, when the free bud requires conjugation with another 
free bud for its development, we have gamogenesis, or sexual reproduction: but cases 
such as those of Daphnia and Apis show that the histological element, which is at one 
time agamogenetic, may at another be gamogenetic. ve 

Time was when the difficulty of the physiologist lay in understanding reproduction with- 
out the sexual process. At the present day, it seems to me that the problem is reversed, 
and that the question before us is, why is sexual union necessary ? Far from seeking for 
an explanation of the phenomena of gemmation in the transmitted influence of the sper- 
matozoon, the philosopher acquainted with the existing state of science will seek, in the 
laws which govern gemmation, for an explanation of the spermatic influence. 


VOL. XXII, > 


[ 221 ] 


XV. On the Agamic Reproduction and Morphology of Aphis.—Part II. By Tuomas 


H. Huxuzv, F.R.S., Professor of Natural History, Government School of Mines. 
Communicated by G. Busk, F.R.S., F.L.S. 


Read January 21st, 1858. 


$ 1. Embryogeny of the external organs of Aphis. 

$ 2. Embryogeny of Mysis as exemplifying the Crustacea. 

$ 3. Embryogeny of Scorpio as exemplifying the Arachnida. 

$ 4. Generalizations regarding the Embryogeny of the Artieulata, and Morpho- 
: logical Laws based on them. 

$ 5. The Embryogeny of Articulata, Mollusca and Vertebrata compared. 


$ 1. Embryogeny of the external organs of Aphis. 


IN the previous part of this paper T sketched so much of the development of the embryo 
of Aphis as was indispensable to the clear understanding of its reproductive processes; 
but it appears to me that the bearings of the embryogeny of this Insect upon morphology 
render it worthy of a more attentive and detailed consideration. 

It would be well worth while, indeed, to trace out the development of all the organs of 
this remarkable animal; but as I shall have for some months no leisure for labours invol- 
ving so great an expenditure of time, I will content myself for the present with a notice 
of some of the leading features presented by the development of the external organs. 

I have already stated, that one of the earliest changes in the germ of the young 
of the viviparous Aphis is the differentiation of its cellular mass into a central portion, 
which takes on the appearance and functions of a yelk, and which I termed “the 
pseudovitellus," and a peripheral coat or layer, the blastoderm. The blastoderm next 
becomes thickened posteriorly ; and in this thickening a division takes place from without 
inwards, so that it is separated into a posterior flap and an anterior portion, which are 
only continuous dorsally. It is the flap which is the rudiment of the abdomen, while 
that portion of the blastoderm against which it is folded stands in the same relation to 
the thorax. In front of this is the rudiment of the head, constituting by far the largest 
portion of the blastoderm. : pe 

Dorsally and posteriorly, the rudiment of the head is originally continuous with the 
thoracico-abdominal thickening; but a separation early takes place at this part, and em 
interval is occupied by the pseudovitellus, which here comes into immediate contact wit 
the pseudovitelline membrane. | à 

In an embryo 745th of an inch in length (Pl. XXXVII. fig. 5), this interval has serre 
so much, that the cephalic blastoderm does not extend on to the dorsal region at all, bu 
lies almost flat under the pseudovitellus, in the anterior half of the ventral region. 


In embryos „15th of an inch in length (Pl. XXXVII. fig. 6), I zei Zi the 


222 PROF. HUXLEY ON THE AGAMIC REPRODUCTION 


cephalic portion of the blastoderm beginning to extend upwards again over the anterior 
face of the germ, so as to constitute its anterior and a small part of its superior wall. 

This portion is divided by å median fissure into two lobes, which play an importa part 
in the development of the head, and will be termed the a procephalic lobes.” I have 
already* made use of this term for the corresponding parts in the embryos of Orustacea. 

. The rudimentary thorax presents traces of a division into three segments; and the 
dorso-lateral margins of the cephalic blastoderm, behind the procephalic lobes, have a 
sinuous margin. 

It is in embryos between this and råsth of an inch in length that the rudiments of 
the appendages make their appearance; and by the growth of the cephalic, thoracic, and 
abdominal blastoderm, curious changes are effected in the relative position of these 
regions. 

In embryos about z$5th of an inch in length (Pl. XX XVIII. fig. 1, la), the proce 
phalie lobes are so completely bent backwards as to lie close against the tergal surface 
of the rest of the cephalic blastoderm, so that no pseudovitelline granules can any more 
be seen in this region of the body. At the same time the lobes have enlarged, and extend 
back as far as the base of the fourth pair of visible cephalic appendages. Their infero- 
lateral angles are rounded and produced, forming an elevation which appears to be the 
rudiment of the eye. 

- Below the anterior extremity of the embryo, the blastoderm is produced on the median 
line into a tongue-like process (Jb), whose inferior part eventually becomes the labrum, 
while superiorly it sends a triangular process (the rudiment of the clypeus) into the 
interval between the procephalic lobes. 

Immediately behind the labrum, the blastoderm curves at first downwards, and then 
sharply upwards and backwards, to a little beyond the line of the posterior edge of the 
procephalic lobes. 

The whole of this portion of the blastoderm belongs to the head. In the re-entering 
angle between it and the labrum the mouth is placed; it is a small aperture, whence the 
cesophagus can be traced ascending and passing backwards with a gradual curve. 

Behind the cephalie region, the thoracie blastoderm passes nearly horizontally back- 
wards, and already presents traces of a division into its three somites. Its upper surface 
is elose to the pseudovitelline membrane, and consequently is covered by but a very thin 
layer of yelk-like granules. 

At the end of the rudimentary thorax the blastoderm is suddenly folded forwards, 8° 
that the sternal surface of the hinder part of the future abdomen is almost in contact 
with that of the thorax. Having come opposite the anterior edge of the thorax, it is bent 
backwards, at right angles to its previous direction, for a short distance,—the extreme end 
being finally folded parallel with this part, and with its apex towards the head. 

- The great mass of the yelk lies over the abdominal blastoderm, in the space left between 
it and the pseudovitelline membrane. The appendages present a singular and beautiful 
uniformity. No trace of the pigment of the eyes is to be seen. The next anterior pair of 
appendages (a £) are more slender and elongated than the others, and are bent inwards near 


* * Lectures on General Natural History,’ Med. Times and Gazette, 1856-7. 


AND MORPHOLOGY OF APHIS. ; 223 


their base so as to form a sort of elbow. In consequence of this, their terminal portions 
are more approximated than their distal ones, and lie close together and parallel. These 
appendages are the antennæ; and it is worthy of remark, that they arise from the proce- 
phalic lobes, or from the point of junction between them and the rest of the cephalic blas- 
toderm above the mouth. 

Behind these and behind the mouth (though the anterior pair are very close to that 
aperture, and might even be described as more or less lateral in relation to it) are three 
pair of short, similar, conical processes. Of these the anterior pair (1v') are the largest, 
and are the mandibles; the two other pairs are nearly equal: the anterior (v') represent 
what are ordinarily termed the maxillæ, but which might be more properly called 
“first maxille," since the second pair (vr), which eventually give rise to the so-called 
“labium,” are precisely like them, and, as Zaddach (2. c. infra) has shown, fully deserve 
the title of * second maxille.”’ 

Three pairs (vir, VIT, IX’) of short processes, unjointed and not much longer than the 
trophi, represent the thoracic limbs. 

The abdomen presents obscure traces of a division into segments. 

In an embryo y5th of an inch in length (Pl. XXXVIII. fig. 1, la, and 2), the pro- 
cephalic lobes have extended so far back as completely to cover the tergal region of the 
head, and even to pass a little beyond the line of the last maxilla posteriorly. The 
fold or depression separating the thorax from the head has become deeper; the antennæ 
have greatly elongated, and are bent downwards and inwards, so as to meet in the 
middle line below, and cover the mandibles. 

The first maxillæ are larger than the mandibles, and somewhat expanded at their ex- 
tremities. The second maxillæ are more slender; and their bases are in a line with those 
of the mandibles, while those of the first maxillæ have taken a more external position. 
Consequently, the bases of the trophi, instead of forming two nearly parallel rows as at 
first, are now arranged as a hexagon, whose outer angles are constituted by the first 

The thoracic members have greatly elongated, the hinder pair being the longest. 

In embryos 7th of an inch in length (Pl. XXXVIII. fig. 5), the blastoderm is found 
to have undergone a wonderful change. Instead of being folded upon itself ventrally 
by the flexure of the abdomen against the thorax, it has become completely extended; 
and so thoroughly has this extension taken place, that the abdomen is now convex in- 
feriorly. At the same time the blastoderm has grown upwards over the sides of the 
body, and roofs-in its tergal region. The head is closed by the union of the procephalie 
lobes, and is now, in consequence of the increased length of the body, proportionally 
much smaller. The pigment of the eyes appears in a few scattered granules towards the 

pig yes app 
posterior margin of the head on each side. 


If the appendages be examined as they become metamorphosed in a succession of spe- 


eimens intermediate in size between roth and „th of an inch, the antennæ are found 
The growth of the mandibles and 


gradually to increase in length and to become jointed. . hort thick 
first maxillæ in length, on the contrary, is suspended; and they remain as short thic 
.. tubercles (Pl. XXXIX. fig. 2), from whose inner surface a long chitinous filament gra- 


224 PROF. HUXLEY ON THE AGAMIC REPRODUCTION 


dually arises. These filaments, thickening and elongating, become the blades of the man- 
dibles and maxillæ. The growth of the second maxillæ makes up, by its excess, for the 
arrest of development of the mandibles and first maxillæ ; for having already approximated, 
their confluent or connate bases elongate as one great process, which extends back in the 
middle line between the thoracic legs, until at length it attains more than half the length 
of the body, and constitutes the well-known proboscidi form “labium ” of the Aphis*. 

The thoracie members or legs have elongated so much, that their terminations are bent 
inwards, to allow of their lying within the pseudovitelline membrane. Their character- 
istie subdivisions are indicated; and the terminal claws are beginning to be formed. 

From this size up to that at which the larvæ are born (Pl. XXXIX. fig. 4) (when they 
are less than 45th of an inch in length), the principal changes are the following. The 
appendages as compared to the body, and the latter as compared to the head, undergo great 
elongation. The anterior pair of thoracic limbs and its somite, the prothorax, come into 
very elose contact with the head, so that the cervical separation becomes obsolete, or is only 
indicated by a groove. The labrum and labium acquire their characteristic form and pro- 
portions; and the mandibular and maxillary setæ elongate, and take their final position. 

The “siphons,” so characteristic of the genus, appear as obtuse tubercles on the dorso- 
lateral region of the fifth abdominal somite. The little larva exhibits unequivocal signs 
of life, but still remains enclosed within its pseudovitelline membrane, to which another 
transparent and structureless envelope, fitting the body of the larva and all its limbs as 
a loose glove fits the hand, seems to have added itself. This second coat is, in fact, 
the embryonie integument, which is now being cast; so that the creature must undergo 
its first ecdysis either before, or immediately after, it is born. 'The head assumes its 
normal proportions. The corneæ become facetted ; and the pigment increases greatly in 
amount, assuming the form of an oval deep-red patch. The clypeus and the procephalic 
lobes unite, but readily give way when the head is crushed, and allow of the exit of the 
cerebral mass, which has in the meanwhile been produced by a differentiation of the inner 
substance of the procephalic lobes, just as the other ganglia are the product of the blasto- 
derm of their somites. 

If the account of the development of the external organs of Aphis which I have just 
given be compared with the statements of Külliker+ and Zaddacht, it will be found that 
there is a close correspondence in all essential respects between the embryogenic pheno- 
mena of at least three orders of Insecta—the Hemiptera, the Diptera, and the Neuro- 
ptera. And, considering the universality of the law that the embryogenic processes of 
members of the same class have a similar fundamental character, I do not doubt that 
eben, of all insects is, in its main features, a process similar to that described 

(phis. 


* Zaddach considers, 
confluent maxillæ, 
labial palpi. I do 
the fact. 

T De prima Insectorum Genesi, 1842. 

I Die Entwickelung des Phryganiden-Eies, 1856. 


from his observations on Phryganea and other Insects, that the labium is the product, not of 
but of an outgrowth of the sternum by which these are supported, the maxillæ remaining a8 the 
not deny that this may be the case in Aphis; but I have been unable to find positive evidence of 


AND MORPHOLOGY OF APHIS. 225 


; $ 2. Embryogeny of Mysis as exemplifying the Crustacea. 

But more than this, if we extend our researches into the embryogeny of the other two 
principal* classes of the Articulata, the Arachnida and Crustacea, we shall see that it 
presents a most remarkable agreement with that of the Insect. 

To illustrate this important truth, I might cite Rathke’s account of the development of 
Astacus as a type of crustacean embryogeny; but I prefer to speak from my own know- 
ledge, and I will therefore describe the development of Mysis, the Opossum-shrimp. 

The fertilized ova of this crustacean have a diameter of #sth to yyth of an inch, and 
consist of a yelk enclosed within a colourless and thin, but strong vitelline membrane. 

The yelk is composed of two elements—small and large yelk-masses, the former having 
about 3560th to zoooth of an inch average diameter, and being usually so closely 
wedged together as to appear polygonal. The latter are large (tooth of an inch or more), 
spherical, and imbedded in the mass formed by the smaller kind of yelk-granules. 

I was unable to detect any trace of endoplasts or cells in these ova, Acetic acid deve- 
lopes neither granules nor endoplasts in the yelk-masses. Upon the yelk thus constituted, 
the blastoderm makes its appearance as a rounded patch, which reflects the light more 
than the yelk, and therefore appears white by reflected, and dark by transmitted light. 
The contrast is greatly heightened by the addition of alcohol}, or of acetic acid. When 
the latter reagent has been employed, or even before, if the examination be very care- 
fully conducted, the structure of the blastoderm is seen to be widely different from that 
of the rest of the yelk. No yelk-granules are visible in it, but it appears to be very finely 
granular; and imbedded within it are numerous close-set vesicular endoplasts, having a 
diameter of +3455th to z555th of an inch. These usually contained many granules, some- 
times only one; but I cannot say I have been able to detect any definite nucleolus in 
them. 

The discoid blastoderm is thickest in its middle region, thinning off gradually on both 
sides, and internally is sharply defined from the substance of the vitellus. In the centre 
it exhibits a more or less marked depression. As development goes on, this depression 
becomes more and more marked, while the disk thickens and increases circumferentially. 
At the same time, the layer of yelk in immediate contact with the disk, and co-extensive 
with it, is found to have a somewhat different constitution from the rest. The globules 
are large, dark, and sharply defined, and acetic acid gives them a granular appearance, 
but developes no endoplast. 

The depression above alluded to now increases, so as to form a fissure which separates 
å small tongue-shaped process from the rest of the blastoderm, to which it nevertheless 
remains closely applied. This process is the rudiment of the abdomen, and in a front 
View it is rendered more distinct by several clear lines, which mark the commencement 


* I have no doubt that the Myriapoda will be found to exemplify the same morphological laws, with the exeeption 
of that relating to the total number of somites in the body, as their congeners ; but I find so much that rige 
in the existing accounts of their development, and so many points in their anatomy requiring re-investigation, 


Prefer for the present to be silent about them. i 
T Rathke, in his numerous embryological researches, appears to have constantly availed himself of this property of 


alcohol in order to render the blastoderm more distinct. 


226 PROF. HUXLEY ON THE AGAMIC REPRODUCTION 


of the future caudal bristles. In front of the end of the abdominal process, two minute 
conical prominences, at first marked by similar, but fewer clear striæ, gradually raise them- 
selves up on each side from the surface of the blastoderm and elongate, their apices being 
directed backwards. They are the rudiments of the antennules and antennæ. 

A delicate structureless membrane is now visible, covering these parts and the adjoining 
portions of the germinal membrane. It is produced into the terminal setæ of the end of 
the abdomen and of the two pairs of appendages, and is the commencement of the first 
skin of the larva*. 

The anterior part of the blastoderm is wider than the posterior, and is produced into 
two great lobes divided by a median fissure. These are the “ procephalic lobes," and have 
the same relation to the anterior division of the head as the corresponding parts in the 
embryo Aphis. 

In this state the embryo becomes a larva, for it bursts its vitelline envelope and lies 
naked in the pouch of the mother. The rudimentary abdomen is at the same time ex- 
tended, so that the little creature is now about 35th of an inch in length, and is very like 
a pear in shape, the stalk being represented by the abdomen, which is terminated by å 
flattened, bifid, spinulose fin. 

The whole larva is covered by a continuation of the delicate membrane already noticed 
on the limbs and abdomen. The blastoderm invests the abdomen almost completely, but 
in front it covers only a somewhat fiddle-shaped area on one face of the yelk. It is still 
more deeply bilobed in front, and the antennules and antennæ are much elongated. 
The larva next begins to grow, being doubtless nourished by the fluid contained in the 
maternal pouch; and at the same time its ventral region assumes a curve, contrary to 
that which it originally possessed, becoming more and more convex. 

The cephalic region is now clearly distinguishable; it occupies nearly one-half of the 
whole length of the body. The procephalic lobes extend upwards over the anterior face 
of the vitellus, and upon each a large rounded elevation, the rudiment of the ophthalmic 
peduncle, has made its appearance in front of the antennule. The latter, like the antennæ, 
elongate greatly, and become divided longitudinally, within the sheath afforded by the 
primitive integument, into their two terminal branches. 

Å slight constriction indicates the boundary between the antennulary and antennary 
sterna, and behind these, similar depressions mark off the surface of the blastoderm into 
seventeen additional segments. 


Attached to them are as many pairs of appendages, which in the youngest larve 
examined had the following form :— 
The first pair were rounded massive elevations, situated one on each side of the pit 
indicating the position of the oral aperture; from their anterior edge a short oval palp 
already projects. These are the rudimentary mandibles. 

The next pair, or first maxillæ, are small rounded elevations meeting in the middle line. 
The second maxillæ succeed, and are more elongated, three-jointed, and bent back parallel 
with one another. The maxillipedes and the thoracic ambulatory legs form one continuous 


* See, for illustrative figures of the development of Mysis, my ‘ Lectures’ above cited. 


AND MORPHOLOGY OF APHIS. 227 


series of similar appendages, all elongated and bent back against the sternal surface of 


the body. 

The abdomen is very short, but is clearly distinguishable from the thorax by its less 
complete segmentation, and by the rudimentary condition of all its appendages save the 
last pair. 

The blastoderm as yet extends only for a little way on the sides of the body. The pri- 
mitive larval integument still invests the whole body loosely, but passes smoothly over’ 
all the appendages, except the antennules and antennæ, which continue to be ensheathed 
by it. 

The larva remains in this general condition until it attains th of an inch in length, 
the principal differences in its later stages being the increased growth of the body as com- 
pared with the head, the completion of the dorsal surface by the upward extension of the 
blastoderm, and the gradual restriction of the yelk to the anterior part of the body. 

I have been unable to determine, as precisely as in Aphis, the exact share taken by the 
procephalic lobes in the composition of the roof of the head in the crustacean; but they 
assuredly extend over a considerable part of its latero-dorsal parietes. 

The carapace appears at first as a ridge-like process developed from the dorso-lateral 
region of the antepenultimate and preceding thoracic and cephalic somites, as far for- 
wards as the bases of the antennules. It is certainly not an extension backwards of the 
terga of any of the anterior cephalic somites, but is from the first continuous with, and 
developed from, the thoracic somites. 

It is needless to trace the history of the larval Mysis further, —what has been said suf- 
ficiently proving the close resemblance of its development to that of Aphis. 


$ 3. Embryogeny of Scorpio as exemplifying Arachnida. 

I have not yet had the opportunity of working out the development of an Arachnidan ; 
but the researches of Rathke* and Heroldt are so full and clear, that the omission is of 
little moment. 

Rathke's observations on the development of the Scorpion show that after, or even 
before, the blastoderm has extended over the whole yelk, a papillary elevation appears at 
one pole. It is the rudiment of the future abdomen, including under that term all the 
segments of the body behind that which carries the last pair of respiratory organs. In 
front of this, eleven pairs of closely approximated thickenings make their appearance; and 
then, at the sides of the sixth to the tenth pair of them, inclusively, counting from the 
rudimentary abdomen, papillary processes are developed. It is clear, from Rathke’s 
figures, that the anterior pair of thickenings are the “ procephalic lobes,” while the succeed- 
ing ones are the sterna of the somites between the mouth and the abdomen. The five pairs 
of processes thrown out by the five anterior of these are the great chelæ and the four pairs 
of ambulatory appendages. The antennæ make their appearance subsequently from the 


procephalic lobes (or their junction with the rest of the blastoderm) in front of the mouth. 


Tt is not expressly stated, but I do not doubt, from Rathke’s figures, that the upper region 


Of the head is formed, as in Insecta and Crustacea, by the union of these lobes. 


* Reisebemerkungen aus Taurien, 1837. + De Generatione summ d 


VOL. XXII. 


228 PROF. HUXLEY ON THE AGAMIC REPRODUCTION 


Rathke’s account of the number of rudimentary post-oral sterna would lead one to sup- 
pose that in the embryo one sternum is wanting. I believe, however, that the truth is, 
that the sterna of the genital and pectiniferous somites were already so much smaller 
than the rest in the embryos which Rathke chanced to examine, as to be regarded by him 
as one. 

I base this conclusion upon the condition of the nervous system, which consisted of 
eleven pairs of clearly distinguishable post-oral cephalo-thoracic ganglia; that is, of just 
the same number as in an embryonic Astacus. Of these, the four posterior were widely 
separated, and lay in the pulmoniferous somites; while the seven anterior pairs extended 
only a little way beyond the ambulatory appendages, and were united into a triangular 
mass. The anterior of these ganglia were the largest, the posterior the smallest. The 
anterior pair gave off the nerves to the chelæ. 

It would be difficult to obtain a more clear and conclusive proof than this, that 
the chelæ of the Scorpion are the homologues of the mandibles of the Crustacean, and 
that the succeeding somites, as far as the last pulmoniferous one, correspond with the fifth 
to the fourteenth somites, inclusively, of the typical Crustacean. The six succeeding 
somites are the homologues of the six abdominal somites of the Crustacean ; the aculeated 
sting corresponds with the telson; and the only difference presented by the pre-oral 
somites is that common to all air-breathing Articulata, viz. the sessile eyes, and the non- 
development of one of the pairs of antennæ. 


$ 4. Generalizations regarding the Embryogeny of the Articulata, and Morphological 
Laws based on these. 


From all these facts of development, I deduce the following morphological laws (some 
of which have already been enunciated for particular classes) for the Articulata (Insecta, 
Arachnida, Crustacea) generally. 

1. The first-formed rudiment of the embryo corresponds with its sternal surface, or 
with the side upon which the great centres of the nervous system are placed. Itisa 
neural rudiment. 

2. In the thorax and abdomen this neural rudiment grows up on each side towards the 
tergal region, or that on which the great centre of the circulation is placed. 

3. In the Articulate embryo, therefore, the neural wall is formed first, and gradually 
extends tergally so as to form the hæmal wall. d: 

4. The cephalic blastoderm very early undergoes a peculiar flexure, a greater or less 
portion in front of the mandibles being bent up at right angles to the rest, and even in 
many instances extending backwards, so as to constitute the entire hæmal region of the 
head. In these cases the top of the head is in reality a sternal, and not a tergal, surface. 

As a consequence of this flexure, the line of attachment of the bases of the eyes and 
antennæ is frequently altogether above that of the other appendages, so that they appear 
to be tergal, and not sternal, appendages. 

5. The anterior extremity of the cephalie blastoderm becomes early divided by ? 
median fissure, each lateral portion being a “ procephalic lobe.” In Insects the line of 
Junction of these procephalic lobes is the epicranial suture. 


AND MORPHOLOGY OF APHIS. 229 


6. In Insecta and Crustacea the head, in the embryo, is easily distinguishable from the 
rest of the body. In Podophthalmous Crustacea it is clearly seen to be composed of six 
somites, each possessing à pair of appendages; of these, the first are the eyes; the second, 
the antennules; the third, the antennæ; the fourth, the mandibles; the fifth, the first 
maxillæ ; and the sixth, the second maxillæ. 

In Insecta, on the other hand, only four pairs of appendages appear in the head, the 
eyes being sessile, and one pair of antennary organs remaining undeveloped. 

In the Arachnida it appears to me to be quite clearly shown by development that the 

anterior pair of appendages are antennæ ; the second pair, mandibles, with a hugely deve- 
loped palpus; the third pair, first maxillæ ; and the fourth pair, second maxillæ, converted, 
like the next two pairs of appendages, into ambulatory legs. 
. It follows, therefore, if we take the number of moveable appendages as the test, that in 
the Articulata never more than six, and never fewer than four somites enter into the com- 
position of the head. But is the number of moveable appendages a just test of the num- 
ber of somites entering into a part? No one will pretend that it is so in the abdominal 
and thoracic regions; and if we consider the head of Crustacea alone, we find the eyes 
becoming sessile, and one pair of antennary organs aborting, without the least reason for 
concluding that the typical structure of the head is altered. It seems to me, then, hardly 
a hypothesis to assume that the sessile eyes of Insects represent the appendages of a 
somite, since it is universally admitted that they do so in Edriophthalmia. But by this 
assumption we arrive at a still closer approximation of the different classes in regard to 
their cephalic structure; for all would, on this supposition, have either five or six cephalic 
somites,—the former number being invariably met with in the true air-breathers (though 
in many purely aquatic forms also), while the latter is found only in those which respire 
by means of gills. 

I repeat, I can see nothing in this generalization but a simple expression of the facts. 
But I would go a step further, and add to this the hypothesis, that in the Artieulata 
the head is normally composed of six somites, which are all fully developed only in 
Podophthalmia, Stomapoda, and some Branchiopoda; while in other Crustacea, some one 
or more of the pre-oral somites is more or less abortive, and in Arachnida and Insecta, 
the appendages of the first somite are sessile, and those of the second or third unde- 
veloped. Admitting this hypothesis, I find further, that of the six cephalic somites, the 
sterna of three (the mandibular and two maxillary) are always situated behind the mouth 
and on the ventral surface of the body. The position of the other three varies 3 but the 
most anterior or ophthalmic is always bent upwards in consequence of the cephalic flexure, 
and not unfrequently, as in Insects, constitutes the greater part, or the whole, of the dorsal 
region of the head. The next two, or antennulary and antennary sterna, may present cray 
variation from approximative parallelism with the axis, in Squilla, to extreme reflexion, as 
in Insecta and many Crustacea. : 

7. Nothing can be more variable than the number of the somites whence appendages 
are developed in the various classes and orders of the Articulata; and in the Myriapoda 
the total number of somites even is susceptible of an extreme amount of variation. But 


in the other classes it appears to me that there is a typical number of se 43 whence 
H 


230 PROF. HUXLEY ON THE AGAMIC REPRODUCTION 


but comparatively few forms depart either by way of excess or defect. Thus, if we leave 
out the Læmodipoda, all Podophthalmous and Edriophthalmous Crustacea have twenty 
somites, of which six are cephalic, eight thoracic, and six abdominal. In a very few 
Branchiopoda, and in Trilobita, there is more than the typical number of sonde but I 
believe that in all other Crustacea, where the number of somites is not twenty, it is less. 

The question of the typical number of somites in the body of the Insecta is one which 
has been much diseussed. But all the theories on this subject with which I am acquainted 
are, in my apprehension, vitiated by the mistaken view which their authors take of the 
composition of the Insect's head. Many seem to consider it to be a simple segment; while 
those who admit a multiplicity of segments, appear to be misled by the position of the 
eyes and antennæ, into regarding them as tergal appendages of the segments over whose 
sternal appendages they lie—as a kind of wings of the cephalic somites, in short. Again, 
it is supposed by many that the labrum and the lingua are the representatives of the ap- 
pendages of distinct somites, a conception which is at once negatived by the study of their 
development. 

Ås I have endeavoured to show, there are certainly five, and hypothetically six, somites 
in the head of Insecta; there are certainly at least three in the thorax; but the number 
in the abdomen has been as much disputed as the number in the head. Zaddach considers, 
as å general rule, ten to be the number of abdominal somites in Insect larvæ; Westwood 
and Newport enumerate eleven in some Hymenoptera, and this last is, I believe, the 
maximum number of somites which has yet been found in the abdomen. Now, if we 
assume the number of somites in the head to be six, the number in the thorax three, and 
the number in the abdomen eleven, we shall arrive at twenty as the maximum number of 
somites in the body of an Insect. 

This conclusion is in remarkably close accordance with the results obtained by M: 
Lacaze-Duthiers from his laborious and remarkable researches into the structure of the 
female genital apparatus of Insecta. M. Duthiers finds that the vulva always opens 
between the eighth and ninth abdominal somites, and that in Neuroptera, in Orthoptera, 


in most Hemiptera, and in Thysanura, three somites intervene between the vulva and ~ 


the anus, which is always placed at the very extremity of the body. There are thus — 
eleven abdominal somites, and, therefore, a total number of twenty, in these four orders. 

Some Hemiptera have the last abdominal somite abortive, and this appears to me to 
be the case in Aphis. In Coleoptera and Hymenoptera, the tenth and eleventh somites 
abort, nine only remaining : in Lepidoptera, finally, all three post-genital somites remain 
undeveloped. M. Lacaze-Duthiers’ researches tend to show that a fundamental unity 
prevails amidst those apparently most diverse apparatuses which we know as stings, 


borers, and ovipositors, and that they are always the result of a modification undergone 
by the ninth abdominal somite. 


I do not consider myself competent to give an opinion as to the details of the investi- 


gations to which I have just alluded, but I cannot refrain from expressing the belief that — 


the labours of future investigators will bring only a confirmation of their general 
accuracy. 


The only adult Insect, besides Aphis, which I have studied with sufficient care in refer- 


AND MORPHOLOGY OF APHIS. 231 


ence to these views, is the common Cockroach (Blatta orientalis), an insect which I 
can recommend as admirably adapted for investigation. Here it is very easy to find the 
eleven abdominal somites, and to satisfy oneself that the vulva is placed between the 
eighth and ninth, and that the two outer elongated pieces of the curious clasping appa- 
ratus for the ovisacs are formed by a modification of parts of the ninth somite. The smaller 
and inner processes, on the other hand, are clearly developed from the sternum of the 
tenth somite, while the lateral anal valves represent the eleventh somite. 

I have found that while the vulva opens between the eighth and ninth somites, the 
aperture of the spermatheca is situated on the sternum of the ninth, and that of the col- 
leterial glands on the sternum of the tenth somite. 

In the male the complex penis is formed by a modification of the tenth somite, and the 
aperture of the vas deferens is on the sternum of this somite, or between it and the 
eleventh. 

Weighing all these facts, the conclusion to which they point seems obvious, viz. that in 
Insecta, as in Crustacea, the typical number of the somites is twenty. 

I have shown above that the development of the Scorpion proves that there are seven- 
teen post-oral somites besides the sting (which is plainly the homologue of the telson in 
the Crustacea) in this Arachnidan. If we make the same assumption for the Scorpion 
as for the Insect, that one of the antennary somites is abortive, we shall have a total of 
twenty somites here also. The anatomy of the adult Scorpion appears to me fully to 
confirm this view. Beginning at the hinder end, we find, including the telson, six seg- 
ments behind those which carry the respiratory apertures. Of these there are four; and 
in the three posterior, the sternum has nearly the same length as the tergum; but in 
the anterior one the sternum is much longer than the tergum. Furthermore, these 
sterna at first seem to occupy the whole space up to the posterior boundary of the cepha- 
lothorax, while, on the dorsal side, two narrow terga lie between the tergum corresponding 
with the anterior sternum and the cephalothorax. 

It appears, therefore, as if there were two more terga than sterna in the abdomen; but 
on more careful investigation, the missing sterna show themselves as the supports of the 
pectines and of the genital aperture in front of these last curious organs. Indications of 
the terga which belong to the two posterior pairs of ambulatory limbs are clearly visible 
on the posterior part of the cephalothorax, and these limbs are strongly distinguished 
from the anterior two pairs by the absence of the triangular processes directed towards 
the mouth, which characterize the bases of the latter. Indeed, the anterior ambulatory 
legs, by means of these processes, take part in the formation of the oral cavity as com- 
pletely as do the maxillæ of any other Articulate animal. ae 

Another exceedingly natural demarcation between the two anterior and two posterior 
pairs of ambulatory limbs is afforded by the so-called ‘diaphragm’ which pees the 
thoracic from the cephalic cavity, and whose attachment corresponds with the interval 


between these two sets of appendages. 
In Galeodes, the two posterior pairs of ambulatory 
Segments. 


There is no necessity to enter into any disquisition upon the homolo 


legs are attached to distinct 


gy of the append- 


232 PROF. HUXLEY ON THE AGAMIC REPRODUCTION 


ages and the general uniformity in plan, so far as the anterior part of the body is con. 
cerned, in all Arachnida. But it may be asked, what becomes of the hinder thoracic and 
the abdominal somites in the Spiders and Mites? Without, at present, giving a positive 
answer to this question, I am inclined to think that the Spiders stand to the Scorpion in 
the relation of Lemodipoda to Amphipoda, and that many of their posterior somites are 
aborted. 

I do not doubt that many minor variations will be detected when the morphology 
of the Articulata is carefully examined; but I venture to think it a result of no small 
moment, if it can be proved that a Lobster, a Cockroach, and a Scorpion are composed 
of the same primitive number of somites; that the head in each consists of the same 
number of parts, and that the great differences are the consequence of the different modi- 
fication of the thoracico-abdominal somites, all fourteen of which bear appendages in 
the Lobster, while only three (or if we consider the genital apparatus in the light of 
appendages, five) are so provided in the Insect, and only two (leaving out of consideration 
the “ pectines”) in the Scorpion. 

8. I have elsewhere* explained at length my views with regard to the nature of the 
carapace in the Crustacea, and I will only repeat here, that there seems to me to be no 
constancy in its composition. The rudimentary carapace of Squilla is assuredly deye- 
loped from not more than four somites, the antennary, mandibular, and maxillary. In 
Apus, I doubt whether more than the six cephalic somites enter into its composition, 
In Cwma it is constituted by the cephalic and three anterior thoracic somites, in Mysis 
by the cephalic and six or seven anterior thoracic, and in ordinary Podophthalmia by all 
the cephalic and thoracic somites. 

9. Lastly, there are certain parts developed singly in the median line in the Articulata. 
Of this nature are the frontal spines of Crustacea, their telson, and the sting of the 
Scorpion, whose mode of development appears to be precisely similar to that of a telson. 
In the same category we must rank the labrum in front of the mouth, which in the 
Crustacea (at least) appears to be developed from the sternum of the antennary or third 
somite, the metastoma (or so-called labium or lingua) of Crustacea, and the lingua of 
Insecta, behind the oral aperture. 

However much these appendages may occasionally simulate, or play the part of, 
appendages, it is important to remember that, morphologically, they are of a very different 
nature, and that the confusing them with true appendages must tend completely to 
obscure the beautiful relations which obtain among the different classes of the Articulata. 


§ 5. The Embryogeny of the Articulata, Mollusca, and Vertebrata compared. 

I find it difficult to conclude this memoir without saying a few words on the resem- 
blances and differences between the embryogenic changes of the Articulata and those of 
the Mollusca and Vertebrata. Absolute and fundamental differences appear to me 2 
separate the members of these three classes almost from the first appearance of the germ. 

As we have seen, it is the neural side of the Arthropod which is first developed, while, 
80 far as I am aware, it is the opposite or hæmal side which is first formed in every 


* “Lectures,” Med. Times and Gazette, 1857. 


AND MORPHOLOGY OF APHIS. ; . 938 


Mollusk. The germ of the Arthropod becomes antero-posteriorly segmented ; the germ 
of the Mollusk never does so. From these two fundamental differences a multitude of 
others necessarily follow. 

The Articulate embryo is no less markedly separated from that of a Vertebrate animal, 
although in the latter, as in the former, it is the neural surface which is first developed ; 
for I know of nothing in the Articulate embryo to be compared with the primitive groove, 
the chorda dorsalis, and the dorsal plates of the Vertebrate*. They, like the amnion and 
the allantois, are, I believe, structures without a representative in the other two sub- 
kingdoms. 

There is perhaps, as Zaddach maintains, a certain analogy between the primitive seg- 
ments of the Articulate animal and the primitive vertebræ (** Urwirbel" of Remak) in the 
Vertebrate, but with the commencing differentiation into tissues the resemblance entirely 
ceases. The appendages of the Vertebrate embryo are more Molluscan than Articulate in 
their primitive mode of development. Notwithstanding all these great and real differ- 
ences, however, there appears to me to be one respect in which a most singular analogy 
obtains between the Vertebrate and the Articulate type :—it is in the construction of the 
head. 

Adopting, in some respects, the views of Prof. Goodsirt, I can recognize at least six 
more or less complete segments in the completely ossified Vertebrate cranium. It is 
clear that the Vertebrate mouth opens like that of the Articulate animal, though on the 
opposite side of the body, between an anterior and a posterior set of cephalic segments. 
In the interior of the cranium a no less natural boundary between the anterior and the 
posterior set of cephalic segments is afforded by the pituitary body and its fossa, when 
the latter exists. 

I find, again, in the cranio-facial bend of the base of the cranium in the Vertebrate 
embryo, something wonderfully similar to the cephalic flexure of the Articulate head, and 
in the cranial trabeculæ (Schädel-balken of Rathke), analogues of the procephalic lobes. 

While fully recognizing the fundamental differences between the Articulate and the 
Vertebrate type, then, I think we should greatly err if we overlooked such singular ana- 
logies as these. Future research will show whether they are or are not the outward 
signs of a deeper internal harmony than has yet been discerned, between the Articulata 
and Vertebrata. 


Since the present memoir was read to the Society, some additional facts of importance | 
have come to my knowledge. In the first place, my friend Mr. Lubbock, having under- 
taken to work out the development of Coccus, was led thereby to search for what I have 
called * ovarian glands" in other insects. His results will be published at length else- 
Where; but he permits me to say that corresponding organs exist in all Lepidoptera, 
Hymenoptera, Geodephagous and Hydrodephagous Coleoptera, Diptera, and most sn 
roptera, while they are absent in Orthoptera, Pulex, Libellulidæ, &c., and are all terminal, 


* I therefore by no means agree with what Zaddach says on this subject, or with regard to the homologue of the 
amnion in Artieulata. 
t As expressed in the Edinburgh New Philosophical Journal, 1857, p. 118 ef seg. 


234 - PROF. HUXLEY ON THE AGAMIC REPRODUCTION 


instead of forming groups between the egg-germs, in the non-geodephagous Coleoptera 
and Hemiptera. They have been figured in Lepidoptera by Herold, Meckel, Thompson, 
and Stein, in Diptera by Stein and Leuckart, and in Coleoptera by Stein. 

Secondly. In September last I received the fourth Part of the fourth volume of Moles- 
chott’s ‘ Untersuchungen,’ which contains a long and remarkable Essay by Leuckart, 
«Zur Kenntniss des Generations-wechsels und der Parthenogenesis bei den Insekten.” 
The first article in the memoir is on the “ Alternation of Generations in the Aphides,” 
The author describes at length, and figures, the female reproductive organs of Aphis Padi; 
and although the arrangement of these organs is somewhat different from what obtains in 
my Vacuna, I am happy to say that his account of the ultimate structure of the ovaries 
essentially coincides with mine. The view which Leuckart takes of the relation of the 
ova and agamic germs (p. 346) is also in close agreement with my own. I lay the more 


weight upon these coincidences because Prof. Leuckart’s observations must have been 


made at the same time with, and were of course wholly independent of, mine. 

Lastly, not having the works of either Kaltenbach or Koch at hand when my memoir 
was read, I abstained from attempting to give the specific names of my Aphides. I haveno 
doubt now that the viviparous form is the Aphis Pelargonii of Kaltenbach, especially 


as my friend Mr. Dallas, who has paid particular attention to the Hemiptera, is of that 


opinion. The oviparous female resembles so much in form and habit the Vacuna dryo- 
phila of Schrank, that I have little doubt it is really that species, though, when carefully 
examined, the antennæ are found to have six unquestionable joints, and seven, if the 
swollen base of the last division of the antenna is to be regarded, as I believe it should 
be, as a distinct joint, The eyes also have a small and inconspicuous tubercle; and the 
promuscis is not nearly so long as either Kaltenbach or Koch states.— Nov. 16, 1858. 


DESCRIPTION OF THE PLATES. 


Tar.. XXXVI. 


Aphis Pelargonii. 


The letters have the same signification throughout. The fractions indicate the measured size, 
in parts of an inch, of the objects. 


Fig. 1. The three anterior chambers of a pseudovarium: a. the apical chamber; 8. the second} 
©. the third. a. Pseudovarian ligament; b. wall of the pseudovarium; c. its epithelium - 
d. periplast or homogeneous matrix of the apical chamber; e. clear vesicle; f. its endoplast, 
the two corresponding with the germinal vesicle and spot of an ovum; g. a pseudovum par 
tially detached, its periplast greatly enlarged; €. its vesicle, whose endoplast is invisible; 


h. blastoderm ; i. pseudovitellus. 


Fig. 2. Terminal chamber of a pseudovarium, with the second chamber beginning to be formed in 


+ 


AND MORPHOLOGY OF APHIS. 235 


consequence of the enlargement of the pseudovum (9), which is about „45th of an inch in 
diameter. 

Fig. 3. The pseudovum is still more enlarged, and the second chamber is nearly distinct. The vesicle,’ 
e, remains, and exhibits certain indistinct granules in its contents. The cells of the blasto- 
derm of c measure about 3-,;th of an inch in diameter. 

Fig. 4. The second chamber is quite distinct from the first, and contains a mass (g) in which no clear 
vesicle could be discovered: this mass became clearer and irregularly areolate by the action of 
water. 

Fig. 5. The cellular germ-mass. The cells or clear cavities have a diameter of about 33 sth of an inch ; 
their endoplasts are hardly more than :5455th of an inch in diameter. 


Tas. XXXVII. 


Aphis Pelargonii. Letters as before. 


Fig. 1. A portion of the blastoderm and pseudovitellus of an unaltered germ, only ;i.th of an inch in 
length, but otherwise like the preceding. The clear vesicles measured „„;sth of an inch; the 
endoplasts r7åzsth. 

Fig.2. A germ extracted from its chamber and treated with acetic acid. It has no pseudovitelline 
membrane. 

Fig. 3a. A germ extracted from its chamber. It is enclosed within a pseudovitelline membrane (4); and 
its pseudovitellus is arranged in obscure spheroids, of which one is represented in 3 4. acted on 
by water. Its granules are about s¢;sth of an inch in diameter. 

Fig. 4. Germ gzl;th of an inch in length. The cells of the posterior end (3) present a sort of break (1), 
and the blastoderm on one side is greatly thickened. The thickened portion offers an indication 
of a division (m). "The anterior end (y) is also somewhat thickened. 

Fig. 5a. Germ „45th of an inch, enclosed within its pseudovitelline membrane: n. rudiment of the 
abdomen; o. of the thorax; p. of the head; /'. gap corresponding with Z, and now filled by the 
pseudovitellus; g. inner layer of the germ; 7. that portion of it which will become the pseud- 
ovarium. 50. Diagrammatic view of the same, viewed from above. 

Fig. 6. Lateral view of a larger germ without its pseudovitelline membrane. "The anterior part of the . 
cephalic blastoderm (p) has extended upwards, and constitutes the procephalic lobe p'. The 
rudiment of the pseudovarium (r) is still more distinct than in the preceding. 


Aphis Pelargonii. 


Fig. 1. Embryo enclosed within its pseudovitelline membrane. The pseudovitellus has aggregated over 
the abdomen, and more or less completely left the thorax. Letters as before, se the 
first larval integument; lb. labrum; at. antenna; 1v. mandible; V. first maxilla; vr. second 
maxilla; vir. first, viri. second, and 1x’. third thoracic leg. 14. The same embryo seen from 


below. ; ld d th 
Fig. 2. Embryo of the same size, viewed from below and the side, the blastoderm unfolded, an » 


appendages separated. : fih iem 
Fig. 3. Highly magnified view of part of the pseudovitellus, and of the rudiment of the pse : 
: in an embryo th of an inch in length. 
Fig. 4. Embryo «th of an inch, enclosed in its pseudovitelline membrane. 
Fig. 5. Embryo 74th of an inch, in its pseudovarian chamber. 
VOL. XXII. 


i 


À. 
236 PROF. HUXLEY ON THE AGAMIC REPRODUCTION AND MORPHOLOGY OF APHIS, 


e 


Tas. XXXIX. 
Aphis Pelargonii. 


Fig. 1. a. Nearly full-grown fœtus, extracted from its investments, and somewhat unfolded: 4. anus, 
whence the alimentary canal is seen taking a curved S-like course to the mouth. B. Terminal 
chamber of one of the pseudovarial cæca of this embryo. 

Fig. 2. The mouth of this embryo seen from below. The “labium” (vr') already appears as a large single 
process bilobed at its free end. 

Fig. 3. Side view of the head of a similar embryo, showing the relative position of the different append. - 
ages and the course of the æsophagus. ; 

Fig. 4. A nearly full-grown fætus in its pseudovitelline membrane: r. the pigment of the eye; s. rudi- - 
mentary siphons. : 

* Fig. 5. A partially diagrammatic figure of the wingless viviparous form of Aphis Pelargonii. The Roman - 
numbers indicate the typical somites of the body and their appendages; the other numbers — 
mark the abdominal somites. A, Anus; c, genital aperture; s. siphon. Lo 


på 
Tap. XL. 
Reproductive Organs of the oviparous Aphis (Vacuna dryophila). 


Fig. 1. The female organs entire. One ovarian cæcum only is represented ; and I have purposely selected — 
one of those, the ovarian glands in whose apical chamber are very similar, at first sight, to ova. — 
A. Anus; B. vulva; c. vagina; p. oviducts j E, F, G, H, 1, K. chambers of the ovary ; L. ovarian Es 
glands; m. colleterial glands; x. spermatheca; 7, 8. seventh and eighth abdominal sterna. — 
Fig. 2. The three anterior chambers of an average ovarian cecum. Letters as before, with the addition — 
of—o. germinal vesicle of the nascent ova in the terminal chamber (x) ; o!. germinal vesicle of 
ovum in 1; and o". of ovum in H ; P. epithelium; -g. cord-like secretion of ovarian gland, l; 
l. inner capsule of ovarian gland. > Ne 
Fig. 3. The end of another ovarian cæcum, showing very distinctly the apparent continuity of the cord, — 
9, with the ovum in the third chamber. The granules of the viscid vitelline mass (which is | 
surrounded by no membrane) are so numerous as to hide the germinal vesicle. a. 
Fig. 4. a. Posterior extremity of the ovum unaltered: r. chorion; s. tubercular elevation; f. append- 
age; lg its gelatinous, investment; v. rod-like bodies imbedded therein ; v. the same more 
tented, B. Anterior end of the ovum after the action of potash; y. papillary elevation ; 
* internal markings of the chorion (r); w. vitelline membrane; +. vitellus. c. Posterior 
extremity treated in the same way: y, micropyle ? 3 
a -o HUE 


Trans Limn. Soo Vol XXI, tab. $4. p 


Trams Lim. $00. Vol. XAIL, tab. 37, fv 


| ex) 
20/9000 


v» 


a : GJarman so.. 
— TH Hud Nat dei? 


Trams Tinm Soo Vol XXT, tab. 38. p. 


GJarmam sc. 


Tinm. Soo. Vol XXII tah. 39. jr. 


Trans. 


Frams. Linn. Soo Vol XXIL tab.40, p. 


G Jarman sc. 


THH ad, Nat del 


à 32 


XVI. On the external Anatomy and Natural History of the Genus of Annelida named 
Palolo by the Samoans and Tonguese, and Mbalolo by the Fijians. By Joux DENIS 
MACDONALD, Assistant-Surgeon of H.M.S. * Herald,’ employed on Surveying Service 
in the South-western Pacific, under the Command of Captain H. M. DexHam, R.N., 
FRS. Communicated by G. Busk, F.R.S., F.L.S. 


Read December 3rd, 1857. 


On the examination of specimens of a remarkable Annelidan, obtained at the Navigator 
Islands (Samoa), and presented to the British Museum by the Rev. J. B. Stair, Dr. Gray 
founded a new genus, which he called Palolo, adopting the native name of the animal. 
It is thus characterized :— 

* Body eylindrical separated into equal joints, each joint with a small tuft of three or 
four spicula on the middle of each side. Head? Last joint ending in a couple of ten- 
tacles. Eggs globular. 

* Palolo viridis, n. sp. Green, with a row of round black spots down the middle of the 
dorsal? surface; one spot on the middle of each joint.— Habitat. Navigator's Islands.” 

Dr. Gray found his specimens so mutilated, that he could not detect à single head 
amongst all the broken pieces; and consequently this essential part remained without 
description. He seems to refer his want of success in this respect to mere casualty. It 
is very remarkable, however, that we at first experienced the same difficulty, and only 
obtained a single head, though we carefully examined considerable numbers of this 
- species in the Fiji Group, where it makes its short annual appearance at a period which 
the natives predict with unerring precision by observing the phases of the moon, as at 
Samoa. The comparison of specimens from both these localities proves them to be speci- 
fically identical; moreover, the Fijian name for the animal is Mbalolo, Mb in that 
language being substituted for the Samoan P. 

The great antiquity of this name is attested by the fact, that the parts of the year 
nearly corresponding with our months of October and November are respectively named 
Mbalolo lailai (little), amd Mbalolo levu (large). The latter, as its name implies,, 16 
distinguished by the appearance of the Mbalolo in such vast numbers, that it is collected 


by the natives as a dainty article of food, and is so much prized that formal presents of 


it are often sent considerable distances, from certain chiefs, to others whose oe 
nions do not happen to be visited by the Möalolo. The Fijians entertain supers’ tb! 


ideas connected with it; but further notice of these would be foreign to our present - 


purpose. ee es Rp = a et | 
Reverting to the separation or absence of the anterior extremity of the Mich (e 
first noticed by Dr. Gray, and subsequently by us, in the specimens including ditteren 


species) at the regular period of visitation, this ra is too remarkable iud upon 


+ 


238 MR. J. D. MACDONALD ON THE ANATOMY AND 


coincidence or casualty. It would rather seem to indicate the fulfilment of some im- 
portant end in the propagation of the species. Mr. F. M. Rayner suggested to me that 
the phenomena of reproduction in the case of Tenia and Bothriocephalus afford a curious 
parallel to the circumstance just alluded to with reference to the Mdbalolo. The trans- 
verse fission in the latter case is evidently connected with the dispersion of the ova, rather 
than the development of new individuals from the pre-existing materials of the animal’s 
body, as in Nereis, &c. 

I had the good fortune to discover a single head of the Mbalolo,—and the only one to 
be found amongst a large bottleful of bodies and tails collected for me by my esteemed 
friend, the Rev. S. Waterhouse, Wesleyan Missionary, Fiji. The joints of the body, to the 
number of about twenty, remaining in connexion with the head, were considerably smaller 
than those that would succeed them were the specimen perfect ; besides which, the aciculi, 
of two sorts, were more numerous in the little bundles springing from the lateral tuber- 
cles. The dark spots and characteristic markings of the dorsal surface were also very 
faint, or scarcely distinguishable. The head itself was very little narrower than the joints 
of the neck, blunt and rounded, with a slight emargination in front. Eyes two, placed one 
on either side of the upper surface, including, in the space between them, three conical 
tentacula, of which the central is the longest, and projects a little beyond the head. 
The mouth was inferior, subterminal, and armed with two pairs of jaws—those of the first 
pair being sickle-shaped and simple, and those of the second broad and jaw-like, having 
a curved external outline and a series of dental points on their opposable border. The 
tissues in the neighbourhood of the jaws appear to be much indurated ; and one structure 
in particular is worthy of notice, that its true nature, if not already known, may be inves- - 
tigated in the neighbouring genera. It consists of two slightly diverging series of scale- 
like plates overlapping one another from before backwards, in which direction also they 
gradually increase in size. The free edges of the plates are directed backwards; and as 
distinct muscular bundles may be traced into their deep surface, it is highly probable 
that they are capable of elevation and depression, acting, so to speak, as a prehensile 
palate, opposable to the jaws. 

The typical elements of the lateral appendages of the body-segments (often so distinctly 
seen in allied Annelida) appear to have become blended together, more or less, so as to form 
a single setigerous tubercle, transmitting, however, as is usual, two characteristic kinds 
of setæ, and bearing a simple papilla-like, dorsal cirrus above, and a somewhat smaller 
ventral cirrus below,—the former lying near the outer extremity of the tubercle, and the 
latter somewhat hearer the base. The repetition of these cirri may be traced backwards, 
cae all = annuli, to the penultimate joint, in which they are quite suppressed ; but 

om reappear In the anal segment, and the ventral cirri in particular, having attained 
"1 considerable length, project posteriorly like those of better-known Nereids. Besides the 
En "i Puig sty lets of the feet (exhibiting so much sameness of character in all the Anne- 
E en, Bere ue = of M balolo, as above noticed, are of two distinct 
ilis Sith où tin dam ane el crate eo Me Like ee oompeomed ONE 
is much stouter and eng and terminating in an exquisitely fine point; while és 
r, with a small claw-like terminal appendage having two 


minute conical teeth on its concave edge. 


NATURAL HISTORY OF PALOLO, OR MBALOLO. 239 


=- 


The bundles of sete generally consist of two 


or three of each of these forms, the latter holding a position inferior to the former. 
Mbalolo has been supposed to exhibit an alliance with Arenicola; but the anatomical 
characters above given refer it, very clearly, to the Nereide. 


Fig. 
Fig. 


Fig. 
Fig. 
Fig. 


Fig. 


Fig. 
Fig. 
Fig. 


= 0. 


QD 


DESCRIPTION OF THE PLATE. 
Tas. XLI. 


. Portion of Mbalolo. Natural size. | 
. Magnified figure of the head, with its three frontal tentacula and eyes, each of which appears to 


consist of two distinct organs closely approximated. The position of the retracted jaws is 
shown in the central dark space behind the tentacula. , 


. The double set of jaws, with their basal plates. 
- The palatal ? erectile plates noticed in the text. 
. Three of the feet, seen from below: a. setigerous tubercle; 5, b, b. dorsal cirri; c, c, c. ventral 


cirri; d. muscles which protrude the aciculi, acting through the basal stylet, e; f. retractor 
muscles, derived from the lateral border of a longitudinal muscular band, 9, running along the 
ventral surface. 


. Posterior extremity of the Mbalolo, dorsal aspect: a. terminal dorsal cirri; 5. terminal ventral 


cirri, or oval styles so called. 


- Basal stylet, or that which affords support to the lateral tubercles. 
. Whip-like aciculus. 
. Clawed aciculus. 


Trans Lian doc Vol. I PLATT 


2 


[1241] 


XVII.—On the probable Metamorphosis of Pedicularia and other forms ; affording pre- 
sumptive evidence that the Pelagic Gasteropoda, so called, are not adult Forms, but, 
as it were, the Larvæ of well-known genera, and perhaps confined to species living in 
deep water. By Joux Denis MACDONALD, Assistant-Surgeon of H.M.S. * Herald,’ 
employed on Surveying Service in the South-western Pacific, under the command of 
Captain H. M. Drennan, R.N., P.R.S. Communicated by Q. Busk, F.R.S., F.L.S. 


Read February 18, 1858. 


IT has been long known that certain genera of Gasteropoda, which are shell-less in the 
adult state, possess both shell and operculum not only while yet within the ovum, but for 
some little time after their liberation, and that ciliated vela precede the more perfect de- 
velopment of the foot. This is especially true of the Nudibranchs; and Janthina, which 
exhibits so near an approach to them in its organization, merely loses the little operculum 
of its embryonic condition, while the spiral shell is retained. But a more striking change 
than this occurs in the case of the genus Pedicularia, if my observations be correct; for 
I believe that I have identified the anatomy of a certain species whose shell presents a 
beautifully cancellated nueleus, with that of one of our little pelagie Gasteropods also having 
à cancellated shell, but presenting an aperture so closely resembling that of Cheletropis as 
to have misled me in naming figures of its labial and lingual dental organs, given in illus- 
tration of a former paper. I am, however, now in a position to prove that the oral 
teeth of Cheletropis are not lateral as in the little Gasteropod just referred to, and that its 
lingual ribbon is triserial and constructed on the type of that of Murex, Purpura, Turbi- 
nella, Ricinula, and such genera,—not septiserial as in Pedicularia and the little animal 
Which I believe to be its fry. In the latter case moreover, it must be mentioned that the 
external series of uncini are often rudimentary, or not at all apparent,—a fact which is 
clearly in aecordance with the common law of development of the lingual ribbon (as 
noticed in a previous paper, with an illustrative figure selected from the fry of Cypræa 
umbilicata). Without reference to the contained animals, the most acute conchologist 
could only regard Oheletropis Huxleyi and its little oceanic ally as distinct species of 
one genus, although we now know that it would be a violation of the simplest gr 
principles to place them even in the same family. Here, to a certain extent, similar con- 
ditions have arisen out of similar necessities in two otherwise very dissimilar beings. 
The final modelling, and thickening of the lip, moreover, afford no proof whatever that 
these shells have attained their adult state; for this change is usual in other cases, as in 
Carinaria, where its further progress is more easily traced, not only as an dona of x 
close of one stage of development, which had been going forward during the ud = | 
of the active life of the being, but as establishing a basis upon which the characteristic 


lines of growth of the future shell are laid. | : 
Macgillivraia pelagica possesses the labial plates of Natica or Triton, and the lingual 


242 MR. J. D. MACDONALD ON THE PROBABLE METAMORPHOSIS 


teeth of Ranella or Dolium, with the spherical otolithes common to all; and it is highly 
probable that it will ultimately prove to be the young of some neighbouring genus. The 
little shell itself is perfectly identical with the summit of the spire of a species of Malea 
which is very common in the South Seas. I regret much that I have not yet had an 
opportunity of examining the animal of the latter for the purpose of more accurate 
comparison; but this much I can say—that the lingual teeth of Doliwm, which cannot be 
far removed from Malea, very closely resemble those of Macgillivraia. 

I formerly figured and described a pelagic Gasteropod with six ciliated arms, and which 
I believe may be safely referred to the Tritomidæ. I now offer the portrait of another 
species, obviously belonging to the same genus, though occupying a very differently 
shaped shell. In the paper alluded to I also noticed a minute Natica, with mentum, 
lateral labial plates, septiserial tongue-strap, and, in fact, only differing from other or at 
least full-grown members of the genus in the possession of a cervical circlet of ciliated 
arms, and a vesicular float; but I am now quite satisfied that I have identified this . 
species with the nucleus of a handsomely spotted Natica occurring in deep water. 

It is very remarkable that the characteristics just mentioned should be present in all 
the little pelagic Gasteropoda, while their internal anatomy unmistakeably refers them to 
very different families. But if, in connexion with these particulars, we take into con- 
sideration the large size of the fatty globules contained in the sacculi of the liver, indica- 
tive of almost incipient development, and the total absence of ova or spermatozoa (at | 
least so far as the matter has yet been investigated), it can scarcely be doubted that these 
little creatures are not perfect, but transitional forms, the further development of whose 
head and proboscis, coincident with the shedding of the ciliated arms and a general adap- 
tation to a new sphere of existence, may, with all propriety, be regarded as a veritable 
metamorphosis. It also appears to me that a bold indication of a developmental change 
such as this, is exhibited in the abrupt transition from the marking of the young shells. 
into the very different and characteristie sculpturing of the adult state of the corresponding 
species, —an occurrence which takes place long subsequent to the period at which, in 
some Instances, a sinistral nucleus merges into a dextral shell. 

Since the above remarks were written, as if to afford them additional support, a suc- 
cessful haul of the towing-net has made me acquainted with another little genus, of which 
I had no previous knowledge. It was invested with a thick and globose cartilaginous 
envelope, with an irregular subterminal aperture, not very unlike that of the cartila- 
gious covering of the Pteropod Burybia. 
| On endeavouring to remove the animal from this singular shell, I found that it was 
xm bes eiii ug ca, probably the egg-capsule itself; for the little creature A 
CINE: se : cupying the true shell, which was yet so membranous and nn 
the tiny occupant ni ee ficte ui Å 
HARAS : à SEREN upon the ocean of life, will be at once apparent on = 
OP Rea "ws rx. Ha entacula were of considerable length, with ocelli at their oiam 

„~ mouth was encircled by a deeply cleft calyx of four segments, with richly ciliated 
a — — that of Atlanta, minus the swimming plate, " 
ough rudimentary creeping dise and a broad three-lo 


OF PEDICULARIA AND OTHER FORMS. 243 


posterior part bearing no operculum. The margin of the mantle, as in Jasonilla, was 
rather thick; and a pectinate gill extended along the dorsal region. The mouth was armed 
on each side with a beautifully set pavement of sharp-pointed labial teeth; and the lingual 
ribbon, supporting 7 series of members, exhibited the closest relationship to that of 
Hyponyæ and Pileopsis. Finally, single spherical otolithes were apparent through the 
transparent pedicle of the foot. Here is an incontrovertible case of a young oceanic 
Gasteropod still undergoing development, but manifesting every essential anatomical 
character present in Macgillivraia and Cheletropis, which we have hitherto regarded as 
perfect forms. 


EXPLANATION OF THE PLATE. 
Tas. XLIT. 


Fig. 1. Little cancellated pelagic shell, closely resembling the minute spire of Pedicularia. 

Figs. 2 & 3. Enlarged back and front views of the same. 

Fig. 4. Small portion of the shell, more highly magnified to show its characteristic sculpturing agreeing 
with that of Pedicularia. 

Fig. 5. The operculum. 

Fig. 6. One transverse row of the lingual ribbon. 

Fig. 7. A species of Pedicularia brought up from a depth of 40 fathoms, on a small Madrepore, 1 mile and 
a half off Elizabeth Reef. Lat. 29° 55' 27" S., Long. 159° 2! 54" E. (The natural size is seen 
to the left.) 

Fig. 8. One transverse row of the lingual teeth of the same, for comparison with fig. 6, which is not so 
fully developed, but in other particulars very similar. 

Fig. 9. Enlarged figures of a minute pelagic Gasteropod, having six ciliated arms and a clawed operculum, 
allied to a species previously figured and described. 

Fig. 9a. Portion of mantle with ciliated bands and processes, more highly magnified. 

Fig. 10. Three-quarter view of the same shell. 

Fig. 11. Lateral labial teeth. 

Fig. 12. Lateral view of small pelagic Gasteropod enclosed in a stout cartilaginous case. 

Fig. 13. Front view of the same. " 

Fig. 14. The animal removed from the outer case and slightly twisted in its rudimentary shell. 

Fig. 15. The auditory capsule. 

Fig. 16. Lateral labial teeth. 

Fig. 17. Portion of lingual ribbon. 


VOL, XXII. er 


Trans Linn soo ol NTP XI 


PEDICULARIA &° 


f 


HO 


ZEA 


WhWest mp 
ETG 14 
TD Macdonald RN Ad nat dei Tuffen West lith 


[0248 «Jo 


XVIII.—On the Anatomy of Eurybia Gaudichaudi, as bearing upon its Position amongst 
the Pteropoda. By Joux Denis MACDONALD, Assistant-Surgeon of H.M.S. * Herald, 
employed on Surveying Service in the South-western Pacific, under the command of 
Captain H. M. Denna, R.N., F.R.S. Communicated by G. Busk, F.R.S., F.L.S. 


Read February 18th, 1858. 


ONE of the most constant products of the towing-net, in the S.W. Pacific, is the little 
Pteropod Zurybia Gaudichaudi. It is often captured in the daytime, which is seldom 
the case with other members of its class. 

The enveloping mantle forms a sac of an oval figure and scarcely exceeding $th of an 
inch in length, with an anterior transverse subterminal slit on the ventral side, giving 
passage to the head with its tentacula, a rudimentary foot, and the swimming-fins. It is 
composed of large spheroidal cartilage-cells, in which the nuclei are distinctly visible, and 
a sparingly interspersed fibrous tissue, opposing an almost insuperable obstacle to the 
study of the internal anatomy of the animal. 

The Eurybia may be said to possess a distinct head and neck, the head bearing on 
either side a large tapering and gently curved tentaculum, with a small nipple-like pro- 
cess at the inner side of the base. All these appendages are richly ciliated, the cilia being 
generally disposed in parallel, longitudinal lines, and exhibiting a dextral, undulatory 


' motion. 


The mouth is a vertical opening, with moderately prominent lateral lips, just within 
the borders of which may be noticed, according to the age of the animal, one, two, or 
three longitudinal series of small square plates, with a delicate eutting edge near the 


outer part of each. 


The lingual ribbon is somewhat more lengthy than that of any other Pteropod with 


which I am acquainted. It presents a single row of simple, lancet-shaped teeth in the 


rachis, and a single row of fang-like uneini in each pleura. The buccal mass is globose in 
figure, and composed of fibro-cartilage with investing museular fibres. + 
Å very distinct though rather small foot springs from the under surface of the ly, 
considerably behind the head, having a flat creeping-dise, with a Ben on 
and a pointed posterior extremity, and in many particulars closely resembling the foot o: 


the true pelagie Gasteropods. i 
The swimming-fins are unquestionably the epipodia of this foot, arising by a subeylin- 


drieal base, just above the lateral border, and near the middle of its eei eg 
at first diminish a little in size, and then gradually expand to form a br y 
compressed paddle, widely emarginated at its extremity. Fe 

The æsophagus holds a dorsal position, and is distinguished en parts of the 
alimentary canal by the thickness of its walls and its richly eiliated lining. 


i i ior of which 
The stomach is large, consisting of several wide sacculated portions, the PE of whic 


2— 


246 MR. J. D. MACDONALD ON THE ANATOMY OF EURYBIA GAUDICHAUDI. 


is beset with large hepatic lobuli containing cells distended with rich brown-red, yellow, 
and green fatty globules, imparting to them a beautifully variegated appearance. No 
gastric plates or teeth are visible, though universally present in the true Thecosomata, 

- Having left the stomach, the intestine soon curves forwards to terminate anteriorly near 
the generative openings, on the right side of the body. 

The salivary secretion is furnished by two irregularly lobulated organs, lined with rather 
large secreting cells, and lying one on either side of the cesophagus as it emerges from the 
buccal mass. 

The heart occupies an antero-dorsal position, and its long axis appears to lie transversely; 
but I have not been able to detect a respiratory organ, or even the actual distribution of 
the blood-vessels. _ 

The main centres of the nervous system are the subæsophageal ganglia, which are well 
developed, and support the auditory capsules containing vibrating otokonia on their 
inferior surface. 

The ovarium consists of a fan-like expansion of lobuli, on the borders and extremities 
of which the ova are clustered within the enveloping sheath. From this expansion a 
number of tubuli arise, and ultimately unite to form a wide oviduct distinguished from 
the spermatic duct and intestine by the large longitudinal and nucleated fibres of its mus- 
cular coat. A large cecal appendage, distended with a bright-orange and plastic fluid, 
appears to open by a short duct at the same external orifice. This organ is probably the 
homologue of the so-called spermatheca of the Pulmoniferous Gasteropods ; and its con- 
tained fluid offers a remarkable analogy to that occurring in the ovarian sac of some 
Cephalopods. 

The lobuli of the testis lie upon and coincide so closely with those of the ovarium, that 
it is difficult to distinguish them; but the stout vas deferens, arising from the point at 
which the divisions converge, soon communicates, above and a little to the right of the 
ovary, with a spheroidal sac or vesicula seminalis, which may be readily seen from 
without, through the cartilaginous mantle. The spermatic duct thence proceeds forwards 
in company with the oviduct, and enters the neck of the external male organ, to termi- 
nate in a perforated dart or rigid point, which occupies the fundus of the inner protrusible 
membrane of a double-walled pyriform sac having an external orifice near that of the 
oviduct, on the right side, behind the corresponding swimming-fin. 

The spermatozoa of this Pteropod are remarkable for their great length and peculiar 
looping and spirally twisting movements, as illustrated in Pl. XLIII. fig. 8 Bd. The fila- 
ments appear to rotate on their long axes, and twirl up, like an over-spun thread unlay 
themselves, and twist up again, with wonderful rapidity while their activity lasts. —— 

Eurybia 18 ovo-viviparous; and the development of the young may be studied with 
much interest While yet within the body of the parent. The simple capsule of the ovum 

zum: ‚little for remark ; but the ineluded embryo in the comparatively advanced stage 

n my points of analogy to that of an ordinary Gasteropod. : It 
‘ e incipient cartilaginous mantle is of large size compared with the embryo itself. 

is globose in figure, with an opening at one end, through which the ciliated vela of the 
young Eurybia may be seen protruding. These latter organs form expanded lobes, lying 


MR. J. D. MACDONALD ON THE ANATOMY OF EURYBIA GAUDICHAUDI. 247 


one on either side of the head, having the borders beset with large vibratile cilia. A 
frontal elevation rises up between them, from which, it would appear, the future tentacula 
spring. At this period of development the rudimentary external parts exhibit a loose 
cellular structure, the basis of the more complex tissues of the full-grown animal. 

The cephalic ganglia show themselves very distinctly, through the transparent enve- 
lopes, as disproportionately large and rounded masses; and the auditory sacs, containing 
a single primordial otolithe, lie quite in contact with their nervous centres. 

The intestine may be seen taking a simple horse-shoe flexure towards the neural 
surface, while the gastric dilatation fills up the hollow of the curve to a considerable 
extent. i 

The organ above described as corresponding with the spermatheca of the Pulmoniferous 
Gasteropods is just visible on the right side; and what appears to be the true vesicula 
seminalis on the left. 

The external surface of the mantle is pretty equally divided into three zones, by two 
ciliated rings disposed transversely, and consisting of a single series of prominent cells 
bearing moderately. large cilia. 

The development of the cartilage-cells of the mantle may be traced at any part, from 
the round transparent nucleus and its acquisition of a membranous investment, to their 
perfect condition as structural elements. 

The fibrous tissue may be also seen in course of evolution, from very thin-walled cells 
formed upon plastic but minutely granular nuclei. Having attained a certain size, the 
cell-wall shoots forth into delicate tubular processes, which either spring from two oppo- 
site ends; or irregularly from several parts of the circumference. As the nutrition of the 
cells goes forward, the simple tubular processes increase in length and in bulk, more 
particularly at the base, and break up into numerous and exquisitely delicate ramusculi, 
by which the ultimate fibrillation of the body of the cell seems to be effected. The 
elongated or fusiform nuclei permanently remain to strengthen the tissue, which now 
begins to assume a reticulated structure in the intercellular spaces. 

The remains of the ciliated rings of this embryo are distinguishable in Burybie of 
considerable size; and from what has been above noticed of the anatomy of the animal 
and the intimate structure of the mantle (which, though answering the purpose of a 
shell, is, strictly speaking, not homologous with such), it is evident that this genus is not 
conformable to the characters of the Thecosomata or first section of the Pteropoda pro- 


posed by De Blainville. 
The principal points of difference may be thus analysed :— 


EURYBIA. 


1. No external shell. 

2, Head and neck quite distinct. 

3. The foot is small, and only on this account 
rudimentary; the tentacula are large, and not 
combined with the fins. 

4. The mouth is not situated in a cavity formed 


by the locomotive organs. à 
5. This last particular may possibly apply. 


THECOSOMATA. 
1. Animal furnished with an external shell. 
2. Head indistinct. 
3. Foot and tentacula rudimentary, combined 
with the fins. 


4. Mouth situated in a cavity formed by the 
union of the locomotive organs. 

5. Respiratory organs contained within the man- 
.. tle-cavity. 


248 MR. J. D. MACDONALD ON THE ANATOMY OF EURYBIA GAUDICHAUDI. 


In the following synopsis of the principal genera of Pteropoda, Eurybia occupies a 


position suggested by general anatomical principles, and which I believe to be the 


natural one*. 


PTEROPODA. 


f Central series of lingual pavement absent. 
Cheek-pouches deep and tentacular 


(Having a lingual pavement, dental 
Gymnosomata. cheek-pouches, and å membra- suckers large and well-developed ....  PNEUMODERMON. 
Animal naked, head | nous envelope .............. | Central series of lingual pavement present. 
distinct. No gas- Buccal pouches shallow, and tentacular 
tric teeth appa- Having a triserial lingual strap, | suckers minute or rudimentary ...... Cr10. 
AP nee numerous and minute lateral 
labial plates cutting longitudi- 
t TT eere eere IRI . Evnvsn. 
( Shell cartilaginous (symmetrical) ; 
Thecosomata. (lod EN NNN CyMBULIA. 
Animal protected by Commissures of the 
a shell, with the ( With lateral slits through aperture closed.. Hyauza. 
head indistinct. which processes of the 4 
DET mantle protrude ...... | Lateral slits com- = 
Having a triserial municating with 
i ribbon, 4 Shell L the aperture. ... DIACRIA. 
large and definite Få eng Parietes angular or com- 
lateral labial erg pressed antero-posteriorly ........ CLIODORA. 
plates ^ cutting Lateral Pil af Subeylindrical, 
transversely, and hoe eon Pon apex decollated. — Cuvizru. 
two or more gas- | Shell calcareous ; entire and smooth or 
tric teeth ...... gastric teeth < crimped. trans. | Acuminate, apex 
| more than 2.. D vermely .... [. entre v i s ÜRESEIS. 
Inoperculate...... LImacIna. 
Shell sinistrally spiral........:... 
Operculate ...... SPIRIALIS. 


It will be seen by the Table, that Eurybia differs still further from the true Thecoso- - 
mata in the peculiar structure of its labial dental organs (whose cutting edges run longi- 
tudinally, not transversely), and in the total absence of gastric teeth. 

The members of the last division, having calcareous shells, are too closely related by 
their internal anatomy to admit of their separation into two distinct families; and I am 
much inclined to believe, though not certain, that Cymbulia ought to be received amongst 
them. The families thus indicated are the Oliide, Eurybiide, and Hyaleide, the principal 
amendment of the commonly received arrangement being the separation of Hurybia frome — 
the Hyaleide, with which it has been improperly associated; and the union of the Zima- — 
cinide with the same family, from which they have been unnecessarily dismembered. 
The oe of Spirialis occurring in these seas may be truly described as the soft parts 
of Creseis occupying a spiral operculate shell aping that of the Gasteropod. 


* Th; : à * >. S. 
ng table is advanced with à little more confidence as, it has nets by the revising hand of Mr. W. 


MR. J. D. MACDONALD ON THE ANATOMY OF EURYBIA GAUDICHAUDI. 249 


EXPLANATION OF THE PLATE. 


Tas. XLIII. 
Eurybia Gaudichaudi. Natural size. 


. Eurybia Gaudichaudi, magnified to show more distinctly its head, tentacula, oral aperture armed 


with lateral dental cells, the swimming-fins, and foot. The abdominal viscera are imperfectly 
seen through the transparent mantle. 

Portion of the fibro-cartilaginous mantle, highly magnified. 

The lateral labial teeth, forming in this case a double series. 

Six of the dental cells belonging to the right side, still further magnified to show the longitudinal 
cutting edge near the external border of each. 


- Three transverse rows of the lingual ribbon. In this view the curvature of the lateral teeth is 


scarcely apparent. 

The auditory capsule containing otokonial particles. 

Generative systems: A a. expansion of the lobules of testis and ovarium lying in juxtaposition ; 
A b. ova seen through the walls of the ovarian lobuli; A c. primary oviducts uniting in A d, 
the common duct; A e. vesicula seminalis; A f. the vas deferens; B a. pointed extremity of 
the penis, composed of indurated cells, and traversed by the ejaculatory canal; m à. sperma- 
tozoa exhibiting their peculiar looping and twisting movements. 


. Embryo of Eurybia within the ovum: a. frontal and lateral ciliated head-lobes; 5, 6. subæso- 


phageal ganglia supporting the auditory sacs with their single primordial otolithes ; c. alimentary 
canal with the rudimentary abdominal viscera; d d, d d. two ciliated rings which gird the exte- 
rior of the incipient cartilaginous mantle. 


Fig. 10. Structural elements of the mantle: a. portion of one of the ciliated rings of the embryo; 


b. stages of development of the fibrous tissue ; c. cartilage-cells in different stages. 


TransLinn.Soc. Vol XX PI XIII 


EURYBIA. 


W.West ımp 


JD Macdonald RN. Ad nat del loffen West lith. 


[ 351 ] 


XIX.— Notes on Phoronis hippocrepia. By F. D. DysrER, Esq., F.L.S. 
Read July 1st, 1858. 


HAVING recently had an opportunity of examining the very remarkable creature, first 
described by Dr. Wright* under the above appellation, I was desirous of contributing 
some details, with respect to its structure, in addition to those contained in that author’s 
observations. 

The colony on which my observations were made appeared in a tank, on a small piece of 
hard limestone which had been in the aquarium for a considerable time (I believe, some 
months), and which, covered with Serpulæ and other tubicular Annelids, and Actinie, was 
clearly derived from between tide-marks. To the naked eye, the Phoronis presents very 
much the appearance of Cycioum, but is rather more robust in build. The animals grow 
closely aggregated together, each individual inhabiting a tube buried in holes in the rock, 
but which does not project beyond its surface, and is considerably larger than the body: 
the tube is membranous and flexible, and appears to be formed by the incorporation of 
mud with an exudation furnished by the worm. In colour they present the silvery white- 
ness of the Polyzoa. They differ greatly in size, varying from fth to Tsths of an inch in 
length (when fully protruded), of which length the tentacula make about 3th ; the breadth 
of the body is from sth to rd of an inch, and the spread of the tentacula from jgth 
to ths of an inch. The body is cylindrical, slightly flattened on the hæmal surface, and 

_Semitransparent, presenting no trace of somites, annulations, setæ, or uneini. Truncate 
obliquely forwards and backwards, with a slight dilatation at the summit, it is crowned with 
a double row of tentacles springing from the two margins of a horseshoe-shaped lophophore, 

round the rami of which they are continued, recalling most vividly the cephalic apparatus 
of the hippocrepian Polyzoa. The outer row of tentacles spread into a cup, while the — 
row arch towards each other, covering the space between the arms of the horseshoe, "— 
lacing at their tips, and forming a cradle for the ova; the tentacles are filiform, silvery 
White, varying in number from 16 to 86, and united for a short distance from their bases by 
a delicate membrane. The motions of the tentacles are individually voluntary, and some- 
what sluggish. The creature itself is timid; and the slightest touch, or any jar commu- 
nicated to the water, causes it to withdraw with great activity: but the tips of the tenta- 
cles always protrude from the tube. The whole body is in constant gentle me 
tion, and is capable of considerable extension as well as protrusion. The animals are 
provided with great power of reproducing lost parts. An abstracted head is renewed 
within 48 hours, not completely developed, but with å serviceable mouth and its covering 
valve and stumpy tentacles which do their work of providing food. The tentacles are 

* Edinburgh New Philosophical Jourual, vol. vii. p. 313, tab. vii. 


2L 
VOL. XXII. 


252 MR. F. D. DYSTER ON PHORONIS HIPPOCREPIA. 


situated as in the Polyzoa, in lines, and not (as in many tubicular Annelids) all over the 
surface. They exhibit no trace of thread cells, and do not appear to be used as organs of 
prehension, but provide the food required by the worm by the current produced by the 
cilia. The body is not ciliated externally. Below and between the rami of the lophophore 
are two somewhat sigmoid ridges, which are the terminations of the oviducts. In some 
individuals, and always in large ovigerous specimens, at the posterior inner margin of the 
concavity of the lophophore, are seen two nearly circular lips, apparently with a perfora- 
tion in their centre. Their use I cannot indicate. They may possibly be the terminations 
of the sperm-ducts; but as when the worm is in a position to exhibit the protrusion of 
the ova, these organs are hidden, I am unable to say whether any action occurs in them 
simultaneously with the deposition of eggs. The integument of the body is composed of 
a very delicate epidermis, beneath which are bundles of longitudinal muscular fibres, 
connected transversely by others, shorter and more delicate. The length of the largest 
specimens dug out from the rock would vary from I to 3ths of an inch. 

Between the two rows of tentacles, on the neural side of the body, but nearer the 
hæmal row, is the mouth, which is somewhat elliptic, ciliated, and surrounded by a mus- 
cular sphincter, and covered by a delicate transparent marginated crescentic lip, attached 
by its concave edge to the convex portion of the lophophore. The mouth opens into a 
delicate expansible (non-ciliated ?) æsophagus, which occupies the middle of the body. I 
have failed to detect the bands of areolar tissue alluded to by Dr. Wright as the stays of 
the alimentary canal. Just below the portion of the body protruded from the tube, the 
cesophagus opens, apparently through a sphincter, into an oblong stomach, richly 
ciliated, in which the food revolves rapidly in pellets, as in Pedicellina. What lies beneath 
the stomach I am unable to state decisively. ‘The lower part of the body is so deeply — 
imbedded in the hard rock, ahd its substance so fragile, while the tube is comparatively 
so tough, that very many hours of effort failed to extract one in perfect integrity ; and 
the lower portion is so opaque that its walls do not permit its contents to be seen. I 
believe, however, that the stomach terminates in a very capacious intestine, which, filled 
with fæces, occupies the lower portion of the body, and which, traversing the whole tube 
and gradually narrowing, ascends again to terminate in a circular anus lying a little 
above, behind, and between the ridges made by the oviducts between the extremities of 
the rami of the lophophore The intestine lies above the cesophagus and the great 
blood-vessel, but beneath the oviduct. It is exceedingly delicate in structure, and can 
ecarealy be made out except by its contents. It is not ciliated. The fæces are voided by 
jerks, in fusiform pellets connected by slender filaments, and frequently equal in length 
the whole exposed portion of the body. 

The only organs to which hepatic functions could be attributed were some coloured 
cells on the walls of the stomach. 
puit Fe yen tas but this part of the organization demands further 
- "Sade ita y die that the two obscure organs mentioned as being present 
no eye-spots ; iu à : the ee hophore may be cesophageal ganglid ee 

Above the œso "e menge N sensibility to the influence of light. 

phagus, and attached to it by one margin, lies the great blood-vessel, 


MR. F. D. DYSTER ON PHORONIS HIPPOCREPIA. 253 


which for convenience may be called the artery. Along it the blood rushes upwards in a 
powerful stream, until it arrives at the base of the lophophore, where it bifurcates, giving 
a branch to each of the rami. These branches open into sinuses which extend all round 
the lophophore ; and a twig is given off to each tentacle. The blood pursues its course to 
the extremity of the tentacles, which are provided with contractile vessels, tied down on 
one side, free on the other. The progress of the blood is not uniform in the tentacles, as 
it will be frequently seen to be ascending one while it is descending another, and some- 
times the stream may be observed to recede from one tentacle and then fill the adjoining 
one instead of falling back into the general circulation. ‘Two venous trunks open from 
the sinuses above and behind the arterial branches, and then proceed downwards, half 
encircling the cesophagus, till they unite in a large vessel on its neural surface. The 
blood moves by pulsations in the artery, at the rate of from twelve to fifteen beats a minute, 
the vessel contracting on it as it passes upwards, and remaining empty in the intervals 
between the beats. The returning stream through the neural vessel is perfectly conti- 
nuous. In the course of the body the neural and hæmal vessels are connected by nume- 
rous capillary loops; and when the upper portion of the body is removed, the circulation is 
quickly re-established through the loops nearest the point of scission, and carried on as 
powerfully as before. 

The blood consists of a colourless liquor sanguinis, densely charged with red globules 
of irregular shape and size, varying from circular to elliptical, flattened and somewhat 
concave on one side. In length they vary from 5355th to røygth of an inch. The thick- 
ness is about zg5gth. All are provided with one, many with two nuclei of granular ap- 
pearance, about 5545s5th of an inch in diameter. They are exceedingly flexible, and turn 
about, double, elongate, and flatten when pressed for room by meeting other globules in 
the capillaries, exactly as globules of human blood do when seen coursing about under 
thin glass. There are no colourless corpuscles; and very careful watching detected no 
- Ameba-like movements. They coagulate in masses which appear homogeneous, the nuclei 
only remaining visible. Treated with acetic acid, the cell-lining contracts, and all the 
globules assume a perfectly spherical form. There seems no ground for supposing that 
any special heart-like organ is concerned in the circulation of Phoronis. At whatever portion 
of the body section was made, after the shock of separation was recovered from, the pulsa- 
tions of the hæmal vessel were renewed with the same vigour as before; and this occurred 
in the posterior extremity of one individual which was dug out to nearly its full extent. 

The ovary lies below the stomach, and is, I believe, single. It is a long cylindrical 
vessel, pyriform at its base, perfectly transparent, and scarcely distinguishable except by 
its contained eggs, which appear to be attached to the inner surface. The ova are slightly 
elliptic, granular, about røsgth of an inch in diameter: the individuals of which the 
-ovaries were examined were all young; and there was no difference in the size or wind 
lopment of the ova. No specimen was observed without an ovary; bat cin conly eed 
spermatozoa were found. The body of these measured from rærsth to rørsth of an inch, 


with a filiform tail of equal length. 
The ova when deposited are white, 
not ciliated. The. oviduct lies above the rectum, on 


spherical, about shoth of an inch in diameter, and 
the hemal surface, immediately 
212 


254 MR. F. D. DYSTER ON PHORONIS HIPPOCREPIA. 


under the integument, and is not ciliated. The ova lie in its upper and visible portion 
for some hours, vibrating backwards and forwards under the influence of the wave of 
blood in the hæmal vessel. They are driven slowly upwards, till they arrive between 
the two ducts which appear as ridges under the rami of the lophophore. The oviduct 
here seems to contract in its dimensions; and the ova assume a cylindrical form. They 
pause here for about half an hour; but at length the upper end of the cylinder dips sud- 
denly downwards, passes into the hollow ridge, and then mounting through it, is dis- 
charged in a spherical form into the space between the inner tentacles, to which the 
ova adhere by a glutinous exudation. They are voided alternately through each ridge, and 
form a compact white mass, separable only with considerable difficulty, on each side of 
the space in the concavity of the horseshoe, shadowed over by the interlacing extre- 
mities of the inner tentacles. They vary in number from 10 to 80. When first extruded, 
they are granular with a clear margin, and show the usual germinal spot on pressure. 
In a few hours, cilia are developed all over the surface; and two depressions appear on the 
circumference, indicating a circular groove. This groove rapidly deepens; and within 
twenty-four hours the young exhibit distinctly a cephalic and an abdominal segment; ante- - 
riorly the line of separation deepens; and the abdominal portion becomes concave on the 
upper surface, alternately receding from and embracing the convex surface of the cephalie 
portion which lies above it. The cilia increase in length and power; and very soon, in 
certain positions, the alimentary canal becomes distinguishable. The cephalic segment 
divides into three lobes, of which the lateral are the longest and anterior, the central 
highest and posterior. The larva has now great power of locomotion, and quits the 
parent-nest when about forty-eight hours old. 

The principal point of interest in the Phoronis is the indubitable presence of blood- 
corpuscles in proper closed vessels of the circulatory system. Von Siebold* is obscure and 
brief on this subject, and simply says—“ the blood of the Annelids . . . is composed of å 
liquid containing globules... which are always colourless and of a spherical form.” 
Milne-Edwards+ says that, in the Vertebrata, “la couleur rouge du sang est due aux glo- 
bules que ce liquide charrie; chez les Vers å sang rouge, c’est en dissolution dans le liquide 
lui-méme, que se trouve la matière colorante. .. . Les globules ne jouent dans cette colora- 
tion aucun róle essentiel, et d'ordinaire ces corpuscules paraissent méme manquer compléte- 
ment dans ce liquide. . . . M. de Quatrefages a été niéme conduit à penser que dans l'im- 
mense majorité des cas, le sang rouge des Annélides est complétement privé des globules quel- 
conques. Il n'a rencontré qu'une exception à cette regle, et elle lui a été fournie par une 
espèce de Glycère des côtes de la Manche, chez laquelle il a trouvé des globules rouges et de 
forme discoïde, nageant dans un liquide incolore. Mais M. Williams t, qui a publié récem- 
ment une série nombreuse d'observations sur le fluide nourricier des animaux invertébrés, — 
affirme que cette exception n'existe pas; que les globules rouges décrits par M.de 
Quatrefages se trouvent dans le liquide de la cavité générale du corps, et non dans les 
vaisseaux sanguins, et que dans aucun Annélide le sang proprement dit (ou sang coloré) 


P å 
Von Siebold, Comp. Anatomy, translated by Burnett, vol. i. p. 168. 
+ Leçons sur la Physiologie, tome i. pp- 106-108 et note. 


+ On the Blood proper and Chylaqueous Fluid of Invertebrate Animals.— Philosophical Transactions, 1852, p. 63°: 


MR. F. D. DYSTER ON PHORONIS HIPPOCREPIA. 255 


ne renferme des éléments *morphotiques' quelconques, c’est-à-dire des globules.” And 
Mr. Huxley* says, “it may be considered an established fact that, whatever the func- 
tions of the varied vascular system and its contents in different classes of the Annuloida, 
they have nothing to do with the blood or the blood-vessels. The latter are entirely 
absent in the Annuloida at present known, the blood being simply contained in the peri- 
visceral cavity and its processes.” Nothing short of the most patient observation would 
have induced me to state a fact which is incompatible with the opinions and observations 
of Mr. Huxley and Milne-Edwards; but while my own investigations leave no room for 
doubt that the proper fluid of the vascular system in Phoromis consists of a colour- 
less liquor sanguinis densely crowded with red corpuscles, I am confirmed in the pro- 
bability of the fact by the discovery of globules in the vascular system of Glycera by 
M. de Quatrefages, against whose accuracy I do not think the sweeping statement of 
Dr. Williams is a sufficient balance. Not only is it easy to define the vessels which con- 
tain the corpuscles in the living worm, but I have several times, under the compressorium, 
succeeded in isolating a capillary loop with its string of globules. 

There are one or two other points in which the Phoronis deviates very remarkably from 
the Annelidan type. In the position of the anus at the anterior extremity in close proxi- 
mity to the mouth, it stands, I believe, alone, though Mr. Busk has reminded me of the 
analogy which this presents to the arrangement in Sipunculus, the annulose form of the 
Echinoderms. The development of the nervous system is very small—indeed at present, 
as before remarked, I cannot do more than guess at the presence of two æsophageal ganglia, 
—while there is no trace of eye-spots, nor does the creature, like Serpula and Sabella, 
exhibit any appearance of sensibility to light. Negatively, Dr. Wright confirms this view 
inasmuch as he makes no allusion to the nervous system, while Professor Allmant distinctly 
says he could perceive none. In all the Capitibranchiate Annelids the pharynx is short 
and muscular, while in Phoronis it is long and presents no appreciable trace of muscular 
structure. In the same division, the alimentary tube has numerous dilatations corre- 
sponding to the somites, while in Phoronis it is a simple canal, and there exist neither 
external segments nor internal septa, and there is no approach to pedal lobes, hooks, 
paleæ or bristles. I believe that in Phoronis there is no perivisceral cavity ; at all events, 
there are no corpuscles such as are present in the perivisceral fluid of other Annelida. 

I am indebted to my friend Mrs. Brett for the figures, which she has translated into 


beauty from my rough diagrams. 


* Lectures on Natural History, Medical Gazette, vol. xxxiv. p. 385. 
+ Freshwater Polyzoa (Ray Society), p- 55. note, 


256 MR. F. D. DYSTER ON PHORONIS HIPPOCREPIA. 


EXPLANATION OF THE PLATE. 
Tas. XLIV. 


n Figs. 1,2, 3. Side, vertical, and front views of Phoronis hippocrepia. 
». Fig.4. a. Lophophore. 
b. Œsophagus. 
c. Hæmal vessel overlying æsophagus. 
d, d, d. Ova overlying hæmal vessel. 
6,6. Branches of hæmal vessel opening into sinus of lophophore. 
Af. Venous branches uniting beneath the cesophagus, and forming neural vessel. 
9,9. Ridges forming terminations of oviduct. 
À, À. Uncertain organs. 
Fig. 5. View of vascular system. 
a. Lophophore. 
b. Venous branches. 
c. Neural vessel. 
d. Hæmal vessel. 
e; €. Capillary loops connecting neural and hæmal vessels. 
J. Œsophagus. 
g. Mouth with crescentic lip. 
h. Tentacula with twigs opening into sinus of lophophore. 
Fig. 6. Diagram of lophophore seen from above. 
a. Lophophore. 
b. Bases of tentacula. 
c. Mouth covered with crescentic lip. 
d. Anus. 
e. Terminations of oviduct. 
Fig. 7. Blood-corpuscles, magnified 460 diameters. 
Fig. 8. Ovary. 
Fig. 9. Spermatozoa, magnified 460 diameters. 
Fig. 10. Ovum just separated. í 
Figs. 11, 12, 13. The same in more advanced stages of development, the last about 48 hours old. 


E 


+ 


G Jarman se. 


MSBrt del. 


[ 257 | 


XX.—Synopsis of the Fructification of the Gros Sphæriæ of the Hookerian 
Herbarium. By FREDERICK CURREY, Esq., MA., ERS., FL.S. | 


Read March 4th, 1858. 


THE title of this paper discloses the source from which the materials for it have been 
derived; and before proceeding with it, I am desirous, in the first place, to record my 
great obligations to Sir William and Dr. Hooker, through whose kindness and liberality 
I have been enabled to carry out those detailed examinations, the results of which, so far 
as they are completed, are embodied in the following pages. 


Amongst the numberless treasures of the unrivalled Hookerian herbarium is a large : 


collection of fungi belonging to the genus Sphæria. This genus, as originally limited, is 
the most extensive in the fungal alliance. i 

Its magnitude may be inferred from the fact, that in the second volume of Fries’s 
‘Systema Mycologicum,’ published thirty-five years ago, no less than 528 species are 
described; and since that time vast numbers of new species haye been added, which are 


to be found in Fries’s * Elenchus Fungorum ;’ in the papers published from time to time in | 


. the ‘Annales des Sciences Naturelles,’ by M. Desmazidres; in the ‘Sylloge Plantarum 
Cryptogamarum’ of Dr. Montagne; in the * Notices of British Fungi,” by Messrs. Berkeley 
and Broome, contained in the different volumes of the * Annals of Natural History ;’ in 
the *Micromycetes Italici’ of De Notaris, and in other scattered and less accessible 
sources of information. 
In so extensive a genus it will necessarily follow that, for the correct determination of 
. Species, many aids must be necessary; and accordingly in the * Systema Mycologicum ’ 
the genus is broken up into two great divisions, viz. the ** Composite,” in which the 
perithecia are united by a common stroma ; and the * Simplices," in which the perithecia 
are solitary. These divisions again are separated into sections, and the sections into sub- 
sections, the details of which, so far as relates to the Composite, which are the subject of 
the present paper, will be given hereafter. — e oae 
These details are necessary, because in the Hookerian herbarium the Spherie are 
arranged in accordance with the ‘Systema Mycologicum ;’ but it should ‘be observed that 
in his subsequent work, the ‘Summa Vegetabilium Scandinavie,’ published in 1849, 
Professor Fries has abandoned his former arrangement, and formed or adopted several 
new genera instead of the subdivisions of the ‘ Systema Mycologicum.’ These new genera 
have been generally adopted by Continental mycologists ; and there is an evident tendency 
to increase their number. The present, however, is an epoch of transition in the classi- 
fication of fungi, as may be inferred from the fact that within the last year two rearrange- 
ments of the family have been proposed—one by Dr. Bail in Germany, the other by Mr. 
Henfrey in this country. Neither of these authors professes to enter into minute details : 
but their systems, as far as they go, have very little in common; and the latter of them is 


258 MR. F. CURREY ON THE FRUCTIFICATION OF COMPOUND SPHÆRIÆ. 


quite subversive of existing notions. Whether either of them will have any permanence, 
it would be premature to speculate, especially in the face of Dr. de Bary’s observations on 
the ascigerous fructification of Agaricus melleus, lately communicated to the meeting of 
German naturalists at Bonn*. If these observations should be confirmed, it is hardly 
likely that the case of Ag. melleus should be a solitary one; and if ascigerous fructification 
should be proved to exist in the Agarics generally, one great line of demarcation in the 
classification of fungi, and upon which all systems are more or less founded, will be almost 
obliterated. 

In the tribe of the Pyrenomycetes, to which the genus Sphæria belongs, the limits of 
genera are far from settled: for the researches of Tulasne and others have gone far to 
show that many of the existing genera are only stages of growth or abnormal conditions 
of other well-known plants of the same tribe; and it remains to be seen whether future 
mycologists will confirm the numerous genera into which the original genus Sphæria has 
been split up, or whether their judgment will not eventually favour the adoption of the 
arrangement of the ‘Systema Mycologicum.’ 

However this may be, there is no doubt that many, if not most, of the later genera are 
exceedingly well defined ; and I have therefore thought it advisable in the following de- 
scriptions, whilst adhering to the earlier arrangement as being that in use at Kew, to 
notice in each case the genus to which, according to more recent views, each particular 
plant would belong. 

The present paper is the commencement of an attempt to render the discrimination of 
species in this extensive and intricate genus more certain and easy than it has hitherto 
been, by means of drawings and detailed descriptions of the fructification of each parti- 
cular plant. ; 

It will have been observed by all who have consulted the * Systema Mycologicum’ that 
no notice is taken of the nature of the fruit as distinctive of species: nor could such 
notice be expected; for at the time of the publication of that work, microscopical appli- 
ances were quite insufficient to render the necessary observations inviting, or even feasible. 
~ The * Summa Vegetabilium Scandinaviæ” contains some general allusions to the nature 

of the sporidia ; but detailed descriptions were not within the scope of that work, which 
professes only to be a Syllabus of Scandinavian vegetation. 

For some time past, however, the importance of the fructification as distinctive of 
species has been fully recognized ; and any details of new plants would at the present day 
be considered imperfect which did not afford full descriptions of the fruit. 

Figures of the sporidia of a number of species are to be found in the pages of the 
‘Annales des Sciences Naturelles, in the works of Dr. Montagne, De Notaris and others; 
and the * Notices” in the ‘ Annals of Natural History’ above referred to, and which 
relate to new species discovered in this country, are illustrated by excellent drawings. 
After all, however, the whole mass of the older Species remain, so far as regards ger 
fructification, almost entirely undescribed ; and I have long thought that good service might 
be rendered to Mycology by any botanist who would undertake the description and illus- 
tration of the fruit of any considerable number of these plants. The opportunity jé 


* See Bot. Zeitung, No, 45, 1857. 


MR. F. CURREY ON THE FRUCTIFICATION OF COMPOUND SPHÆRLÆ. 259 


doing so having been kindly accorded to me, I have done my best to avail myself of it, 
and have now the honour of laying before the Linnean Society a * Synopsis of the fructi- 
fication of the compound Sphæriæ of the Hookerian herbarium.” In describing the nature 
of the fruit, the points to be particularly attended to are, 1st, the number of sporidia in 
each ascus; 2ndly, their mode of arrangement in the ascus, i. e., whether in one row, in 
two rows, or in a crowded and irregular manner; 3rdly, the shape of the sporidia ; 
4thly, their colour; 5thly, their structure, i. e., whether unicellular and simple, or divided 
by septa, which latter may be transverse, or transverse and longitudinal, or even oblique ; 
and 6thly, the length of the sporidia. The shape of the asci varies so much, not only in 
the same species, but in the same specimen, that I do not think it a character to be relied 
upon. | 

In the great majority of cases the number of sporidia is eight, very rarely it is less, in 
a few instances the number is sixteen, and in a few others the number is unlimited. 

The mode of arrangement in the ascus, although tolerably constant, is not a very cer- 
tain characteristic. "There are many species in which uniseriate and biseriate sporidia 
may be found even in the same perithecum. 

The shape and colour of the sporidia are very constant, and sure marks of distinction. 
With regard to colour, however, it must be remarked that it is different at different 
periods in the growth of the plant. For instance, in a very common species, S. stercoraria, 
Sow., the sporidia are colourless at first, they then assume a rich sea-green sort of colour, 
and eventually become quite black and opaque. It might be supposed that the darker 
colour evidenced the maturity of the plant, but I doubt if this is so, for I have seen the 
sporidia of S. stercoraria when in their colourless state germinate as actively as those of 
the darker shade. j 

The structure of the sporidia when perfect is also a very safe guide, but it is to be 
observed that there are many species in which the sporidia rarely attain their perfect 
state. This is particularly to be seen in some species of the division “ Villosæ,” where 
the sporidia when perfect are of a dark brown colour, and divided by numerous septa, 
but where the sporidia (although attaining their full size) more frequently remain simple, 
continuous, and colourless. Sphæria Corticis one of the Obtectæ is another striking 
instance of this peculiarity. The sporidia in this species are almost always colourless 
and simple, whereas in the perfect state (to which as far as my observations go they 
seldom attain) they are of a dark rich brown colour and divided by several septa. 

I may here observe, that it is important to distinguish between real and apparent septa; 
it is not uncommon in unicellular sporidia to find the endochrome divided into two or 
more portions, and if these portions of the endochrome touch one another, the line of 
contact often appears so like a real partition that it is not always easy to ascertain 


whether in fact any septum exist or not. 


The presence of nuclei is a point of structure which should be noticed in describing - 


sporidia, but it is a characteristic of no great certainty, the sporidia in the same species 

being often found indifferently either with or without nuclei, and the number of them 

also is variable. In some plants, however, as in the sporidia of S. verrucæformis, and of 

some Hypoxyla, they are invariably to be met with. 
VOL. XXII. 


2M 


ke né nt: gm 


260 MR. F. CURREY ON THE FRUCTIFICATION OF COMPOUND SPHJERLE. 


Micrometric measurements of length should always be given. The size of sporidia is 
of course subject to variations dependent upon the circumstances of growth of the plant 
producing them, but as a general rule these variations take place Within narrow limits. It 
not unfrequently happens that the sporidia increase in size after their escape from the 
ascus, so that measurements taken from specimens in which the asci have disappeared, 
will often be found to exceed the mean lengths of the sporidia when measured in the 
interior of the ascus. 

I will conclude these introductory observations with a caution, superfluous to practised | 
microscopists, but which may not be without its use to others who may consult the 
measurements here given; viz., that it is indispensable for correctness of observation to 
ascertain with accuracy the magnifying power of their object-glasses. Opticians, as is 
well known, always furnish tables giving estimates of the magnifying powers, but the 
tables are not (nor do the opticians profess that they are) minutely accurate. They ex- 
press, in fact, the magnifying power aimed at, not that actually attained; and it will be 
found in practice that in object-glasses of the same focal length, no two will magnify 
exactly the same number of diameters. In the measurement of objects so minute as the 
sporidia of many of the Spherie, great accuracy is indispensable for avoiding erroneous 
conclusions. I need hardly add that the correct determination of the value of the micro- 
meter divisions is equally necessary. The arrangement of the “ Sphæriæ composite” 
according to the * Systema Mycologicum’ is subjoined, with the essential characters of the 
genus, sections and divisions. 


SPHÆRIA, Hall. 


CHAR.—Perithecia rounded, entire, perforated at the apex. Asci mixed with paraphyses, convergent, 
deliquescent. Sporidia various. 


A. COMPOSITÆ. 


SECTION I. PERIPHERICZ.—Perithecia more or less divergent, situated in the periphery of the 
stroma; ostiola even, destitute of a neck. 

Div. 1. Corpycers.—Stroma club-shaped, erect, simple or branched, stipitate. 

Div. 2. Poronta.—Siroma marginate, cupshaped, open. Perithecia ovate, situated in the disk of 
the cup only, destitute of a neck ; ostiola even and prominent. x 

Div. 3. PuLvinatæ.—sStroma sessile, convex, more or less hemispherical, immarginate. Perithecia 
in the periphery of the stroma. 

Div. 4. Connat#.—Stroma widely effused, indeterminate, immarginate, plane, surrounding the 
perithecia, or arising from their confluence. Perithecia destitute of a neck, immersed in the 
stroma, or protruding. 


SECTION II. Hyroruericæ.—Perithecia vertical, immersed, covered with the stroma, and having an 
attenuated neck. : 
Div. 5. GueBosa.—Stroma more or less effused, determinate, glebose, distinct from the matrix, 
at length rigid and brittle. Perithecia ovate, large, at first buried and mouthless, at length 
attenuated into a short neck ; ostiola even. 


Div. 6. Licnosæ.—Stroma effused, thin, plane, more or less determinate, connate with the matrix; 


circumscribed with a black line. Perithecia sunk down to the bottom of the stroma, crowded, 
with prominent ostiola. 


Div. 7. VERSATILES.— Sfroma immersed, at length emergent, determinate, but confluent with the 


MR. F. CURREY ON THE FRUCTIFICATION OF COMPOUND SPHÆRLÆ. 261 


matrix, not circumscribed (as in the Lignosæ) with a black line. Perithecia vertical, irregularly 
scattered through the stroma, their necks at first included in it, but at length exserted. 

Div. 8. CoNCRESCENTES.—Stroma thin effused, indeterminate, never cireumscribed, innate, not 
erumpent, formed from the matrix or from the confluence of the perithecia. Perithecia subglo- 
bose, vertical, irregularly aggregated, at first solitary, then confluent, emergent, attenuated into 
a short neck. No erumpent disk. 


SECTION III. AMPHIPHERICÆ.—Perithecia with elongated necks, convergent, circinating, surrounded 
by a spurious pustulate stroma. 

Div. 9. Cıroumsckıpr2.—Stroma formed from the matrix, more or less rounded, included in a 
proper black ventricose conceptaculum which is compressed at the apex. Perithecia scattered in 
the stroma, irregularly circinating, decumbent, with rather long, converging, ultimately umbili- 
cate necks, bursting out from the conceptaculum. 

Div. 10. Incusa:.—Stroma formed from the matrix, rounded, included below in a proper open 
dimidiate conceptaculum, covered above with the subcomate epidermis, through which it bursts, 
forming a somewhat waxy, more or less flat disk. Perithecia collected in the centre of the 
stroma, irregularly circinating, their necks perforating the disk, but less protruded than in the 
former tribe. 

Div. 11. OBVALLATÆ.— Stroma cortical without any proper conceptaculum. Perithecia immersed 
in the inner bark, collected in a circle; ostiola collected into a disk. 

Div. 12. CIRCINATÆ.— Stroma none, or formed of the corroded matrix. Perithecia covered, simple, 
aggregated, arranged in a circle, more or less decumbent. Necks of the perithecia elongated, 
united, and perforating the epidermis, at length free. No conceptaculum or heterogeneous 
disk. 

SECTION IV. Erirnericæ.—Perithecia naked, destitute of a neck, collected upon a stroma (which is 
often spurious) at first covered by the matrix. 
= Div. 13. Cæsprrosæ.—Stroma rounded, determinate, convex. Perithecia superficial, simple, free, 
without elongated necks. 

Div. 14. CoxrLuENTES.—S/roma thin rounded or effused, indeterminate, innate, arising principally 
from the confluence of the perithecia. Perithecia simple, connate, at first innate, then erum- 
pent. 

Div. 15. SgR1ATA.— Sfroma thin, effused, indeterminate, formed from the corroded parenchyma of 
the matrix, sometimes altogether wanting. Perithecia seated on the stroma, covered at first 
with the adnate epidermis, at length almost naked, disposed in parallel rows, often connate ; 
ostiola short. ; 

Div. 16. Conrerræ.—Stroma when present effused, formed from the parenchyma of the leaf, more 
often wanting. Perithecia aggregate, nestling under the epidermis of dead or dying leaves. 


sm. the ‘Summa Vegetabilium Scandinaviæ,’ the Sphæriæ included in the above divisions 
are thrown into distinct genera, the main characters of which, such of them at least as 
include the plants to which this paper relates, are given in that work as follows :— 


1. Corpycers.—Stroma vertical, clavate or capitate, fleshy; perithecia pale-coloured ; sporidia very 
numerous, arranged in moniliform rows. e. i 
2. XYLARIA.—Stroma vertical, clavate, between fleshy and corky, or leathery ; perithecia horny, black; 


at first immersed in the stroma. Sporidia eight in an ascus, usually uniseptate*. 


* This last character is by no means to be relied on; the sporidia are very frequently not septate. They have 
sometimes two nuclei, and when these latter are large and close to one another, there may be an appearance of a 


septum where none really exists. 
TES 2x2 


262 MR. F. CURREY ON THE FRUCTIFICATION OF COMPOUND SPHÆRIÆ. 


3. Poron1A.—Stroma between vertical and horizontal, suberoso-fibrous; perithecia collected in edid 
ginate disk. Sporidia black. 

4. Hypocrea.—Stroma horizontal, sessile. The genus is intermediate between Cordyceps and Nectria, 
and differs from Hypoæylon precisely as Cordyceps does from Xylaria*. 

5. HypoxyLon.—Stroma horizontal, sessile, homogeneous, discrete from the matrix, covered at first 
with a floccose furfuraceous veil.  Sporidia opaque. Intermediate between Xylaria and Spheria, 

6. DiatryPe.—Stroma formed (in part at least) from the matrix: perithecia sunk in the stroma; necks 
of the perithecia elongated and often prolonged into a beak. Sporidiat pellucid, simple, not opaque 
or bilocular, as is common in the Hypoxyla. 

7. VaLsa.—Perithecia circinating with long converging necks; ostiola connate or united into a disk. 

8. Nzorria.—Siroma none. Perithecia pallid, free, but often seated on a tuberculate mycelium of a 
variable nature, membranous, flaccid, brightly coloured. Sporidia 8 in an ascus, pellucid (very 
often, I may add, uniseptate). 

9. SPHÆRIA.—Perithecia carbonaceous, black, superficial and bicorticate, or immersed and then of a 
thinner texture; papillate, sometimes beaked. Asci octosporous, mixed with paraphyses, sporidia 
normally septate or cellular, but often simple, ejected like powder. 


Div. I. Corpycers. 


. B. (CORDYCEPS) Gunn, Berkel. Decades. The asci produce long filamentous sporidia, 
which probably eventually break up into joints, as is the case with other Sphæriæ of 
this division: the sporidia have an alga-like appearance, with an undulating or 
horned outline; the transverse divisions are sometimes very indistinct; each division 
measures about 0-0002 inch. Tag. XLV. fig. 1 represents an ascus X 225 diameters; 
and fig. 2 a somewhat smaller ascus ruptured, with the sporidia protruding, x 450. 
The undulating outline may be caused by the shrinking of the fruit in drying. 
Robin, in his * Végétaux Parasites,’ describes the spores as “courtes, tronquées, 
cylindriques,” 

2. 8. (CORDYCEPS) micrrants, L. ; Fr. S. M. p.323. The asci produce long sporidia, which 
break up into minute joints. Ta». XLV. fig. 3a shows an ascus x 220; and b a 
portion of some of the filaments with the joints into which they divide, more highly 
magnified, _ 

8. 8. (Xyzarns P) PILEIFORMIs, Berkel. Tas. XLV. fig. 4, ascus and sporidia, and free 
sporidia, x 325. Sporidia uniseriate, dark reddish brown, irregularly almond-shaped, 
0:0003 inch long. CUR E 

* 8. (XYLARIA) PEDUNCULATA, Dicks, ; Sow. t. 437. Tas. XLV. fig. 5, ascus with sporidia, 
and free sporidia, x 225. Sporidia biseriate or uniseriate, clear brown and granular 
at first, eventually quite black, almond-shaped or elliptical, at or before maturity sur- 
rounded by a gelatinous envelope, 0-0015 to 0-0018 inch long. 

5. 8. (ConDYozps) FNTOMORRHIZA, Dicks.; Fr. S. M. p. 324. Tas, XLV. fig. 6, X 22. 


P 


* * 
ue era (C. c)» that Hypocrea is hardly distinguishable from Cordyceps, except by its stroma, but that d 
globose nd m er I may add, that in many species of Hypocrea the sporidia consist of squarish or su 

i : a 6 in each ascus, exhibiting a marked departure from the fruit of any other division. j 


MR. F. CURREY ON THE FRUCTIFICATION OF COMPOUND SPHÆRLÆ. 263 


The asci produce colourless jointed sporidia, which divide at the joints. The latter 
are rod-shaped or cylindrical, 0*0003 inch long, and are shown separately at 5, a, 
being an ascus with its contained sporidia. 


6. S. (CORDYCEPS) OPHIOGLOSSOIDES, Ehrh.; Fr. S. M. ii. p. 324. Tan. XLV. fig. 7 a, 


an ascus X 225, b and c joints of the filaments more highly magnifled. The asci 
produce jointed sporidia, which divide at the joints, each joint measuring about 0°0001 
inch inlength. Ifind them cylindrical, not oval or elliptical as figured by Robin and 
Berkeley. When free they often appear quite circular (see fig. 7 c), and to consist 
of an outer membrane and a very highly refractive nucleus. This circular appearance 
may arise from a change in the line of sight, although I think it probable that the 
joints, when freed from their mutual pressure, may assume a globular form. 


7. S. (XYLARIA) INVOLUTA, Kl. Tas. XLV. fig. 8, ascus with sporidia, x 325. Sporidia 


uniseriate, dark brown, elliptical or sub-reniform, 0:0007 inch long. The surface of 
this plant, when dry, breaks away from the flesh like a shell, carrying the perithecia 
with it, and the same thing occurs in S. Guianensis, post. 


8. S. (XYLARIA) GUIANENSIS, Mont. Syll. p. 202. Tas. XLV. fig. 9, ascus and sporidia, 


x 325. Sporidia uniseriate, brown, rather dark, elliptical or slightly curved, 0°0006 
inch long. 


9. S. (CoRDYCEPS) CAPITATA, Holmsk. Fr. S. M. p.324 Tas. XLV. fig. 10a, ascus with 


10. 


11. 


12. 


13. 


14. 


15. 


. inch long. Hypoxylon, Fr. 


sporidia; 5, free joints, x 225. The asci produce jointed colourless sporidia, as in S. 
militaris, &c. "The joints are rod-shaped or cylindrical, 0-0006 to 00008 inch long. 
S. (Hyroxyton) Canopus, Mont. Ta». XLV. fig. 11, sporidia, x 325, sporidia clear 
brown, curved, elliptical or plano-convex, 0:001 inch long. This plant is from Guiana, 
from Leprieur and Montagne's collection. I doubt if it is distinct from S. deusta, 
Hoff. It should, I think, be arranged in the Glebosæ. 
S. (XYLARIA) POLYMORPHA, Pers. Fr. S. M. p. 326. Tas. XLV. fig. 17, ascus with 
sporidia, and a free sporidium, x 325.  Sporidia uniseriate, dark reddish brown, 
curved slightly, 0:0003 to 0:0004 inch long. 
S. (Hxroxvrow or Ponox1A) HELISCUS, Mont. Syll. p. 209. Ta». XLV. fig. 13, spori- 
dia, x 325. Sporidia uniseriate, I think (but there were no complete asci), dark brown, 
irregularly almond-shaped or pyriform, 0:0003 to 00004 inch long. This plant was 
described by Montagne in Cent. ii. No. 44. t. 10. f. 5, under Hypoæylon; but in the 
‘S ' it is transferred to Poronia. 
= en Mont. Tas. XLV. fig. 14, ascus with sporidia, x 325. Sporidia 
uniseriate, dark (rather reddish) brown, elliptical, but irregular in shape, 0°0003 


S. (HYPOXYLON) CRENULATA, Berkel. Tas. XLV. fig. 15, sporidia, x 325. ee 
biseriate, greenish, slightly curved, endochrome usually divided into four po = 
giving a triseptate appearance, 0:0007 to 0:001 inch. long. Sporidia oon y 
uniseriate and overlapping, endochrome sometimes divided into 3 portions only. — 
6. Tas. XLV. fig. 16, ascus with sporidia, 


S. (XYLARIA) DIGITATA, Pers. Fr. S. M. p. 32 i 
ae ar uniseriate, dark brown, curved, 0°0007 inch long. When unripe, 


transparent with one or two nuclei. — 


264 MR. F. CURREY ON THE FRUCTIFICATION OF COMPOUND SPHÆRIÆ. 


16. S. (XYLARIA) HYPOXYLON, Linn.; Fr. S. M. p. 327. Tas. XLV. fig. 17, ascus and 
sporidia, X 325. Sporidia uniseriate, dark brown, elliptico-acuminate, but often 
subeymbiform, 0-0004 inch long, usually with two nuclei, BOM with one, 

17. S. (XYLARIA) MICROCERAS, Mont. Tas. XLV. fig. 18, ascus with sporidia, x 325, 
Sporidia uniseriate, brown, elliptical, rather irregular, 0:0004 inch long. 

18. S. (XYLARIA) MULTIPLEX, Kze. Linn. vol. v. p. 536. Tas. XLV. fig. 19, ascus with 
sporidia, x 325. Sporidia uniseriate, dark brown, plano-convex or irregular elliptical 
0:0004 inch long. | 

19. S. (XYLARIA) CORNIFORMIS, Mont. Tas. XLV. fig. 20, ascus with sporidia, X 325. 
This plant is marked in Hook. Herb. * Hypoxylon corniforme, an Sphæria corniformis, 
Fr." The sporidia are uniseriate, almond-shaped, double- or plano-convex, rather dark 
brown, 00004 inch long. In fig. 21 I have drawn an ascus with sporidia (x 325) of 
S. corniformis, Fr. The sporidia are considerably larger than the above, and are 
furnished with one or two nuclei in each. There are many species, however, in which 
the size of the sporidia varies even to as great an extent as is shown in the difference 
between figs. 20 and 21; and the existence or non-existence of nuclei depends upon 
age and other circumstances to such an extent that they afford no safe characteristic, 
I think S. corniformis, Fr. and Hypoxylon corniforme, Mont., must be the same. Mr. 
Berkeley thinks they may possibly be distinct, the former being European, the latter 
tropical. Dr. Montagne's plant is rather a Xylaria than a Hypoxylon. 

20. S. (XYLARIA) IANTHINO-VELUTINA, Mont. Syll p. 204. Tan. XLV. fig. 22, sporidia, 
X 225. Sporidia uniseriate (I think, but there were no perfect asci), brown, irregularly . 
almond-shaped or pyriform, 0:0005 long. The perithecia are rather larger, with.vel- 
vety hair between them. 

21. S. (XYLARIA) CARPOPHILA, Pers. ; Fr. S. M. ii. p.328. Tap. XLV. fig. 23, ascus with 
sporidia, x 325. Sporidia uniseriate, clear, light brown, elliptical or slightly reni- 
form, with one or two nuclei, 0:0004 inch long. 

22. S. (CoRDYCEPS) ALUTACEA, Pers. The fructification of this plant is not distinguish- 


able from that of S. rufa (see post). It is a-Oordyceps with the fructification of : T 


Hypocrea. 


23. S. (HYPOXYLON) saGRÆANA, Mont. Tan. XLV. fig. 24, sporidia, x 450. Sporida — 
obtusely elliptical, sometimes slightly incurved in the middle, clear (rather dark) 2 
brown, obtuse or almost square at the ends, 0-0004 inch long. | ee 

24. 8. (Corpycers) PURPUREA, Fr. Ta». XLV. fig. 25 a, ascus with sporidia, X 325; b ta 
free sporidia more highly magnified. Sporidia colourless, filiform, attenuated at each — 

_ end, variable in length, 0:002 to 0-008 long, or even more. It is difficult to count the — 
sporidia, but I should guess them at eight; I found them always entirely filling vd 


ascus, which is very long and narrow, and in my specimens not so clavate as the — 


figures in the Micrographic Dictionary, which are reduced from Tulasne's figures m 


the ‘Ann. des Sciences.”—This and the following plant are placed in a distinct genus — - 


(Cavicens) by Tulasne. They differ altogether in fructification from the moniliform 2 à 
fruit of Cordyceps. E 


25. 8. (CogpvcE»s) MICROCEPHALA, Tul. The fruit in my specimens differs in no respect | ; 


MR. F. CURREY ON THE FRUCTIFICATION OF COMPOUND SPHJERLE. 9265 


from that of S. purpurea, except that the hyaline knob at the apex of the aseus is 
very distinct in each ascus; one ascus which I measured was 0-002 inch, another 
0:0034 inch long; so that their length is very variable. The sporidia entirely fill the 
ascus, as in S. purpurea, but I think they sometimes, if not always, overlap one 
another, so that their length is not always equal to that of the ascus. 

26. S. (Corpycers) Roszrtsu, Hook. Ta». XLV. fig. 26, ascus with fruit, x 425. 
The fruit here, as in the other true species of Cordyceps, consists of long colourless 
sporidia, which break up into joints. At least, the sporidia are divided by septa 
into a number of small portions, which probably separate from one another at 
the partitions, as in S. militaris, capitata, &c. The septa are sometimes very 
indistinct. 

26a. S. (CORDYCEPS) TYPHINA, Pers. Fr. S. M. ii. p. 553, under Dothidea. The fructifi- 
cation is almost identical with S. militaris, although the asci are smaller. 


Div. 2. Poronna. 


27. 8. (PORONIA) PUNCTATA, Sow. Tas. XLV. fig. 27, ascus with sporidia, x 825. Spo- 
ridia uniseriate, brown when young, and then slightly granular, quite black and 
opaque when ripe, elliptical, varying in length from 0-0008 to 0:0012 inch. 


Div. 8. PULVINATÆ. 


28. S. (HYPOXYLON) MULTIFORMIS, Fr.! S. M. ii. p. 334. Ta». XLV. fig. 28, ascus with 
sporidia, x 450. Sporidia uniseriate, rich yellowish-brown, elliptical, but usually 
slightly curved, 0:0004 inch long. 

29. S. (HYPOXYLON) DURISSIMA, Sz.; Fr. S. M. ii. p. 335. Ta». XLV. fig. 29, ascus with 
sporidia, x 325.  Sporidia uniseriate, clear dark brown, elliptical, but frequently 
slightly curved, 0:0003 to 0:0004 inch long. This plant is marked with a ?; and it 
seems doubtful whether it is not a form of S. multiformis. The perithecia are blacker 
and rather more dome-shaped than in my specimens of S. multiformis, and they have 
a decided nipple-shaped ostiolum. The sporidia have usually one good-sized nucleus, 
sometimes two, or even more. 

30. S. (HYPOXYLON) FRAGIFORMIS, Pers.; Fr. S. M. ii. p. 332. Ta». XLV. fig. 30, spo- 
ridia, x 325. Sporidia uniseriate?, dark opaque brown, elliptical or areuate, with 
sometimes one or two nuclei visible, but mostly too opaque for the nuclei to be seen; 
0:0006 inch long. | 

31. S. (HYPOXYLON) ANNULATA, Mont. (8. depressa) Mont. Syll p. 213. Tas. XLV. 
fig. 31, sporidia, x 450. Sporidia light brown, elliptical, 0°0003 inch long. ^ 

32. S. (HYPOXYLON) CONCENTRICA, Bolt. Fr. S. M. ii. p. 331. Tas. XLV. fig. 32, sporidia, 
x 325. Sporidia uniseriate, dark brown, sometimes with a large elliptical or reni- 
form nucleus, elliptical, but sometimes of irregular shape, 0-0005 inch long. 

33. S. (HYPocREA) GELATINOSA, Tode ; Fr. S. M. ii. p.336. Tas. XLV. fig. 33, ascus with 
sporidia, x 325. Sporidia uniseriate, colourless, squarish in the ascus, more nearly 
round when free, with one or many, or with no nuclei, 0-0002 inch in diameter when 
free. Almost precisely similar in fructification to S. rufa (see post), and like that, 


266 


34. 


35. 


36. 


37. 


38. 


39. 


40. 


41. 


MR. F. CURREY ON THE FRUCTIFICATION OF COMPOUND SPHARIA. 


probably not a true Spheria. In specimens in my own herbarium from Tunbridge 
Wells, each pulvinulus is seated upon a dense white byssoid subiculum, which 
becomes brown with age. 
S. (HYPOXYLON) VERNICOSA, Schwein. Tas. XLV. fig. 34, ascus with sporidia,.x 315. 
Sporidia uniseriate, dark brown, elliptical, 0-0005 to 00006 inch long. Perhaps only 
a form of S.concentrica; but Mr. Berkeley considers it distinct in the structure 
of the stroma. 
S. (HYPOXYLON) ARGILLACEA, Fr. Obs. i. t. 2. f. 5. Tas. XLV. fig. 35, ascus with 
sporidia, x 325. Sporidia uniseriate, opaque black-brown, irregularly elliptical ;. 
sometimes one, two, or three nuclei, or a number of small nuclei, are visible. 
S. (HYPOCREA) RUFA, Pers. Fr. S. M.ii.p.335. Ta». XLV. fig. 36, ascus with sporidia, 
x 325. The sporidia are squarish colourless bodies usually 16 in each ascus, quite 
unlike the usual fructification in the Pulvinatæ. This plant agrees in its fructifica- 
tion with S. citrina and S. lobata, and, like S. gelatinosa and some others, comes 
very near to Dothidea; for the perithecia, as far as I can make out, have no walls 
distinct from the flesh of the stroma; so that the fructifying mass is in fact a 
* nucleus" in the sense in which that word is used in Dothidea. 
S. PRUINATA, Kl. Linn. vol. viii. p. 489. Ta». XLV. fig. 37, ascus with sporidia, x 325. 
Sporidia uniseriate, dark brown, elliptical, with 1 to 4 nuclei, or with a dark curved 
band across, arising, I think, not from a septum, but from the approximation of the 
nuclei; varying slightly in length, but on an average 0:001 inch long. Perithecia 
crowded, dome-shaped, with mammillate ostiola, mostly, but not always covered with 
a white farinaceous powder. 
S. (HYPOXYLON) FUSCA, Pers. Fr. S. M. ii. p.332. Tas. XLV. fig. 38, asci with - 
sporidia, x 325. Sporidia elliptical or almond-shaped; when young pale brown, with 
one or two large nuclei, and sometimes small nuclei also; when ripe very dark and 
opaque, normally uniseriate, but occasionally erowded as shown in fig. 39, which 
represents the fruit (x 225) of a Sphæria not differing materially from Sphæria 


fusca, except that the sporidia are larger and more irregular than usual. De, 


latter plant occurred at Eltham, in Kent, in J une, 1855. The ordinary length of the 
sporidia in S. fusca=0-0005 inch. | 
S. (HYPOXYLON ?) PARMULARTA, Berkel. Tar. XLV. fig. 40, sporidia x 225. Sporidia 
rather dark brown, clear, with mostly one nucleus, irregularly elliptical or almond- 
shaped, 0-0004 inch long. The specimens are in a bad state; and I had some doubt 
whether the plant ought not to be referred to the Connatæ or Confluentes. There B 
however, some appearance of the débris of a Pulvinulus. i 
S. (Hrroxyuon) COHÆRENS, Pers. Fr. S. M. ii. p. 935. Tap. XLV. fig. 1 
with sporidia, x 325. Sporidia uniseriate, rather dark brown, irregularly elliptical, 
ee rather curved, often with one or two nuclei, 0:0003 to 0:0004 inch 
ong. 
S. (HYPOXYLON) RUBRICOsA, Fr. E]. 2. p.63. Tap. XLV. fig. 42, ascus with sporidia, 
x 325. Sporidia uniseriate, dark brown, eventually opaque, elliptical, uniseptate, 
with frequently a nucleus in each division, 0:0004 to 0-0006 Bid long. 


MR. F. CURREY ON THE FRUCTIFICATION OF COMPOUND SPHÆRLÆ. 267 


Div. 4. CoxNATX. 


42. S. (HYPOXYLON) RUBIGINOSA, Pers.; Fr. S. M. ii. p. 340. Ta». XLV. fig. 43, ascus 
with sporidia x 325. Sporidia uniseriate, dark brown, elliptical, but slightly irre- 
gular, usually with one, sometimes with two or even three nuclei, 0:0004 inch long. 

43. S. (HYPOXYLON) ATRO-PURPUREA, Tode; Fr. S. M. ii. p. 340. Tan. XLV. fig. 44, 
sporidia x 325. Sporidia dark brown, paler when young, usually with two, some- 
times one nucleus, 0:0004 to 0:0005 inch long. 

44. S. (HYPOXYLON) PERFORATA, Sz.; Fr. S. M. ii. p. 340. TA». XLV. fig. 45, sporidia, 

. X 325. Sporidia I think uniseriate, like those of atro-purpurea, but rather lighter- 
coloured, 0:0004 inch long. The plant is easily known by the rusty-coloured peri- 
thecia, and the white flat ostiola. 

45. SPHÆRIA BOTRYOSA, Fr. S. M. ii. 342. Tas. XLVI. fig. 46, ascus with sporidia, x 225. 
Asci variable in size, clavate, with the inner membrane very distinct, the latter filled 
with innumerable multitudes of very minute granules; granules colourless, about 
0:0001 inch, endowed with Brownian motion. I should have doubted these granules 
being true sporidia; but I find the fruit of the plant in the Hookerian herbarium 
coinciding exactly with that of authentic specimens from the Scleromycetes Sueciæ. 
Fries considers the plant a Sphæria with confluent perithecia, and not an Hypoxylon. 
See ‘Summa Veg. Scand.’ p. 383, note 3. : 

46. S. (HYPOCREA) LATERITIA, Fr. S. M. ii. p. 338. Tas. XLVI. fig. 47, ascus with spo- 
ridia, x 325. Sporidia uniseriate elliptico-acuminate, colourless, 00006 to 0:0007 
inch long. I had some doubt whether the plants were ripe, and whether the sporidia 
may not become brown in age. The specimens in Hook. herb. are from Mr. Stephens's 
herbarium. I have seen no other specimens of this species. 

47. S. (HYPOXYLON) SERPENS, Pers.; Fr. S. M. ii. p. 841. Tas. XLVI. fig. 48, ascus 
with sporidia, x 325. Sporidia similar in colour to those of S. atro-purpurea, and 
of about the same size. " 

48. 8. (Hyrocrea) HYALINA, Schwein.; Fr. S. M. ii. p. 339. Tas. XLVI. fig. 49, ascus 
with sporidia and free sporidia, X 325. Sporidia uniseriate, colourless or greenish, 
elliptico-acuminate but rather irregular, 0:0006 to 00008 inch long. 

49. S. (HyroxynLon) SASSAFRAS, Sz. ; Fr. S. M.ii.p.343. Tam. XLV I. fig. 50, ascus wäh. À 
sporidia, x 325. Sporidia uniseriate, elliptical, ent dark, 0-0004 : 
long. Very like S. atro-purpurea ; but the aperi are of. a m darker colour, 
and the ostiola obtuse or depressed, not mammillate. 2 

50. S. (Hyrocrea) cITrINA, Pers.; Fr. S. M. ii. p. 337. Tas. ass = 51, ET 
with sporidia, x°S25: --Bporidia “squarisb; irregulat in shape, colour es, Norma y 
16 in an ascus. Hardly distinguishable in fruchfoneun from S. rufa, &c., = € 
known by the yellow colour, and effused condition of the stroma. It may be only a 
peculiar condition of S. gelatinosa, Tode. 


51. S. (HyPOCREA) LOBATA, SZ. Wormsk.; Fr. 8. M. ii. P- 348. Tap. XLVI. fig. 52, 
nåde with sporidia, x 325. Not distinguishable in its fruit from the last, except 


that the asci and sporidia (in the Kew specimens at least) are smaller. : 
N 
VOL. XXII. | 


268 MR. F. CURREY ON THE FRUCTIFICATION OF COMPOUND SPHÆRLÆ. 


52. S. (HyPOCREA) LUTEO-VIRENS, Fr. S. M. ii. p.339. Tas. XLVI. fig. 53 a, ascus with - 
sporidia; 6, free sporidia ; c, stylospores, all x 325. Sporidia uniseriate ; overlapping, 
colourless, very narrowly almond-shaped, 0:0008 to 0-0010 inch long.  Perithecia 
dull-yellow orange colour, imbedded in a woolly subiculum which produces the yellow 
stylospores on the left of the figure. 


Div. 5. GLEBOSÆ. 


53. S. (HYPOXYLON) REPANDA, Fr. S. M. ii. p. 346. Ta». XLVI. fig. 54, ascus with spo- 
ridia X 325. Sporidia uniseriate, dark brown, elliptical, 0:0004 to 0:0005 inch long. 
Stroma rather undulating in surface, ostiola depressed, the whole plant very like 
S. stigma, except in its fruit, which is quite different. 

54. S. (HYPOXYLON) TUBULINA, À. and S. ; Fr. S. M. ii. p.946. Tas. XLVI. fig. 55, ascus 
with sporidia and free sporidia, x 325. Sporidia usually, I think, uniseriate, but 
sometimes biseriate, clear, rather dark brown, elliptical, 0-0002 to 00003 inch long. 

55. S. (HYPOXYLON) LUTEA, A. and S. ; Fr. S. M. ii. p.947. Tas. XLVI. fig. 56, ascus with 
sporidia, x 325. Sporidia minute, uniseriate, brown, elliptical, but rather irregular, 
generally with two nuclei, which sometimes give an appearance of a septum; some- 
times there is one nucleus, sometimes none. Sporidia 0:0002 to 0:0003 inch long. 

56. 8. (HYPOXYLON) DEUSTA, Hoffm. ; Fr. S. Mai. p.345. Tas. XLVI. fig. 57, sporidia, X 
325. Sporidia biseriate I think (the asci were imperfect), rather dark brown, usually 
somewhat curved, with an inner membrane sometimes visible, 0:0012 to 0:0016 inch 
long. a 

57. S. (HYPOXYLON ?) FUSCOSPORA, Schwein. Tas. XLVI. fig. 58, sporidia, x 325. Spo- Lo 
ridia biseriate I believe, at first colourless, then clear dark brown, arcuate, 0:0014 — 
inch long. au 

58. S. (HYPOXYLON) NUMMULARIA, Bull; Fr. S. M. ii. p.348. Tas. XLVI. fig. 59, asci 
with sporidia, x 325. Sporidia uniseriate, brown when young, black when ripe, 
round or elliptical, 00004 to 0-0006 inch long. : 

59. 8. (HYPOXYLON) marçrnara, Fr. El. ii. 69. Tan. XLVI. fig. 60, ascus with sporidia, — 
X 925. Sporidia uniseriate, with one, two, or no nuclei, elliptical or subreniform, — 
00002 inch long, pale clear brown. xn 


| | Div. 6. Lienosa. — Eo 
60. 8. HYPOXYLON) una, Pers.; Fr.8.M.ii. p.358, Tam XLVI. figs. 61 and 62, asci with — 
sporidia, X 825. Fig. 63, free sporidia, x 895. Sporidia at first of a rich olive — 
green colour, eventually dark, clear, transparent brown, sometimes with nuclei, some 
times obtusely elliptical (fig. 61), sometimes almond-shaped (fig. 62). 0:0006 fo 
0:001 inch long. | : 
61. S. (DraTRYPE) DISCIFORMIS, Hoffm.; Fr. S. M. ii. p.353. Tas. XLVI. fig. 64, sporidia, — 
highly magnified. Sporidia biseriate or crowded, yellowish in a mass, almost colour Rå 
less when detached, slightly curved ; 0:0002 to 0-0003 inch long. a 
62. S. (Drarryrn) FLAVOVIRENS, Hoffm.; Fr. S.M. ii. p.357. Tas. XLVI. fig. 65, aset 
with sporidia, x 450. Sporidia crowded, yellowish, curved, 0:0003 inch long. 


MR. F. CURREY ON THE FRUCTIFICATION OF COMPOUND SPHÆRLÆ. 269 


63. S. (DIATRYPE) PLATYSTOMA, Schwein.; Fr. S. M. ii. p.351. Tar. XLVI, fig. 66, spo- 
ridia, X about 450.  Sporidia colourless or with a slight yellow tinge, less curved 
and rather smaller than in Sphæria stigma; about 00003 inch long. The plant is 
distinguishable from S. stigma by its rimose stroma, and by its larger, more protru- 
ding Pezizæ-form ostiola. 

64. S. (HYPOXYLON) VOGESIACA, Pers. in Litt. Ta». XLVI. figs. 67 and 68, sporidia, x 
325. Sporidia uniseriate (I believe), at first pale brown, eventually almost opaque 
with granules and nuclei, elliptical, but rather irregular, sometimes almost pyriform, 
0:0007 to 00008 inch long. * Ad truncos emortuos Aceris pseudo-platani ; affinis 
Sphæriæ serpentis sed distincta," is the note attached. 

65. S. (HYPOXYLON) viRGULTORUM, Fr. S. M. ii. p. 351, El. xi. 71. Ta». XLVI. fig. 69, 
sporidia, x 450. Sporidia sometimes uniseriate, sometimes biseriate, lageniform, 
colourless, or with an almost imperceptible yellowish tinge, 0°0004 inch long. 

66. S. (DIATRYPE) ATRO-PUNCTATA, Schwein. ; Fr. S. M. ii. p.951. Tas. XLVI. fig. 70, 
ascus with sporidia and free sporidia, x 325. Sporidia uniseriate, overlapping, brown, 
with granular nucleate contents, broadly almond-shaped, 00010 to 0:0012 inch long. 
Easily known by the small punctiform ostiola penetrating the white upper surface 

` of the stroma. : 

67. S. (DIATRYPE?) raors, Schwein. ; Fr. S. M. ii. p. 356. Ta». XLVI. fig. 71a, sporidia ; 
b, stylospores. The specimens are in bad condition; but one perithecium produced the 
sporidia a, which are yellowish like those of Sphæria verrucæformis ; another produced 
the stylospores or spermatia b, which were colourless and produced on sporophores 
attached to the wall of the perithecium. Length of a about 0:0003 inch, of å from 
0:0010 to 0:0014 inch, measured on the chord of the arc. 

68. S. (DIATRYPE) LIRIODENDRI, Schwein. ; Fr. S. M. ii. p.356. Ta». XLVI. fig. 72, asci 
with sporidia and free sporidia, x 325. Sporidia very crowded, quite filling the asci, 
brown, just like the sporidia of Sphæria favacea and S. aspera, 0:0002 inch long, or 
rather more. Possibly the plant is only a state of Diatrype BPM idis 

69. S. (DIATRYPE) cAPNODEs, Berkeley. Tas. XLVI. fig. 73, ascus with sporidia and free 
sporidia x 325. Sporidia uniseriate, at first a clear brown, ultimately quite opaque, 
uniseriate, elliptical, 00004 to 0:0005 inch long. This species is allied to Sphæria 

stigma; but the stroma is of a blackish grey colour, and the ostiola more prominent. 
The fruit, as will be seen, is totally different from that of S. stigma. 

70. 8. (Hyroxxzon) mELANASPIS, Mont. Sylloge, p. 215. Tas. XLVI. fig. 74, ascus with 
sporidia and free -sporidia, x 325.  Sporidia uniseriate, overlapping at the ends, 
almost colourless, but with a faint tinge of yellow or rather straw colour, fusiform, 
but wider at one end than at the other, 0001 inch long. Forming round black stains, 
with the ostiola slightly protruding in the middle of the stain. amet 

71. S. (DIATRYPE) DRYOPHILA, n.s. Ta». XLVI. fig. 75, asci with sporidia an p 
ridia, x 325. Sporidia uniseriate, narrowly elliptical, subacuminate å - - ae 
first of a pale greenish brown, then darker, and eventually almost black, w så er 
with two or more nuclei in each sporidium, 00004 to 0-0005 inch long. Perithecia 


ovate or sub-globose surrounded by a dirty green stroma, rather pd buried, 


270 


72. 


73. 


74. 


75. 


76. 


TL 


- 78. 


MR. F. CURREY ON THE FRUCTIFICATION OF COMPOUND SPHÆRLÆ. 


piercing the bark by their long converging necks, forming compact scattered black 
pustules on oak-branches. On dead and decaying branches of oak at Weybridge, 
September, 1856. Much resembling SpA. verrucæformis in its external form, and in 
the very glutinous nature of the contents of the perithecia. 

S. (DIATRYPE) NUCLEATA, n. s. Tas. XLVI. fig. 76, ascus with sporidia, x 395. 
Sporidia linear-acuminate, but constricted in the middle, colourless, 0:0007 to 00008 
inch long. Perithecia ovate or globose, with rather short ostiola, collected in elon- 
gated irregular patches surrounded by a dark line. Weybridge, J anuary, 1856, on 
furze. 

S. (DIATRYPE or VALSA) VARIANS, n. s. Tas. XLVI. fig. 77, ascus with sporidia, x 
325. Sporidia biseriate, obtuse, constricted in the middle, colourless, with granular 
endochrome, uniseptate, 0-0006 inch long. Perithecia subglobose; ostiola conical, 
sometimes umbilicate at the apex. The masses of perithecia penetrate the bark in long 
parallel lines. This plant unites the characteristics of the Circumscriptee and Lig- 
nose, some specimens having a conceptaculum, and others being scattered, with a 
black line and without a conceptaculum. Eltham, June 1855. 

S. (DIATRYPE or VALSA) DENIGRANS,n.s. TAB. XLVI. fig. 78, ascus with sporidia, x 
325. Sporidia uniseriate, often partly overlapping, dark olive-brown, elliptical, bi-, tri-, 
or multi-nucleate, sometimes quite simple, 00005 inch long.  Perithecia conical or - 
depressed, with long ostiola, deeply set in the wood; ostiola penetrating the bark and 
raising the wood into minute but prominent black tubercles. When a nest of perithecia 
Is cut transversely, there is a black line round them which is very well defined. 

8S. (DraTRYPE) INÆQUALIS, n. s. Tas. XLVI. fig. 79, ascus with sporidia, x 325. 
Sporidia uniseriate, almost colourless, but rather of a green tinge, obtuse, constricted 
in the middle, uniseptate, 0:0006 inch long or a little over; contents granular and 
nucleate. Perithecia subglobose ; ostiola short and rugose. The surface of the wood 
(beneath the epidermis) under which the perithecia lie is blackened and rugged; the 
perithecia are sometimes scattered. Masses of perithecia surrounded by a black 
line. Nearer to the Lignosæ than to any other division. On furze, Weybridge, 
January 1856. 

S. (DraTRYPE) BaDHAMI, n. s. Tar. XLVI. fig. 80, ascus with sporidia and free spo- 
ridia, x 325. Sporidia biseriate, colourless, usually 4-nucleate, with a minute hyaline 


appendage at each end (which, however, is invisible in the asus), narrowly almond- — — 


shaped, 0°0005 to 0:0006 inch long without the appendages. The perithecia occur - 
we and in masses ; they are deeply imbedded in the wood ; the surface of the inner 
bark is much blackened. The plant in habit and appearance resembles S. inequalis, 
supra; but the sporidia are altogether different. | 


S. (DrarRYpz) VERRUCÆFORMIS, Ehr.; Fr. S. M. ii. p.355. Tap. XLVI. fig. 81, ascus 2 _ 
with sporidia and free sporidia, x about 225. Sporidia very numerous, crowded, ae 


yellowish in the mass, almost colourless when 


S. (DraTRYPE) FAVAOEA, Fr. S. M. ii. p. 354. 


MR. F. CURREY ON THE FRUCTIFICATION OF COMPOUND SPHÆRLÆ. 271 


— 


79. S. (DIATRYPE) aspera, Fr. S. M. ii. p. 354. The only difference between the fructifi- 


cation of these plants and that of S. verrucæformis is that their sporidia are somewhat 
smaller and without nuclei, and their asci are clavate, not fusiform. 


80. 8. (DIATRYPE) stiema, Hoffm.; Fr. S. M. ii. p. 350. Tan. XLVI, fig. 82, sporidia, x 


81. 


82. 


83. 


84. 


85. 


86. 


450. Sporidia yellowish in the mass, almost eolourless when single, slightly curved, 
0:0004 inch long. 

S. (DIATRYPE) BULLATA, Ehr.; Fr. S. M. ii. p. 349. Sporidia just like those of 5. 
platystoma (see fig. 66), 00002 to 0:0003 inch long. 

S. UNDULATA, Pers.; Fr. S. M. ii. p. 350. Sporidia biseriate, yellowish, slightly 
curved, rounded at each end, 0:0004 inch long. The perithecia break through 
the bark in an undulating manner. The sporidia are just like those of Sphæria 
stigma (see fig. 82), of which this species is probably only a variety, as was long 
since remarked by Fries. 

S. (Hyrocrea) LENTA, Tode ; Fr. S. M. ii. p. 349. Ta». XLVI. fig. 83, ascus with spo- 
ridia, X 325. The fructification of this plant is precisely similar to that of S. rufa; 
and I would refer to the remarks appended to that plant. The square-shaped, irregu- 
lar, colourless sporidia, after escaping from the ascus, become round or nearly so, their 
average diameter being less than 0:0002 inch. I have retained the plant in this 


. division, as it is so arranged in the Herbarium, and by Fries in the ‘Syst. Myc.’ 


In the ‘Summa Veg. Scandinaviæ ” it is placed (strange to say) in the genus Diatrype ; 
according to the arrangement in that work, it ought certainly to be a Hypocrea. 


Div. 7. VERSATILES. 


S. (DrATRYPE) scABROSA. Dec.; Fr. S. M. ii. p. 860. Tas. XLVI. fig. 84, ascus with spo- 
ridia and free sporidia, x 450. Sporidix uniseriate, clear brown (one had a hyaline tip 
at each end), subcylindrical, rounded at each end, triseptate, slightly constrieted at the 
septa, 0:0006 inch long. In this specimen the perithecia are very slightly immersed, 
and look like large specimens of Sphæria pulvis-pyrius, which, although belonging 
to the Denudatæ, is often subcuticular in its growth. The fruit also is the same as 
in S. pulvis-pyrius, except as regards size. The plant, however, being marked 5. sca- 
brosa by Mr. Berkeley, I have described it as such, but with some doubt. i 
S. (DIATRYPE) PODOIDES, Pers. Syn. p. 22, excl. syn. Ta». XLVI. fig. 85, ascus with 
sporidia and free sporidia, x 225. Sporidia biseriate or crowded, clear, rather pale- 
brown, 7-septate, with a hyaline joint at each extremity, often slightly curved, 0:0024 
to 0003 inch long: Quite distinct from S. scabrosa, Dec., under which it was placed 
by Fries as a variety. id 
S. PODOIDES, var. LEVIS, does not differ from the above, as far as the fruit is concerned, 
but only in the smoothness of the tubercles, arising from the ostiola of the perithecia 


not penetrating the stroma. 


87. S. (DiarRYPE?) PETIGINOSA, Fr. in litt. Tas. XLVI. fig. 86, ascus with sporidia and 


88. S. (DIATRYPE) LEPROSA, 


free sporidia, x 450. Sporidia biseriate or crowded, yellowish, rounded at each end, 


narrow, slightly curved, 0-0004 to 0-0006 inch long. i: 
iron: Pers.; Fr. S. M. ii. p. 365. Tas, XLVI. fig. 87, ascus with 


972 


89. 


91. 


92. 


93. 


94. 


95. 


MR. F. CURREY ON THE FRUCTIFICATION OF COMPOUND SPHÆRIZÆ. 


sporidia and free sporidia, x 325. Sporidia biseriate or crowded, colourless or 
yellowish, slightly curved, 0:0004 to 0:0005 inch long, rounded at each end, 
narrow. 

S. (DIATRYPE) STRUMELLA, Fr. S. M. ii. p. 365. Tas. XLVII. fig. 88, sporidia, x 450. 
Sporidia biseriate, colourless, elliptico-acuminate, triseptate, or, I think, pseudo- 
triseptate from the division of the endochrome into four parts, 0:0005 to 0:0006 inch 
long, sometimes slightly curved and slightly constricted in the middle, and frequently 
only biseptate. 


. S. (DIATRYPE) RADICALIS, Fr. El. ii. p. 73. Tap. XLVII. fig. 89, sporidia highly mag- 


nified. Sporidia biseriate or crowded, elliptical or subturbinate, colourless, hyaline 
very minute, 00002 to 0:0003 inch long, always, I think, with one, sometimes with 
two or three septa. The perithecia are deeply buried and surrounded by an orange- 
coloured stroma, through which the ostiola penetrate in the form of little, round, 
black, dise-like spots. : 

S. (DIATRYPE) QUERCINA, Pers. Tap. XLVII. fig. 90, ascus with sporidia and free 
sporidia, x 325. Sporidia biseriate, endochrome 4-, sometimes 6-partite, oleaginous, 
but sometimes granular, 0:0020 to 0-0028 inch long. This plant has been much 
misunderstood. "The above description is from a specimen lent to me by Mr. Berkeley. 
I doubt whether any of the specimens at Kew are the true species. Some of them 
are certainly S. leiphemia; and the others are not in a sufficiently good condition 
to decide upon. 

8. (DIATRYPE) LANCIFORMIS, Fr. S. M. ii. p. 362. Tap. XLVII. fig. 91, aseus with 
sporidia, x 325. Sporidia usually biseriate, but occasionally (when the sporidia are 
small, I think) uniseriate, pale clear brown, when ripe flatly elliptical, rather obtuse 
at the ends, the extreme tips frequently, if not always, pellucid, and the sporidium 
of a darker colour in the part immediately adjoining the pellucid tips. Length very 
variable, ordinarily 0-002 inch. | 

S. (DIATRYPE) nysrRIX, Tode; Fr. S. M. ii. p. 364 Tas. XLVII. fig. 92, asci with 
sporidia, X 325. Sporidia uniseriate or biseriate, elliptical, slightly constricted at the — 
septa, 3- or 4-septate with frequently longitudinal and oblique septa, pale clear brown ——— 
0:0006 to 0:0008 inch long. Described from a specimen out of Mr. Berkeley's herba- > 
rium, the plant at Kew not having perfect fruit. g or 
8S. (DraTRYPE) CERATOSPERMA, Tode; Fr. S. M. ii. p.364. Ta». XLVII. fig. 92, ascus — 
with sporidia, x 325. Sporidia biseriate, colourless, simple, curved, 0:0003 to 0:0004 — — 


inch long. Described from a specimen of Mr. Berkeley's, the plant at Kew not å å 


being the true species. 


S. (DIATrYrE) FERRUGINEA, Pers.; Fr. S. M. ii. p. 363. Tap. XLVIL fig. 94, asi —— 


with sporidia, x 395. Sporidia very long, linear, acuminate at the ends, colourless, 2 


. with many nuclei. 


+ S. (DraTRYPE) NIGERRIMA, Bloxam, MSS. Sporidia biseriate, closely packed, "s : 


irregular in shape (arcuate, subelliptical or lozenge-shaped), with a number g 
circular nuclei, colourless, but with a greenish tinge, 3-, 4-, or 5-septate. Pertha 


ir u : on g^ nie 
regularly ovate or conical, sometimes arranged in tiers, raising the matrix intg 


MR. F. CURREY ON THE FRUCTIFICATION OF COMPOUND SPHJERLE. 273 


black pulvinate tubercle, through which the densely crowded ostiola penetrate and 
cause a scabrous appearance; ostiola somewhat elongated and rounded. Communi- 
cated by Mr. Bloxam, from Leicestershire. 

97. S. (DIATRYPE) IRREGULARIS, Sow.=S. gastrina, Fr. S. M. ii. p.979. Tas. XLVII. 
fig. 96, ascus with sporidia, x 450. Sporidia at first colourless or pale green, at 
last dark brown, uniseriate; average length a little over 00004 inch. The sporidia 
have one, two, or three nuclei, and sometimes the nuclei are so close as to give a 
septate appearance to the sporidium; but I do not think there is ever any real 
septum. i 

98. S. (DrATRvPE) Movexortt, Pers. in litt. Tab. XLVII. fig. 97, ascus with sporidia, 
x 325. Sporidia biseriate, yellow (the same colour as in S. herbarum and S. siparia), 
irregularly oblong, constricted in the middle, multi-septate, with frequently longitu- 
dinal septa (or rather pseudo-septa; for all the septa are, I think, only apparent, 
arising from divisions of the endochrome), 0:0012 inch long. Perithecia with deci- 
duous (?) ostiola. Arranged in the herbarium with the Cespitose, but clearly belong- 
ing to the Versatiles. On dried branches of Acer Pseudo-platanus. 


Div. 8. CONCRESCENTES. 


99. S. FIBROSA, Pers.; Fr. S. M. ii. p. 384. Tas. XLVII. fig. 98, ascus with sporidia, x 
420. Sporidia uniseriate, obtusely elliptical, but slightly constricted in the middle, 
colourless; endochrome bipartite, 0-0005 inch long. 

100. S. DEPRESSA, Fr.; Montagne, Sylloge, p. 232. Ta». XLVII. fig. 99, ascus with 
sporidia, x 325; detached sporidia, x 450. Sporidia curved, of a pale yellowish 
colour, each ascus containing a great number. Asci with their sporidia much 
resembling SpA. favacea; sporidia 00004 inch long. 

101. S. PARALLELA, Fr. S. M. ii. p. 373. Tas. XLVII. fig. 100, sporidia highly magni- 
fied. Sporidia pale clear brown, straight or slightly curved, rounded at each end, 
0:0004 to 0:0005 inch long. | | 

102. S. (DiATRYPE) LEIOPLACA, Fr. S. M. ii. p. 370. Tas. XLVII. fig. 101, sporidia 
highly magnified. Sporidia biseriate, curved, of a pale yellowish tinge, just like the 
colour in S. stigma, 0:0004 inch long. Very like Sp. stigma, only there is no dark- 
coloured stroma. | | er 

103. S. FIMETI, Pers, var. EQUINA. Ta». XLVII. fig. 102, ascus with sporidia, and 
a free sporidium, x 325. Sporidia uniseriate, broadly ovate, sometimes slightly acu- 
minate, sometimes obtuse at the ends, of a deep brown colour, or almost black, quite 
opaque when ripe, 0:0008 inch long. When immature, the sporidia are pale brown, 
with large nuclei. ie : 

104. S. (DreLop1a) DrosPvnr, Schwein.; Fr. 8. M. ii. p. 872. This specimen is not in 
good condition. I could find no asci, and am doubtful whether or not the sporidia 
are produced on sterigmata as in Diplodia. The sporidia are dark brown, unisep- 
tate, obtuse, slightly constricted in the middle, 0:0006 to 0-0007 inch long. åå 

105. S. vELATA, Pers.; Fr. S. M. ii. p. 375. Ta». XLV. fig. 104, ascus with sporidia, 


x 325, and two free sporidia more highly magnified. Sporidia crowded, quite colour- 


274 MR. F. CURREY ON THE FRUCTIFICATION OF COMPOUND SPHARIA. 


less, pseudoseptate by division of the endochrome into 2 or 4 partitions, narrowly 
almond-shaped, 00005 inch long. 

106. S. (DrATRYPE) DISCINCOLA, Schwein.; Fr. S. M. ii. p. 368. Tap. XLVII. fig. 105, spo- 
ridia, X 325. Sporidia uniseriate, I think (at least it seemed so from what appeared 
to be the remnants of the asci), dark opaque brown, almost black, globular or ellip- 
tical, 00006 inch long. = 

107. S. (DiATRYPE) MILLIARIA, Fr. S. M. ii. p. 370. Tap. XLVII. fig. 106, sporidia, 
x about 450. Sporidia crowded, curved, colourless, 00003 inch long. Resembling 
S. eutypa both in habit and fructification ; but the ostiola are more prominent, and 
the perithecia more crowded. | 

108. S. (SPHERIA) LIMÆFORMIS, Schweinitz; Fr. S. M. ii. p. 869. Ta». XLVII. fig. 107, - 

- sporidia, x about 450. Sporidia, crowded, colourless when separate, yellowish when 
the asci are in a mass, curved, usually rounded at the ends, 0-0002 inch long. On 
oak, very black, very crowded, well-named * limæformis.” Fries considers it a form 
of Sphæria spinosa. 

109. S. (SPHÆRIA) spınosa, Pers.; Fr. S. M. ii. p. 368. Tap. XLVII. fig. 108, sporidia, 
X 325. Sporidia yellowish in the mass, almost colourless when single, curved, 
rounded or acute at the ends, 0-0003 inch long. 

110. S. zuevara, Berk, Tas. XLVII. fig. 109, asci with sporidia, and free sporidia, x 325. 
Sporidia numerous, crowded, closely packed, dull yellowish brown in a mass, almost 
colourless when separate, curved, rounded at the ends, 0:0006 to 0:0007 inch long. 
From Swan River, Mr. Drummond. 

lll. S. (DraTRYPE) LATA, Pers.; Fr. S. M. ii. p. 369. Tag. XLVII. fig. 110, sporidia, 
X 325. Sporidia crowded, yellowish in a mass, almost colourless when single, 
curved, obtuse at each end, 0:0002 to 0:0004 inch long. 

112, 8. (Sparta) sPICULOSA, Pers. ; Fr. S. M. ii. p. 369. Tas. XLVII. fig. 111, sporidia, - 
x 325. Sporidia narrowly almond-shaped, acuminate, quite colourless, with the 
endochrome divided into four portions, 0-0004 to 0-0005 inch long. Easily known 
by its black appearance and hair-like ostiola. | 


113. S. (VALSA) MONADELPHA, Fr. S. M. ii. p. 382. Fructification not distinguishable — 


from that of S. stigma, The contents of the peritbecia are glutinous, as in that . 
species. | 

114. S. (DraTRYPE or VALSA) ÆQUILINEARIS, Schwein.; Fr. S. M. ii. p. 374 In this 
plant the fructification is not distinguishable from that of the last. 

115. S. (Spars) Drscumrens, Berk. Engl; Fr. p. 245. Ta». XLVII. fig. 112, sporidia, 
X highly. Sporidia biseriate or crowded, 1-2- or 3-septate, or apparently so from 

. division of the endochrome, colourless, almond-shaped, 0:0005 inch long.  Perithecia 
deeply buried; ostiola long and protruding. 

116. SPHÆRIA STIPATA, Currey; Phil. Trans. 1857. Tas. XLIX. fig. 197, asci with spo- 
ridia, x 225. Sporidia biseriate, pale clear brown, very transparent, slightly curved, 
sometimes with å nucleus at each end, but usually without; 0:0008 to 0:001 inch 
long. Perithecia ovate, crowded ; ostiola sulcate, often exactly resembling those of 
S. stellulata. This plant might be placed either here or in the Confluentes with 


MR. F. CURREY ON THE FRUCTIFICATION OF COMPOUND SPHÆRLÆ. 275 


almost equal propriety; but the ostiola are somewhat elongated, and the perithecia 
immersed in the matrix. Eventually, however, the bark is thrown off, and the peri- 
thecia form rough, naked, scabrous masses. A dark line often surrounds the masses 
of perithecia ; but the area it includes is very irregular, and the line itself not always 
present. It might be taken for a coarse abnormal form of S. stellulata ; but the spo- 
ridia are more than double the size of those of S. stellulata, and of a different colour. 
It is very common in this country, principally on elm. 


Div. 9. CIRCUMSCRIPTÆ. 


117. S. (VALSA) ENTEROLEUCA, Fr. S. M. ii. p. 381. Tas. XLVII. fig. 113, ascus with 
sporidia, x 325. Sporidia biseriate, colourless, oblong, acuminate at each end, con- 
stricted in the middle, often slightly curved, 4-nucleate, uniseptate (I think, but the 
septum very difficult to make out), 0:0006 to 0:0007 inch long. On Robinia Pseuda- 
cacia. Perithecia very numerous in each pustule, deeply imbedded in a very white 
stroma; when cut across horizontally, a very well-defined black line is seen, which 
arises from the wood surrounding the perithecia being blackened, thereby causing the 
appearance of the existence of a conceptaculum ; but there is not really any. I have 
received a plant from Mr. Bloxam marked as S. enteroleuca, Fr., of which fig. 114 
represents an ascus with sporidia x 325, and free sporidia more highly magnified. 
The sporidia are very numerous, crowded, brownish yellow in the mass, almost colour- 
less when single, curved, 0:0003 inch long. I suspect, from the habit of the latter 
plant upon its fruit, that it is a form of Diatrype aspera. 

118. S. (Varsa) ANOMIA, Fr. S. M. ii. p. 381; El. ii. p. 77. Sporidia (spermatia ?) 
colourless, slightly curved, very minute, about 0:0002 inch long. A very doubtful 
specimen ; but the fruit does not differ very materially from that figured in Sturm's 
* Deutschland's Flora.’ 

119. S. (VALSA) EXTENSA, Fr. S. M. ii. p. 381. Tas. XLVII. fig. 116, ascus with 
sporidia, x 325. Sporidia uniseriate, colourless, obtuse, slightly constricted in the 
middle; endochrome bipartite, smooth, and refractive, but doubtless sometimes 
granular. 

120. 8. (Varsa) DETRUSA, Fr. S. M. ii. p. 382. Tas. XLVII. fig. 117, ascus with spo- 
ridia, x 325, and a free sporidium, x 450. Sporidia biseriate, colourless; endochrome 
divided into four, sometimes apparently into only two portions, elliptie, subobtuse or 
subacuminate at the ends ; 0:0006 inch long. 

121. S. (VALSA) PmvasrRI, Pers.; Fr. S. M. ii. p. 980. Sporidia biseriate, almost 
colourless, but brownish yellow in the mass; minute, curved, 0:0003 inch long. 
Hardly distinguishable from the fruit of Diatrype stigma. 

122. S. (Vasa) CARPINI, Pers.; Fr. S. M. ii. p. 384 Tas. XLVII. fig. 118, sporidia, 
x 325. Sporidia curved, colourless, highly refractive, 00006 to 0-0008 inch long. 
There is another specimen in the Herbarium, marked SpA. Carpini; but as the 
sporidia are just like those of Spå. syngenesia, it probably belongs to the latter 
Species. 

123. S. (VALSA) SYwGENEsIA, Fr. S. M. ii. p. 382. Tas. XLVII. fig. 119, sporidia, x 

' VOL. XXII. 20 


276 MR. F. CURREY ON THE FRUCTIFICATION OF COMPOUND SPHÆRLÆ. 


325. Sporidia biseriate or crowded, colourless, elliptic-acuminate; endochrome 4. 
partite, sometimes only 2-partite; 0:0005 to 0:0006 inch long. 

124. 8. (VALSA) CELLULATA, Fr. Syst. Mye. ii. p. 880. Tas. XLVII. fig. 120, sporidia, 
x 450. Sporidia biseriate, very pale yellowish brown, simple, slightly curved, rounded 
at both ends, 0:0004 inch long. | 

125. S. (DIPLODIA ?) JUGLANDICOLA, Schwein. ; Fr. S. M. ii. p.385. Tas. XLVII. fig. 121, 
fruit, x 325. Sporidia (?stylospores) uniseptate, dark brown, 00009 to 0.001 inch 
long. To the naked eye the plant resembles S! pulvis-pyrius. 


Div. 10. Inousz. 


126. S. (Varsa) ANGULATA, Fr. S. M. ii. p. 890. Tas. XLVII. fig. 122, ascus with spori- 
dia, and free sporidia, x 225. Sporidia uniseriate, very seldom biseriate, eolourless 
or pale sea-green, obtuse, constricted in the middle, furnished with 4 (? sometimes 5) 
cilia, one proceeding from each pole and one from the middle of each side; endo- 
chrome usually granular; 0:0008 to 0-0012 inch long. The above is the usual form 
of the sporidia; but sometimes, when the endochrome is oleaginous and not granular, 
the appearance of the sporidia is so different that they might almost be taken for a 
second form of fruit. A second form of fruit does exist in this species, as to which I 
would refer to my paper “On the Fructification of certain Sphæriaceous Fungi,” 
read before the Royal Society, in June 1857. See Phil. Trans. 1857. 

127. S. (VALSA) MELASPERMA, Fr. S. M. ii. p.989. Tap. XLVII. fig. 123, ascus with 
sporidia, and free sporidia, x 325. Sporidia very numerous, slightly curved, of a 
pale-yellowish tinge, 0:0002 to 0:0003 inch long. FE 

128. S. (Vasa) TALEOLA, Fr. S. M. ii. p.990. Tag. XLVII. fig. 124, ascus with sporidia, 
and free sporidia, x 450. Sporidia biseriate, colourless or very pale green, elongate- 
acuminate, frequently or usually slightly curved, 0:0007 to 00008 inch long. I think 
there is no real septum. I find not the slightest difference between the sporidia of 
Sphæria taleola and those of S. leiphemia, and have little doubt that the two plants 
are not distinet. I feel sure that the conceptaculum which is said to distinguish 
S. taleola is a characteristic not to be relied upon; for there are some species of 
Spheria which sometimes have a conceptaculum and sometimes not. The plants 
marked §. ¢aleola at Kew have no conceptaculum ; and I have seen other specimens 
marked S. taleola also without a conceptaculum. | 

129. S. (Varsa) FULVO-PRUINATA, Berk. Tas. XL VII. fig. 125, asci with sporidia, x 325. . 
Sporidia uniseriate, sometimes slightly acuminate at each end, but sometimes obtuse 
at the ends, slightly constricted in the middle, uniseptate, dark brown, paler at first 
(and then an inner membrane and sometimes nuclei are visible), 0-0006 to 0-0007 inch 


eh Easily recognized by the tawny colour of the protruded bark surrounding the 
ostiola. 


130. S. (Varsa) NIVEA, Hoffm. ; Fr. S, M. ii. p. 386. Tas. XLVII. fig. 126, asci with 
sporidia, and free sporidia, x 325. A sei crowded with sporidia; sporidia colourless, 
slightly curved, 0:0002 to 0-0003 inch long. 

131. B. (VALSA) FORAMINULA, Pers. in litt. Tan. XLVII. fig. 127, ascus with sporidia, and 


MR. F. CURREY ON THE FRUCTIFICATION OF COMPOUND SPHÆRLÆ. 277 


free sporidia, x 325. Sporidia biseriate, elliptical, sometimes acuminate, sometimes 
obtuse at the ends, colourless, 0:0011 to 00016 inch long. In some sporidia the 

, endochrome was broken up into two, three, or six portions, but, I think, only from 
being dry. 

132. S. (VALSA) PROFUSA, Fr. S. M. ii. p. 392. Ta». XLVII. fig. 128, sporidia, x 450: 
a, à young sporidium almost colourless or very slightly greenish; 5, a sporidium 
more advanced; c, a ripe sporidium. The sporidia when young have a gelatinous 
envelope, which disappears in age, the asci frequently containing only four sporidia 
arranged in one continuous series; I believe, however, the normal number to be 
eight, and that they are sometimes arranged biseriately. "The asci are absorbed at a 
very early period, so that usually only free sporidia are to be found. "These when 
ripe are of a clear rich olive-brown colour with a very slight greenish tinge, oblong- 
elliptic, 0:001 to 0:002 inch long. One of the most beautiful Sphæriæ in point of 

. fructification. 

133. S. (DIPLODIA ?) RUDIS, Fr. El. ii. p. 98. Tas. XLVII. fig. 129, fruit, x 325. No 
asei; sporidia (?stylospores) rather dark brown, broadly elliptical, but slightly con- 
stricted in the middle, 0:0010 to 0:0011 inch long. 

134. S. (VALSA) MICROSTOMA, Pers.; Fr. S. M. ii. p. 388. Ta». XLVIL fig. 130, ascus 
with sporidia, and free sporidia, x 325. Sporidia biseriate, colourless, transparent, 
curved, obtuse at the ends, 0:0005 to 0:0007 inch long. 

185. S. (VALSA) DISSEPTA, Fr. S. M. ii. p.392. Tas. XLVIII. fig. 131, asci with sporidia, 
and free sporidia, x 325. Sporidia biseriate, colourless, linear-acuminate or almond- 
shaped; endochrome bi- or quadripartite; 0*0004 to 00005 inch long. 

136. S. (VALSA) CIROUMSCRIPTA, Fr.; Mont. Syll. p. 220, under “ Valsa.” Tan. XLVIII. 
fig. 132, sporidia, x 325. Sporidia biseriate, colourless, linear, pointed at the ends ; 
endochrome bipartite, sometimes quadripartite ; 0:0006 inch long. This agrees with 
Montagne's description, except that he calls the sporidia obtuse. = 

137. S.(VaLsa) Kunzen, Fr. S. M. ii. p.388. Tas. XLVIII. fig. 133, ascus with sporidia, 
and free sporidia, x 325. Sporidia biseriate, brownish yellow in the mass, almost 
colourless when single, curved, 0:0004 inch long. Described from a specimen of 
Mr. Berkeley’s, that at Kew having no fruit. | 5 bi | be 

188. S. (VALSA) CONCAMERATA, n.s. - TAB. XLVIII. fig. 134, ascus with sporidia, x 825. 
Sporidia crowded, colourless, curved, 0:0004 inch long ; perithecia raising the inner 
bark into a dome-shaped conceptaculum, nests of perithecia united by white woolly 

- fibres. On oak; quære, a form of S. ceratosperma ? MR 
139. S. crxcra, n.s. Tas. XLVIII. fig. 185, ascus with sporidia, x 325. Sporidia unise- 
riate, dark rich brown, obtuse, constricted in the middle, pes - long, och ; 
* Ke ] : 1 eathery conce uium ; 
ger st i RE PE se gist gm ne en lens the 
ostiola bursting through the bark and forming ro p 
ostiola seem surrounded with a dirty-olive-green stroma. Blackheath Park, March 
1855. | j *3* . * H 
140. S. (Varsa) SorBI, Schmidt; Fr. S. M. ii. p. 380. Sporidia not distinguishable from 


those of S. stigma. | 
202 


278 MR. F. CURREY ON THE FRUCTIFICATION OF COMPOUND SPHÆRLÆ. 


141. S. (Varsa) SACCULUS, Schwein.; Fr. S. M. ii. p.378. In the sphæropsoid state, 
spermatia colourless, delicate, slightly curved, obtuse or subobtuse at the ends, 
0:0002 to 0-0003 inch long. — | 

142. S. ENTEROXANTHA, Berk. Fructification exactly similar to that of S. stigma, as well 
in the glutinous nature of the contents of the perithecia as in the size, shape, and 
colour of the asci and sporidia. From British Guiana. 

143. S. (Varsa) Crarzer, n.s. Tas. XLVIII. fig. 135 a, sporidia, x 425. Sporidia bise- 
riate, oblong or elliptical, sometimes curved; endochrome 4-, sometimes 2-partite, 
colourless, 00006 to 0:0010 inch long; perithecia irregularly globose; ostiola rather 
short. Very common on thorn near Blackheath and elsewhere. 

144. S. (VALSA) DRYINA, n.s. Ta». XLVIII. fig. 1355, ascus with sporidia, and free 
sporidia, X325. Sporidia biseriate or crowded, colourless, strongly curved ; both asei 
and sporidia very delicate and hyaline; 0:0003 inch long. On dead oak-branches at 
Weybridge. Perithecia pyriform, imbedded in a dirty-brown stroma; ostiola rather 
shorter than the body of the perithecia, and somewhat thickened towards the apex. 


Div. 11. OBVALLAT A. 


145. S. (Varsa) coronama, Hoffm. ; Fr. S. M. ii. p.395. Tas. XLVIII. fig. 136, ascus with 
sporidia, x 325. Sporidia colourless, biseriate, elliptic-acuminate, constricted in 
the middle, 0:0007 to 0-0008 inch long. 

146. S. (Varsa) LEIPILEMIA, Fr. Syst. Myc. ii. p. 399. Tas. XLVIII. fig. 137, sporidia, 
X 450. Sporidia biseriate, colourless or pale green, elongate-acuminate, slightly 
curved ; endochrome refractive or granular, consisting of two distinct portions with a 
clear space (not, I think, a septum) between them; the outer membrane is so delicate 
that it is frequently not visible. I have found specimens in which the sporidia have 
oozed out of the perithecia and formed round rose-coloured tubercles at the apices 
of the ostiola, looking like Tubercularia vulgaris (see remarks under S. taleola). 
Length of sporidia about 0:0007 inch. | 

147. 8. (Varsa) AMBIENS, Pers. Syn. p.44. ; Fr. S. M. ii p.403. Ta». XLVIII. figs. 188, 
138 a, asci with sporidia, and free sporidia, x 325. Sporidia colourless, crowded, 
curved, rounded at each end, 00006 inch long. 

148. 8. (Varsa) TURGIDA, Pers.; Fr. $. M. ii p. 400. Tas. XLVIII. fig. 139, ascus 
with sporidia, and free sporidia, x 325. Sporidia uniseriate, at first a pale clear 
greenish brown with two or more nuclei, eventually a very dark opaque brown 
without (or with hardly visible) nuclei, elliptical, subacuminate, 00003 to 0:0004 
inch long. 

149. S. (Varsa) srILBOSTOMA, Fr. S. M. ii. p.403. Tap. XLVIII. fig. 140, ascus with 
sporidia, and free sporidia, x 325. Sporidia biseriate, colourless, acuminate-elliptic 
but constricted in the middle; the endochrome divided into two portions (not 
I think, septate), sometimes smooth, glossy, and highly refractive, sometimes 
granular; 0-0007 inch long. The variety Platanoides (see fig. 140 a) has triseptate 


or pseudo-triseptate sporidia, which are 0-0010 to 0:0011 inch long, and sometimes 
mucronate. ; 


1 


MR. F. CURREY ON THE FRUCTIFICATION OF COMPOUND SPHÆRIÆ. 279 


150. 8. (Y ALSA) QUERNA, n. S. Tas. XLVIII. fig. 141, ascus with sporidia, x 325. Spo- 
ridia slightly curved, simple, linear, colourless, subhyaline, crowded at the apex of 
the ascus as in many Pezize, 0-0002 to 0-0003 inch long. Weybridge, 1856. Not 
in a sufficiently good condition to describe the perithecia accurately. The fruit 
differs from that of any of the allied species with which I am acquainted. 

151. S. (VALSA) BICONICA, n. s. Weybridge, January 7, 1856. Ta». XLVIII. fig. 142, 
ascus with sporidia, and free sporidia, x 220.  Sporidia biconical; endochrome gra- 
nular or oleaginous, greenish; 0'0011 to 0:0012 inch long; perithecia globose or 
depressed, few together, arranged in circles, penetrating the bark with their rather 
short ostiola and forming small pustules. 

152. S. (VALSA) PULCHRA, n. s. Tap. XLVIII. fig. 143, ascus with sporidia, x 225. 
Sporidia biseriate, elliptical but slightly constricted in the middle and slightly 
acuminate at each end, uniseptate, greenish ; perithecia imperfect. 

153. S. (Vausa) SUFFUSA, Fr. S. M. ii. p. 399. Ta». XLVIII. fig. 144, sporidia, x 225. 
Sporidia variously curved from their great length, colourless, granular, biseriate or 
crowded. 

154. S. (Varsa) JUGLANDIS, Schwein., non Fries. Tas. XLVIII. fig. 145, sporidia, x 
450. Sporidia biseriate, yellowish, as in S. stigma, 0-0003 to 00004 inch long. 

155. S. (Varsa) ABIets, Fr. S. M. ii. p. 398. Tas. XLVIII. fig. 147, ascus with sporidia, 
and free sporidia, x 325. Sporidia biseriate, colourless, curved, 0:0003 to 0:0006 
inch long. This I believe to be the true S. Abietis of Fries. Fig. 146 represents 
the fruit of a Spheria also occurring on Fir and resembling S. Abietis in general 
appearance. The sporidia in this latter plant are biseriate, colourless, subhyaline, 
curved, acuminate at each end, 1-septate or pseudo-septate, 0-0007 inch long. It is 
possible that the species may not be really different from S. Abietis, and that the 
latter may, in its perfect state, have septate sporidia. There are, as stated above, 
several Sphæriæ in which the normal state of the fruit is septate, but in which that 
state is often not attained. 

156. S. (VALSA) TETRASPORA, n. $s. Weybridge, January 1856. Tas, XLVIII. fig. 148, 
ascus with sporidia, and free sporidia, x 325. Sporidia simple, slightly curved, rounded 
at each end, colourless, only 4 in each ascus, biseriate, 0:0008 inch long; perithecia 
globose; ostiola surrounding or scattered through a white disk. The plant hardly 
differs from S. ambiens, except in its tetrasporous fructification. : 

157. S. (VALSA) SALICINA, Pers.; Fr. S. M. ii. p. 401. TAB. XLVIII. fig. 149, ascus with 
sporidia, and free sporidia, x 325. Sporidia biseriate, colourless, uniseptate, with gra- 
nules and small nuelei, the septum often very difficult to make out, elliptical, but 
usually slightly curved, and often slightly constricted in the middle, 0-0009 inch long. 

158. S. (P VALSA) INTEXTA, n.s. TAB. XLVIII. fig. 169, ascus with sporidia, x 325. 
Sporidia interwoven, generally nearly as long as > ascus, filiform and flexuous. 
This plant was not in a state to enable me to describe the perithecia. It appears to 


belong to the Circinatæ, and occurred on oak at Weybridge. The paraphyses were 


very long and numerous, and septate. I know of no species to which it can be 


referred, although the sporidia somewhat resemble those of S. suffusa, Fr. 


280 


159. 


160. 


161. 


162. 


163. 


164. 


165. 


166. 


MR. F. CURREY ON THE FRUCTIFICATION OF COMPOUND SPHÆRLÆ. 


Div. 12. CIrCINATÆ. 


S. (VALSA) PULCHELLA, Pers.; Fr. S. M. ii. p. 406. Tas. XLVIII. figs. 150-153, 
asci with sporidia, and (in fig. 152) free sporidia, X 325. Sporidia biseriate or 
crowded, colourless, slightly curved or nearly straight; normally uniseptate, but the 
septum sometimes not distinguishable; varying much in size, 0-0005 inch in length 
being about the average. The sporidia are sometimes rounded at the ends, some- 

times slightly acuminate. 

S. (VALSA) FURFURACEA, Fr. S. M. ii. p.409. Tas. XLVIII. fig. 154, sporidia, 

x 450. Sporidia colourless or pale green, uniseriate, 00008 inch long. 

S. (VALSA) CONVERGENS, Tode; Fr. S. M. ii. p. 410. Ta». XLVIII. fig. 155, fruit, 

x 325. Sporidia dark brown, subeymbiform but irregular in shape, frequently con- 

stricted in the middle; length variable. On Platanus occidentalis, the perithecia 

being under the bark so as not to be seen above, except by the undulations of the bark. 

I found that some of the perithecia produced, in lieu of the regular sporidia, numbers 

of the small diplodioid bodies shown in the figure, below the sporidia. One of the 
sporidia has commenced germination. 

S. (VALSA) HYPODERMIA, Fr. S. M. ii. p.407. Tas. XLVIII. fig. 156, asci with spori- 

dia, x 325. Sporidia colourless, granular, narrowly oblong, obtuse or slightly acu- 

minate at each end, sometimes with two or more large globose nuelei, biseriate. 

S. (VALSA) THELEBOLA, Fr. S. M. ii. p.408. Tas. XLVIII. figs. 157, 158, and 159, 

asci with sporidia, and free sporidia, x 325. Sporidia biseriate, greenish or colourless, 
slightly arcuate, obtuse at both ends ; usually with a cilium at each end, which is not 

visible whilst the sporidia are in the ascus. These cilia require care, to be rendered 

visible even when the sporidia are detached ; they are sometimes absent. Length of 
the sporidia and size of the asci variable, the sporidia sometimes reaching 0:0017 

inch. These figures show in a marked manner the difference in the appearance of 
the sporidia when the endochrome is oleaginous and refractive, and when it is 

granular. | 

S. (VALSA) XANTHOSTROMA (or CHRYSOSTROMA), Mont. Sylloge, p.221. Tas. XLVII. — 
fig. 160, ascus with sporidia, and free sporidia, x 350. Sporidia uniseriate or bise- — 


riate, more often the former, colourless or greenish, almond-shaped, pseudo-septate — 


from the division of the endochrome, 0-0007 inch long. Perithecia small, hardy — 
visible above the bark, but (like all the Circinatæ) very visible when the epidermis is 
stripped off. Easily known by its yellow stroma. Endochrome smooth and refractive. 
In another specimen of the same species I found the sporidia not septate, and reach- 
ing nearly 0:0009 inch in length. 

S. (Varsa) vesrira, Fr. S. M. ii. p.410. Tas, XLVIIL fig. 161, ascus with sporidia, 
X 425. Sporidia clear brown, multicellular, having transverse, longitudinal, and 
oblique septa, 0:0006 to 0:0009 inch long. As to the fructification of this species, 8 
my paper in the ‘Phil. Trans.’ 1857, above referred to. En. 
S. (DrPLoprA?) Meur, Fr. El. ii. p. 85. Tap. XLVIIL fig. 102, sporidia, x 325- 
Sporidia clear dark brown, some margined, some (not many) with a central nucleus, 


MR. F. CURREY ON THE FRUCTIFICATION OF COMPOUND SPHARIA. 281 


elliptic, rather irregular, varying much in length, the average being about 0:0008 
inch. I could find no asci. 

167. S. (Varsa), Psp. Tas. XLVIII. fig. 163, ascus with sporidia, x 225. This plant is 
one of the * Fungi Pyrenei,” and is marked “ S. cohærens, Pers." in that collection. 
It has, however, no affinity with S. cohærens, but belongs to the Circinatæ. The 
fruit differs from that of all other species of the division Cireinate with which I am 
acquainted. The sporidia are uniseriate, of a rich brown colour, obtusely elliptical, 
uniseptate, 00014 inch long. 

168. S. (VALSA) QUATERNATA, Pers. Syn. p. 45. Ta». XLVIII. figs. 164 and 165, asci 
with sporidia, and tree sporidia. In fig. 164, a is x 220, b about 325 diameters. 
Fig. 165 isx450. Sporidia biseriate, almost colourless when separate, but in a mass 
of dull-brownish yellow, like the colour in S. stigma, S. stellulata, &e., curved, 0:0004 
to 0:0006 inch long. 

169. S. (VALSA) INNESI, n. s. Tas. XLVIII. fig. 166, sporidia, x 325. Sporidia 
biseriate, colourless, acute at each end, and constricted three times; endochrome 
4-partite ; 0°0010 to 0-0012 inch long. Sporidia frequently, if not always, with a 
delicate appendage at each end. Perithecia irregularly globose; ostiola elongated and 
frequently thickened at the apex. Externally much resembling S. pulchella, but 
smaller and differing altogether in fructification from that species. 

170. S. (VALSA) ARCUATA, n.s. Tas. XLVIII. fig. 167, asci with sporidia, and free spori- 
dia, x 325. Perithecia globose, arranged in circles; ostiola penetrating the bark and 
forming a dark-coloured, sometimes dirty-white disk. Sporidia biseriate, pale green, 
4-5-septate, very flexible, sometimes constricted at the articulations, sometimes not, 
variable in length, somewhat resembling 5. quercina, Pers. in its sporidia; but S. 
quercina belongs to the Versatiles, and this plant to the Circinatæ. 

171. S. (VALSA) FAGINEA, n. s. Eltham Grove, October 1856. Tan. XLVIII. fig. 168, 
sporidia, x 420. Sporidia biseriate, colourless, elliptic-acuminate, but constricted in 
the middle, and irregular, 0:0005 inch long. Perithecia conical; ostiola penetrating 
the bark, normally long and protruding, but mostly broken off; when the long ostiola 
are rubbed off, the plant looks just like S. quaternata or S. turgida. On beech. 


Div. 15. CÆSPITOSZ. 


172. SPHÆRTA CUPULARIS, Pers. ; Fr. S. M. i.p. 416. Ta». XLVIII. fig. 170, asci with 
sporidia, and free sporidia, X 325. Sporidia biseriate, colourless, curved, 0°0004 inch 
long, sometimes rather longer. 

173. S. PsEUDo-BOMBARDA, Mont. Sylloge, p. 228. Ta». XLVIII. fig. 171, ascus with 
sporidia, and free sporidia, X 225. Sporidia biseriate or erowded, colourless, irre- 
gular, 0:002 inch long. This plant was in a young state ; the spores when ripe are 
described by Dr. Montagne as 6-septate and * olivaceo-fuliginosas.” sj 

174. S. (NzorRIA) EXAMINANS, Berk. Tap. XLIX. fig. 172, ascus with sporidia, and free 
sporidia, x 325. Sporidia uniseriate or biseriate, clear dark brown, elliptical, sub- 
acuminate, 0-0008 inch long. Perithecia forming long black lines. The contents of 
the sporidia were mostly clear, sometimes multinucleate, and sometimes granular. 


282 MR. F. CURREY ON THE FRUCTIFICATION OF COMPOUND SPHÆRIÆ. 


175. S. (NECTRIA) COCCINEA, Pers. ; Fr. S. M. ii. p. 412. Ta». XLIX. figs. 174 and 175, 
ascus with sporidia, and free sporidia, X 325. Sporidia almost always uniseriate, 
colourless, uniseptate, frequently with a nucleus in each septum, elliptical, subacu- 
minate, 0:0005 inch long. In the variety “ sanguinella”” I did not find any difference 
in the fruit, except that the average size of the sporidia was somewhat smaller, 
Fig. 175 represents free sporidia of the variety * cicatricum,” Desm., x 450. 

176. S. (NECTRIA) CINNABARINA, Tode; Berk. Engl. Flora, No. 77, under * Sphæria.” 
Tas. XLIX. fig. 175, sporidia, x 450. Sporidia sometimes uniseriate, sometimes 
biseriate, normally I believe uniseptate, frequently if not always constricted in the 
middle, rather pointed at each end, colourless or pale sea-green, 0:0004 to 0:0006 
inch long. I found some sporidia attached to long threads; but whether they were 
in the nature of stylospores, or whether the threads were the product of germination, 
I cannot say. Several of the sporidia were biseptate. See a, fig. 175. 

177. 8. (NECTRIA) DECOLORANS, Pers. Syn. p. 49. Tas. XLIX. fig. 176, ascus with spo- 
ridia, and free sporidia, x 325. Sporidia uniseptate, colourless or greenish, 0:0005 
inch long. Placed by Fries under S. cinnabarina, from which it differs in its more ` 
compact habit and in the more broadly elliptical sporidia ; the perithecia, however, 
are rugged as in that species, with which perhaps it ought to be united. 

178. S. BERBERIDIS, Pers.; Fr. S. M, ii. p. 415. Tas. XLIX. fig. 177, sporidia, X 325. 
Sporidia uniseriate, but so overlapping as to be almost biseriate, yellow when young, 
brown when mature, constricted in the middle, usually acuminate at the ends, 0:0011 - 
to 00016 inch long. Quiere if distinct from S. Laburni ? 

179. S. (NECTRIA) CUCURBITULA, Tode; Fr. S. M. ii. p. 415. Tap. XLIX. fig. 178 a, 
sporidia ; b, spermatia ; and c, caudate irregular bodies : all x 325. Sporidia colour- 
less, irregularly elliptical, 0:0004 inch long. Two membranes very visible in the 
sporidia. Besides the normal asci and sporidia, the perithecia produce asci containing 
the spermatia-like bodies (5) on the right: and the caudate bodies on the left also 
appeared to proceed from the same perithecia; but of this I am doubtful. The 
length of the spermatia is 0-0001 to 0-0002 inch, of the caudate bodies 0:0006 

- inch; what the latter may be I cannot say. 

180. S. LABURNI, Pers. Syn. p. 50. Tas. XLIX. fig. 179, ascus, x 225, and free sporidia, 
x 450. Sporidia, when perfect, multicellular, dark clear brown. I find in different 
perithecia in the same stroma every gradation from colourless minute unicellular 
spores (mostly with a nucleus at each end), through septate, biseptate, and triseptate 
forms, up to the multicellular. The colour varies by degrees, being at first colourless, 
then green, and eventually dark clear brown. 

181. S. PULICARIS, Fr. S. M. ii. p. 417. Tas, XLIX. fig. 180, ascus with sporidia, and 

free sporidia, x 325. Sporidia biseriate, colourless, elliptical, sometimes pyriform, 

pseudo-triseptate from division of the endochrome, 00006 to 0:001 inch long. - 

= (Nzorria) Purton, Grev. Tas. XLIX. fig. 181, ascus with sporidia, X "n 

poridia uniseriate, colourless, acuminate, elliptical, 00004 inch long. Fries Jo 


this species with S. Abietis; I do not know why, as it is quite distinct in habit and 
fructification, 


182. 


x 


MR. F. CURREY ON THE FRUCTIFICATION OF COMPOUND SPHÆRLÆ. 283 


183. S. (NECTRIA) ACERVALIS, Moug.; Fr. El. ii. p. 88. Ta». XLIX. fig. 182, ascus with 
sporidia, X 325. Sporidia uniseriate, rarely biseriate, colourless, elliptical, uniseptate 
or pseudo-uniseptate, 0:0005 inch long. 

184. S. (NEcrRIA) AQurronn, Fr. El. ii. p.82. Tas. XLIX. fig. 183, ascus with sporidia, 
and free sporidia, x 325. Sporidia uniseriate, colourless, elliptical, pseudo-uniseptate 
by division of the endochrome, which is oleaginous or granular; 00005 inch long. 

185. SPHÆRIA ACERVATA, Fr. S. M. ii. p. 416. Ta». XLIX. figs. 184, 185, ascus with 
sporidia, and free sporidia, x 325. Sporidia biseriate, colourless, hyaline, curved, 
0:0003 to 0-0005 inch long. Very like (? different from) S. cupularis. Mr. Bloxam 
informs me that he has sometimes found the sporidia elliptical, uniseptate, or with 
several nuclei. 

186. SPHÆRIA CONGLOBATA, Fr. The Hookerian herbarium contains several plants 
marked S. conglobata, Fr.; but they are all specimens of S. pulvis-pyrius, some being 
subcuticular, some bursting transversely, some longitudinally, others forming a 
cæspitose mass, others in the usual scattered condition. They form a very instruc- 
tive series of specimens, showing the Protean habit of S. pulvis-pyrius. The speci- 
men of S. dioica, Fr., at Kew, is also a subeuticular form of S. pulvis-pyrius, a form 
by no means uncommon in this country. 

187. S. Bux1. Milton, Northamptonshire, Berkeley. Ta». XLIX. fig. 186, ascus with 
sporidia, highly magnified. Sporidia uniseriate, ? colourless, acuminate, elliptical, 
0-0005 inch long. Perithecia light yellow, rather longer than broad, with a small 
mamillate ostiolum. I doubt if the sporidia are not sometimes uniseptate. The 
fruit was hardly ripe, although a good many sporidia were visible. 


Div. 14. CONFLUENTES. 


| 188. SPHÆRIA SCORIADEA, Fr. El. ii. p. 87. Ta». XLIX. fig. 187, sporidia, x 325. Spo- 
ridia reddish brown, opaque, lageniform, when young with a gelatinous envelope. 'The 
tips of the sporidia are paler than the body ; they are irregular in length and breadth, 
varying from 0-0016 to 0:0028 inch in length. There is some doubt whether - 
plant be not a Verrucaria. See Annals of Nat. Hist. vol. vi. p. 360, ren s 
* British Angiocarpous Lichens,” p. 39, under * Verrucaria conferta, Tayl. 

189. SPHÆRIA RHIZOGENA, Berk. Ta». XLIX. fig. 190, — with sporidia, X 325. 
Sporidia uniseriate, rarely biseriate, colourless or greenish, probably brown when 
ripe; elliptical, sometimes slightly constricted in the middle, uniseptate or biseptate 
or (?) sometimes cellular, 0:0005 to 00006 inch long. On the roots of Gleditschia 
triacanthos, washed bare by the Ohio freshets. Perithecia seated on or immersed in 
a ferruginous woody or leathery stroma, producing, when young, Very minute sub- 

indrical spermatia. 

190. er et, Ness; Fr. S. M. ii. p. 424. Tas. XLIX. fig. 189, ascus with 
sporidia, x 325. Sporidia uniseriate, frequently (if not usually) slightly overlapping, 
dark brown, cellular, acuminate, constricted in the middle, 0:0011 to 0:0012 inch long. 
This species — S. elongata, Fr. 

191. Sez ERrA ? Dorurea, Fr. (? Mont.), 

VOL. XXII. 


S. M. ii. p.423. Ta». XLIX. fig. 190, ascus 
2P 


284 MR. F. CURREY ON THE FRUCTIFICATION OF COMPOUND SPHÆRLÆ. 


with sporidia, x 325. Sporidia crowded, colourless, broadly almond-shaped, 0:0007 
to 0:0009 inch long. 

192. SPHÆRIA MUTILA, Fr. S. M. ii. p. 424. Tas. XLIX. fig. 191, sporidia, x 325. 
Sporidia crowded, colourless, but with a greenish tinge; endochrome divided trans- 
versely, and frequently also longitudinally; sporidia irregular in shape, variable, 
0:0008 to 0:0012 inch in length. 

193. SPHÆRIA MEGALOSPORA, Mont. Sylloge, p.229. Tas. XLIX. fig. 192, ascus with 
sporidia, X 225. Sporidia uniseriate, rich dark brown, obtusely elliptical, constricted 
in the middle, frequently slightly curved, 0:0016 inch long. 

194. SPHÆRIA INSIDENS, Schw.? Tag. XLIX. fig. 193, sporidia highly magnified. I could 
find no asci. Sporidia pale clear brown, irregularly elliptical or subturbinate, 0:0003 
inch long, or rather less; on rough corky elm-bark, blackening the furrows with its 
small perithecia. 

195. SPHÆRIA MELOGRAMMA, Pers.; Fr. S. M. ii. p.420. Tap. XLIX. fig. 194, sporidia, 
x 450. Sporidia biseriate, arcuate, acuminate at both ends, triseptate, pale greenish 
brown, probably ultimately clear dark brown, 0:0014 to 0-002 inch long. 

196. SPHÆRIA CALYCANTHI, Fr. (sed qu. Schwein.), S. M. ii. p.421. Ta». XLIX. fig. 195, 
ascus with sporidia, x 325. Sporidia crowded, colourless, hyaline, with one or many 
nuclei, or none; broadly almond-shaped, 0:0008 to 0:0011 inch long. 

197. 8. (DIATRYPE) DECIPIENS, Dec.; Fr. S. M. ii. p.371. Tag. XLIX. fig. 196, sporidia 

. highly magnified. I could find no asci. Sporidia elliptical, varying much in length 
and breadth, sometimes slightly curved, rather dark brown, 0:0003 inch long, crowded. 
. Ostiola stellate, just as in S. stellulata. 

198. S. (DIPLODIA) SUBSOLITARIA, Schwein. ; Fr. El. ii. p. 86. Tas, XLIX. fig. 198, spo- 
ridia, x 325. This plant might be placed in the genus Diplodia, the sporidia being 
borne on sterigmata, not produced in asci. Sporidia of a clear brown colour, 
irregularly elliptical or subpyriform, pseudo-uniseptate from the division of the 
endochrome. ! 

1984. SPHÆRIA CHRYSENTERA, n.s. Perithecia rounded, without any neck, crowded, 
situated underneath the outer bark, which is thrown off sometimes irregularly, some- 
times in rings. Contents of the perithecia yellow. Sporidia biseriate, bright yellow 
when young, then a pale clear light brown, eventually darker brown; irregular in 
shape, usually somewhat acuminate at each end, always constricted. in the middle, 
the outline sometimes wavy, nof (T think) septate, although the endochrome is always 
divided into two distinct portions, frequently into four, and occasionally into three. 
Length of sporidia, 0-0008 to 0-0009. inch. Weybridge, September 1856. Easily 
known by the bright yellow colour of the contents of the perithecia, and by its pecu- 
liar fruit. The perithecia a 


re sometimes seated on a byssoid subiculum, which made 
me doubt to which division it should be referred. 


1 Div. 15. Serrarx. 
99. J P Fr. Tas. XLIX. fig. 199, sporidia, x 325. Sporidia biseriate, yellowish- 
own, linear-ac te, 3-septate, 0:0012 inch long, hardly distinguishable from the 


MR. F. CURREY ON THE FRUCTIFICATION OF COMPOUND SPHJERLE. 285 


fruit of S. arundinacea. S. Junci is placed, in the * Summa Veg. Sc., after Dothidea, 
with the remark “ Incerti generis, priori affinis." 

200. SPHÆRIA ARUNDINACEA, Sow. Tas. XLIX. fig. 200, sporidia, x 326. Sporidia 
biseriate, yellowish-brown, linear, acuminate at one or both ends, 3-5-septate, 0*0010 
to 0:0016 inch long. 

201. SPHÆRIA NEBULOSA, Pers.; Fr. S. M. ii. p. 430. Tap. XLIX. fig. 201, ascus with 
sporidia, x 325. Sporidia biseriate, colourless, subacuminate, straight or slightly 
curved, uniseptate, 0:0003 to 0:0005 inch long. 

202. SPHÆRIA GODINI, Desm. Ta». XLIX. fig. 202, ascus with sporidia, x 325. Sporidia 
biseriate, linear-acuminate, uniseptate, colourless, 0:0010 to 0-0012 inch long. Messrs. 
Berkeley and Broome state (Ann. Nat. Hist. s. 2. vol. vii. p. 187) that this species 
is identical with S. arundinacea, Sow.; but in the Kew specimens the species differ 
in the colour of the sporidia, and in the number of the septa. "This, however, would 
not prove them distinct, as the number of septa varies in many species, and sporidia 
which are colourless when young frequently become brown in age. 

203. SPHÆRIA CLARA, n. s. Tas. XLIX. fig. 203, ascus with sporidia, x 325. Sporidia 
biseriate, colourless, subfusiform, 4-nucleate, sometimes also with some smaller 
nuclei ; slightly constricted in the middle, 0*0010 to 0-0012 inch long. On some reed. 
This fruit is described from a specimen which I had in my own herbarium, but 
which I have mislaid. "The species is not in the Hookerian herbarium. 

204. S. PARDALOTA, Mont. This plant produced only colourless, rather narrowly ellipti- 
cal stylospores 0:0002 to 0:0004 inch long. The true sporidia are described as cym- 
biform and biseptate. See Mont. Syll. p. 235. 

205. SPHÆRIA ? PANTHERINA, Berk. The specimen of this plant produced no asci, but 
fusiform stylospores 0:0008 to. 0:001 inch long, and extremely narrow. 

206. SPHÆRIA LONGISSIMA, Pers. This species forms long dark spots studded with the 
perithecia. 'The specimens at Kew are imperfect; buf in two of the perithecia I 
observed elliptical or slightly curved bodies with two nuclei, and sometimes with a 
septum in the middle, greenish or yellowish in colour, 0:0004 to 0*0006 inch long 
and about half as wide; but whether true fruit or not, I cannot say. I have seen no 
other specimens. 


Div. 16. CoNrERTE. 


207. SPHÆRIA BIFRONS, Schm. and Kunze; Fr. S. M. ii. p. 438. Tas. XLIX. fig. 204, 
ascus with sporidia, and free sporidia, x 325. Sporidia biseriate, colourless, curved, 
0:0005 inch long. toe 

207 a. SPHÆRIA GRAMINIS, Pers.; Fr. 8: M. ii. p. 434. Tas. XLIX. fig. 211, asci with 
sporidia, and free sporidia, x 450: Sporidia colourless, broadly elliptical, frequently 
with a single large globose nucleus. 

208. 8. Birte P) ROR, Berk. Engl. Flora, vol. v., “ Fungi,” p.257. Tas, XLIX. 
fig. 205, stylospores, x 325. Stylospores greenish, margined; contents granular; 
0:0012 inch long. 


209. S. GIGANTEA, Mont. Syll. 230. Tas. XLIX. figs. 206 and 207, ascus with sporidia, 
2rP2 


286 MR. F. CURREY ON THE FRUCTIFICATION OF COMPOUND SPHARIA. 


and free sporidia, x 325. Sporidia usually biseriate, but sometimes uniseriate, dark 
brown, multiseptate (always, I think, 7-septate), and also divided longitudinally in 
each division except the two end ones; curved, cymbiform, surrounded by a gelati- 
nous envelope not perceptible in the ascus, at least in the Kew specimens; 0:0018 
inch long. On Agave. A large, black, subhemispherical mass + inch and more wide. 
The fruit is very striking. 

210. S. (HENDERSONIA ?) YUcoæ-GLORIOSÆ, Schwein. ; Fr. S. M. ii. p. 437. . TaB. XLIX. - 
fig. 208, stylospores, x 350. Stylospores colourless, linear, 0:0010 to 0:0018 inch 
long. Not a true Spheria, or, if so, only in a secondary state of fructification. 

211. SPHÆRIA RHYTISMOIDES, Bab. in Proc. Linn. Soc. vol. i. p. 32. Ta». XLIX. fig. 209, 
ascus with sporidia, X 325. Sporidia crowded, colourless, hyaline, obtuse, 0:0005 
inch long. 

212. S. (DrATRYPE?) INSULARIS, Berk. MSS. Ta». XLIX. fig. 210, ascus with sporidia, 
and free sporidia, x 325. Sporidia biseriate, colourless ; narrowly almond-shaped, or 
flat on one side ; 0:0005 inch long. Placed in the Herbarium with the Confertæ, but 
belonging, I think, to the Lignosæ. The black line is very distinct. On Aucuba 
Japonica. The endochrome is sometimes 1-, 2-, or 4-partite. It is, I think, very 
common on Aucuba Japonica, but has not, as far as I am aware, been described, 
unless, as Mr. Berkeley has suggested to me, it may be S. controversa, Desm. 

213. SPHÆRIA FIMBRIATA, Pers. Syn. p. 36; Fr. S. M. ii. p. 436. The sporidia are, I 
believe, colourless and curved, similar to those of Yalsa ambiens. Fig. 212 represents 
asci and sporidia, x 225; but the specimens are not quite ripe. 

214. SPHÆRIA CEUTHOCARPA, Fr. S. M. ii. p.439. Tas. XLIX. fig. 213, ascus with spori- - 
dia, and a free sporidium, x 425. Sporidia lying side by side, extending in length all 
along the ascus, colourless, divided by numerous septa, which are not always visi- 
ble except under a high power. 


INDEX. 
| [The figures refer to the numbers afixed to each Plant, and not to the pages.] | 

geo Pe arundinacea, 200. Calycanthi, 196. clara, 203. 
acervalis, 183. aspera, 79. capitata, 9. coccinea, 175. 
st atro-punctata, 66. capnodes, 69. coenopus, 10. 
equiliiearis, 114. atro-purpurea, 43. Carpini, 122. cohærens, 40. 
BAR, carpophila, 21. concamerata, 138. 
ment 147. Badhami, 76. ceratosperma, 94. concentrica, 32. 
angulata, 126. Berberidis, 178. ceuthocarpa, 214. conglobata, 186. 
annulata, 31. biconica, 151. chrysentera, 198 a. convergens, 161. 
pare E bifrons, 207. cincta, 139. corniformis, 19. 
ms diu botryosa, 45. cinnabarina, 176. coronata, 145. 
aui 25 bullata, 81. circumscripta, 136. Cratægi, 143. 

N Buxi, 187. citrina, 50. crenulata, 14. 


MR. F. CURREY ON THE FRUCTIFICATION OF COMPOUND SPHÆRIZÆ. 


cucurbitula, 179. 
cupularis, 172. 


decipiens, 197. 
decolorans, 177. 
denigrans, 74. 
depressa, 100. 
detrusa, 120. 
deusta, 56. 
digitata, 15. 
Diospyri, 104. 
disciformis, 61. 
discincola, 106. 
discutiens, 115. 
dissepta, 135. 
Dothidea, 191. 
dryina, 144. 
dryophila, 71. 


durissima, 29. 


elevata, 110. 
elongata. See 190. 
enteroleuca, 117. 
enteroxantha, 142. 
entomorrhiza, 5. 
examinans, 174. 
extensa, 119. 


faginea, 171. 
farinosa, 37. 
favacea, 78. 
ferruginea, 95. 
fibrosa, 99. 
fimbriata, 213. 
fimeti, 103. 
flavo-virens, 62. _ 
foraminula, 131. 


fragiformis, 30. å 


fulvo-pruinata, 129. 
furfuracea, 160. 
fusca, 38. 
fuscospora, 57. 


gastrina. See 97. 
gelatinosa, 32. 
gigantea, 209. 
Godini, 202. 
graminis, 207 a. 


Guianensis, 8. 
Gunnii, 1. 


heliseus, 12. 
hyalina, 48. 
hypodermia, 162. 
hypoxylon, 16. 
hystrix, 93. 


ianthino-velutina, 20. 
inæqualis, 75. 
Innesii, 169. 
insidens, 194. 
insularis, 212. 
intexta, 158. 
involuta, 7. 
irregularis, 97. 


juglandicola, 125. 
Juglandis, 154. 
Junci, 199. 


Kunzei, 137. 


Laburni, 180. 
lanciformis, 92. 
lata, 111. 
lateritia, 46. 
leioplaca, 102. 
leiphæmia, 146. 
lenta, 83. 
leprosa, 88. 
limæformis, 108. 
Liriodendri, 68. 
lobata, 51. . 
longissima, 206. 
lutea, 55. 
luteo-virens, 52. 


Malorum, 208. 
marginata, 69. 
megalospora, 193. 
melanaspis, 70. 
melasperma, 127. 
Meli, 166. — 
melogramma, 195. 


microcephala, 25. 


microceras, 17. 


microstoma, 134. 
militaris, 2 
milliaria, 107. 
monadelpha, 113. 
Mougeotii, 98. 
multiformis, 28. 
multiplex, 18. 
mutila, 192. 


nebulosa, 201. 
nigerrima, 96. 
nivea, 130. 
nucleata, 72. 


nummularia, 58, 
ophioglossoides, 6. 


parallela, 101. 
pantherina, 205. 
pardalota, 204. 
parmularia, 39. 
pedunculata, 4. 
perforata, 44. 
petiginosa, 87. 
pileiformis, 3. 
platystoma, 63. 
podoides, 85. 

„ var. levis, 86. 
polymorpha, 11. 
profusa, 132. 
Prunastri, 121. 
Pseudo-bombarda, 173. 
pulchella, 159. 
pulchra, 152., 
pulicaris, 181. 
punctata, 27. 
purpurea, 24. 
Purtoni, 182. 


rhytismoides, 211. 
Robertsii, 26. 
rubiginosa, 42, 
rubricosa, 41. 
rudis, 188. 

rufa, 36. 


saceulus, 141. 
sagræana, 23, 
salicina, 157. 
Sassafras, 49. 
seabrosa, 84. 
scoriaden, 188, 
serpens, 47. 
Sorbi, 140, 
Spartii, 190. 
spienlosa, 112. 
spinosa, 109. 
stellulata, 124. 
stigma, 80, 
stilbostoma, 149. 
stipata, 116. 
strumella, 89. 
subsolitaria, 198. 
suffusa, 153. 
syngenesia, 123. 


taleola, 128. 
tetraspora, 156. 
thelebola, 163. 
tubulina, 54. 
turgida, 148. 
typhina, 27. 


uda, 60. 
undulata, 82. 


varians, 73. 
velata, 105. 
verrucæformis, aes 
vestita, 165. 
virgultorum, 65. 
vogesiaca, 64. 


xanthostroma, 164. 


Yuecæ-gloriosæ, 210. 


287 


GEN. SPHÆRIA. 
(Figs 1 to 45) 


GE | | ER 
FCusrey del Tuffen West se. | : 
| All the figures magnified 325 Diamevers,enogt where otherwise stated. 


Trans Linn. Soo. Vo XXI PL ZLVI 


GEN. SPHÆRIA. 
(Fig! 46m 87) 


E Currey del Tuffen Wast se. — RE 5 Lic. KI cu ce Hu T m 
HE er AU the Figures magnfeå 325 Diar ters, coven where otherwise stated. Nu uiu. | 


Fur: 


Trans. Linn. Soc. Vol HIT PLALVE 


D 
Ye 


Cr ee Q g FIN 
LOE EL NGC 7 


| 


| | AU the figures magnified: 25 Diamaters, omegn where otherwise Stated, 0 


T LJ YY DI PT FT 
Trans Lun Soc. Vol ANT PLALVZL 


GEN. SPHÆRIA 
[Fig? 181 to 174) 


F Currey del. Tuffn West se. 


E V TOU B m ias 
| Trans Linn. Se LIT PIL 


GEN. SPHÆRIA. 
(Fig? 172 w 28) x i 


X. 


I 


THE 


TRANSACTIONS 


OF 


THE LINNEAN SOCIETY 


OF 


LONDON. 


VOLUME XXI. 
PART THE FOURTH. 


LONDON:. 
PRINTED BY TAYLOR AND FRANCIS, RED LION COURT, FLEET STREET: 


SOLD AT THE SOCIETY'S APARTMENTS, BURLINGTON-HOUSE ; 
AND BY LONGMAN, GREEN, LONGMANS, AND ROBERTS, PATERNOSTER-ROW. 


M.DCCC.LIX. 


[ 289 ] 


XXI. Observations on the Structure of the Stem im certain Si the Natural Orders 
Caryophylleæ and Plumbagineæ. By DANIEL OLIVEX, Jun., F.L.S. 


- Read December 10th, 1858. 


CARYOPHYLLEÆ. 


THE internal structure of the stem in certain suffruticose species of the genus Acantho- 
phyllum presents some interesting anomalies which appear to me to be undescribed. I 
venture, therefore, to communicate to the Linnean Society this brief memorandum, as a 
further contribution to an already accumulated store of material in our possession relating 
to what we are accustomed to regard as abnormal modes of arrangement in the elements of 
Dicotyledonous wood, furnished to us by species often widely removed in the order of their 
natural affinities. The time may not be distant when a careful correlation of all our facts 
may be seasonable, and likely to render good service, not only to systematie botany, but 
also in enabling us to attain to a more comprehensive perception of vegetable structure 
generally. At the present time, how isolated, as well as fragmentary, are the items of 
our knowledge upon this subject ! 

The points of peculiarity in the species of Acanthophyllum to which I wish to direct 
attention at this time are—1st, the general relative arrangement of the vascular and cel- 
lular tissues or ‘systems’ of the stem; 2nd, the histological character of some of these 

tissues ; and 3rd, the occurrence in remarkable abundance, in some species, of frequently 
| large concretions of oxalate of lime in the parenchyma of the stem. To the difficult and 
time-absorbing study of the relations subsisting between the vascular bundles and the 
leaves, scapes, adventitious roots, or other axial appendages, I cannot at present address 
myself. Their connexion, so obscure and imperfectly understood even in succulent plants 
with lengthened internodes, is much complicated by a dense habit and dividing woody 
stem, apart from the difficulties attending the examination of specimens which have been 
dried many years. In regard, however, to this point, I may say that I am quite unable 
to attribute the differences in internal structure, presented by the different species, to a 
varied disposal of lateral organs; the mode of arrangement of the leaves is, upon the 
whole, very uniform in the plants which I have had through my hands. 

In this notice, I speak of the perennial, leaf-hearing, woody, and frequently branching 
axis of the genera Acanthophyllum, Arenaria, and Dianthus, as the stem: in the present 
terminology of these parts, there is, by the way, I think, somewhat ar a vagueness and 
laxity, which, without multiplying technicalities, or further burdening our glossaries, 
might be removed. : 

With regard to the arrangement of the tissues of the stem :—In some of the species of 
Acanthophyllum, as also in Dianthus and Arenaria, a manifest, though often very wise 
trie pith occurs ; in the first-named genus, however, often deranged, as are also the woody 

27 2Q 

VOL. XXII. # 


290 MR. D. OLIVER, JUN. ON THE STRUCTURE OF THE STEM 


bundles to a great extent, by radial processes of cellular tissue, which may, in some cases, 
be regarded as projections from the medulla, probably in connexion with lateral organs; 
in others, as induplications of the cortical parenchyma. In Acanthophyllum spinosum it 
is often impossible, from the excessively dislocated character of the vascular bundles, to 
determine satisfactorily a true pith. In this species, the entire stem, almost to the young 
and leafy annual shoots, is partitioned into numerous vascular masses by tortuous in- 
truded plates of cellular tissue, presenting at first sight, in a transverse section, a striking 
similarity to the irregular structure of some species of Combretaceæ, Malpighiaceæ, or 
Bauhimie. The sole relation of the parts in these stems to a common centre is indicated 
more or less distinctly by some of the peripheral vascular bundles, which are laterally 
bounded and separated by lines of cellular tissue exhibiting a radial disposition. If we 
trace the structure by transverse sections from the young and yet green shoots, internode 
by internode downwards, we may observe the rapid accession of an irregular relative 
arrangement of the parenchyma and vascular tissues of the stem. Some of these changes 
are sketched in their consecutive order under fig. 3 (PL L.) ; figs. 4, 5, 6 and 7 represent 
also this curious and complicated structure. In the very young internodes the pith is 
found to be much elongated transversely, extending nearly or quite through the vascular 
zone, as shown by fig. 2. When dividing the wood entirely, as in fig. 4, the narrow ring 
of small and delicate cells, which in fig. 2 is represented as a normal * cambium" zone 
: surrounding the wood, penetrates the fissure and encloses separately the divided portions. 
Whatever may be the first determining cause of a deviation from the usual structure in 
this species, it is, I consider, obviously to the penetration of this belt of young and active 
cells that the subsequent and anomalous arrangement of the wood is due. These cells 
are doubtless capable of continuing the process of division and growth as under the 
ordinary conditions of the * cambium" layer, though probably to a limited extent. It 
may be observed, that in the very young shoots the direction of the transverse elongation 
of the pith may be found to alter materially from one internode to another. (See refer- 
ence to fig. 3.) . | 
The surrounding, intropenetrating *cambium zone’ is more or less apparent, not 
only in the earlier shoots, but also in portions of the stem which exhibit a highly dis- 
torted vascular system. In Acanthophyllum (Griffiths Affghan Coll. no. 1562) the 
dislocation of the wood is by no means so marked as in 4. spinosum, yet in this plant 
broad wedges of parenchyma are found to divide the vascular tissues into most unsymme- 
trical sectors, and sometimes extend quite across them. In À. laxiflorum these invasions 
are accompanied even to the apparent pith by cells containing the highly-coloured contents 
characteristic of the outer cortical layers. How far these wedges of cellular tissue may 
gen replace the medullary rays of Exogens generally, it would be interesting 
Fue. Singularly enough, although it was probably in Acanthophyllum (Gr. no.1562) 
that an irregular structure first caught my attention, I did not note or appreciate the 


IN CERTAIN SPECIES OF CARYOPHYLLEÆ AND PLUMBAGINE E. 291 


phyllum, in Dianthus (D. hispanicus) and Arenaria (A. laricifolia), as also in Armeria in 
Plumbagmee. Schacht* records their absence in the ‘rhizome’ of Viola (F. mirabilis 
‘and odorata), and A. Brongniart in certain Orassulaceæt. It is probable that a wood 
destitute of these processes may prevail through other natural orders in which a similar 
perennial, depressed axis is found. 

In very young internodes of Acanthophyllum spinosum, and in the stem of Acantho- 
phyllum (Gr. no. 1562), more or less regular concentric zones are apparent in the woody 
mass. These vary in number and width. I am unable to satisfy myself entirely as to 
the occasion of these alternating zones. They may, in these species, result, as might be 
expected, from the annual alternation of seasons; but upon this head further evidence is 
required. In the younger branchlets of A. spinosum they are often distinctly perceptible ; 
in the older portions, from the advanced distortion of the tissues, they are nearly or 
. quite lost. 

The change, from whatever cause it may result, in the character of the vascular 
bundles which gives rise to these concentrie rings, is of much interest, and I think im- 
portant to the phytotomist. I feel tolerably satisfied that in Acanthophyllum (Gr.no.1562) 
and A. spinosum, the vascular mass, which consists in great measure of very numerous 
‘ slit-marked’{ vessels of various calibre, traversing a prosenchymatous tissue, is inter- 
rupted by the formation of narrow annular belts of spiral vessels of small diameter 
repeated at definite (?) intervals, and that, in the young shoots of A. spinosum at least, the 
concentric zoning is due to these. In Acanthophyllum (Gr. no. 1562) numerous cords 
of a thick-walled prosenchyma occur along with the vessels, and these probably assist 
in determining the annular formations: in A. spinosum I have not observed any of the 
tissue to become thus thickened. This recurring deposition of spiral vessels is, I be- 
lieve, a repetition periodically of that ring in which they occur, and which is so generally 
recognizable, in Dicotyledons, immediately, and only, around the pith, and which, from 
the peculiar character which the presence of these vessels confers, is usually distin- 
guished by the special term ‘medullary sheath.’ It remains yet to be ascertained, and I 
have not leisure to follow up the investigation myself except to the neglect of other 
studies, whether an annular formation of spirals thus repeated is common to all species 
destitute of medullary rays. The inquiry is an important one, and pilen arr Bon 
logical value. In the older stems of A. spinosum, in which no concentrie disposition of 
the elements of the vascular masses is apparent, I should observe that apaapitels seem 
to be almost indiscriminately scattered amongst the wider and * slit-marked vessels. In 
the other species of Acanthophyllum examined, and in Arenaria laricifolia, lam not sure 
that true spirals reeur in the wood, in the manner described; nor am I quite clear that 
they do so in Dianthus. 

In the case of Acanthophyllum (Gr. no. 1562), allusion has been made to the presence 
of cords of much-thickened prosenchymatous cells, which traverse, with the vessels, the 
thinner tissues of the woody masses. In this plant these thickened cords are, I think, 


* «Die Pflanzenzelle,’ p. 280. f Arch. Mus. i. 437 (Lindl. V. K. p. 344). 
. $ I use the term ‘slit-marked’ as best expressing the form presented by their pits, which are transversely more or 


less lengthened. 202 


292 MR. D. OLIVER, JUN. ON THE STRUCTURE OF THE STEM 


sometimes considerably larger than I have represented in fig. 13. This remarkable 
thickening of the prosenchyma attains a maximum, however, in Acanthophyllum (Gr. 
no. 1570), in which its extensive development is the conspicuous feature of the wood, 
Here it is not disposed in cords or isolated masses, but is nearly continuous through each 
annular zone, although with many sinuosities in its cross section, traversed by the vessels, 
which are not very numerous, and chiefly confined to the inner portion of the rings, here 
and there à few of them being radially approximated. The thinner tissue which separates 
these annular zones is extremely narrow, and with difficulty resolvable from its ready 
rupture in cutting the adjacent cells, the cavities of which are nearly obliterated by 
secondary hardened deposits. In A. laxiflorum the arrangement of the thick-walled 
tissue is perhaps yet more curious. It is here remarkably dense, and is disposed, not in 
continuous belts, but in large cords, which are singularly lobed and divided in their hori- 
zontal section. The tissue filling up their winding interstices is almost entirely made up 
of ‘slit-marked’ vessels often of conspicuous diameter. The vertical continuity of these 
vessels through so dense a prosenchyma is interesting and readily observable in a thin 
longitudinal slice, the transverse septa of the cells of which they were originally consti- 
tuted frequently remaining quite distinct. The irregular form of these masses does not 
interfere with the regular arrangement of the annular zones within which they are 
confined. | | 

In respect to this thick-walled tissue, it will be apparent, from what has been stated, 
that Acanthophyllum (Gr. no. 1562), 4. laxiflorum, and Acanthophyllum (Gr. no. 1570), 
exhibit their development in a sort of graduated series, while in 4. spinoswm they are, so 
far as I have observed, totally absent. The singular prevalence in some of the species of 
Acanthophylium of an unusually large deposition of crystals of oxalate of lime is an addi- 
tional item of interest worthy of remark. I am indebted to my kind friend Daniel Han- 
bury for the accurate determination of these crystalline concretions from, I think, Acan- 
thophyllum (Gr. no. 1562). They oceur abundantly scattered through the parenchyma 
of the stem, both in the inner cortical layers and its inversions, the irregular cellular 
plates which radially traverse the wood, and also in the pith. | | 

In Acanthophyllum spinosum they are small, and do not probably much inerease be- 
yond the boundary of the cell in which they originate. They are very numerous, accom- 
panying the parenchyma in its twistings through the stem-structure,—in the vertically 
elongated cellular tissue being often also lengthened, or rod-like. In Acanthophyllum 
(Gr. no. 1570) they are comparatively few, but chiefly very large, without doubt obli- 
terating several cells in their increase. The concretions of Acanthophyllum (1562) are 
very variable in size and extremely numerous,—the parenchyma, laid bare by removal of 
the outer cortical layers, being quite gritty from their presence. 


PLUMBAGINEA. 


The stems of Armeria maritima, Acantholimon diapensioides, and, I think also, Statice 
arborea, present a wood destitute of medullary rays. Parenchymatous processes which 
may be accompanied by a few vessels are found very irregularly traversing the vascular 
bundles radially, from three or four of them to a considerable number sometimes occur- 


IN CERTAIN SPECIES OF CARYOPHYLLEÆ AND PLUMBAGINEÆ. 293 


ring in one horizontal plane. These processes, which I consider to be in connexion often, 
if not at all times, with the lateral appendages of the stem, are frequently in Armeria 
maritima of considerable width, and distort the generally more or less central and readily 
recognizable pith materially. In the laxer parenchyma of the medulla of this plant indi- 
vidual cells of very irregular form are found. In Statice arborea, the pith, the cells of 
which are marked by transversely lengthened pits, is vertically traversed by cords of a 
rather thick-walled elongated tissue. In Acantholimon diapensioides the rather compact 
woody mass is singularly lobed in some older stems, as represented in fig. 61, cortical 
inversions penetrating almost to the centre; in Acantholimon (sp. À. tomentello affinis, 
Gr. no. 1589) and Acantholimon (no. 1579), also, the wood is more or less dislocated by 
parenchymatous radial plates or cords ; in the latter species these processes contain many 
much-thickened apparently * sclerogen’ cells. The older wood in various species which I 
have examined presents more or less of a tolerably thick-walled prosenchyma. In Acan- 
tholimon (No. 1589) this tissue occurs in irregular dense masses, which exhibit a manifest 
disposition in concentric belts, as also a decided radial arrangement, as represented by 
fig. 24 (PL LI). The tissue intervening between the conspicuous cords abounds in vessels 
of considerable diameter, which, as in the other species examined, present * slit-marked * 
walls similar to those observed in Caryophylleæ, &e. 

Perhaps the most interesting point in the histology of the wood of these plants is the 
occurrence of minute, apparently intercellular cavities in the tissue traversed by the 
vessels in Statice arborea and Acantholimon diapensioides. In the latter species I have 
more minutely examined these. In this species the cells of the wood in which the small 
and rather sparingly distributed vessels are immersed, are elongated, presenting a reti- 
culated or spiral arrangement of their secondary deposits; it is between these cells, or 
between them and the vessels, that the very minute slit-like spaces are visible, in a suffi- 
ciently thin section of the wood, when examined with a magnifying power of 300 or 400 
diameters. On the nature of these interspaces I scarcely feel myself competent to offer 
a positive opinion, believing it possible that the eyes of a more experienced phytotomist 
might differently interpret it. I regard them as either very minute intercellular verba 
corresponding in some measure to those of Conifere; or as the much-widened blind ex- 
tremities of the pore-canals which traverse the thickening layers of the enclosing cells,— 
the primary cell-walls between each pair of opposing canals becoming absorbed. 


Note.—The specimens belonging to the genera Acanthophyllum (Caryophylleæ) and 
Acantholimon (Plumbagineæ) which I have examined, have been derived from Griffith 5 
extensive Affghan collections. The names of those which have been clearly determined 
I have obtained from the Hookerian Herbarium, in which some of the species have 
been examined and labelled by Boissier. ; 

To my friend Thos. Atthey, of Cramlington, Northumberland, I desire to acknowledge 
myself much indebted for a valuable series of sections illustrative of these structures, 


which he kindly prepared for the microscope at my request. 


294 ON THE STEM IN CERTAIN SPECIES OF CARYOPHYLLEÆ AND PLUMBAGINEZÆ. 


EXPLANATION OF THE PLATES. 


Tas. L. 


Acanthophyllum spinosum, C. A. M. 

Fig. 1. Portion of a young leafy shoot. | 

Fig. 2. Transverse section from a young internode: a. cortical parenchyma; b. zone of small and delicate 
cells (cambium region) ; c. wood which abounds in vessels of various calibre ; d. transversely 
lengthened pith-cavity, the cells partly obliterated. 

Fig. 3. a. From a young internode; 6. from the next below; c. from the third—the dividing fissure has 
changed to an opposite direction; d. traces of further lobing become apparent; e. about 4th of 
an inch lower; f. point from which a branchlet is given off. 

Fig. 4. Semi-diagrammatic section from a young shoot of the same, showing the division of the woody 
mass, and a ‘generative zone? surrounding each portion. 

Fig. 5. Transverse section from lower portion of twig, fig. 1. 

Figs. 6 & 7. Sections from older stems. 


Fig. 8. Semi-diagrammatic and enlarged ; the vessels often cut the horizontal plane obliquely. 


Fig. 9. Vertical section showing the vessels obliquely traversing the thin-walled tissue forming the mass 
of the older stems. 


Fig. 10. Diagram from one of the upper branchlets of the twig, fig. 1. In the wood, which is very full 
of vessels—more so, I think, than I have represented—four nearly equal zones are to be counted; 
through these the vessels are irregularly distributed. "The zones are separated by lines of very 
narrow vessels, which, according to my observations, are true spirals. 


Acanthophyllum (sp. dub., Griffiths Affghan Coll. no. 1562). 
Fig. 11. Thin sections, successively removed from an older stem. 
Fig. 12. wu section enlarged : a. cortical parenchyma, with numerous concretions of oxalate of 
e. i 
Fig. 13. Portion more highly magnified, with cords of thickened prosenchyma. 


Figs. 14 & 15. Vertical sections showing * slit-marked” and spiral vessels with prosenchyma. _ 
Fig. 16. Pith, with oxalate of lime. 


Fig. 17. Tangential section dividing a radial parenchymatous cord. 


Ta». LI. 


; Acanthophyllum laxiflorum, Boiss. " 
Fig. 18. Transverse and, fig. 19, vertical sections of the stem. i 


Acanthophyllum (sp. dub., Griffith’s Affghan Coll. no. 1570). 
Fig. 20. Transverse section. Fig. 21. Same more highly magnified. 


Acantholimon diapensioides, Boiss. 
Fig. 22. Transverse section. Fig. 23. Same enlarged. ; 


Acantholimon (sp. A. tomentello aff.). 
vertical sections of portions of the stem. 


Armeria maritima, Willd. 


Fig. 24. Transverse and, fig. 25, 


Fig. 26. Transverse section from lower portion of the stem (passing into the root?). Fig.27. From about jth 
of an inch higher up. Fig. 


j : 28. Section showing numerous radial processes in the same plane. 
1g. 29. Magnified transverse section showing the distribution of the vessels and cords of thick-walled 
prosenchyma. " | 


Fig. 30. Tangential section dividing-a radial cord. 
Fig. 31. Isolated cells from the pith. — 


Ao Val 


UU. . 


Trans L 


‘tab. 50 294. 


ef 


D.Oliver del Tuffen West se. 


294 


Ap 


Soc. Vol KUM tab. 


um. 


2 Me 


he tn 


1 


[ 295 ] 


XXII. On the genus Henriquezia of Spruce. By GEORGE BeNTHAM, Esq., V.P.L.S. 


Read February 3rd, 1859. 


THis genus is one of the many interesting new ones discovered by Mr. Spruce in 
Northern Brazil and Venezuela. Some years since, I published the first species received 
from him, in the 6th vol. of Sir W. Hooker’s Kew Journal of Botany; but at that time 
the fruit was not known. Since then Mr. Spruce has transmitted fine flowering specimens 
of a second species, and fruits of two more, which, from their foliage, appear to be distinct 
from the flowering ones. I am thus enabled to complete the generic character, and to lay 
before the Society the accompanying illustrations from the artistic and accurate pencil of 
Mr. Fitch. 

It will be observed that these trees have stipules, an inferior ovary, and five perfect, 
nearly equal stamens; and yet they undoubtedly belong to the family of Bignoniacee. 
The affinity which I formerly alluded to, with Platycarpum of Humboldt and Bonpland, 
is fully confirmed by these specimens, as well as by one or two fruiting specimens of the 
true Platycarpum gathered on the Orenoco by Mr. Spruce, which show that that genus 
also has stipules, although, as in the first-received specimens of Henriquezia, they had 
not been observed, being very deciduous. In Platycarpum the ovary is figured as supe- 
rior. I have not seen the flowers; but on the fruit there is an external scar at the base, 
which shows that there had been a partial adherence to the calyx. In Henriquezia the 
ovary at the time of flowering is entirely below the adherent part of the calyx. After 
fecundation it gradually emerges, and the ripe fruit shows about its centre a transverse 
convex line or scar left by the free part of the calyx on falling off. The fruit is nearly 
flat and hard as in Platycarpum, but much larger and more woody ; and instead of open- 
ing out vertically in four valves which remain attached by the central line formed by the 
axial entire dissepiment, it opens horizontally to a little below the middle in two entire 
valves, down the centre of each of which, in the inside, is a raised line, being the remains 
of the dissepiment. In each cell are four large flat seeds, remarkable, at least in Mr. 
Spruce's specimens, for their tendeney to germinate whilst the capsule is yet Ravi i" 
the tree,—the radicle lengthening out, twisting round the seeds, and forming un : 
network, the origin of which was very puzzling till I had carefully soaked the an 


traced it out. : po 
The genus, as I stated in the above-mentioned article, was dedicated by Mr. Spruce to 


Senhor Henriquez Antonij, a native of Leghorn, but for more than thirty years settled at 


the Barra do Rio Negro, where he has constantly rendered every csi rt 
and other travellers during that period. The species are all described as t gr 


L Th 2 4 h 
beauty, , and some of them of considerable size. The following are the technical cha- 
racters :— : 


296 MR. BENTHAM ON THE GENUS 


HENRIQUEZIA, Spruce. Benth. in Hook. Kew Journ. Bot. vol. vi. p. 338. 
(Genus e familia Bignoniacearum.) 


Calyæ basi turbinatus ovario adhærens, limbo supero, 4-fido v. 4-partito, circumscisse deciduo. Corolla 
oblique infundibularis, fauce ampla, limbo 5-lobo, laciniis subæqualibus, æstivatione bilabiata, 
Stamina 5, corolla breviora, 2 inferiora cæteris paullo breviora ; antheræ subæquales, lineari-oblongæ, 
omnes fertiles. Ovarium per anthesin inferum, disco carnoso breviter cupulato v. annulato coro- 
natum, biloculare, post anthesin semisuperum. Ovula in quoque loculo 4, axi collateraliter affixa, 
horizontalia v. subadscendentia. Stylus filiformis, apice bilobus, lobis seu lamellis lineari-oblongis, 
complanatis, intus stigmatosis, mox revolutis. Legumen compressum, lignosum, bivalve, valvulis 
integris, dissepimento lineari valvis contrario, maturitate diviso. Semina in quoque loculo 4, magna, 
plano-compressa, hilo lineari affixa. Testa tenuis, laxiuscula. Cotyledones subearnosæ, basi profunde 
bilobæ. | ES 

Árbores Brasiliæ borealis et Venezuela. Folia 3—5-natim verticillata, coriacea, integerrima, pennivenia. 
Stipule lineares v. lanceolatæ, rigidæ,caducæ. Flores speciosi, in paniculas densas terminales dispositi. 


l. HENRIQUEZIA VERTICILLATA, Spruce, J. ¢.; foliis subquinis anguste oblongis sub- 
acuminatis basi acutis, venis primariis valde obliquis subtus prominentibus, calycis 
4-fidi laciniis lanceolatis, corolla extus cano-sericea. 


Arbor pulcherrima, 50 ad 100 pedes alta, trunco 4 pedes crasso, ramis summis fastigiatis subquinatim 
verticillatis, ramulis inflorescentiaque ferrugineo-tomentellis mox glabratis. Folia pleraque 5-natim 
verticillata, petiolata, in speciminibus suppetentibus 8-10 poll. longa, 2-21 poll. lata, inferiora vero- 
similiter majora, obtusiuscula v. breviter et acute acuminata, integerrima, basi cuneato-acuta, glaber- 
rima, coriacea, supra nitidula, subtus costa media venisque primariis obliquis valde elevatis percursa, 
rete venularum vix conspicuo. Petiolus 4-1-pollicaris, basi dorso sub insertione cicatrice stipularum 
deciduarum signatus. Stipulas ipsas non vidi. Panicula terminalis, intra folia summa subsessilis, 
late thyrsoidea, ferrugineo-tomentella, ramis crassis subcompressis verticillatis et dichotome cymi- | 
feris. Calyx incurvus, crassus, circa 6 lin. longus, extus ferrugineo-tomentosus, intus sericeus, limbus 
ad medium divisus in lacinias 4 lato-lanceolatas acutas æstivatione vix imbricatas, quarum infima — 
parum latior. Corolla sesquipollicaris, extus tomento sericeo appresso in sicco canescens, in vivo — 
rosea, intus alba, glabra, exceptis laciniis puberulis et serie pilorum flavicantium secus lineam | 
mediam labii inferioris, necnon pilis nonnullis ad basin faucis. Stamina glabra, infra medium tubi — 
inserta. Anthere longæ, medifixæ, biloculares, longitudinaliter dehiscentes. Stylus basi glaber, — 
apice in lamellas oblongas birto-stigmatosas divisu 
Capsula non visa. 


One of the finest trees met with by Mr. Spruce in his voyage up the Rio Negro in 
December 1851. It was frequent in the Gap6, from above Barraroá to San Gabriel do 


s. Discus epigynus brevis, crassiusculus, pubescens. — 


Cachoeiras. 


2. HENRIQUEZIA OBOVATA, Spruce, MS.; foliis quaternis obovato-oblongis obtusis retu- 


sisve basi rotundatis cordatisve, venis primariis subtransversis subtus valde promi- 
nentibus, capsula transverse oblonga. 


Arbor 40-60-pedalis. Stipule lanceolatæ, juniores 4-5 lin. longæ, a foliis adultis jam deciduæ. Folia ; 
(sec. Spruce) constanter quaternatim verticillata, 4-5 poll. longa, 21-3 poll. lata, venis primaris | 


a costa multo magis divergentibus quam in H, vertici E 1 long 
. icillata. Flores ignoti. Capsula 2 poll. longi 
4 poll. lata. Semina 13 poll. lata. oe Fr 


. 


HENRIQUEZIA OF SPRUCE. — | Eo 297 


Å single specimen was gathered by Mr. Spruce, in June 1854, in woods along the 
Guiainia, or Upper Rio Negro, above the confluence of the Casiquiare. 


3. HENRIQUEZIA NITIDA, Spruce, M$.; foliis longe oblongis obtusis crasso-coriaceis 
nitidis, venis vix prominulis, calycis 4-partiti laciniis subovatis acutis imbricantibus, 
corolla extus sericeo-villosa. i 
Arbor 30-pedalis, ramosissima. Folia (an constanter ?) ternatim verticillata, 8-10 poll. longa, 2-3 poll. 
lata, basi angustata, utrinque viridia et nitida, costa utrinque prominula ; venæ tamen primariae parum 
conspicuæ et non prominentes.  Sfipulæ lineari-lanceolatæ, pollicares, rigidæ, persistentes. Paniculæ 
rami ternatim. verticillati, dichotome cymosi, cano-tomentosi. Calycis limbus usque ad ovarium 
divisus; lacinie 5 lin. longæ, tomentosæ, multo latiores quam in H. verticillata, marginibus imbri- 
cantibus. Corolla sesquipollicaris, rosea maculis nonnullis cinnabarinis notata, intus prater lineam _ 
pilorum flavicantium glabra, extus mollissime sericeo-villosa, fauce latiore quam in H. verticillata. 
Fructus ignotus. ` ; 


Collected by Mr. Spruce in October 1854, on the banks of the Casiquiare, and distri: 
buted under the number 3690. | | 


4. HENRIQUEZIA OBLONGA, Spruce, MS.; foliis anguste oblongis obtusis subcoriaceis 
opacis subtus pallidis, venis vix prominulis, capsula suborbiculari. 

Possibly a variety of the H. nitida; but the specimens of the one being in flower only, 
of the other only in fruit, I do not think it safe to unite them, as the leaves do not quite 
agree. In the H. oblonga they are scarcely 6 inches long, although with longer petioles 
than in H. nitida, not so thick, and not nearly so shining, and much paler underneath. 
The stipules are nearly the same. The capsule is of the same woody consistence as in 
H. obovata, but not broader than long, being about 3 inches each way; it is marked with 
a similar slightly-arched transverse line, being the scar left by the limb of the calyx. 

. À small tree of about 18 feet in height, with few spreading branches; frequent on the 
-Rio Atabapo, an affluent òf the Orenoco near the mouth of the Casiquiare. There were 
only two or three specimens gathered by Mr. Spruce in June 1854, and numbered 3702. 


V EXPLANATION OF THE PLATES. - 
TAB. LII. | 


l ' Henriquezia verticillata. 
Fig. 1. -Calyx and style. 7 
Fig. 2. Corolla cut open. d 
Fig. 3. Ovary, transverse section. 
Fig. 4. Ovary, vertical section. 
Fig. 5. The same shortly after flowering. . ; oR 
C VOL. XXII, 


298 


Fig. 
Fig. 
Fig. 
Fig. 
Fig. 
Fig. 
Fig, 


- Fig. 


WAAR we 


. Å seed. En 
. The same seen edgewise. 

. Embryo with the growing root. - 
- The same with one cotyledon removed, showing the growing plumula, 


. Calyx and style, — | 


MR. BENTHAM ON THE GENUS HENRIQUEZIA OF SPRUCE. 


Tam. LIII. 
Henriquezia obovata. 


Ripe fruit seen from one side. 


"The same, end view. 


'The same with one valve removed, showing the seeds entwined by the young roots. 


LE 


Tas. LIV. 
Henriquezia nitida. 


Corolla cut open. 


. Calyx and base of the style after the calyx-lobes have fallen off. 


Ovary, transverse section. 


: Ovary, vertical section, ` 


Trans, Linn So Vol II rad 58, ju 298. 


WE Fam, del. 


296. 


“x 


59 


vs 


Vol JAIL bab. 


200. V 


c. 


Brans Lin. 


^, del. 


WI, 


LIU tab 55. 298. 


å 


29] 


XXIII Remarks on Gnetura. By the late WILLIAM bs ELS., Madras. 
. Medical Service. Communicated by A. HENFREY, Esq., F.R.S., F.L.S., Professor of 
_ Botany, King’s College." iy = | è 


Read April 21st, 1859. 


: PREFATORY Nore BY Pror. HENFREY. | 
THE following paper is the original from which were derived the particulars communi- 


cated by Dr. Lindley, in the article Gnetacee, in his * Vegetable Kingdom ;’ and it is now 


brought forward under the following circumstances. 

My curiosity was excited by Mr. Griffith’s account, quoted by Dr. Lindley, of the exist- 
ence of a long convoluted suspensor in the ovule of Gnetum. It appeared to me that this 
indicated an additional affinity between the Guetaceæ and the Conifere and Cycadaceæ, 
and that this would be still more striking if it were accompanied by the phenomena of 
polyembryony, such as are met with in the undoubted Gymnosperms. On examination of 
some specimens, I found that the lower end of the long convolutéd suspensor does divide 
into à number of distinct processes, as in Conifers, and that the embryo is developed at 
the end of one of these. I now became anxious to examine some flowers in an early stage 
of development with a view to ascertain if Gnetum produced corpuscula ; which I thought 
might have been overlooked by Mr. Griffith, his Memoir having been written before the 
publication of Mr. Brown's celebrated Memoir on the Plurality of Embryos in the 
Conifere. rd MEET Fr SER us 

Through the kindness of Dr. Hooker, I obtained from the Kew Museum a supply of 
specimens of Gnetum in various stages of growth. From the same friend I learnt that 


the original Memoir of Mr. Griffith was in the hands of the Secretary of the Society, v +: 
with the consent of Dr. Lindley, Mr. Bennett placed the paper in my hands. The study 


of this paper, under the light of my own observations, has led me to attach great im- 
portance to it, and I have recommended its publication before communicating the results 
_ of my investigations, on account of the Aa 
| of the facts important in the history of developmen genus, and rendered it un 
cessary for ae do more than sp nett his observations in a few points, rx enter- 

‘ing upon the general conclusions I have drawn from his and my own studies. | 
I hope shortly to offer a paper on this subject for 


S La i; s S € j * 8 rste- 
. favourably upon the opinion expressed by Prof. J. der ec syke Is Loren 
matis Plantarum,' that the Gnetacee are related even more el | y o gata 
than to the Coniferæ. Å em M te á 


London, January 2, 1800. SET | og 


ving forestalled me in the greater part 
of this genus, and rendered it unne- 


the consideration of the Linnean 


+— Society, and shall merely say at present, that my investigations lead me to look very . 


de 


300 MR W. GRIFFITH ON THE GENUS GNETUM. 


Tuıs paper owes its existence to Dr. Lindley, who with his usual kindness pointed out to : : 
me this, among several other genera, as an interesting subject for study. In glancing over | 
the history of this genus, we find that very little additional information had been acquired — 
upon its structure from the period of its original publication by Linnæus, from whose . 
character all the subsequent ones have been more or less. derived, until the appearance of — 
Dr. Brown's: Memoir relative to the character and description of Kingia appended to 
King's Voyage*. Previous to the appearance of this important publication, all botanists | 
who had noticed this genus had ascribed to the female flower the ordinary structure, It — 
‚will hereafter be seen that all those who have formed this opinion of the structure of . 
Gnetum have examined the female flowers at a rather late period. Tr 
Dr. Brown, in the Memoir above referred to, p. 23, gives it as his opinion, that in 
Cycadee, Conifere, Ephedra, and Gnetum, the ovarium is either altogether wanting, or — 
so imperfectly formed that the ovulum itself becomes directly exposed to the action of the . : 
pollen. He further states.that “the similarity of the female flower in Cycadeæ and — 
Conifere to the ovulum of other phænogamous plants, as I have described it, is indeed 
sufficiently obvious to render the opinion here advanced not altogether improbable; but : 
the proof of its correctness must chiefly rest on a resemblance, in every essential point, - 
being established between the inner body in the supposed female flower in these tribes, 
and the nucleus of the ovulum in ordinary structures, not only in the early stage, but 
also in the whole series of changes consequent to fecundation. Now, as far as I have yet 
examined, there is nearly a complete agreement in all these respects.” 


After repeated examinations of Cycas, Gnetum, and Agathis, in their growing states, it — 3 


may perhaps not be considered presumptuous in me to add my feeble testimony to the — å 
extreme accuracy of this statement of Dr. Brown t. E 0 
Professor Lindley, in his valuable Introduction to the Natural System of Botany, — 


while he admits to the fullest extent Dr. Brown’s notions on Cycadeæ and Conifere, : 


appears to entertain the old opinion as to the structure of Gnetum; and here it is again — 
evident that this truly philosophical botanist has only examined somewhat advanced | 
female flowers. I find, however, that in his remarks on Garrya (Bot. Register, vol. vi. — 
new series, t. 1686), this author has adopted the opinion of Dr. Brown, and speaks of E 
Gnetaceæ, a naked-seeded order. — — d 
: Having thus briefly alluded to the opinions entertained on the structure of this 
singular genus, I shall now proceed to the consideration of the structure and develop- 
ment of the female flower, which consists of a single naked ovulum. B 
At a period long before the exsertion of the anthers, the ovula, which lie upon the male — 


flowers, are generally of an oblong form, and consist of a central cellular solid body, : ; 
enclosed in two envelopes. The outermost of these is fibro-cellular, and divided longitudi- s 
nally on the upper face, or that nearest the axis, the fissure extending nearly to the base — 


of the ovulum?. The inner or second envelope is cellular, and is divided irregularly — 


* ls Meno 
Of this | oir I have seen only an 8vo copy, in the possession of Dr. Wallich. 


T At the same time, Dr Brown's observati : : 
; ; . t addi 
membrane, * (See Kingia, Apes rvations refer only to a period subsequent to the appearance of the 


? This division is perhaps similar to that which Dr. Brown states to take place in Dacrydium. 


" 
4^ 
. 
NUES 


MR. W. GRIFFITH ON THE GENUS GNETUM.  . | 301 


towards its apex. This envelope does not at this period entirely enclose the nucleus: the 
points of some of the laciniæ or divisions project occasionally beyond the apex of the outer 
envelope. The nucleus is an oval or oblong cellular body, rounded off at its apex, which 
is composed of lax cellular tissue. The next change consists in the commencement 
of the obliteration of the longitudinal fissure existing along the posticous face of each 
outer envelope, and of an extension of the inner coat over the nucleus, the apex of which 
becomes more or less depressed,—the centre of the depression, however, projecting in the 
form of a cone of a very slight elevation. At the time of flowering, or of the exsertion 
and dehiscence of the anthers, the fissure originally existing along the upper face of the 
outer coat has disappeared, with the exception of a small portion at the apex of the ovulum 
which remains unclosed throughout : the ovula are at this period in some species oblique. 
The inner envelope is generally entirely enclosed within the outer ; the points of its lacinise 
reach, however, to the opening existing in the apex of this latter, and occasionally, but 
by no means universally, project beyond it to a short distance. This coat has undergone 
scarcely any change, and corresponds in shape to the cavity of the outer envelope. The 
nucleus is completely covered by both integuments, and its apex, which continues of the 
same form, is occasionally tinged with brown. Within its substance, which is entirely 
cellular, and towards its centre, there exists a small cavity, lined with a membranous sac, 
attached apparently to the apex of the cavity, and containing a number of minute 
grumous-looking brown masses, arranged without any obvious regularity. This sac* exists 


+ at a rather early period, and is developed within a cavity formed by some excavating 


process. 

A short time after the fall of the male flowers an extraordinary change will be found to 
have occurred, consisting of the very rapid and apparently sudden development of a now 
membrano-cellular envelope between the second coat and the nucleus. This new forma- 
tion, which I may term the additional coat, envelopes the nucleus pretty closely, and is 
continued upwards beyond the apex of the nucleus into a cylindrical tubular — ho 
mouth of the tube being laciniate or fimbriated. At the period now referred to, its apex 


barely projects beyond the outer envelope. During its development, no particular change 


igi i | hat later 
has taken place either in the original integuments or nucleus. At a somewhat | 
å w +), hitherto concealed by the invo- 


period, the ovula (except in the instance quoted belo | o" 
lucrum, will be found exposed, and the outer coat to have become of a green colour: 


* . * - | g elo à 
th i i is distinct, and its direction vertical. The second envelope 
the opening through its apex is distin ook the additional or third envelapeiiów 


continues unchanged. The tubular prolonga | 

- projects through the openings in the original coats toa considerable unde itn em 

of the tube is also rather dilated, and the fimbriæ of its margin sp aa irregular | ud 
and to various extents. The whole of the tubular prolongation has me tinged ` 


brown, in some cases approaching to black. 
It is to this stage or period that the descriptions of those tinker 
a style and stigma to this genus apparently refer. Both Dr. Brown an ; : 
Trail Linda tbe du gue, Vii Mid AMO NOM MEA Lo 
+ In one species, G. Brunonianum, the ovula are at an early 
late involucrum, 


302 MR. W. GRIFFITH ON THE GENUS GNETUM. - 


must likewise advert to this period when they state the nucleus to be surrounded with 
three envelopes. | i 

The nucleus, which up to the present time continues free from adhesion, has now — 
undergone a slight alteration in figure, consisting in a tendency to constriction towards 
its apex, which is now invariably more or less brown: its cavity has become enlarged, 
This constriction may possibly be only apparent, since it may originate in the greater 
growth of the elevated part of the depression, stated to exist at the apex. : 

- As the young seed progresses in its development, the two outer coats increase in size, - 
The base of the additional membrane adheres to the corresponding portion of the nucleus, | 
the adhesion continuing to increase in extent until it has reached upwards to the com- : 
mencement of the constriction. The projecting tubular prolongation does not appear to 
have any regular term of existence; its coloration or sphacelation increases, and at a 
variable period the exserted portion becomes detached at the situation of the opening of 
the outer coat. The cavity existing in the nucleus undergoes a corresponding enlarge- - 
ment, and its walls are more irregular. It will be found to contain, about the period of 
detachment of the exserted portion of the tubular process, a pendulous body of a similar — 
form, attached by a rather broad cellular band to the apex of the cavity. This is the — 
rudimentary amnios, developed originally within the sac, stated to line at a prior period - 
the cavity of the nucleus. As the development proceeds, this body increases in size; it — 
appears to be developed from below upwards, and as it enlarges, the lax cellular attach- 
ment undergoes a corresponding diminution, chiefly, I imagine, by pressure. As this 
body, in which the albumen becomes deposited, increases, the substance of the nucleus d 
gradually disappears, and subsequently merely forms a thin covering to the large and — 
fleshy albumen. The last change which I have traced in the albumen consists in the for — 
mation of a cavity within its substance, commencing at its apex, and gradually extending — 
downwards. The corresponding changes consist in the development of pulp and increase — 
of the fibres of the outer coat; in an induration (rather in an induration of the second 
envelope depending upon the development of fibrous tissue) and development of fibrous 
tissue in the second envelope, which subsequently becomes brown, subosseous, and fragile. 
The originally large and distinct openings through both these become narrowed, but never. 
completely filled up—not, at least, by an extension of their own substance. The additional 
envelope is now membranous and thin, united throughout the greater part of its extent 
to the thin remains of the nucleus. The included (and permanent) portion of its ori in» 
ally tubular prolongation has become filled up, and is embraced closely within the con 
stricted openings existing through the two outer coats. The remains of the nucleus, 
owing to the pressure this has undergone during the growth of the albumen (or rather of 
the amnios and deposition of albumen), finally assume the form of a membrane, united, 
except towards its apex, which is brown or black ånd apiculate, to the inner paries of the 
third coat. : is — 

Mi h regard to the development of the embryo, I can add nothing. Although I hav 
examined abundance of fully-formed fruits of two species, I have never observed anything 
towards the development of the embryo beyond the formation of the cavity in - 
albumen, and the very rare and partial development of the funiculus, which, moreover, i 


MR. W. GRIFFITH ON THE GENUS GNETUM. 308 


the only two cases in which I observed its partial formation laid loose in the cavity. 
Excepting the want of this essential portion, the seeds were, as I have since ascertained 
by comparison with seeds with perfect embryos, fully and completely formed. 

The following description of the mature seeds is taken from exembryonate specimens, 
‘and from the examination of some embryonate seeds of Gnetum scandens, brought from 
Arracan, and communicated, with many other interesting productions, to Dr. Wallich by 
his late friend Captain R. R. Margrave ;— | 
_ The mature seed is more or less ovate, obtuse, or acute, and of a reddish-orange colour, 
and either entirely smooth, or covered with appressed, peltate, silvery cellular scales. Its 
. coats occur in the following order :— 

l. Outer and baccate. Origin: outer envelope of ovule.—Of this the outer half is 
entirely cellular ; the inner composed of several layers of fusiform fibres, sharp at both . 
ends, of a yellow colour and shining appearance. They are often punctated, and occa- 
sionally marked with incomplete spiral fibres ; they are pungent, and occasion considerable 
irritation, 

2. Drupaceous : originating from the second envelope of the ovule.—This is externally 
sulcate, the innermost fibres of the outer coat being lodged in these sulci; it is of a brown 
colour and nearly osseous texture, composed of cells, the innermost series of which are 
arranged transversely, and a great number of longitudinally-disposed fibres, which are 
longer and of much less diameter than those of the outer baccate coat. 

3. Fibro-cellular coat. Origin: the additional envelope, and chief part of the nucleary 
membrane.—The external part, which is composed of fibres similar to those of the drupa- 
ceous coat, arranged longitudinally, is terminated by the persistent and now woody p 
tion of the originally tubular prolongation, the apiculus being lodged in the opening 
existing even at this period in both the outer coats: its apex is discernible exteriorly at 
the corresponding end of the seed, but does not, however, project beyond the level of the 
outer coat. The internal portion forms the | 

4. Cellular and membranous envelope. Origin: the circumferential tissue of the 
nucleus, — This adheres to the preceding throughout the greater part of ws extent. Towards 
the apex, where it is thinnest, most membranous and sphacelated, it is free, and it is 
terminated by a conical point (the original elevated portion of the centre of the depressed 
_ apex of the nucleus) : it is free from adhesion with the former coat. The point, however, 
alwa tes with the albumen, the upper portion of which it covers somewhat in 

ys separates wi 
the manner of a cap. | x 
5. Albumen. Origin: a deposition in the tissue of the amnios.—This - eom en 
cavity of the last-mentioned envelope; it is of considerable size, fleshy, and nn 
feculent granules, Around its apex an indistinct areola is visible, ar zZ : 2% 
"m ør Jara ; 
apex are attached the remains of the originally lax, cellular mn d zs the 
Wise marked with a depressed, indistinct areola, Along its contre, m aa ha 
: ; ; í ity of considerable size, which tapers off in 
upper half* of this portion, there is a cavity of co 


: : T ios to the apex of the 
* With regard to the axis at least; otherwise the base of the original amnios corresponds 
- nucleus, 


. the pollen of anthers belonging to p 


304 | : MR. W. GRIFFITH ON THE GENUS GNETUM. 


feriorly, terminating towards the middle of the long diameter of the albumen ; the tissue, 
however, on the same line, and between it and the apex of the albumen being more dense, 
and having a peculiar appearance. The walls of this cavity are rugged and irregular. To 
the upper portion, and to one side of the cavity, is attached the embryo by means of an 
enormously long, tortuous, and spirally but irregularly-twisted cellular funiculus, the: 
cells being much elongated and twisted. Its length varies, when moderately pulled out, 
from 34 to 5 inches, the length of the fruit being 1inch: This funicle, as well as the. 
extremely similar one of Cycas, has the property of contracting when immersed in water. 


When in situ it is tolerably closely packed ; it is dilated towards its attachment with the 


embryo. The embryo itself is more or less ovate, its radicle tapering off superiorly, and 
being completely continuous with the apex of the funiele. The cotyledons are equal, very 
small, and mutually applied by their plane contiguous faces. The plumule is incon- 
spicuous, and only indicated by a rounded, entire, very minute elevation. . The lower half 
of the embryo is lodged in lax, apparently dislocated tissue. ay bøn | 
[Roxburgh * describes the seed of G. scandens as having, besides the two outer coats, . 
which he refers to the fruit, “two integuments : the exterior one thin and fibrous ; the l 
inner one less distinct, and adhering to the perisperm very firmly. Perisperm conform 
to the seed, cartilaginous. Embryo in the apex of the perisperm, straight. Cotyledons 
two, ensiform, unequal. Radicle superior.” In his drawing of G. scandens the embryo is 
represented with a short obtuse radicle, two unequal cotyledons, and an immense plumule, 
consisting of a filiform stalk equalling in length the larger cotyledon, bearing at its — 
apex two small leaflets. N othing is said of the funiculus. ] . us 
From the foregoing account, I trust it will be. seen. that, with the exception of the 
subsequent and sudden appearance of the additional membrane t, the development of the 


ovula of Gnetum presents scarcely a single peculiarity worthy of notice. The changes s 


that occur in the two outer coats from an early period up to the time of the maturity of 
the seed, I consider as of secondary importance, and as the result of modifications de- 
pending upon their anomalous situation. The sudden appearance of the additional mem- — 
brane does not seem to me to be capable of any explanation: hitherto I have not met — 
with it in any stage of development prior to that when its apex reaches to the opening T 
existing in the outer coat. I can assert with tolerable confidence, that this coat does not 
originate in a separation of the cireumferential tissue of the nucleus, and that hence it is 
not analogous to the fercine, I am unable to state what functions it performs, although - 
its exsertion, its free communication with the apex of the nucleus, its dilated laciniate - 


apex, and its subsequent sphacelation would lead us to suppose that it is connected with — 


| : spikes at an earlier period of growth. | ROI 
The complete agreement in development of the nucleus and albumen with that of the 2x 
' * "Flora Indica, vol. iii. p. 519. 


. + Roxburgh, in his figure of G, 
the existence of the male flowers. 


scandens, represents the exsertion of this membrane or coat as occurring dunng | 


* 


MR. W. GRIFFITH ON THE GENUS GNETUM. | 305 
e» 


same parts in ordinary ovula, precludes the necessity of any remark ; neither is there any 
feature in the embryo, excepting its enormously long funiculus, that would lead me to 
suspect any material deviation from the usual structure. ` Å 

The only genera of Comiferæ with which I have been able to compare Gnetum are 
Agathis and Thuja, in both of which the nucleus has only one envelope, its cavity being 
in Agathis lined with a very distinct amnios. The males of neither of these genera, how- 
ever, exist in the Botanical Garden at Caleutta; and neither of them appears to possess 
that power, which exists to such a degree in Oycas and Gnetum, of continuing the deve- 
lopment of the ovulum independently of fecundation. With Cycadeæ I have had ample 
opportunities of comparison, so far, at least, as regards the young ovula and perfect fruit. 
The mutual resemblance is, indeed, so strong, that I have little hesitation in affirming 
that the fruit of Cycas differs only materially from that of Gnetum in the absence of 
an additional envelope, and perhaps in the presence of the remains of the amnios. With 
regard to the ovula of Cycas, the nucleus has, as Dr. Brown has stated in the Memoir 
referred to, p. 24, only one envelope; but I have no doubt, both from the difference in 
its tissue, in which there is, indeed, an obvious line of demarcation, as well as from the 
disposition of its vessels, that it consists of two, united throughout their whole extent. 
The nucleus is likewise united by its lower portion to the envelope,—its apex, which is 
conical, being free, and prolonged into a membranous point, which is engaged within 
the lower part of the canal formed by the opening that exists through the coat. This 
membranous portion is tubular, the cavity extending some distance within the actual 
substance of the nucleus : its walls are cellular, and not lined by any membrane. 

The body of the nucleus contains a large globular cavity, in which a sac of a corre- 
sponding size and form exists. This sac (the amnios) is, at the period referred to, mem- 
branous, and is rendered tense by an abundant gelatinous fluid; it is easily detached, and 
appears to be merely applied to the walls of the cavity. 

I hope to be H Bone future period to enter into more detail on the subject of the 
development of the ovula of Cycas, which appears to promise several yr al and na 
interesting facts, particularly as regards the first appearance of cellular Hesne E; 
the sac of the amnios, while at the same time its great size renders it peculiarly fitted Ar 
observation. ge ER | Li 

It is probable that the fruit represented by Dr. Hooker (Bot. Mag. gi em 
t. 2827) as that of Cycas circinalis, belongs to another ne The true MEG of 
has, as Richard has stated, a fungous envelope surrounding the albumen, \ iini 

considerable thickness towards the base of the fruit, or rather seed ; a a ye ae 
probably the remains of the amnios, being interposed, and OaE 9 ES har 
velope, except towards its apex. In this species the cotyledons ifii i AER 
likewise stated, united, except at their bases, where å ipu deep z 


coats, agrees much 
ge intion, particularly as regards the coats, : 
on both sides. Dr. Hooker's description, particularly apex of the albumen, first pointed 


better with C. sphærica, Roxb. The sacs existing in the | 
| rman in whic! "yo is develo 
out by Dr. Brown *, are in C. circinalis permanent ; that in which the embryo ped 
* Prodr. Flore Nova Hollandie, vol. i. p. 347. A 


VOL. XXII. 


EEE EEE GNOME, TES pe. SNE 


306 MR. W. GRIFFITH ON THE GENUS GNETUM. 


becoming necessarily much enlarged, and adhering firmly to the walls of the cavity it 
lines. 


* Although I have repeatedly examined the ovula of Gnetum with a view to the nature 
of their impregnation*, I am not in possession of a single fact relative to its performanee. 
I must, however, mention, that the ovula of Gnetum scandens and lepidotum do not 
appear to be ever submitted to the action of the pollen derived from their proper male 
flowers. In G. Brunonianum this is obviated by the extreme smallness of the annulate 
involucrum. My residence at the Botanical Garden of Calcutta, and the great and un- 
varying kindness of Dr. Wallich, have put me in possession of some, I think, interesting 
facts relative to the impregnation of Cycas, which it may not be amiss to state. I may 
here observe, that the consideration of the mode of application of the pollen-tubes to the 
apex of the nucleus in some plants possessing the ordinary structure of ovarium, had led 
me to suppose that these productions are merely organs of communication, developed on 
account of the distance that necessarily exists in these plants between the stigmatic 
surface and ovulum, and that hence, in Cycas and plants of a similar simple structure, 
in which actual application of the pollen itself to the apex of the nucleus can obviously 
take place, no pollen-tubes would be produced. 

In forming this view, I had not lost sight of the apparent penetration of the pollen- 
tubes into the ovula of Asclepiadeæ, first observed by Dr. Brown; but I supposed that it 
might be the result of the anomalous formation of the nucleus in these cases. Repeated 
observation has since, however, taught me, that not only does application of the pollen- 
granules to the apex of the nucleus of Cycas take place, but that pollen-tubes are like- 
wise generated, although, as might be expected, much shorter than usual. Indeed, the 
tubular membranous portion of the apex of the nucleus becomes actually crammed with 
pollen-granules, from the lower and outer of which pollen-tubes are pretty generally 
produced. | 

The orifice of the envelope of Cycas has a callous and shining appearance; and — 
although I have often examined pollen-grains which had been in apposition with it for — 
some time, I have never seen any production of boyaux, except in the cavity of the apex 
of the nucleus. E 

Although I have by no means proved the necessity of the production of the pollen- — 
tubes in Cycas to ensure fecundation, I consider the fact of their production a strong — 
argument in favour of the idea that actual penetration does occur in every case in which E 
the application of the tubes to the apex of the nucleus can be conceived. The fact of the. ; 
production of the tubes likewise seems to me to put the nature of the bodies from which > 
they originate out of all doubt, and to prove the truth of Dr. Brown's remark (Memo — 
cited, p. 30), that it would be quite gratuitous, on the grounds stated, to consider the par — 
ticles contained in the thecæ to be analogous to the fovilla. E. 


Dr. Brown, in his account of microscopical observations on the particles contained in i : 
the pollen of plants, published in the * Edinburgh Journal of Science,’ vol. ix. p. - 2 


* The opening of the outer coat never presents a shining appearance. 


MR. W. GRIFFITH ON THE GENUS GNETUM. 307 


says, after alluding to his want of success in tracing the particles contained in pollen- 
grains through the tissue of the style, “ Even in those families in which I have sup- 
posed the ovulum to be naked, namely Cycadeæ and Coniferæ, I am inclined to think 
that the direct action of these particles, or of the pollen containing them, is exerted 
rather on the orifice of the proper membrane than on the apex of the included nucleus; 
—an opinion which is in part founded on the partial withering confined to one side of 
the orifice of that membrane in the Larch,—an appearance which I have remarked for 
several years.” It is, however, most probable that, from the late rapid increase of our 
knowledge of the process of fecundation in Phænogamous plants, chiefly indeed owing to 
the beautiful observations of this distinguished botanist, the opinion above cited has been 
modified by its author. 

With regard to the two principal objections urged by Dr. Brown * against the opinion 
of the female flower of Cycadeæ and Conifere being a naked pistillum, the first, viz. that 
arising “from the perforation of the pistillum, and the exposure of that point of the 
ovulum where the embryo is formed to the direct action of the pollen,” still holds good. 
The second, viz. the too great simplicity of structure of the supposed ovulum, I look 
upon as in a great measure destroyed by the reduction of the usual number of the 
constituent parts of this organ in Loranthacee}. 

Respecting the male flower, I have to add, that Linnæus was correct in referring the 
genus to Monæcia Monadelphia. Roxburgh adopts the same view in his MS. Synopsis ; 
but in his ‘Flora Indica’ he refers it to the Monæcia Monandria. The correctness of 
Linnzeus’s view is proved by the fact that there are two filaments at an early period, or at 
least that their union is only partial, and by the number, situation, and distribution of 
the vascular fascicles. 


GNETACEE, Lindley, in Bot. Register, vol. vii. N. S. t. 1686, sub Garrya. 


DraGnosis.— Plante gymnospermæ, dicotyledoneæ, aquosæ. Rami articulati. Folia op} ; 
venis anastomosantibus reticulata. iis ie d 

CHAR. gssENTIALIS.—Flores mohoici, in spicis amentiformibus verticillatim dispositi ; qum = 
lus involucro annulato integerrimo (rarius obsoleto) suffultus distans. MASCULI pluriseriati, in Mrd 
cillis inferius dispositi. Perianthium tubulosum, inverse subulato-conicum — a 
pressione angulatum, apice planiusculo vel depresso, rimá transverså | erg ses thesi ; 
mentum unicum, monadelphum (e 2 nempe coalitis formatum), hypogynum, , gere T 
per rimam transversam exsertum. Anther@ duæ, uniloculares, — E Mara a ple b 
nino discreti longitudinaliter et centraliter secus latus utrumque — ; m red 
oblongum.—Fæmrnan in verticillis superius dispositi, 1- niat deis pos ea E ix. 
ovata, sepe obliqua, transverse sita, orthotropa. (Junius.) Tegum — A ]lulosus å 
losum, interius cellulosum, sæpius inclusum, utrumque T apertum. Nucleus = a a 
mento conformis, liber. (Maturius.) Tegumenta terna; tertium T oda » e 
ejus apicem in tubum cylindricum styliformem longe exsertum ore ma pac ec us, 
niatis productum. Fructus (semen maturum) omnino exsertus, oo ve à le ig Auge 
lævis vel lepidotus, ovatus, indehiscens. Sarcocarpium extus cellulosum, intu 


i ium fibrosum 
utrinque acutæ pungentes, diametro magno, coloratæ), apice caper vcre = 
fragile, subosseum, sarcocarpio admatum, e fibris longitudinaliter cellulisque 

| my Paper on this subject. 
* * Kingid, p. 28. + See my Pape ds 


308 MR. W. GRIFFITH ON THE GENUS GNETUM. 


formatum, apice apertum. Tegumentum tertium fibroso-cellulosum albumen amplectens, apice in 
apiculum subulatum induratum endocarpium et sarcocarpium perforantem sed vix exsertum pro- 
ductum. Albumen semini conforme, carnosum, copiosum, apice membranä apiculatå lacerå spha- 
celatå (nuclei reliquiis) incomplete tectum, intus excavatum. Embryo inversus, in excavatione albu- 
minis reconditus, ope funiculi longissimi (3-5 uncialis) cellulosi, spiraliter torti, albuminis excavationi 
lateraliter et apicem versus affixi sustentus. Radicula conica, supera, cum texturå apicis funiculi 
continua. Cotyledones parvæ, æquales. Plumula inconspicua.—Arbuscule, fruticesve sæpius scan- 
dentes succo aquoso fæti. Truncus conicus, ramosus. Lignum fibroso-vasculosum, zonatum ; Zonis con- 
centricis maxime evolutis, e radiis medullariis inter se lateraliter confluentibus formatis ; e vasis mazi- 
mis glanduloso-punctatis (more Coniferarum) fibrisque ligneis crebre punctatis constans. Rami articu- 
lati ramulique virides ad articulos tumidi. Folia opposita, petiolata ; petioli superne plani inferne con- 
vezi, basibus in annulum. intrapetiolarem (fere ut in Potaliaceis) connati. Limbus indivisus, ovato- 
vel lanceolato-oblongus, plus minus acuminatus, sepius repandus, integer, penninervis, nervis venis 
secundariis et tertiariis arcuatim coalitis, Supra secus nervum centralem et bases secundariorum 
stomatibus donata ; infra. per totam superficiem venis venulisque exceptis organis iisdem donata.—(See 
Note 1.) Folia novella per vernationem paginis superioribus planis mutuo et arcte approximata. In- 
florescentia paniculatim spicata, axillaris vel sepius terminalis. Spice cylindrice, 3 in quáque divi- 
sione panicule ; laterales opposite pedunculate ( pedunculi ad bases annulo obsoleto cincti) ; terminalis 
longius pedunculatus, pedunculus supra medium bracteis 2 ovatis alte connatis stipatus. Involuera 
annuliformia, primo approximata demum discreta vel distantia ; infima spicarum presertim lateralium 
biapiculata. Flores pilis cellulosis articulatis quasi moniliformibus immizti presertim feminei ; 
utriusque sexis arcte verticillati, radiatimque patentes. Masculi inferiores, et in seriebus pluribus 
dispositi. Foeminei superiores, I vel 2 seriales, per anthesin involucro pilisque sepius omnino obtecti ; 
Jloribus masculis delapsis cito exserti, et demum omnino nudi. 


- 


I shall conclude with a synopsis of the species I have ascertained to be indigenous to 
the Tenasserim Provinces. 


GNETUM, Linn., Mantissa, p. 18, no. 1278. 


Character ordinis. 


Sectio I. Zrecta. 


G. BRUNONIANUM, mihi; fruticosum ; foliis lanceolato-oblongis (membranaceis), involu- _ 
cris obsoletis. 


Hab. in sylvis ad Banlau, urbem antiquam anglice Tenasserim dictam prope, florens ab Aprile usque ad - 
Februarium. Fructus non vidi. Frutex humilis, erectus ; ramis divaricatis. Folia breviter petio- 


lata, subrepanda. Spice axillares et terminales, sæpius solitariæ ; flores fœminei ezserti, 1-seriati, 
virides, pilis cellulosis albis immixti. 


Sectio II. Scandentia. 


G. APICULATUM, mihi; foliis lanceolato-oblongis, fructibus sessilibus acutis lævibus. 


Hab. in Silhet, Roxb. Arracan, Capt. Margrave. Legi in sylvis circa Mergui, ore Tenasserim. Frutex 


longe scandens ; folia. coriacea, subacuminata. Flores pilis cellulosis brunneis immixti, foeminei per 


anthesin, marium omnino obtecti. Fructus (lanceolato-ovati) apiculati, omnino læves. 


MR. W. GRIFFITH ON THE GENUS GNETUM. 309 


G. SCANDENS, Roxb.; foliis ovato-oblongis vel ovalibus, fructibus breve stipitatis obtusis 
lepidotis. Ula, Hort. Mal. vol. vii. p. 41. t. 22 opt. Gnetum scandens, Roxb. MSS. 
Synopsis; ejusdem Icones pictæ in Horto Botanico Caleuttense asservate, Suppl. 
vol. iv. t.78; Flora Indica, vol. iii. p. 518. 

Hab. in sylvis ore Tenasserim. Legi ad Moulmein, Amherst, et Mergui. Floret Decembre, Januario. 
Frutex longe scandens. Folia coriacea, obtuse acuminata; supra atro-viridia, raro pallida. Paniculæ 
terminales. Flores utriusque sexås pilis cellulosis albidis immixti. Fructus (ovato-oblongi), præ- 
sertim juniores, pulcherrime argenteo-lepidoti. 


Under this species Roxburgh has confounded two distinct plants. In the drawing 
quoted above he has figured the fruit of @. apiculatum and of @. scandens as the pro- 
duce of the same plant. In his description of the fruit of G. scandens, he seems to advert, 
although obscurely, to some peculiarity of the superficies, at least before maturity. This 
species is at once known by the silvery scales of the fruit, which are very conspicuous 
before it begins to assume its orange colour; these scales are peltate, closely appressed, 
and composed of cells radiating from the situation of the attachment. 

I have not quoted Rumph, whose figure does not at all resemble our plant. Bu- 
chanan’s synonym in Rees’s Cyclopædia, founded on the supposed identity of Rumph's 
plant with the above, is therefore not to be taken without great hesitation. 


Note 1.—(Stomata.) The arrangement above referred to is not uncommon, but pe 
haps limited to those leaves in which the parenchyma is continued over the veins. It is 
remarkable that in these cases the stomata differ considerably in size ; those of the upper 
surface I have hitherto found to be the largest. As good instances of such distribution, 
I may mention Costus speciosus, and perhaps all species of Nymphæa. A curious, and I 
believe hitherto unnoticed singularity occurs in the distribution of these "arii m 
Nelumbium speciosum, in which the callous spot in the centre of the leaf, and opposite to 
the termination of the petiole, is crowded with stomata of a large size, to the pov of 
which the unusual colour of the spot appears to be partly owing. These a" - 
plant certainly open into cavities, through which they communicate indirectly with the 
cavities in the petiole, from the apex of which they radiate into the ims un ser så 
of the vast limb is minutely papillose, the Menace Sk ncm 
stomata are very indistinct, and indeed almost o m 

Note 2.— Dr. Lindley, in the * Botanical Register’ quoted above, states Ts CUIR 
of Gnetum is zoneless. In the climbing species the zones are highly dev LE : í 
G. Brunonianum no zones existed; this is, however, in all probability ais E eA 
specimens that I examined being the growth of Uno pe t em ro RR A 
confirming the statement of the above author respecting the Api TM E ve 
least in the young parts. I may add, that the proportion of v —-— y 


considerable. omale flowers reduced to naked 


Finally, the three families characterized by having the fe a rir stile pari 
ovula, agree in the following remarkable points: VIZ. peculiar p | sr 
presence of at least two, often 


of their tissue, unisexuality, orthotropous ovula, and in the 
more, opposite cotyledons. 
Aug. 4th, 1835. 


310 


MR. W. GRIFFITH ON THE GENUS GNETUM. 


EXPLANATION OF THE PLATES. 


Tas. LV. 


Fig. 1. Very young ovule of Gnetum scandens viewed on its upper or posterior surface, which is observed 


to be cleft longitudinally. 
Fig. 2. Second envelope, showing its divided apex, the laciniæ of which are connivent. This coat is 


visible at a in fig. 1. 


Fig. 3. Nucleus of the same, both coats being removed. | 
Fig. 4. Posterior view of an ovule (long before the dehiscence of the anthers), showing the longitudinal 
cleft, the margins of the upper part of which are in this case, and indeed generally at a period 


Fig. 7. 
Fig. 8. 
Fig. 9. 
Fig. 10. 


Fig. 11. 


Fig. 12. 


Fig. 13. 
Fig. 14. 


Fig. 15. 


Fig. 16. 


Fig. 17. 


Fig. 18. 


Fig. 19. 


Fig. 20. 


Fig. 21. 


rather later than that to 


which figs. 1-3 refer, in apposition ; part of the second coat is visible 


at a, and another portion projects through the apex at 5. 
Fig. 5. Inferior or anterior view of the same ovule, along which no fissure is visible. 
Fig. 6. View of the inner or second coat, showing its irregularly-divided apex. 


Nucleus highly magnified (the coats removed), showing the depression at the apex and the 


central conical elevation. 


Apex of the nucleus; the conical elevation appears emarginate. 
Ovule of G. Brunonianum ; no fissure through the outer coat except at the apex. 


Longitudinal section ; the apex of the inner coat is visible on a level with that of the outer coat. | 
Å central cavity enclosing the remains of a sac is visible within the nucleus. 
Transverse section of the same. 
The inner coat, with part of the outer remaining at the base. This figure shows the narrow, - 
deeply-divided apex of this coat. | 
Nucleus, with half the bases of the coats remaining, | 
Ovule of G. Brunonianum some time after the fall of the male flowers; at its apex are visible — 
several projecting processes of cellular tissue. v 
Vertical section of the same, showing that a new coat has been developed between the inner (now = 
second) coat and the nucleus. This coat is prolonged beyond the nucleus into a cylindrical 2 
tube, the mouth of which is divided, and projects slightly beyond the opening through the RB 
of the outer envelope. The second coat is now tinged with yellow. E 
Ovule with outer coat detached; the tubular prolongation is seen to project considerably beyond 


the opening in the apex 
yet become expanded. 


of the second coat. The fimbriæ of the extremity of the tube have not 


The same with the second coat removed, showing the third coat throughout. 2 
Vertical section of an ovule of G. Brunonianum, showing the parts in situ. The apex of h 


second coat is preserved 


entire : the correspondence of the apex of the nucleus to the commence- 


ment of the tubular prolongation, the expanding fimbriæ of this, and the adhesion with the base — 
of the nucleus, are represented. The tube is not much sphacelated at this period. The termi- - 
nations of the vascular fascicles at the base of the nucleus are shown. fe 

Transverse section of the ovule near its apex, showing the two envelopes and a portion of the a 


tubular prolongation. 


Vertical section of the whole of the third envelope and the nucleus. At the base of the nucleus a 
appear the terminations of the vessels ; but this is only apparent, the vessels being cut through as- 


they diverge outwards to pass into the envelopes ; none exist in the nucleus. m 


Vertical section of the i 


mmature fruit of Gnetum apiculatum, some time after the fall of the : 


Fig. 22. 


MR. W. GRIFFITH ON THE GENUS GNETUM. 311 


exserted portion of the tubular prolongation. The prolonged apex of the outer coat (a) is distinct, 
as well as the formation of the large yellowish fibres. The second coat (5) has become thicker 
towards its apex, and somewhat indurated. The inner or additional coat (c) is now adherent to 
the nucleus throughout the greater part of the extent of this latter organ. That part of it which 
corresponds to the constricted apex of the nucleus is thickened ; its prolongation is considerably 
dilated after it has passed through the narrow, and now cylindrical, tubular perforation of the 
second coat. The excavation is large, and its margins are rugged and more or less blackened. 

Vertical section of an ovule of G. scandens ; the nucleus is surrounded inferiorly by the two outer 
coats ; the notch on either side (a, a) indicates the place of separation of the additional (third) coat. 
The cavity is much enlarged, and occupied by a clavate body, of which the outline is given in 


- the adjoining figure 23. 


Fig. 25. 


Fig. 26. 


Fig. 27. 
Fig. 28. 


Fig. 29. 


Fig. 30. 


Fig. 31. 
Fig. 32. 


Fig. 33. 
Fig. 34. 


Body occupying the nucleary cavity ; this is the rudiment of that which subsequently becomes 
the very copious fleshy albumen. 

Vertical section of a full-grown fruit of G. apiculatum, in which, however, no embryo had been 
developed: a. outer baccate coat, the inner half of which is composed of very large fibres ; 
these are of a yellowish colour, shining appearance, and imbedded in a reddish-yellow pulp ; 
b, b. drupaceous covering; c, €, c. inner or additional coat, with its woody, fragile, enclosed and 
prolonged apex (to this the fourth coat adheres firmly, except at its sphacelated apex, d); 
e. albumen, now copious and fleshy, by the great development of which the original nucleus has 
become reduced to a thin membrane, of which the free apex is seen at d; f. excavation at the 
apex of the albumen occupied by broken-up tortuous tissue, the rudiments of a funiculus. The 
direction of the subsequently-extended excavation may be traced lower down, and is indicated by 


a greater density of texture than occurs elsewhere. 
Albumen of the same detached, viewed on its entire surface : at d. is seen the apex of the nucleary 


membrane with its sphacelated apiculus. 
Vertical section of the same, showing the cap which the adhering apex of the fourth or nucleary 


membrane makes for it. The tortuous tissue is visible in the cavity. 


Tas. LVI. 


Considerably advanced ovule of G. Brunonianum ; vertical section. 
Vertical ender of a mature and, with the exception of the embryo, "d : - ead 
dens: a, a, a. the outer coat, which has become pulpy and baccate, m lying rear LA fe 
the pulp; it is perforated by the prolonged persistent apex of ME es x Dan de 
second or drupaceous coat, shown to consist of two layers = eee: deg Ed (RES 
third membrane, composed of the additional and nuclear mem ge : dn bien. and 
at their apices,—the apex of the nucleary membrane always separating with 


: > Er ; e. the central apical cavity, empty. 
ng Ge e NE : sse a. outer coat; 5. second coat; c. nucleary 


Vertical section of an immature seed of G. apiculatum m 
membrane of albumen ; d. albumen ; e. sphacelated apex of albumen ; f. apical cavity; g- 
suspensor lodged in ditto. : 

Tortuous suspensor (g) in fig. 29 extracted and highly magnified. 


Vertical section of the separated albumen of a similar seed. 


Vertical section of the albumen with its cellular attachment ; 


lateral attachment, : 3 MR 
Embryo and funiculus extracted and highly magnified, showing the ey sqm apt 
Embryo with the apex of the funiculus ; the cotyledons are seen 


equal. 


showing the embryo in situ and its 


312 


Fig. 35. 
Fig. 36. 
Fig: 37. 
Fig. 38. 


Fig. 39. 


Fig. 40. 


Fig. 41. 


Fig. 42. 
Fig. 43. 
Fig. 44. 


Fig. 45. 
Fig. 46. 


Fig. 47. 


MR. W. GRIFFITH ON THE GENUS GNETUM. 


The same, the embryo varied in shape. 

The same, with one of the cotyledons removed, showing the rudimentary plumule. 

Portion of the centre of the funiculus, showing the spiral disposition. 

Portion of inflorescence of G. apiculatum; the ovules are totally concealed and surrounded by 
very dense brown hairs. Exsertion of the stamina has taken place. | 
Portion of inflorescence of G. scandens ; the ovules are totally concealed, and the filaments are 
exserted to a considerable length; the transverse opening through the male perianth is re- 
presented. 

A verticil viewed from above, showing the ovules in situ, and partially surrounded with hal 
In this species the ovules are arranged in about two series, and the fissure is visible at the 
apices of some. 


Portion of inflorescence of G. Brunonianum, including one verticil with its obsolete ring (invo- id 
lucre). In this species the ovules are completely exposed at this period, their bases being sur- Pide 


rounded with hairs; they are arranged in one series : some of the stamens are exserted. 


Portion of the spike of G. scandens, representing the ovules fully exposed, and the exsertion of | = 


the tubular prolongation of the additional or third membrane. 
Stamen of G. Brunonianum (?) at an early period ; the filaments are still distinct at their bases ; 
the number, direction, and termination of the vascular fasciculi are shown. 

Male flower of G. Brunonianum (?) before dehiscence. 

Stamen of the same, the base of the perianth partly remaining; the anthers have dehisced. 
Pollen of the same, 

The same after immersion in water. 


& 


* [ 313 | 


XXIV.—Synopsis of the Fructification of Jf Simple Sphæriæ of iia Hookerian 
‘ Herbarium. By FREDERICK Currey, Esq., M.A., F.R.S., F.L.S. 


&. ; 
Read May 5th, 1859. 


Ina paper published in the last Part of the‘ Transactions’ of this Society, I have figured 
and described the fructification of all the species of compound Sphæriæ contained in the 
Hookerian herbarium, that is to say, of all those species whose fruit was sufficiently 
perfect. Sir William and Dr. Hooker having kindly afforded me the same facilities for 
the examination of the other great division of the genus, that is the simple Sphæriæ, I 
am now enabled to lay before the Society the result of my examination of the remaining 
species contained in their herbarium. I adopt the course pursued in the former paper, 
of prefixing a short account of the characters på f the sections and divisions, as given in the 
‘Systema Mycologicum.’ ^ 

Most of the plants described in the ox: paper kake to thé genus Sphæria as 
limited in Fries’ ‘Summa Vegetabilium Scandinaviz,’ but a few are referrible to some of 
the new genera proposed in that work, and I have noted such of the species as belong to 
the new genera, and have given the characters of such genera in the notes, as they occur. 

The simple Sphæriæ commence with the 5th Section, * SUPERFICIALES.” 


SECTION V. SUPERFICIALES.—Perithecia free, bicorticate, seated on an effused villous subiculum, 
or altogether superficial. - a " 
Div. 17. Byssisepx —Perithecia free, glabrous, with a short subpapillæform ostiolum, seated on 
a tomentose subiculum, formed of densely interwoven threads. 


Div. 18. ViLLosæ — Perithecia. ovate or globose, clothed with simple persistent down; en 


even, subpapillæform, rarely elongated or obsolete. | 
Div. 19. Denupatæ.—Perithecia naked, ovate or globose, glabrous, without kg Fr 


© z 
ostiolum short, subpapillæform. , us På 


Div. 20. PERTUSZ. ‚—Perithecia : naked, glabrous, flattened at the base, adnate o: 
` by the falling off of the ostiolum. iind 
Section VI. SUBIMMERSÆ.—Perithecia immersed, often erumpent ; ostiolum conspicuous, i 
or elongated into a neck. a 


Div. 21. PLATYSTOME. «—Perithecia at first covered, then more or less exposed ; — some- 
what compressed, very broad, opening bya longitudinal fissure. 
Div. 22. Czratostomæ.—Perithecia at first covered, often surrounded with down, then emerging, 


naked, free, black, terminated by a beak eylindrig ostiolum generally longer than the. 
af 


Ee 
> perithee ER 
Div. 23. DN V EN immersed in the. perennial parts of plants, with a mas erumpen 
neck, which is often dilated at the apex. — : um 2 


of the matrix. 
Secrion VII. Susinnaræ.—Perithecia innate in the epidermis l * 
Div. 24. OBTURATÆ.—Perithecia at first innate, covered by the epidermis, then Er naked, 


almost superficial, glabrous; ostiolum naked, pa] il form or rimosely dehi 
VOL. XXII. 


Ll 
x2 


314 MR. F. CURREY ON THE FRUCTIFICATION OF SIMPLE SPHÆRILÆ. 


Div. 25. Susrecræ.—Perithecia innate, covered, at length erumpent, concrete with the matrix, 


often without a prominent ostiolum. | 
Div. 26. CavticoL.z.—Perithecia subinnate, at first covered, at length exposed by the separation 


of the epidermis, discrete from the matrix. 
Div. 27. Forucozæ.—Perithecia innate, covered, connate with the matrix, very rarely erumpent — 


or free. ; 
Div. 28. Depazea.—Perithecia simple, innate, seated on an arid spot on leaves which are still 


green. 


Div. 17. BYSSISEDÆ. 


215. S. Guss, Berkel. and Currey MS. Tas. LVII. fig. 1, ascus with sporidia, x 220. 
Sporidia biseriate, pale brown, simple, slightly curved, 0:0009 to 0-001 inch long, 
rounded at each end. Perithecia rather large, round and very flat, seated on a dense 
subiculum, usually entirely hidden beneath the bark, not erumpent, but raising the 
bark into smooth, rounded, or elongated swellings; the perithecia and subiculum 
are usually of a dark dirty green tinge. I do not know where to classify this plant, 
if not with the Byssisedæ, although it is quite anomalous to find a Spheria of this 
division entirely hidden by the bark. The presence of the fungus is only indicated 

_ by the smooth swelling of the small branches of oak, on which it occurs. The wood 
of these branches is generally somewhat decayed. The Spheria is common in the 
neighbourhood of Weybridge, and I have also found it near Tunbridge. It grows, as 
far as I know, only on oak, and I am not aware that it has been hitherto described. 
Some of my correspondents have received it from me under the manuscript name of 
5. tomentosa; but Mr. Berkeley, to whom I sent it at first without a name, marked 
it in his herbarium as Spheria Glis, and at his request I have adopted that name. 

216. S. Desmazrert, Berk. and Br.; Ann. & Mag. Nat. Hist. 2 ser. vol. ix. p. 318. pl. 9. 
fig. 1. Tan. LVII. fig. 2, ascus with sporidia, and free sporidia, x 325. Sporidia - 
uniseriate, overlapping, dark brown when ripe, with globular and irregularly -shaped 
nuclei, subeymbiform, somewhat flexuous, 0:0013 inch long. 

217. S. (NECTRIA) ROSELLA, A.and S.; Fr. S. M. ii. p.441. Tas. LVII. fig. 3, sporidia, 

X 325. Sporidia fusiform, slightly constricted in the middle, colourless or greenish, 

_ with the endochrome bipartite, 0-0010 to 00012 inch long. 

218. 8. AQUILA, Fr. 8. M. ii. p.442. Tam. LVII. fig. 4, sporidia, x 325. Sporidi& 
dark brown, almost opaque, almond-shaped or subeymbiform, or oblong and slightly 
curved, sometimes with one, two, or three nuclei, 0:0006 to 0:0008 inch long. 

219. S. TRUNCATA, Sz.; Fr. S. M. ii. p. 442. Apparently not distinct in any material 
respect from S. aquila, Fr. ; | 

220. S. FULVA, Fr. EL ii. 90. Tas. LVIL fig. 5, asci with sporidia, and free sporidia, 
X 325. Sporidia biseriate or crowded, oecasionally uniseriate, colourless, broadly 
almond-shaped, sometimes subeymbiform, 0-0004 to 0-0005 inch long; endochrome 
bipartite, sometimes continuous. d 

221. 8. (Nzcrrna) AURANTIA, Pers.; Er. S. M. ii. p.440. Ta». LVII. fig. 6, ascus with 
Sporidia, and free sporidia, x 395. Sporidia uniseriate, colourless or greenish, 
almond-shaped or flexuous, 0:0006 to 0-0008 inch long ; endochrome bipartite. On 


MR. F. CURREY ON THE FRUCTIFICATION OF SIMPLE SPHÆRIÆ. 315 


Polyporus versicolor. I have in my herbarium a Sphæria on Polyporus hispidus, 
not apparently distinct from N. aurantia, Pers., but producing the fruit shown in 
fig. 7, where the sporidia are elliptical, not acuminate or flexuous, 00005 inch long, 
and very different in appearance from those of the Kew plant. 

222. S. THELENA, Fr. S. M. ii. p.441. Ta». LVIL fig. 8, ascus with sporidia, x 825. 
Sporidia uniseriate, black-brown, opaque, elliptical or slightly curved, sometimes 
with a large nucleus, 00009 inch long. Not distinct, I should say, from $8. aquila, 
Fr., although the sporidia are rather narrower. 

223. S. LANATA, Fr. 8. M. ii. p.442. The specimens of this plant are in bad condition. 
I could find very few sporidia, and those hardly differed from the sporidia of S. the- 
lena, except perhaps in being slightly more acuminate. Perithecia somewhat col- 
lapsed, without a manifest ostiolum ; subiculum loose, hairy. 

224. S. SUBICULATA, Sz.; Fr. S. M. ii. p.443. Tas. LVII. fig. 9, sporidia, x 325. Spo- 
ridia broadly elliptical, with sometimes one, sometimes two nuclei; the two occa- 
sionally approaching so close as to give a septate appearance, dark brown, 00004 
inch long. The subiculum is wanting in the Kew specimens. The plant is like 
S. aquila, but the perithecia are blacker, flatter, and much smaller. 

225. S. ALLIGATA, Fr. S. M. ii. p. 445. Tas. LVII. fig. 10, ascus with sporidia, x 325. 
Sporidia uniseriate or biseriate, yellowish, oblong, obtuse or somewhat pointed at 
the ends, constricted in the middle, 0:0007 inch long. Erumpent, but furnished 
with a subieulum ; endochrome bipartite. ae 

226. S. TRISTIS, Tode ; Fr. S. M. ii. p.444. Ta». LVII. fig. 11, sporidia, X ww. Sporidia 
biseriate, linear, acuminate, usually with four nuclei, but sometimes with only two 
large ones, producing a spurious uniseptate appearance, colourless, 0:0003 to 0.0004 
inch long. 

227. 8. POENE D. R. and M.; Fl. Alg. t. xxvi. f. 2. TaB. LVII. fig. 12, am 
x 450. Sporidia triseptate, colourless, or almost so, at each end, clear ee th 
middle, with or without nuclei, 00014 to 00016 inch long. This plant is the 


S. tristis B of the * English Flora.’ 


Div. 18. VILLOSÆ. | ; 
228. S. scABRA, n. s. TAB. LVII. fig. 13, sporidia, x 325. Sporidia — | 
constricted in the middle, with many (usually 6) nuclei, t gts ota On 
inch long. Perithecia very hairy, seated on à dense subi deir B. and Br. 
furze, Weybridge, Oct. 1857. The fruit is like that of S. — sr N 
229. S. CANESCENS, Pers.; Fr. S. M. ii. p.448. Tas. LVII. fig. 14, 


: p à mes more pointed at one 
Sporidia colourless, straight or curved, subacuminate, T s aga m os 


end than at the other, 0-0011 to 0-0014 inch long. See | 
plant. . 15, ridia, X 220. 
230. S. Racoprum, Pers.; Fr. S. M. ii. p. 449. Tas. Pe SINE not distin- 
Sporidia biseriate, in the early state (as m the brown, and 7-septate, 0:002 
guishable from the fruit of S. hirsuta, eventually pale 9T9 


316 MR. F. CURREY ON THE FRUCTIFICATION OF SIMPLE SPHÆRIÆ. 


to 0:0026 inch long. I suspect that the sporidia of S. hirsuta, and perhaps also those 
of Spheria ovina and Sphæria canescens, are multiseptate when perfect. 

231. S. ovina, Pers.; Fr. S. M. ii. p.446. Fructification not distinct from that of $ hip. 
suta (post). I have found, however, on one or two occasions specimens which I be. 
lieve to be S. ovina, in which the sporidia exhibit under the microscope a peculiar 
pink tinge, each sporidium being also furnished with a clear, round, colourless spot. 
Tas. LVII. fig. 16, represents the fruit of this form x 325 diams. 

232. S. CJESIA, Carm. Ta». LVII. fig. 17, ascus with sporidia highly magnified, Sporidia 
(? always) uniseriate, colourless, elliptical, or subpyriform, about 0-0002 inch long. 
Perithecia white, very hairy. This specimen is marked “S. ovina statu juniori;” but 
whoever so named it, had clearly not compared the fruit of the two species. 

233. S. HIRSUTA, Fr. S. M. ii. p. 449. Tas. LVIL fig. 18, ascus with sporidia, x 220, 
Sporidia biseriate or crowded, pale brown, long, flexuous, 0:002 to 0:0026 inch long. 
See remarks under §. Racodium (supra). 

234. S. RHODOCHLORA, Mont. Syll. p.227. Tas. LVII. fig. 19, ascus with sporidia, x 395. 
Sporidia crowded, colourless, broadly elliptical, with a reticulated appearance, but 
whether from the spores being multicellular, or from a number of nuclei touching 
one another, I cannot say; 0-0005 to 0:0007 inch long. 

235. S. MUTABILIS, Sz. Tas. LVII. fig. 20, ascus with sporidia, x 325. Sporidia bise- 
riate, colourless, curved, acuminate, endochrome 4-partite, 0°0008 inch long. Very 
like S. pulvis-pyrius to the naked eye, but under a power of 50 diams. the perithecia 
exhibit a very slight brownish hairiness, principally at the base. — | 

236. S. PILOSA, Pers.; Fr. S. M. ii. p. 440. Tas. LVII. fig. 21, ascus with sporidia, X 
about 325. Sporidia uniseriate, colourless, elliptical, uniseptate, or with the endo- 
chrome bipartite, 0:0003 to 00004 inch long. A small Sphæria, like S. pulvis- 
pyrius, but decidedly hairy. 

237. S. (CERATOSTOMA *) CHIONEA, Fr. S. M. ii. p. 446; EL ii. p.92. Tas. LVII. fig. 22, 
ees X 925. Sporidia elliptical or subglobose, dark brown, 0:0008 to 00005 inch 
ong. | oo) 

238. S. STRIGOSA, A. and S.; Fr. S. M. ii p.448. In my opinion not distinguishable — 
from S. canescens, Pers. Fries Says, “ Å præcedente ” (that is S. canescens) “non 
facile dignoscitur." 

259. 8. Brassrom, KL; Eng. FL p. 261. “Tan. LVII. fig. 28, asus with sporidia, x 290 
Sporidia uniseriate, at first colourless, then pale brown, ultimately dark opaque — 
IM rei. elliptical, with pointed ends, or lozenge-shaped, 00014 to 0-002 inch long. — 
On Brassica oleracea. Perithecia large, hairy at the bottom, hairs white or brown; 
sporidia sometimes with two large nuclei, sometimes with a dark line not extending 
quite across the sporidium. | | 

ma. TVSVEULOG Berks Lak TAURO TE sporidia, x 325. Sporidia irregular ™ — 
shape, multiseptate, ? multicellular, very dark brown, varying much in size, from — 

T Crratostoma, Fr. Obs. 


Nucleus gelatinous, asci soon di 


ssolving (in some speci | : i mpent, surroun 
the apex of the peritheeium. pecies not yet discovered) ; spores simple, erump | 


Perithecium membranaceous, soft ; ostiolum subulato-rostrate, with å penicillate b ig 5 


MR. F. CURREY ON THE FRUCTIFICATION OF SIMPLE SPHÆRLÆ. 317 


0:0010 to 00014 inch long. Perithecia roundish, rather pulvinate, very hairy, but 
the hairs are very short. On wood from Swan River. 

241. S. SUPERFICIALIS, n. s. Tas. LVII. fig. 25, asci with sporidia, x 420. Sporidia 
uniseriate, overlapping, elliptical, or subturbinate, rarely slightly curved, colourless, 
2-nucleate, 0-0003 to 0:0004 inch long. Perithecia hairy, subglobose, very small, 
seated on a hairy subiculum. Like some other Sphæriæ, combining the charac- 
teristics of the Villose and Byssisede. 

242. S. CAPILLIFERA, n.s. Tas. LVII. fig. 26, ascus with sporidia, X 325. Sporidia 
| uniseriate, slightly overlapping, rather dark brown, 1-2 nucleate, broadly elliptical, 
0:0003 to 00004 inch long. Perithecia globose, clothed with very short, rather stiff 
black hairs, seated on a pale subiculum, and furnished with a mammillate ostiolum. 
Quere, if distinct from S. crinita, Pers. 


Div. 19. DENUDATZ. 


243. S. INSPERSA, Berk. Tas. LVII. fig. 27, ascus with sporidia, x 325. Sporidia mostly 
(? always) uniseriate, colourless, subhyaline, elliptico-acuminate, 00005 to 00006 inch 
long. On wood from Swan River. Not distinguishable from S. pulvis-pyrius, except 
by the sporidia. 

244. S. RHODOMPHALOs, Berk. Ta». LVII. fig. 28, ascus with sporidia, and free sporidia, 
x 325. Sporidia biseriate, colourless or greenish, curved, subfusiform, uniseptate 
or with a bipartite endochrome, 00007 to 0:0008 inch long. On wood. Perithecia 
globose, with a distinct red circle round the ostiolum, which is frequently slightly 
depressed. Sporidia sometimes slightly constricted at the division, and with a 
nucleus in each partition, as in the right-hand figures. -— 

245. S. Bomparpa, Batsch; Fr. S. M. ii. p. 456. Tas. LVII. fig. 29, sporidia, x 325. 
Sporidia crowded, colourless, intertwined, 0-0016 to 0:0020 inch iuit imd. vor 
variable, frequently, if not usually, with a division in the middle. qr eg 
ing much in the length of the ostiolum. I have seen once or twice a bead- er 
pearance in the sporidia, arising from the breaking-up of the endochrome " ivi- 
sions. The fruit of this plant should be particularly observed, - sed 
sometimes be multiseptate, as in S. Racodium, corticis, &e. in 30 i 

246. S. morrrormis, Tode; Fr. S. M. ii. p. 458; mu S — - etui 
with sporidia, and free sporidia, x 325. Sporidia crowded, u. nlar 
with a greenish tinge, linear, but slightly curved; endochrome usually grann ar 
sometimes nucleate; asci usually broad as in the figure, but 
elongated. des 202 Kinn 

247. en n.s. Tar. LVII. fig. 31, ascus with es ie, 
colourless, or greenish, subelliptical, but slightly — ; t in the sporidia. 
endochrome bipartite. On wood. Very like S. pulvis-pyrius, excep 


Bunga , Mr. Stock. ae e | å a 

248. 8. seid. Pers. ; Fr. S. M. ii. p. 458. Ta. LVII. fig. x oam = RPR 
and a free sporidium, X 325. Sporidia straight, er Mq c the * Disoductiol 
slightly constricted at the septa, pale brown, 00005 inch long. | 


318 MR. F. CURREY ON THE FRUCTIFICATION OF SIMPLE SPHJERLE. 


to Oryptogamic Botany,’ p. 281, Mr. Berkeley figures the sporidia of S. pulvis-pyrius 
with as many as five septa, and of a more elongated form than in my figure. In all 
the specimens (and they are very numerous) which I have examined, I have never 
found more than three septa, nor have I ever seen the sporidia so elongated as in — 
Mr. Berkeley’s figure. Mr. Berkeley himself mentions the sporidia as being trisep- 
tate in ‘ Ann. and Mag. of Nat. Hist.’ ser. 2. vol. vii. p. 189. 

249. S. CONGLOBATA, Fr. S. M. ii. p. 444. Tas. LVII. fig. 33, ascus with sporidia, and free 
sporidia, x 325. The specimens of this Spheria in Hook. herb. show clearly that 
this species is only a crowded, subcuticular, erumpent form of S. pulvis-pyrius. Some 
of the perithecia are bursting transversely, some longitudinally, and others form 
cæspitose masses; others again have the ordinary scattered habit of S. pulvis-pyrius. 
They form a very instructive series of specimens. 

250. S. DIOICA, Fr. S. M. The specimens of this plant in Hook. herb. show clearly that 
(like S. conglobata) this species is only a subcuticular form of S. pulvis-pyrius. 

251. S. MOROIDES. n.s. Tab. LVII. fig. 34, ascus with sporidia, and free sporidia, x 325. 
Sporidia biseriate, greenish brown at first, eventually brown, subhyaline, elliptical, | 
0:0004 to 00005 inch long. Perithecia rugose, small, like very small specimens of 
S. moriformis, from which species the present differs altogether in the nature of its — 
sporidia. 

252. S. PLATEATA, Pers. in litt. Tas. LVII. fig. 35, ascus with sporidia, x 325. Sporidia 
crowded, yellowish brown, multipartite, subelliptieal, 00008 to 0:0010 inch long. 
On wood. Very like S. pulvis-pyrius, except in the sporidia—possibly only a small - 
form of S. Spartii, Nees, which latter does not differ essentially from S. elongata, Fr. 

253. S. SPERMOIDES, Hoffm.; Fr. S. M. ii. p.457. Tas. LVII. fig. 36, ascus with sporidia, 
and free sporidia, x 325. Sporidia biseriate, colourless, curved, endochrome some- 
times bipartite, 0-0008 inch long. 

254. S. MAMMÆFORMIS, Pers.; Fr.S. M. ii. p. 455. Tan. LVIL. fig. 87, sporidia, x 450. 
Sporidia dark clear brown, subcymbiform, varying much in size, from 0:0008 to 
00016 inch long. e 

255. S. srERCORARIA, Sow.; Fr. S. M. ii. p. 455. Tan. LVIL fig. 38, ascus with sporidia, 
X 220. Sporidia uniseriate, brown, eventually quite opaque, at first colourless, ellip- — 


tical, or almond-shaped, 0-0016 to 0-002 inch lono. B 


256. S. STERCORARIA, Sow. var.? I subjoin here a description of a Sphæria which I ha 
. found growing on horse-dung, and which is probably the same species as the last, 
notwithstanding its smaller sporidia and curious ostiola.  Perithecia solitary, or few 
together; ostiola formed of a number of processes arranged in a penicillate manner, 
each consisting of a single row of irregularly-shaped cells, the upper cell being 
pointed. Sporidia mostly uniseriate, but sometimes biseriate, greenish at first, then 
darker, probably eventually black, elliptical, 0-0010 inch long. Tas. LVII. fig. 39 (a) 2 
represents an ostiolum broken off, x 325 diameters, and fig. 39 (5), an ascus vid 
sporidia similarly magnified. | 
257. S. STERCORARIA, Fr. S. M. ii. p. 455; S. stercoris, El. ii. p.104. Ta». LVIL fig 40 
sporidia, x 325. Sporidia biseriate, dark opaque rich brown, consisting of four 


MR. F. CURREY ON THE FRUCTIFICATION OF SIMPLE SPHJERLE. 319 


joints (? at first continuous), Which frequently separate when the sporidia escape 
from the ascus, 0:0018 inch long. 

258. 8. BIFORMIS, Sz., non Fries. Ta». LVII. fig. 41, fruit, x 325. I could find no asci. 
The perithecia are filled with balloon-shaped bodies, with granular contents, colourless, 
and of irregular size. There is a reddish tinge on the upper part of many of the 
perithecia, as in S. rhodomphalos, Berk. If the plant be a Sphæria, it would belong 
to the present division, but I am doubtful as to the existence of asci. 

. 259. S. VERRUCOSA, Grev.; Fr. Index Alphabeticus=8. moriformis, Tode. 

260. S. PULVERACEA, Ehrh.; Fr. S. M. ii. p. 459. Tas. LVII. fig. 42, ascus with sporidia, 
x 325. Sporidia uniseriate, clear dark brown, elliptical, subglobose, or subturbinate, 
0:0008 to 0-0004 inch long. 

261. S. SORDARIA, Fr. S. M.ii.458; EL ii. p.94 Ta». LVII. fig. 43, ascus with sporidia, 
x 325. Sporidia uniseriate, dark brown, elliptical, 0:0006 to 0-0007 inch long. 

262. S. OBDUCENS, Schum. =S. plateata, Pers. supra. 

263. S. POMIFORMIS, P.= 5. pulois-pyrius. 

264. S. (NECTRIA) Przıza, Tode; Fr. S. M. ii. p. 452; El. ii. p. 92. Ta». LVII. fig. 44, 
sporidia, x 450. Sporidia elliptical, uniseptate (or with the endochrome bipartite), 
colourless, 0-0004 to 0:0005 inch long. Perithecia fawn-coloured or brown. 

265. S. (NECTRIA) SANGUINEA. Tas. LVIL fig. 45, ascus with sporidia, x 395. Sporidia 
elliptical, colourless, uniseriate or overlapping, uniseptate, 0-0004 to 0-0005 inch 
long. Fig. 46 represents sporidia of the same species, X 420, at a more advanced 
period, after the asci have become dissolved. 

266. S. (NECTRIA) EPISPHÆRIA, Tode; Fr. S. M. ii. p. 454; El. ii. p. 93. Tas. LVII. fig. 47, 
sporidia, x highly. Sporidia uniseriate, or overlapping, colourless, elliptieal, — 
nate or round at the ends; endochrome bipartite, or with two nuclei, 00002 to 
0:0004 inch long. ré | 

267. S. RUBICOLA, n.s. Tam. LVIL fig. 48, sporidia, x 420. Sporidia biseriate, colourless, 
subeymbiform, or narrowly almond-shaped, with 2 or 4 nuclei, sometimes with the 
endochrome divided into two parts as in the lower figure, 0-0006 to ee 
Perithecia small, globose, or subglobose, with a mammillate ostiolum 4 — 
with a strong lens, they are seen to be closely surrounded å dd | 

i : age ithecia, and similar hairs cover the bark 
hairs, sometimes springing from íhe perithecia, am i ber 1857. Almost 
Where no perithecia are visible. On bramble, Wey bridge, Octo bel N to the 
as nearly allied to the Villose as to the Denudate, but ape PRES ae 


latter by preference. | | 4 Ta 

268. 8. (Gnana *) Vaconen, Som. t. 878. £ 1; gn eR en 
LVIII. fig. 49, ascus with sporidia, X 325. Sporidia UE adit ord in 
colourless, but yellow in a mass, almost almond- haped, femme ly Hook. herb. 
the middle, uniseptate, 0-0006 inch long, or slightly less. — to the Denudate. It 
with the Cæspitosæ, but quere, if not more properly belonging 
has quite the habit of S. moriformis. 

* Grppera, Fr. Perithecium ceraceo-corneous, free, — 
Which is at length ejected ; asei linear, fixed to the base of the perithecium. 


always closed, filled with a waxy nucleus, 


320 MR. F, CURREY ON THE FRUCTIFICATION OF SIMPLE SPHÆRIÆ. 


269, S. CAUDATA, n. s. Tas, LVIII. fig. 50, ascus with sporidia, x 220. Sporidia bi- 
seriate, consisting of a pointed subelliptical brown head, and an elongated colourless 
tail; length of the sporidia (including the tail) 0:002 inch, of the head alone 0:0008 
inch. Perithecia small, scattered, or few together, conical or subglobose, with a 
conical ostiolum. On rotten wood, near Twyeross, Leicestershire. This Sphæria, 
communicated to me by Mr. Bloxam, is very peculiar in the form of its sporidia. 
The perithecia are sometimes naked, sometimes almost buried in the soft rotten 
wood, so that I have had great doubt as to its proper division. Its sporidia resemble 
those of Podospora fimicola, Ces., figured in * Hedwigia,’ tab 14. fig. A. 

270. S. COLLABENS, n. s. Tas. LVIII. fig. 51, sporidia, x 325. Sporidia biseriate, fusi- 
form, swollen or constricted in the middle, with several nuclei, colourless, 0:0014 to 
0:0016 inch long. Perithecia subglobose, with an impressed ostiolum, but the osti- 
olum is often furrowed, or rimose. The perithecia and sporidia agree very nearly 
with those of S. macrotricha, B. & Br., but the perithecia have no hairs. Its habitat 
is wood, whilst that of S. macrotricha is dried leaves and beech mast. The sporidia 
also resemble those of S. scabra; but, besides being smooth, the perithecia of S. col- 
labens are four times the size of those of S. scabra. i 

271. S. Curreyn, Blox. MSS. I have a Spheria from Mr. Bloxam under this name, 
which is possibly the same as the plant last described. In Mr. Bloxam's plant, how- 
ever, the sporidia rarely, if ever, exceed 0:0010 inch in length, and the contents of 
the perithecia have a rose-red tinge. In both this and the Sphæria last described, 
the perithecia with rimose ostiola might easily be mistaken for those of a Hysterium. 

272. S. PULVISCULA, n. s. Tas. LVIII. fig. 52, ascus with sporidia, and free sporidia, 
X 325. Sporidia biseriate, curved or cymbiform, colourless or greenish, when perfect 
with three septa, or at least with the endochrome divided into four portions, giving 
an appearance of three septa; sometimes the endochrome is only once divided, and 
in a young state the sporidia are continuous. Length of the sporidia variable, from 
0:0008 to 0:0012 inch. Perithecia very small, black, rather shining, conical or sub- 
globose, crowded or scattered, with a minute mammillate ostiolum. This species is 


not distinguishable by the naked eye from S. pulvis-pyrius, but its fruit, as will be 
seen, is quite different. 


Div. 20. PERTUSÆ. 


273. S. PERTUSA, Pers.; Fr. S. M. i. p.464. Tap. LVIII. fig. 53, sporidia, x 220. Spo- 
ridia biseriate, I believe; brown, usually 4-septate, broad and curved, 0:0016 to 
0:002 inch long. The specimens are not in good condition. 

274. S. PICASTRA, Fr. S. M. ii. p. 463. Ta». LVIII. fig. 54, sporidia, x 220. Sporidia 
dark brown, multicellular, very irregular in size and shape. I suspect that this plant 
ought to be placed in the genus Hendersonia*. Some of the sporidia were elongated 
as into a foot-stalk. There was no trace of asci. 


* * * i 
* nn Berk. Perithecium carbonaceous, subinnate, emergent, almost mouthless, bursting with a pore 
ü ; m 
an arly. Nucleus, compact, then diffluent ; sporidia erect, elongate, pedicellate, multiseptate. 
IPLODIA, Fr. Sporidia clavate, uniseptate ; otherwise as Hendersonia. 


MR. F. CURREY ON THE FRUCTIFICATION OF SIMPLE SPHÆRIÆ. 321 


275. S. (DIPLODIA) LECYTHEA, Schwein.; Fr. S. M. ii. 460. Ta». LVIII. fig. 55, fruit, 
x 220. Spores colourless or greenish, elliptical, 0:0010 to 0:0014 inch long. Re- 
sembling externally the Sphæriæ of this division; but I think it is a Diplodia, although 
there is not any septum in the sporidia, 

276. S. MERDARIA, Fr. El. ii.100. Ta». LVIII. fig. 56, ascus with sporidia, x 420. Spo- 
ridia uniseriate, dark black-brown, elliptical or subglobose, 00008 to 0:0004 inch 
long. Arranged with the Pertusæ at Kew, but placed by Fries in the Obtectæ. 
I should rather consider it as belonging to the Denudate, judging from the speci- 
mens in the Hookerian herbarium, which are authentic. The perithecia are very 
small. | | 

277. S. OLEARUM, Cast. Tas. LVIII. fig. 57, ascus with sporidia, X 325. Sporidia uni- 
seriate or biseriate, rather pale brown, elliptical, 5-septate, with a nucleus in each 
septum, 0:0010 to 0:0012 inch long. Peritheeia large, mammillate, subinnate. 
Apparently belonging to the Pertuse. 

278. S. ULMICOLA, n.s. Ta». LVIII. fig. 58, sporidia, x 450. Sporidia oblong, brown, 
uniseptate, 0-0009 inch long, slightly constricted in the middle. I could find no asci; 
but, there being no trace of sporophores, the asci had probably become dissolved : if 
ascigerous, the plant is certainly a Sphæria, belonging to this division. Perithecia 
large, globose, more than half-buried in the wood, furnished with a mammillate 
ostiolum, which drops off, leaying a large round aperture. 

279. S. micrasprs, Berk. Tas. LVIII. fig. 59, represents the fruit of this plant, which 
is probably not a Spheria, but a Pertusaria. n 

280. S. PUTAMINUM, Schwein.; Fr.S. M.461. Ta». LVIII. figs. 60 & 61, sporidia, x 220. 
Sporidia uniseriate or crowded (I could not determine whether there were mom 
than four in any of the asci), dark brown, 0-004 to 0-005 inch long. Tam uncertain 
whether the bodies in fig. 60 are sporidia with gelatinous envelopes, or asci with a 
single sporidium. À 

281. 8. Re n.s. Ta». LVIII. fig. 62, ascus with sporidia and free toig 
X 325. Sporidia biseriate, very broadly fusiform, 00024 to 0:0030 inch net : ad 
slightly constricted in the middle, with a median septum, and from one to tare 

idium: colour of the sporidia 

other septa close together at each end of the spor 4 hyaline tip at each 

greenish, becoming brown in age; the sporidia have usually a på a ag 
end. Perithecia large, almost globose, with a small mammilla 


Lie , Kid- 
piece of old broken paling, the surface of which was q AP Oh imm 
brooke, Blackheath, March 12, 1859. Quære if distinct from . putaminus 


Div. 21. PLATYSTOME. 
282. S. BARBARA, Fr. S. M. ii. 468. Tas. LVIII. fig. 65, ard elg a 
Sporidia linear, packed side by side along the vun nd | a Fructification 
‚great length, but probably breaking up into short n i 
. resembling that of many plants in the division or ee ^d 4 sporidia, x 425. 
283. S. MACROSTOMA, Tode; Fr. S. M. ii. 469. TAB. LVIII. fig. 04, vår 


VOL. XXII. 


322 


284. 


285. 


286. 


287. 


MR. F. CURREY ON THE FRUCTIFICATION OF SIMPLE SPHÆRLÆ. 


Sporidia biseriate (? sometimes uniseriate), yellow at first, eventually brown, 5. 
septate, rarely with 6 or more septa, frequently with longitudinal divisions, rendering 
the sporidia multicellular, 0*0010 to 0:0012 inch long. I have a plant from Mr. 
Bloxam marked S. macrostoma, Tode, the fruit of which is drawn in fig. 65, x 220 
diameters. These sporidia attain a length of from 0:0020 to 0:0026 inch. 

S. LIGNIARIA, Grev. Sc. Cr. Fl. tt. 82. Tas. LVIII. fig. 66, ascus with sporidia, 
X 425. Sporidia uniseriate, at first pale brown, then dark brown, elliptico-acuminate, 
0:0005 inch long. Perithecia somewhat flask-shaped, with a rugose, or rather velvety 
appearance, but not at all shining. 

S. CRISTATA, Pers. = S. crenata, Fr. S. M. ii. 469. Tas. LVIII. fig. 67,sporidia, x 395. 
Sporidia almond-shaped, brown, 5-7-septäte, 0:0016 to 0-0018 inch long. Quiere if 
distinct from S. macrostoma, Tode. 


Div. 29. CERATOSTOM X. 


S. BREVIROSTRIS, Fr. S. M. ii. 474. Tas. LVIII. fig. 68, ascus with sporidia, x 325. 
Sporidia biseriate or uniseriate, pale brown, 0:0006 to 0:0007 inch long, elliptical, 
rather pointed at the ends. i 

S. LONGISPORA, n. s. Ta». LVIII. fig. 69, ascus with sporidia and free sporidia, 
(a) x 325, (b) x 425. Sporidia filiform, arranged side by side, very variable in length 
and in the number of septa, yellowish brown. Perithecia flattened, with a short 


_ ostiolum, mostly covered by the bark, excepting the tip of the ostiolum. This plant 


288. 


289. 


290, 


291. 


292. 


is marked in the herbarium « S. rostellata," but it differs from that species in the 
ostiolum. 

S. (OvrisPORA) MICULA, Fr. El, ii. 101. Taz. LVIII. fig. 70, fruit, x 325. No asci; 
stylospores variable in length, colourless, fusiform, very pointed; ostiola fuscous. 
An imperfect state of some Spheria. 

S. (SPHÆROPSIS ?) PILIFERA, Fr. S. M. ii. 472. Not ascigerous, the perithecia pro- 
ducing only a mass of minute colourless spermatia. Remarkable for its beautifully 
fine, hair-like ostiolum. Perithecia small and globose. 
S. CIRRHOSA, MS. Tas. LVIIT. fig. 71, ascus with sporidia, x 425. Sporidia bi- 
seriate, elliptical, colourless, 0-0004, to 00005 inch long. Perithecia ovate or turbi- 
nate, quite buried in the wood, but piercing the surface by their very long, almost 
filiform ostiola. There is another plant marked S. cirrhosa, the fruit of which 1s 
shown in fig. 72, x 325 diameters. Here the sporidia are uniseriate or biseriate, pale 
greenish brown, elliptical, frequently with one, two, or more nuclei, 0:0008 to 
00004 inch long. Perithecia subovate, ostiola about as long as the perithecia. 

S. BOSTRATA, Fr. S. M. ii. 499. Sporidin Bitte, elliptical or slightly curved 


0*0003 to 0-0004 inch long, colourless, or with a tendency to pale brown. Probably 
not distinct from S. cirrhosa, supra. 


Div. 23. OBTECTÆ. 


S. BUTTPA, Fr. S. M. ii. 478. Man. EVIIL. fig. 73, ascus with sporidia and free — 


MR. F. CURREY ON THE FRUCTIFICATION OF SIMPLE SPHARLA. 323 


sporidia, x about 450. Sporidia biseriate, colourless, slightly curved, 0:0002 to 
0:0003 inch long, much resembling those of §, stigma. — 

293. S. VIBRATILIS, Fr. S. M. ii. 482. Authentic specimens of this Sphæria exist in the 
Hookerian herbarium, but the fruit is not ripe. As far as I could make out, the 
sporidia appear to differ very little from those of S. stellulata. There is another 
Sphæria in the collection marked by Mr. Berkeley “ 8. vibratitis, Fr.” in which the 
fruit is very different. Ta». LVIIL fig. 74, represents the fruit of the latter plant, 
X 325, of which the following is a description :—Sporidia uniseriate or overlapping, 
pale brown, oblong-elliptic, but slightly curved, endochrome bipartite, 0°0005 to 
0:0006 inch long. 

294. S. OPERCULATA, A. & S.; Fr.S. M.ii.479. Tan. LVIII. fig. 75, ascus with sporidia, 
X 925. Sporidia biseriate, brown, curved, 00005 inch long. 

295. S. (HALONIA) CUBICULARIS *, Fr. S. M. ii. 477; El. ii. 97. Ta». LVIIL fig. 76, 
sporidia, x 325. Sporidia uniseriate, dark brown, elliptieal, often acuminate at the 
ends, 0-0006 to 0:0007 inch long. Easily known by the ostiola being surrounded by 
a white tubercle formed from the wood. The sporidia, when free, are often surrounded 
by a narrow border. I have a specimen from Dr. Montagne in which the sporidia 
reach 00010 inch in length. It will be observed that the sporidia differ entirely 
from the generic characteristies given by Fries. LE ee 

296. S. LrvmDA, Fr. S. M. ii. 479. Tas. LVIIL fig. 77, sporidia, x 825. Sporidia uni- 
seriate, brown or yellowish-brown, normally I think triseptate, but frequently with 
longitudinal partial septa, and sometimes (from the breaking up of the endochrome) 
having a cellular appearance, elliptical, 0-0005 to 0:0007 inch se Ne 

297. S. (SPHÆROPSIS) PRUINOSA, Fr. S. M. ii. 486. Not ascigerous, the A 7 
dueing only a quantity of minute eurved spermatia, colourless ve eas d 
brown in the mass. There are two specimens in the herbarium, one from 
Suec. without fruit, and the other containing the spermatia, just mentioned. 

k ji 489. Ta». LVIII. fig. 78, fruit, x 325. 

298. S. (SpHaropsis ?) OLES, D.C.; Fr.8.M.ii.489. Tas. UE cda enia, 
There appear to be no asci ; stylospores cylindrical, enm adei. | 
0:0008 inch long, sometimes with a border, and sometimes A ^ AB. LVIII, fig. 79, 

299. S. (Hencosrora) RHODOSTOMA t, A. & 8.; Fr. 8. M. ii. DH with constrictions at 
sporidia, x 325. Sporidia dark brown when perfect, — of the sporidia 
the septa, 0-0009 to 0-001 inch long. In the Kew specimens many 


are very small and uniseptate. oe | i long and narrow ; 

300. S. CErasarum, MS. Ta». LVIII. fig. 80, es ita ie slightly con- 
sporidia biseriate, rather dark brown, with the = "t inch long. The sporidia are 
stricted in the middle, acuminate at each end, 0 0009 


* Harowia, Fr. Perithecium entire, covered, membranaceous ; Mer A pet. 
coloured heterogeneous disk ; nucleus gelatinous ; asci delicate ; spores sé ar open, 
t Hercospora, Fr. Perithecium discoloured, subcarbonaceous, ver mixed with paraphyses; spores dark- 
by the bark, breaking through a heterogeneous tubercle; asci € 
coloured, opaque, septate. 2v2 


324 MR. F. CURREY ON THE FRUCTIFICATION OF SIMPLE SPHÆRIZÆ, 


— drawn from the specimen marked “a” in the herbarium, the other marked * $$. Ce- 
rasarum, b," not being in good condition. : 

301. S. TAMARISCINIS, Grev. Tas. LVIII. fig. 81, ascus with sporidia, x 325. Sporidia 
biseriate, rather dark brown, triseptate (or ? the endochrome 4-partite), usually slightly . 
eurved, 0:0008 inch long. Occasionally the sporidia have four septa, and rarely none. 
There is another plant in the herbarium, marked by Mr. Berkeley $.- Zamariscinis, 
Grev., the perithecia of which contain no asci, but a multitude of minute, elliptical, 
turbinate or irregular yellowish spermatia, varying from 0°0005 to 0:0003 inch in 
length. It may be, and probably is, an imperfect state of the true S. Tamariscinis. 

302. S. PrNAsTRI, Dec.; Fr. S. M. ii. p.488. Ta». LVIII. fig. 82, ascus with sporidia, 
x 325. Sporidia crowded, colourless, elliptical, often acuminate at the ends, 0:0003 
to 0:0004 inch long. 

303. S. CLYPEATA, Nees; Fr. S. M. ii. p.487. Ta». LVIII. fig. 83, sporidia, x 325, 
Sporidia uniseriate, rather dark brown, subelliptical or slightly curved, continuous, 
or with the endochrome divided into two, three, four, or even five portions, 0:0005 to 
00008 inch long. Easily known by the rounded shining shield-like patches of the 
blaekened epidermis which cover the perithecia. The sporidia are sometimes a good 
deal broader than those in the figure, and slightly constricted at the septa. 

304. S. XYLOSTEI, Pers.; Fr. S. M. ii. p.487; El.ii.99. Tap. LVIII. fig. 84, sporidia, 
x 325. Sporidia uniseriate, dark brown, bordered, elliptical, 00006 to 0:0007 inch 
long. In many of the sporidia I observed a faint indication of a septum in the 
middle, but I could not satisfy myself that it really existed, or that the endochrome 
was truly bipartite. | 

305. S. (HENDERSONTA) MAMMILLANA, Fr. S. M. ii. p.487. Ta». LVIII. fig. 85, sporidia 
on stylospores, x 325. Sporidia not contained in asci, but borne upon pedicels, 
bringing the plant within the genus Hendersonia, brown, triseptate, or with the 
endochrome 4-partite, irregular in shape and length, from 0-0005 to 0:0007 inch long. 
The sporidia are almost exactly the same as in Sphæria pulvis-pyrius. 

306. S. (HENDERSONIA) HIRTA, Fr. S. M. ii. p.483. Fruit not contained in asci, and 
differing In no respect from the fruit of S. mammillana, except perhaps in being 
slightly smaller. Slight variations in size, however, form no material distinction. 
In the ‘Syst. Myc.’ it is noted as a Cytispora, but would now be ranked with Hen- 
dersonia. | 

307. S. SEMI-IMMERSA, Grev. This plant does not differ in fructification from 8. Æylostei, 
Pers,, nor, as far as I can see, is it capable of being distinguished from the latter 
species. | 

308. s. OCELLATA, Fr. S. M. ii. p.480. Ta». LVIII. fig. 86, sporidia, x highly. Spo- 
ridia biseriate, colourless, slightly curved, rounded or acute at the ends, 0°0004 inch 

long. This specimen is marked ß, as if it were a variety, but I do not find any 
variety noticed in the * Syst. Myc.’ 

309. S, ToxICUM, Lev. Ann. d. Sc. Nat. 1848. Tap. LVIII. fig. 87, sporidia, X 450. 
Sporidia uniseriate and overlapping, or biseriate, dark (almost opaque) brown, sub- 
cymbiform, frequently with two large nuclei, 0-0004 to 0:0006 inch long. 


MR. F. CURREY ON THE FRUCTIFICATION OF SIMPLE SPHÆRLÆ. 325 


310. S. CONFORMIS, B. and Br. Ann. and Mag. Nat. Hist. ser. 2. vol. ix. p. 325. pl. 11. fig. 19. 
Tas. LVIII. fig. 88, ascus with sporidia, x 220. Sporidia biseriate, elliptical, often 
slightly curved, colourless, pellucid, endochrome 4-partite, 00007 to 0:0008 inch 
long. There are not, I think, any real septa. 

311. S. (HaLoNtA) DITOPA, Fr. S. M. ii. 381. Tas. LVIII. fig. 89, asci with sporidia and free 
sporidia, x 450. Sporidia crowded, very numerous, colourless, oblong, narrow, rounded 
or somewhat pointed at the ends, 0:0006 to 0:0007 inch long. Fries, in the * Summa 
Veg. Scand.,’ describes the sporidia as septate. It is possible they may sometimes 
be so, but I have never seen them otherwise than continuous, with a nucleate or 
granular endochrome. They are figured so, moreover, by Messrs. Berkeley and 
Broome, in Ann. and Mag. N. H. ser. 2. vol. ix. pl. 10. fig. 15*. Messrs. Berkeley and 
Broome, however (1. c.), mention Dr. Roussel's specimens of S. ditopa as having uni- 
septate sporidia. | ; 

312. S. QUADRI-NUCLEATA, n.s. TAB. LVIII. fig. 90, sporidia, x 450. Sporidia biseriate, 
very closely packed, colourless, narrowly oblong, pointed or rounded at the ends, 
each sporidium with four nuclei, 0-0006 inch long. Perithecia small, subglobose, with 
a mammillate ostiolum, which pierces the outer bark, making a circular hole or a 
rimose transverse fissure in the bark. On a stick with S. pulvis-pyrius, Weybridge. 
Surrey, September 7th, 1856. 

313. S. RUBI, n.s. Tas. LVIII. fig. 91, (a) asci with sporidia not quite ripe, X 325; 
(5) ripe sporidia, x 450. Sporidia biseriate, colourless, each with four large nuclei, 
subfusiform, but wide in the centre, with the sides flexuous, and mostly elongated 
at each end into a hyaline mucronate appendage. Perithecia very small, posco, 
just penetrating the bark with their minute ostiola. On bramble, Weybridge, 
Surrey, September 12th, 1856. ibo cm 

314. S. rxQuiLINA, Wallr.; Fr. EL ii. p. 100. Tas. LVIII. fig. 92, sporidia highly mag- 
nified. Sporidia biseriate, colourless, subfusiform, constricted in the middle, "i 
cleate, 0-0004 to 0-0005 inch long. On dead stems of Smyrnium Olusatrum. vg 
near Lewes, August 24th, 1858. I have no doubt about the species, eam 
contents of the perithecia in these specimens is not black, but colourless. Å té 
examined under a lens, without extracting the contents, they sii black; but ı 
picked out on the point of a needle, they will be found to be colour: d AR 

315. S. acus, Blox. MS., n.s. Tas. LVIIL. fig. 93, sporidia highly magnified. pori 

Fes : indri ith rounded ends, or acu- 
biseriate or crowded, colourless, narrowly cylindrical, with r 1 dochrome 
minate at the ends and then almond-shaped, 0:0005 to 0:0004 er. re id 255 
2-4-partite. Perithecia small, subglobose, flattened, concealed by ep | 


Which is pierced by the sharp-pointed minute ostiola. ; L ix. p. 380. 
316. S. PuowaTosPORA, Berk. and Br. Ann. and Mag. Na, Be d dri Sporidia 
pl. 11. fig. 33. Tas. LVIII. fig. 94, ascus with per a a nucleus at each 

uniseriate, elliptical, colourless, 0-0003 to 00004 inch long, * 

extremity, as in the spores of the genus Phoma. .. 999. pl. 10. fig. 9. 
:317. S. ARGUS, Berk. and Br. Ann. and Mag. Nat. Hist. ss aped citis dan or vl 

Ta». LVIII. fig. 95, ascus with sporidia, X 220. Sporidia biseriato 


326 MR. F. CURREY ON THE FRUCTIFICATION OF SIMPLE SPHÆRIÆ. 


dark olive-green, divided into two unequal parts, in the smaller of which the endochrome 
is 3-partite, and in the larger 3- or 4-partite. Sporidia surrounded by a gelatinous 
envelope, which is not always visible in the ascus, especially where the sporidia touch 
one another. Length of sporidia 0:0020 to 0-0024 inch. I have ventured to place 
this plant in the Odfeete, contrary to the opinion of Messrs. Berkeley and Broome, 
who consider its division to be the Subtectæ. Very like S. lanciformis in the form 
and nature of its sporidia. 

318. 8. APICULATA, n. s. TAB. LVIII. fig. 96, (a) ascus with sporidia, x 325; (b) free 
sporidia, x 450. Sporidia uniseriate, olive-brown, straight or very slightly curved, 
biseptate, each furnished with a hyaline tip, which is shut off from the rest of the 
sporidium by one of the septa. The hyaline tip is frequently, if not generally, invi- 
sible in the ascus, owing to the overlapping of the ends of the sporidia. The sporidia 
are often nucleated; the endochrome is somewhat granular. Length of the sporidia 
00010 inch.  Perithecia large, subglobose, deeply buried in the wood, above the 
surface of which the rather wide, circular, somewhat gaping ostiolum just protrudes. 
On a dry, old (? deal) fence of the South-Western Railway, not far from the Wey- 
bridge Station, 1856 and 1857. A very curious plant both in habit and sporidia. 
The perithecia are completely and deeply buried in the wood, and are sometimes 
scattered, sometimes in circles or groups of as many as five together. The perithecia 
seem eventually to throw off the wood above the ostiola, leaving deep depressions in 
the surface of the wood. | 

319. 8. APPENDICULOSA, Berk. and Br. Ann. and Mag. Nat. Hist. ser. 2. vol. vii. p. 189. 
Tas. LVIII. fig. 97, sporidia, x 325. Sporidia uniseriate, overlapping, colourless, 
subfusiform, with a caudate appendage, 0:001 inch long with the appendage. 

320. S. SIPARIA, Berk. and Br. Ann. and Mag. Nat. Hist. ser. 2. vol. ix. p. 321. pl. 9. fig. 8. 
Ta». LVIIL fig. 98, ascus with sporidia, x 220. Sporidia biseriate, at first golden- 
yellow, eventually clear brown, 00020 to 0:0024 inch long, without measuring the 
outer gelatinous envelope in which they are enclosed, and which disappears as the 
sporidia advance in age. | 

321. S. (MASSARIA) AMBLYOSPORA *, Berk. and Br. 1. c. p.322. pl. 10. fig.10. Tas. LIX. 
fig. 99, ascus with sporidia and a free sporidium, x 220. 8 poridia biseriate, greenish- 
brown, eventually brown, biseptate, lageniform, the apicular cell rather lighter- — 
coloured than the others, 0:002 to 0-0024 inch long, without measuring the gelatinous — 
coat in which they are enveloped. In describing the fruit of this plant, I have — : 
adopted the name used by Messrs. Berkeley and Broome, who described it as å € 
species. Iam quite convinced, however, that S. amblyospora, Berk. and Broome, 18 
the Sphæria fædans of the * Syst. Myc.,’ the Massaria fædans of the * Summa ye 
Scand.,’ and the Splanchnonema of Corda, in Sturm’s * Deutschland's Flora,’ t. 54. 5 

322. 8. GIGASPORA, Desm. Tap, LIX. fig. 100, ascus with sporidia, x 220. Spor zi 
biseriate, oblong or almond-shaped, with the sides often incurved at the centre, dark 


* SE : a 
Massarıa, Notar. Perithecium subcarbonaceous ; ostiolum papillate, attenuated. Nucleus gelatinous, m 


with paraphyses and asci, which dehisce a besdi : ; ine, W 
d t the > in gelatine, 
form a dirty black stain, apex, ejecting dark septate spores immersed in ge 


MR. F. CURREY ON THE FRUCTIFICATION OF SIMPLE SPHJERLE. 327 


brown, triseptate, ör with the endochrome 4-partite, surrounded by å narrow gela- 
tinous border. Perithecia small, conical or somewhat flattened, sometimes only just 
piercing the bark, sometimes protruding considerably. Sporidia ejected in vast 
numbers, forming wide black stains round the ostiola, 0-0026 inch long. On dead 
branches of (I think) maple, Blackheath Park, 1855 and 1856. This plant is the 
type of the genus Saccotheciwm of Fries, the characters of which seem to me most 
unsatisfactory, and to have been founded upon imperfect microscopical observations. 

323. S. INQUINANS, Tode; Fr. 8. M. ii. p. 486. Tas. LIX. fig. 101, sporidia, x 220, 
Sporidia dark brown, with a granular and nucleated endochrome, very variable in 
size, sometimes reaching 0:0080 inch in length. 

324. S. BUFONIA, Berk. and Br. Ann. and Mag. Nat. Hist. ser. 2. vol. ix. p. 323. pl. 10. 
fig. 13. Tas. LIX. fig. 102, ascus with sporidia, x 220. Sporidia uniseriate, clear 
rich brown, eventually becoming very dark brown, oblong, constricted in the middle, 
surrounded by a broad gelatinous envelope, 0:0008 to 0-0011 inch long. Not un- 
common on dead branches of oak. One of the finest Spherie in point of fructifica- 
tion. The figure in the ‘ Annals’ conveys no idea of the beauty of the sporidia, being 
hardly more than an outline. 

325. S. FUSCELLA, Berk. and Br. l.c. p.325. pl. 11. fig. 20. Tan. LIX. fig. 108, ascus 
with sporidia, x 825. Sporidia uniseriate, colourless at first, eventually pale brown, 
oblong-elliptic, rarely very slightly curved, or incurved or constricted in the middle, 
0:0007 inch long ; endochrome 4-partite, giving a triseptate appearance. a 

326. S. Tırız, Pers. Syn. p.84; Fr. S. M. ii. p.485. Ta». LIX. fig. 104, sporidia, 
X 325. Sporidia biseriate, dark brown, lageniform, uniseptate, 00018 inch long, 
without measuring the gelatinous coat in which each sporidium is enveloped. — 
not quite certain whether this plant is the true S. Tilia of Persoon. It is Dot un- 
common, and cannot be mistaken for any other if attention be paid to its fruit, and 
to its growing upon lime. In fig: 105 and fig. 106 I have represented asci with T 
sporidia and free sporidia of a Sphæria on lime, communicated to à by n 
Berkeley as S. Tiliæ of Mougeot and Nestler. The sporidia of this gna ex "d 
seriate or biseriate, colourless, acuminato-elliptical, 00006 inch long, with bipart 
endochrome. : | ag 

327. S. ASHWELLIANA, n. s. TAB. LIX. fig. 107, ascus with sporidia, X - 7 

seis nyde ‚reenish, elliptical, but mostly pointed at each 
uniseriate or biseriate, colourless or greenish, d | 
. . . : 1-4-septate, hyaline, 0-0010 to 00014 
end and slightly constricted in the middle, p us 
: : bridge, Surrey, October 1857. Perithecia * 
inch long. On small branches of fir, Weybridge, lant 
ti d I have some doubt whether the p 
The specimens were in bad condition, an se me | 
Seni ut I know of no species in either of those 
belongs to the Obtecte or the Circinate ; but I know er uni di SÜD ide. 
divisions with similar fruit, and have therefore proposed the p 


Div. 24. OBTURAT E. 


Ys : from Å icis, Sow.; and 
328. S. MILLEPUNCTATA, Grev. I cannot distinguish this plant em 


1 . e s € > ii. 98). 
this seems to be Fries’ opinion (see ‘ Elenchus, 1. #9, —. L oon 
329. 8. FRAxINI, Fr. This also seems to me to be identical with S. corticis, 


328 
330. 


331. 


332. 


MR. F. CURREY ON THE FRUCTIFICATION OF SIMPLE SPHÆRIÆ: 


S. corrıcıs, Sow. t. 372.£.5; Fr.S. M. ii. 481; El. ii. 98. Tas. LIX. fig. 108, aseus 
with sporidia and free sporidia, X 325. Sporidia biseriate or crowded, pale brown, 
but of rather a rich colour, eurved, variable in length, but in good ripe specimens 
about 0:0008 inch long. Intermixed with the Sphæria, and exactly resembling it, I 
find other perithecia containing the bodies on the right of the figure, which were of 
a darker brown, and 3-5- or 7-septate, varying from 00006 to 0:0012 inch long, 
which I doubt not are perfect, free sporidia, the asci having deliquesced. 

S. TRANSVERSALIS, Schw.; Fr. El. ii. 94. Tas. LIX. fig. 109, ascus with sporidia, 
x 325. Sporidia uniseriate, dark brown, elliptical, subglobose, or even quite globular, 
0:0004 inch long. Arranged with the Obturate, but belonging, I think, to the Denu- 
date, and in my opinion not distinct from S. pulveracea, Ehrh. Fries (El. ii. 94) con- 
siders S. transversalis identical with S.myriocarpa. I have seen no specimens of the 
latter Spheria in fruit, but it is possibly not distinct from S. pulveracea. 

S. JUGLANDIS, Fr. S. M. ii. 493. Tas. LIX. fig. 110, sporidia, x highly. Sporidia 
biseriate, colourless, almond-shaped or narrowly elliptical, frequently curved, usually 
4-nucleate, sometimes with the endochrome bipartite (? sometimes 5-partite), 0-0004 
to 0°0005 inch long. I believe this to be the perfect form of S. Juglandis, Fr. The 
Hookerian herbarium contains another form, next described, which is an authentie 
specimen from the Scl. Suec., but which is not a true Sphæria, and is perhaps an 


imperfect form of the species just described. 
S. JUGLANDIS, Fr. S. M. ii. 493. Tag. LIX. fig. 111, fruit, x highly. No asci; but 


. the perithecia contain a quantity of minute, colourless or slightly yellowish, narrowly 


334. 


235. 


336. 


337. 


almond-shaped stylospores, borne on rather long pedicels, which, when broken 
look like the fruit of a-Cytispora, but are in reality only the fulcra of the stylospores. 
Stylospores 0:0002 to 0:0003 inch long, sometimes with two nuclei or with a bipar- 
tite endochrome. 

S. INSPERSA, Sz. Tas. LIX. fig. 112, ascus with sporidia, x 325. Sporidia biseriate 
or erowded, with the endochrome 4-partite, dark brown, constricted at the partitions, 
usually rather more pointed at one end than at the other, 0:0010 to 00011 inch : 
long. Not to be confounded with SS. inspersa, Berk., which has very different å 
sporidia. 
S. (SPHÆROPSIS ?) UBERIFORMIS, Fr. S. M. ii. 491. Tan. LIX. fig. 113, fruit, X 925. — 
I could find no asci. Perithecia filled with elongate, acuminate, colourless spores — 
having the endochrome 4-partite, 0-0005 to 0:0008 inch long. Easily known by m 
udder-shaped perithecia ; but it is, perhaps, not a true Sphæria. à 
S. (HenDersonn ?) orPrLATA, Fr. S. M. ii. 493. Tan. LIX. fig. 114, fruit highly — 
magnified. No asci. Perithecia filled with almond-shaped colourless spores wi 
bipartite endochrome, 00003 to 0-0004 inch long. This would be a Sphæria or : 
Hendersonia, according to whether it has or has not asci. 
S. (HENDERSONIA) PALINA, Fr. S. M. ii. 494. The fruit of this plant is not distin: — 
guishable from that of $. oppilata, but the spores rarely exceed 00008 inch in length. m 
The endochrome also is sometimes continuous instead of bipartite, and the spores ee : 
borne on long peduncles, as in S. Juglandis, but which I did not observe in S. oppilata: søn 


MR. F. CURREY ON THE FRUCTIFICATION OF SIMPLE SPHÆRLÆ. 329 


338, S. (HENDERSONIA) STROBILINA, Holl & Schm. The fruit in this species is just like 
that of S. palina, but larger, being 0:0005 inch long. 

339. S. Lonicer#, Fr. S. M. ii. 492, Tag. LIX. fig. 115, ascus with sporidia, x 325. 
Sporidia uniseriate, colourless, or with a greenish tinge, elliptical, 0*0004 to 0-0006 
inch long ; endochrome tripartite. 

940. S. PISIFORMIS, Pers. in lit. Incorrectly arranged in this division ; it belongs to the 
Obvallate, and is probably the Sphæropsoid form of 5. leiphæmia. 

341. S. (SPHÆROPSIS) OBTUSATA, Fr. S. M. ii. 495. Spermatia colourless, curved, about 
0:0004 inch long. à 
342. S. (HENDERSONIA) SYRINGZ, Fr. 8, M. ii. 492, Ta». LIX. fig. 116, fruit, x 325. 

Fruit borne on stalks, not contained in asci, irregular in size and shape, dark brown. 
In the later arrangements it would be placed in the genus Hendersonia, or perhaps 
in Diplodia, 
Div. 25. SUBTECTE. 

349. S. (DrrLopra) Inicis, Fr. S. M. ii. 501. Tas. LIX. fig. 117, fruit, x 325. No 
asci. Fruit borne, I think, on short peduncles, elliptical, oblong, subglobose or tur- 
binate, irregular in size and shape, almost colourless, but with a yellowish-green 
tinge. | 2 

344. S, (DIPLODIA) SARMENTORUM, Fr. S. M. ii. 498. Tap. LIX. fig. 118, fruit, x 325. 
No asci. Fruit borne on peduncles, dark brown, uniseptate when ripe, with sometimes 
a large globose nucleus in each division, irregular in size, about 0:0007 inch long 
on an average. "d 

345. S, (SPHEROPSIS ?) ATROVIRENS, B. Buxt, Fr. No asci. Stylospores exactly like those 
of S. Juglandis, but not septate or nucleate. Peduncles rather long. E 

346. S. EPIDERMIDIS, Fr. S. M. ii. 499. Tas. LIX. fig. 119, ascus with sporidia, x 325; 
Sporidia uniseriate or slightly overlapping, rather light brown, elliptical, st, PT 
or with the endochrome bipartite, hardly 0:0004 inch long. There zu. 15 sr 
in the herbarium, marked by Mr. Berkeley 5. epidermidis, Fr., m whioh ww ilo 
have no asci, but contain a mass of fusiform, straight m lightly a pies å 
spores, faintly 1-, 2-, or 3-septate, or 1-, 2-, or 3-nucleate, — 119 
0:0006 inch. These spores are drawn to the right sia us > T ight or curved, 

347. 8. Ruscr, Wallr. Tas. LIX. fig. 120, sporidia, x 450. Sporidia s each division 
normally 4-septate, occasionally 3- or 5-septate, constricted a MER — ; 
usually with one or more nuclei; colour of the sporidia a y å 
length 0-0006 to 0:0010 inch. | i RR | | 

348. "as n.s. Tas. LIX. fig. 121, sporidia, x s: pe ge c qaam 
slightly curved, with many nuclei, fusiform, 00008 inch long. 


i | of the wood. 
globose, bursting in somewhat parallel lines through the surface 


Div. 26. CAULICOLE. | | ie 
| | 4 idia, x 325. Sporidia 
349. S. Dourorvm, Pers.; Fr. S. M. ii. 509. TAB. LIX. m ipa Ben ost straight, 
_ biseriate, or uniseriate and overlapping, y pg ie i more. 
3-5-septate, constricted at the septa, 0001 inch long, ©! 9 x 


VOL, XXII. 


330 MR. F. CURREY ON THE FRUCTIFICATION OF SIMPLE SPHÆRIÆ. 


350. S. DEMATIUM, Pers.; Fr.S. M. ii. 505. Tas. LIX. fig. 123, sporidia, x 325. This 
plant was not, I think, quite ripe. Most of the sporidia were curved and colourless, 
as in the right-hand one of the three; but septa were just visible in one or two of 
the sporidia, and nuclei in others. I have little doubt that the sporidia, when ripe, 
are brown or yellowish, and three or four times (or even more) septate. There are 
three specimens of this species in the herbarium, marked a, B, and y. The above de- 
scription is from («); I could find no fruit in (8) and (y). 

351. S. ARUNDINIS, Fr. S. M.ii.510. Tas. LIX. fig. 124, sporidia, x 425. Sporidia bi- 
seriate, at first yellowish, then brown, 3-5-septate, sometimes with six septa, slightly 
curved, somewhat pointed at each end, frequently with nuclei in the septa, 0:0010 to 
0:0016 inch long. 

352. S. (SPHÆROPSIS ?) CogN1 Suzcicæ. Notatrue Spheria. Perithecia producing only 
minute, straight, cylindrical, colourless spermatia, 0:0002 inch long. 

353. S. (SPHÆROPSIS ?) ACUTA, Hoffm. I believe S. acuta, Hoffm., to be only a sphærop- 
soid state of some common Caulicolous species; but whether of S. complanata or 
S. herbarum, or of any other species, it is impossible to say. 

354. S. COMPLANATA, Tode. It is difficult, without authentic specimens, to be certain as 
to the true S. complanata of Tode. The sporidia shown in Ta». LIX. fig. 125, x 325 
diameters, are those which I have always supposed to belong to the true species. 
These sporidia are biseriate, or uniseriate and overlapping, colourless, or yellow, 
2-4-partite, sometimes apparently triseptate; they vary at different ages of the 
plant; (a) represents the young state, (b) the more advanced ; (c) is a form equally 
common (in the same specimens) with (5); (d) is, I think, the perfect and typical 
form of fruit. ; | 

355. S. CONIFORMIS, Fr. S. M. ii. 508. Tas. LIX. fig. 126, sporidia, x 325. Sporidia bi- 
seriate, yellow, slightly curved, 5-11-partite, varying much in length, as is shown by 
the figure. See the remarks on the fruit of this species as compared with S. acumi- 
nata, Sow., in Ann. and Mag. Nat. Hist. ser. 2. vol. ix. p- 328. | 

356. S. IMBERBIS, MS. This specimen contains three different plants—lst, Sphæria 
herbarum; 2ndly, perithecia containing curved colourless stylospores, like the sper- 

. matia of S. verruceformis ; 3rdly, perithecia containing minute, colourless, straight, 
or slightly curved cylindrical spermatia or stylospores. : 

357. S. pHmosticta, Berk. TAB. LIX. fig. 127, ascus with sporidia and free sporidia, 
X 325. Sporidia uniseriate or biseriate, very dark brown, subeymbiform, 0:0004 
inch long. Perithecia minute, punctiform. On Hierochloe Brunonis, from the Auck- 
land Group and Campbell Islands. 

we seen ea ‚Fr. S. M. ii. 527. Tan. LIX. fig. 128, we 
ul o. Fi. a m containing the narrowly cylindrical or irregularly- 
her deck bo er - These bodies have a bipartite endochrome, poo 
Perge cpi) ae E s ims of Zea Mays. If there be no asci, this plant 5 1 

; sci may have dissolved, and then it would be a Spherw. — 


There is another specimen in the herbarium marked S. Zee, Sz., which is, I think, 
identical with Hendersonia arcus, B. & Br. 


MR. F. CURREY ON THE FRUCTIFICATION OF SIMPLE SPHÆRLÆ. 381 


359. S. PELLITA, Fr. S. M. ii. 508. Tas. LIX. fig. 129, sporidia, x 325.  Sporidia, . 
x 325. Asci clavate ; sporidia crowded, multiseptate, fusiform, yellow, with a 
swollen joint, which is sometimes the third, sometimes the fourth from the end, 
0-0015 to 0:0016 inch long. There is another specimen in the herbarium, marked 
S. pellita, Fr., in which the fruit is not distinguishable from that of S. coniformis, 
supra. 

360. S. NIGRELLA, Fr. S. M. ii. 512. Tas. LIX. fig. 130, sporidia, x 325. Sporidia bi- 
seriate, colourless or greenish, fusiform, 00008 inch long; endochrome bipartite. 

361. S. Pısı, Sow.; Fr. S. M. ii. 509. Tas. LX. fig. 131, ascus with sporidia, x 325. 
Sporidia biseriate, yellow or yellowish brown, multicellular, 0:001 inch long. On 
Asparagus officinalis. I think not distinct from S. herbarum. 

362. S. ASTRAGALI, n. s. Tas. LIX. fig. 132, sporidia, x 325. Sporidia biseriate, subfusi- 
form, varying at the extremities from acute to obtuse, colourless; endochrome uni- 
partite, 0:0006 to 0:0008 inch long. Perithecia minute, globose, shining, half-hidden 
by the epidermis. On petioles of Astragalus. Dr. Richardson, Arctic expedition, 
1827 (? =S. nigrella, Fr.). 

363. S. ACUMINATA, Sow.; Fr. S. M. ii, 506. Ta». LIX. fig. 133, ascus with sporidia, 
and a broken aseus with sporidia protruding, X325. Sporidia filiform, almost as long 
as the ascus, yellowish brown in a mass, but of a very pale colour when separate, 
arranged side by side. See the remarks of Mr. Berkeley on the fructification of this 
species in Ann. and Mag. Nat. Hist. ser. 2. vol. ix. p. 828. I could not clearly make 
out any septa in the specimens in the Hookerian herbarium, although some of the 
asci when full of sporidia seemed marked with very faint transverse lines. Mr. Berkeley 
says there are about twenty articulations. 

364. S. CARICINA, Desm. The perithecia contain only minute, acuminate, colourless 
spermatia, without septa or nuclei, just like those of 5. Juglandis, but slightly longer. 

365. S. (DIPLODIA) CALVESCENS, Fr. S. S. 401. This specimen has perithecia containing 
the brown uniseptate fruit, not contained in asci, shown in Tas. LIX. fig. 154, 
x 325. It should be classed, 1 think, with Diplodia. In company with p apt 
are some other perithecia with asei and sporidia, the latter differing little, if a a, 
from the sporidia of S. herbarum. : 

366. 8. aes Berk. & Br. Ann. and Mag. Nat. Hist. ser. 2. vol. ix. P. Dee dock 
fig. 29. Tan. LIX. fig. 135, ascus with sporidia, X 220. Sporidia = poste : 
brown, oblong or slightly curved, triseptate, constrieted at the sep 
0:0008 inch long. i + a 

367. 8. RUBELLA, PA Fr. S. M. ii. 506. Tas. LIX. fig. 136, ascus with sporidia, X 220. 


id 
Sporidia filiform, quite filling the ascus, and of the same length, arranged side by 


i i h of the asci 
side, colourless when detached, but yellowish brown in the mass. Lengt 


variable, about 0:007 to 0:008 inch long. : :dia uniseriate, over- 
368. S. Heen, n. s. Tas. LIX. fig. 137, sporidia, X uns eii with the endo- 
lapping, brownish yellow, curved, 3-septate, or said Perithecia conical, truncate. 
chrome 4-partite, or rarely 5-partite, 0-001 inch long: of the perithecia are in the 


This species is very near S. complanata, Tode, but many 


332 MR. F. CURREY ON THE FRUCTIFICATION OF SIMPLE SPHÆRIÆ. 


form of truncated cones. It may, however, be merely a variety of the latter, and I 
propose it with doubt as a new species. 

369.. S. HERBARUM, Fr. S. M. ii. 511. Taz. LIX. fig. 188, asci with sporidia, x 220. 
Sporidia oblong-elliptie, at first yellow, eventually brown, multicellular, varying in 
size and in the mode of arrangement in the asci, as will be seen by referring to the 
figures. | 

Div. 27. FoLrcoLz. 

370. S. RHIZOMORPHE, Kunze. Tas. LIX. fig. 139, ascus with sporidia and free sporidia, 
X 325. Sporidia crowded, brown, of rather a dark colour, but clear and transparent, 
0:0012 to 0-0016 inch long. From Surinam. | 

371. S. TUBÆFORMIS, Tode; Fr. S. M. ii. 516. Tas. LIX. fig. 140, ascus with sporidia, 
x 325. Sporidia biseriate or crowded, colourless, or in age with a dirty-brown tinge, 
elliptical or subeymbiform, 0:0005 inch long. | 

372. S. (SPHÆROPSIS ?) MACULÆFORMIS, Pers.; Fr. S. M. ii. 524. This is not a true Spheria. 
The perithecia produce only exceedingly minute, staff-like, colourless spermatia, not 
so much as 00002 inch long. 

373. S. (SPHÆROPSIS?) OLES, D.C.; Fr. S. M. ii. 489. No asci. Tag. LIX. fig. 141, 
fruit, x 325. Fruit consisting of subcylindrical, colourless, highly refractive spores, 
070007 to 0:0009 inch long. I could not make out the mode of attachment; a few 
of the spores had a sort of string, as in the right-hand figure; but I am doubtful 
whether it were a true peduncle, or a string of a kind of mucus in which the spores 
were imbedded. This appears to be the same plant as No. 298 in the Obtectæ. 

374. S. (SPHÆROPSIS P) LEPTIDEA, Fr. S. M. ii. 522. N ot a true Spheria. Fruit con- 
sisting of minute, colourless, straight or very slightly curved, subeylindrical spores, 
barely 0:0002 inch long. 


375. S. (SPHÆROPSIS ?) DUPLEX, var. Nari, Fr. S. M. ii. 520. Not a true Spheria. 
Fruit the same as in S. leptidea, Fr. 

376. S.? PUNCTIFORMIS, Pers.; Fr. S. S. 86. The perithecia contain colourless saccate 
bodies, grouped together as in Ta». LIX. fig. 142; but whether they are spores or im- 
perfect asci I cannot say, probably the latter. 

377. S. (SPELEROPSIS) Høprrz, Sow.; Fr. S. M. ii. 521. Fruit consisting of colourless 
cylindrical spores, 00008 to 0-001 inch long; not distinguishable from the fruit of 


S. Oleæ, D.C., excepting that the spores are rather narrower and not so refractive, 
and I observed no peduncles. 


378. S. (SPHÆROPSIS) Æeoronn, Pers.; Fr. S. M. ii. 526. The fruit is the same as in 
S. leptidea, Fr., supra. 

379. S. (Discosta*) Artocrzas, Tode; Fr.S.8. 151; S. M. ii 523. Tap. LIX. fig. 149, 
spores, X 325. The fruit consists of straw-coloured, curved, or almost straight spores, 
having an extremely delicate filiform appendage at each end. The average length of 
the spores without the appendage is 0:0007 inch. The spores are probably produced 


* Discosta, Lib. Phlyctid. Notar. 


Perithecium innate, sube icate ; ostio- 
; A i arbonaceo lapsed and plicate ; 
lum pièrced. Spores fusiform, : -— —— z 


prolonged at each end into a filiform appendage. 


* 


MR. F. CURREY ON THE FRUCTIFICATION OF SIMPLE SPHÆRLÆ. 333 


in asci, which deliquesce at an early period. I can i 
rating this plant and the tilovin n from Pra ss NE Een 

380. S. (DiscosrA) ALNEA, Fr. S. M. ii. 520. This plant has asci; but the sporidia in 
the specimens in the Hookerian herbarium, which are from the Scl. Suec., are not 
formed. 

381. S. GLAUCO-PUNCTATA, Grev. Fl. Edin. Tas. LIX. fig. 144, sporidia, x 325. Sporidia 
biseriate, rather dark brown, slightly curved, but sometimes straight; endochrome 
3-6- or more partite, 0-0007 inch long. On Ruscus aculeatus. Not distinct, I think, 
from S. Rusci, Wallr. 

382. S. (SPHÆROPSIS) PALUSTRIS, Fr. in lit. The fruit is the same as in 8. leptidea, Fr. 

383. S. (SPHÆROPSIS) Vinca, Fr. in lit. The fruit is precisely the same in form as in 
S. Hedere, supra. Length=0-0006 to 0:0007 inch. | 

384. S. SETACEA, Pers.; Fr.8.M.ii.518. Tas. LIX. fig. 145, sporidia, x 450. Sporidia 
biseriate, colourless or greenish, arcuate, usually tapering to a point at each extre- 
mity, but sometimes (though rarely) rounded at the ends, 0:0006 inch long, apparently 
septate; but the apparent septum is, I think, only the line of contact of the bipartite 


endochrome. 
Div. 28. DEPAZEA 


385. S. (HENDERSONIA) CORNICOLA, Dec.; Fr. S. M. ii. 530. Tas. LIX. fig. 146, fruit, 
x 825. No asci. Fruit consisting of elliptical, triseptate, pale-brown spores, 0.0004 
to 0-0006 inch long, borne on long peduncles. 

386. S. (DEPAZEA*) vacans, Fr. S. M. ii. 532. No asci. The perithecia contain long 
filiform or fusiform bodies, precisely like those figured under 5. Juglandis, where, 
however, the bodies are not spores, but peduncles. Whether this is so in the present 


case, I am uncertain. 

387. S. (DEPAzEA) Uvarız, Berk. Tas. LIX. 
tained in asci, colourless, elliptical, subglobose or sub i 
long. On leaves of Uvaria triloba. Spots round or irregular, uy in the centre, 
becoming much darker towards the edge. Perithecia F “th à pae i Tar, LIX 

388. S. (DEPAZEA) FRAXINICOLA, MS. ; marked also “Sp iuit "s wege | 
fig. 148, fruit, x highly. Fruit not contained in asci, irregular in epe, NUE 
elliptical or subturbinate, or even subeymbiform, ne p dispu ag er 
round or irregular, brown, with a dark margin. pee idia bi ER colour- 

389. S. Dnvwrpis, Berk. Tas. LIX. fig. 149, sporidia, x 325. €: ieu: pel 
less, uniseptate, the two portions of unequal width, peng BOET ts. This is 
small, black, seated upon quite white, somewhat impressed, rag ee ie 
a true Sphæria with the habit of a Depazea. On aur ; 
Juan Fernandez, May 1830. | GT . ‚vi. p. 862. pl. 11. 

390. 8. (DEPAZEA) PALLOR, Berk. Ann. and Mag. yte leiden = with 


fig.2. Tas. LIX. fig. 150, fruit, x 325. Fruit not " 
mouthless, opening in a circumscissile manner or dehiscing 
spores acrogenous, simple. 


fig. 147, fruit, x 325. Fruit not con- 
turbinate, 0:0002 to 0*0004 inch 


* Derazea, Fr. Perithecium carbonaceous, subinnate, 
at the apex. Nucleus consisting of thread-like sporophores ; 


334 MR. F. CURREY ON THE FRUCTIFICATION OF SIMPLE SPHJERLE. 


a yellow tinge, sometimes straight, but usually more or less curved, 0-0006 to 00008 
inch in the chord of the are. 

391. S. (DEPAZEA) GRAMINICOLA, Berk. Ann. Nat. Hist. series 1. vol. i. p.207. Tas, LIX, 
fig. 151, fruit, x 325. Fruit not contained in asci, colourless, fusiform, straight or 
curved, 0:0008 to 0:0009 inch long ; endochrome sometimes bipartite, sometimes not. - 
I think it not improbable that this is an imperfect form of S. graminis, Pers. | 

392. S. (DEPAZEA) Brassrcæ, Pers. in lit. Fruit not contained in asci, similar to that 
of S. leptidea, Fr., but sometimes slightly thicker at each extremity than in the 
middle ; spots rather large, roundish, with an irregular outline, brown in the centre, 
greenish towards the margin. Perithecia small, black, and numerous. 

393. S. (DEPAZEA) DrANTHI, Fr. S. M. ii. 531. Fruit not contained in asci; similar to that 
of S. vagans, Fr., supra. ; 

394. S. (DEPAZEA) PrsicoLA, Berk. Tas. LIX. fig. 152, fruit, x 325. Fruit not contained 
in asci, colourless, of irregular shape, but mostly elliptical; endochrome bipartite or 
continuous ; length from 0:0004 to 00005 inch. [I find a minute fungus hardly dif- 
fering from this, on dead stems of Smyrnium Olusatrum. In my plant, however, the 
perithecia are not seated on an arid spot, and the fruit is rather larger and narrower, 
and generally more incurved at the sides, giving the spores a sort of dumb-bell shape.] 
Spots pale, round, margined, often confluent. Perithecia small, black. 

395. S. (DEPAZEA) RıBIcoLa, Fr. S. M. ii. 530. Tag. LIX. fig. 153, fruit, x 325. Fruit 
not contained in asci, colourless, bluntly and irregularly crescent-shaped, 0:0006 to 
0:0007 inch between the horns. 

396. S. (DEPAZEA) FUSCELLA, Berk. Fruit not contained in asci; similar to that of 
S. vagans, supra. Spots brown, paler in the centre. Perithecia very small On 
Convolvulus arvensis. King's Cliffe, Northamptonshire. | 

397. S. (DEPAZEA) ALISMATIS, n. s. Tas. LIX. fig. 154, fruit, x 325. Fruit colourless, 
bordered, elliptical, subglobose or turbinate, with sometimes the apparent remains of 
a peduncle, 0:0004 to 0:0005 inch long. On Alisma Plantago. Suffolk, Mr. Stock. 
Perithecia subglobose, chestnut-coloured, at least in the dry plant. 


INDEX. 
[The figures refer to the numbers affixed to each Plant, and not to the pages.) 
acuminata, 363. aquila, 218. biformis, 258. capillifera, 242. 
acus, pe Argus, 317. Bombarda, 245. caricina, 864. 
AG : Årtocrens, 378. Brassicæ, 239, 392. caudata, 269. 
Nes lii, 78. Arundinis, 351. brevirostris, 286. Cerasarum, 300. 
ismatis, 397. Ashwelliana, 327. bufonia, 324. - chionea, 237. 

| vage 225. Astragali, 362. cirrhosa, 290. 

nea, 380. atrovirens, 845. cæsia, 232. Clivensis, 366. 
321. aurantia, 22]. callicarpa, 281. clypeata, 303. 
apic = 18. calvescens , 865. collabens, 270. 
appendiculosa, 319. barbara, 282, 


canescens, 229. complanata, 354. 


MR. F. CURREY ON THE FRUCTIFICATION OF SIMPLE SPHJERLE. 


conformis, 310. 
conglobata, 249. 
coniformis, 355. 
cornicola, 385. 
Corni Suecicæ, 352. 
corticis, 330. 
cristata, 285. 
cubicularis, 295. 
Curreyii, 271. 


Dematium, 350. 
Desmazieri, 216. 
Dianthi, 393. 
dioica, 250. 
ditopa, 311. 
Doliolum, 349. 
Drymidis, 389. 

. duplex, 375. 


epidermidis, 346. 
episphzeria, 266. 
eutypa, 292. 


Fraxini, 329. 
fraxinicola, 388. 
fulva, 220. 
fuscella, 325, 396. 


gigaspora, 322. 


glauco-punctata, 381. 


Glis, 215. 
graminicola, 391. 


Hederæ, 377. 
Helenæ, 368. 
herbarum, 369. 
hirsuta, 233. 
hirta, 306. 


Theis, 343. 


imberbis, 356. 
inquilina, 314. 
inquinans, 328. 
inspersa, 243, 334. 


Juglandis, 332, 333. 


lanata, 223. 
lecythea, 275. 
leptidea, 374. 
ligniaria, 284. 
livida, 296. 
longispora, 287. 
Loniceræ, 339. 


macrostoma, 283. 
maculæformis, 372. ° 
mammæformis, 254. 
mammillana, 305. 
merdaria, 276. 
micraspis, 279. 
micula, 288. 
millepunctata, 328. 
moriformis, 246. 
moroides, 251. 
mutabilis, 235. 


nigrella, 360. 


obducens, 262. 
obtusata, 341. 
ocellata, 308. 
Olex, 298, 373. 
Olearum, 277. 
operculata, 294. 
oppilata, 336. 
ovina, 231. 


palina, 337. 
pallor, 390. 


palustris, 382. 
pellita, 359, 
pertusa, 273. 
Peziza, 264. 
phæosticta, 357. 
phæostroma, 227. 
Phomatospora, 816. 


| picastra, 274. 


pilifera, 289. 
pilosa, 236. 
Pinastri, 302. 
Pisi, 361. 
Pisicola, 394, 
pisiformis, 340. 
plateata, 252. 
pomiformis, 263. 
pruinosa, 297. 
pulveracea, 260. 
pulvinulus, 240. 
pulviscula, 272. 
pulvis-pyrius, 248. 
punctiformis, 276. 
pustula, 247. 
putaminum, 280. 


quadri-nucleata, 312. 


Racodium, 230. 
Rhizomorphæ, 370. 
rhodochlora, 234. 
rhodomphalos, 244. 


rhodostoma, 299. 


rosella, 217. 
rostrata, 291. 
Rubella, 367. 


Rubi, 313. 


rubicola, 267. 
Rusci, 947. 


sanguinea, 265 
sarmentorum, 344. 
scabra, 228. 
semi-immersa, 307. 
seriata, 348, 
setacea, 384. 
siparia, 320, 
sordaria, 261. 
spermoides, 253. 


stercoraria, 255, 256, 257. 


strigosa, 238. 
subiculata, 224. 
superficialis, 241. 
Syringæ, 342. 


Tamariseinis, 301. 
thelena, 222. 
Tiliæ, 326. 
tomicum, 309. 
transversalis, 331. 
tristis, 226. 
truncata, 219. 
tubæformis, 371. 


uberiformis, 335. 
Ulmicola, 278. 
Uvariæ, 387. 


rå 
US 


Ke 


ESC 
DEN 


LT 
GS 


ke? 
N 


> s 


= urey del. Tuffen West sc 


All the figures 


magnifisd 325 Diameters, ext Where 


otherwise saia 


P Currey del DE Då 


AU the figures magrafied 325 Diameters, exopt where otherwise stated. 


ri I7 TIIT tah 59 2.335 
Trans Linn. Soc. Vol XU tal åå, p.425. 


GEN. SPHÆRIA 
(Fig? 99 to Il.) 


P Currey del. Tuffen West c. — 


arwise stated 
All the figures magnified. 325 Diameters exogrt where ohur 


[ .887..] 


XXV. Synopsis of the Genera Camellia and Thea. By BERTHOLD SEEMANN, Ph.D., F.L.S. 


Read May 5th, 1859. 


GREAT diversity of opinion exists as to whether the genera Camellia and Thea ought 
to be merged into one, or regarded as distinct. The advocates of the union argue that 
hitherto no difference of generic importance has been pointed out, which, on being put to 
the test, has not broken down, and that, notwithstanding great authorities have pro- 
nounced in favour of keeping them separate, the generic union of Camellia and Thea is 
dictated by the stern laws of systematic botany, whilst their opponents have as yet not 
been able to disprove, with any degree of satisfaction, those sweeping assertions. In order 
to arrive at an independent opinion, it will be necessary to examine the various generic 
characters that have from time to time been proposed by authorities on the subject, and 
to add such critical remarks as a careful study of these plants enables me to make. 

Linnæus, who was acquainted with only two species of Camellia and one of Thea, thus 
defines them, in his sixth edition of the * Genera Plantarum ° of 1764 :— 


CAMELLIA, Linn. Tura, Kempf. 
Monadelphia Polyandria. : Polyandria Monogynia. 

Cal. Perianthium polyphyllum, subrotundatum, Cal. Perianthium 6-partitum, minimum, planum, 
imbricatum ; squamis subrotundis, obtusissimis, foliolis rotundatis, obtusis, persistentibus. Cor. 
internis sensim majoribus, concavis, deciduis. Petala 6, subrotunda, concava, æqualia, (— A 
Cor. Petala 5, obovata, basi coalita. Stam. Fi- Stam. Filamenta numerosa (ducenta circiter), 
lamenta numerosa, erecta, coalita in coronam filiformia, corolla breviora. Anthere simplices. 


stylo ampliorem, superne libera, corolla bre- Pist. Germen globoso-trigonum. Stylus subu- 
viora. Antheræ simplices. Pist. Germen sub- | latus, longitudine staminum. Stigma triplex. 
rotundum. Stylus subulatus, longitudine sta- Per. Capsula ex tribus globis coalita, Sen 
minum. Stigma acutum, reflexum. Per. Cap- laris, apice trifariam dehiscens. Sem, solitaria, 
sula turbinata, lignosa, sulcis aliquot exarata. globosa, introrsum angulata. 

Sem. Nuclei tot quot striæ capsulæ, subrotundi, = ; 
seminibus minoribus sæpe repletis. 


It is unnecessary to dwell much upon the imperfections of these characters.  Linnæus, 


by placing Camellia in Monadelphia Polyandria, and Thea in Polyandria Monogynia, 
undoubtedly laboured under the belief that in Thea all the stamens were free. The 
greater number of them are, however, as firmly united at the base as those of the typical 
. Camellia Japonica are, and the genus ought therefore to have been placed in Honadelphia 
Polyandria, as has been done by subsequent writers of the Linnean school. But Linnæus 
had very imperfect specimens of Thea at his disposal even when he publis ed the sixth 
edition of his «Genera Plantarum.’ After the publication of that edition, however, he 
seems to have had access to better materials; for in one of his copies of that work pre- 
served in our library we find several marginal notes in his own handwriting, + ER the 
VOL. XXII. 


338 DR. SEEMANN’S SYNOPSIS OF THE 


generic character of Thea: for instance, the words * Perianthium 6-partitum,..... 
planum ...." are altered into “Per. 5-partitum;” the number “6” assigned to the 
petals is struck out; to the description of the stamens is added “filamenta corollæ 
basi inserta,” &c.; so that, if he had brought out a seventh edition, these corrections 
would have been introduced, and they would have tended to improve the limits of the two - 
genera. But even giving Linnæus credit for all these intended improvements, his generic 
characters of the two contain little that either Camellia and Thea do not share with other 
Ternstræmiaceæ, or that more ample materials have shown to be correct. There are, how- 
ever, two characters upon which he relies to distinguish these genera which have never 
been shaken—the calyx of Camellia is termed polyphyllous and deciduous, whilst that of 
Thea is described with a definite number of persistent sepals. Morphologists may possibly 
object to attach much importance to them, as the calyx of Thea is clad with deciduous 
bracts, and what is termed a polyphyllous calyx in Camellia may also be explained as a 
simple calyx surrounded by bracts. This view of the subject would reduce the whole 
difference to one of time,—the sepals remaining longer attached to the plant in Thea than 
in Camellia. Systematic botanists, however, are compelled to take a practical as well as 
a theoretical view of such questions, and that is the course here pursued. 
DeCandolle, who wrote in 1824, and who looked upon Camellia and Thea as forming å 

distinct natural order, chiefly differing from Ternstremiacee by the absence of albumen, 
thus defines (Prodr. i. pp. 529, 530) the genera in question :— 


CAMELLIA, Linn. Tua, Kempf. 

Calyx imbricatus, nempe bracteis sepalisve non- Calyx 5-6-sepalus. Petala 6-9 ima basi subco- 
nullis accessoriis cinctus. Stamina basi poly- hærentia 2-3-serialia. Stamina basi sublibera. 
adelpha aut monadelpha. Antheræ ellipsoideæ. Antheræ subrotundæ. Capsula 3-cocca, septis 
Capsula valvis medio septiferis, axim triquetrum valvaribus, nempe a valvularum marginibus zs 
liberum post dehiscentiam relinquentibus. troflexis formatis. 


: DeCandolle places under Camellia one species which has no right to be there, viz. Camel- 
lia axillaris, Roxb. (=Polyspora axillaris, Sweet), while under Thea he ranges Loureiro's 
Thea oleosa, which I hold to be the old oil-seeded Camellia Sasanqua of Thunberg; and it 
s possible that in drawing up his generic characters the presence of those plants may have 
influenced him in some degree, compelling us to make some allowance in criticising them- 
He terms the stamens of Camellia polyadelphous : that term must, I think, be suppressed, 
as it cannot be applied to those of any true species of that genus, unless we are prepared 
to apply it tothe result of the mechanical rupture of the tube of the monadelphous stamens, 
witnessed in Camellia Sasanqua when the petals begin to fade, bend back, and become 
detached from the stamens. In Thea he describes the stamens as nearly free, but the 
outer series of them is always monadelphous, as in Camellia; and, following Gærtner, he 
ee "m cells of the eapsule, unlike those of Camellia, to be formed by the edges of the 
épris bent inwards. The latter are, however, exactly formed as those of Camellia, 
pie e Fare on the middle of the valve. DeCandolle has therefore not discovered 
= racter by which Camellia and Thea could be distinguished from each 


We now come to W. B. Booth, who in 1830 published an able history of the gne? 


GENERA CAMELLIA AND THEA. 339 


Camellia and Thea in the * Transactions of the Horticultural Society of London," vol. vii. 
p. 519. The essential characters which he assigns to them respectively are as follows :— 


CAMELLIA, Linn. Tura, Kempf. 

Flowers axillary, sessile. Calyx inferior, of 7, 8, Flowers axillary, stalked. Calyx inferior, of 5 
or more deciduous imbricated scales, the inner deeply divided permanent roundish segments. 
ones the largest. Corolla of 5 petals. Styles Corolla of 5, 6, or 9 petals. Styles cohering at 
united nearly their whole length. Capsule fur- the base, dividing towards the point into three 
rowed, having as many cells as furrows, and one distinct bodies. Capsule 3-lobed, 3-celled. 
or two seeds in each. Seeds solitary. 


Here another characteristic feature of the two genera is pointed out, viz. that in 
Camellia the flowers are always sessile, while in Thea they are stalked, though it can 
hardly be admitted in the essential characters of the two genera. The styles in Camellia 
are said to be united nearly their whole length : that is certainly the case in all those 
species with which Booth was acquainted in 1830; but in my Camellia Hongkongensis 
they are entirely free. Applying his generic character to the Camelliacee known at that 
time, Booth followed Lindley’s suggestion, and transferred Camellia euryoides, Lindl., to 
Thea. According to that rule, he ought to have done the same with Camellia maliflora, 
Lindl. The fact that the latter was only accessible to him in a double state may partly 
be considered the reason why he failed to do so. 

The last author who has attempted to frame a character for the two genera under con- 
sideration is Choisy, in one of the most elaborate papers that have as yet appeared on the 
Whole order of Ternstræmiaceæ. It was published in 1855 in the ‘Transactions of the 
Natural History Society of Geneva.’ He defines Camellia and Thea thus :— 

CAMELLIA, Linn. l Tuza, Kempf. : 
Calyce à éstivation imbricative, sépales sur plu- Calyce simple, å un seul rang. Capsule déhiscente 
sieurs rangs se recouvrant comme des tuiles. dans tout sa hauteur. (Cloisons non séparées 

Capsule semi-déhiscente ; cloisons séparées en du placenta central. 

haut du placenta central. 

It does not appear to me that we are justified in saying the capsule of Camellia is semi- 
dehiscent, and its septa do not separate from the central placenta, and that the capsule of 
Thea is quite dehiscent, and the septa do not separate from the central placenta. The 
fact is, that in Thea the septa do at one time separate from the central placenta, and at 
another they do not. In normally developed capsules they generally do separate, Hence 
it follows that the dehiscence or semi-dehiscence cannot be regarded as a character of 
generic distinction between the two genera. : 

There is, however, one point which has been quite overlooked by all who have written 
on this subject, and which seems the more important as it is not one of degree. E 
examining the stamens, I was struck with the fact that there was m all Camellias m 
Theos, besides the outer series of monadelphous stamens of indefinite number, > ne 
series of free stamens, definite in number. With a little practice the outer ak of t = 

ens is easily removed, and the inner one, closely surrounding the ovary, lai vg Fe 
Camellia I found them to be double in number to that of the normal number of petals, 


å agreed i t I 
and in Thea equal to it. I may add, that in the latter respect Thea i es vn 


340 DR. SEEMANN'S SYNOPSIS OF THE 


discovered in Pyrenaria. On the other hand, the structure now discovered in Camellia 
(a definite number of free stamens surrounded by a series of monadelphous ones) was 
known to exist in Calpandria of Blume, placed by Endlicher (Genera Plantarum, n. 5546) 
among the “ genera dubia ” of Meliaceæ, and removed to Ternstræmiaceæ by Choisy, who, 
however, did not suspect that it was so intimately related to Camellia as to render a 
generic separation impossible. True, in Camellia the filaments of the monadelphous 
series of stamens are more free towards their upper extremity than those of Calpandria; 
but it must be admitted this is a mere matter of degree; and no one would like to keep up 
Calpandria as å separate genus on that account only, especially as it agrees in habit and 
all other respects with Camellia. 

The number of styles in Thea I have always found to be three, whilst in Camellia there 
are normally five, or by abortion four or three. I here use the term ‘styles’ for what 
some authors have called stigmas ; and I justify this application by pointing to Camellia 
Hongkongensis, where the styles are quite free to their very base. There is, besides, 
another feature, which, though it cannot be admitted into the technical characters of the 
two genera, serves to distinguish them by their habit; I mean that in Camellia the 
flowers are always erect, whilst in Thea they are nodding. Thus, if we sum up the 
distinguishing peculiarities of the two, we have the following :— 


CAmELLIA, Linn. Tura, Kempf. 
Calyx polyphyllus, sepalis deciduis. Stamina in- | Calyx bracteatus, 5-sepalus, sepalis persistentibus. 
teriora duplo petalorum numero. Styli 5 (ab- Stamina interiora petalorum numero æqualia. 
ortu 4 v. 3).—Flores sessiles, erecti. Styli 3.— Flores pedunculati, declinati. 


When we apply these characters to the aggregate body of Camellias and Theas as it now 
stands, we are enabled to retain all the older typical species under their respective genera. 
Thus Camellia Japonica of Linnæus, together with C. reticulata, Lindl., C. Sasanqua, 
Thunb., and drupifera, Lour., are kept under Camellia, whilst Thea Chinensis, Linn., 
remains undisturbed under Thea. Only four modern species of Camellia are placed with 
Thea; and that is all the change required. A-few other species, which have nothing to do 
either with Camellia or Thea, are referred to their proper systematie position: Camellia 
avillaris, Roxb., is admitted to be the same as Polyspora axillaris, Sweet; C. integrifolia, 
Chois., proves itself to be a Laurinea (Actinodaphne Chinensis, Nees), according to sped 
mens kindly transmitted by Prof. Choisy himself; C.? Scottiana, Wall., is held to be Andi- 
nandra dumosa, Jack; whilst a plant which had been considered to be allied to the Wild 
Tea of Assam, and had been distributed by Wallich under the name of Freziera? atte- 
nuata, I found to be a species of Pyrenaria*. | 

I now proceed to give the generic characters of Camellia and Thea, and a synopsis of 


* Pyrenaria attenuata, 


; Seem. in * Bonplandia,” vii. p. 49 [1 859] (name only) ; foliis obovatis acuminatis serratis 
basi attenuatis glabris, 


; floribus solitariis pedunculatis, pedunculis medio bracteis lunulatis obtusis alternis instructis, 
sepalis (5) suborbiculatis obtusis bracteisque ciliolatis, petalis (5) cohærentibus obovatis v. oblongis obtusis, 3 "E. 
— paulo majoribus, omnibus dorso sericeis, staminibus interioribus liberis 5, exterioribus monadelphis numerosis 
yp glabris, stylis 5 liberis glabris, stigmatibus capitellatis, ovario ovato acuminato 10-suleato hirsuto, fructu - + + ' 
Affinis P. masocarpe, Korth.— Freziera ? attenuata, Wall, an Camellia?, Wall. Cat. n.1451!—Thea Assam 
affinis sp., Chois. in Mém. Soc. Gen. xiv. i. p. 156 (1855).—Thea viridis, var. Assamica ?, Planch. in Herb. Hook. 
Chois, J. e. p. 179.—Hab. in Tavoy (Gomez! in Wall. Cat. n. 1451 ). ; 


GENERA CAMELLIA AND THEA. 341 


their species. I have considerably reduced the number of species, and only describe a single 
new one; but I trust the smallness of the number of species that remain will be regretted 
the less, as we may anticipate ere long a considerable increase from China and Japan, 
since both these countries are now open to å more unrestricted intercourse with Europe. 
I must also beg to apologize for the many imperfections to be found in my enumeration ; 
but it must be borne in mind that, although the great herbaria of London, Paris, Berlin, 
and Vienna contain treasures which were carefully examined, yet these materials are in 
many cases not so complete as could have been wished ; and, as all the species are not repre- 
sented in our gardens, much is left for future investigation. Nor must it be forgotten that 
the plants under consideration have a tendeney to form double or semi-double flowers, and 
that their normal type is traced out only after a great deal of close study and observation. 


CAMELLIA, Linn. 


CHAR. Gen, EMEND.—Calyx polyphyllus, foliolis imbricatis, interioribus sensim majoribus, deciduis. 
Corollæ petala 5 (vel in floribus semiplenis et plenis 6-plurima), hypogyna, imbricata, interiora 
majora. Stamina hypogyna biserialia, exteriora numerosa petalis adhærentia in tubo concreta, inte- 
riora petalorum numero dupla, libera; filamenta subulata, anthere incumbentes, biloculares, ob- 
long, connectivo crassiusculo, loculis longitudinaliter dehiscentibus. Pollen sphæricum vel ovato- 
oblongum. Ovarium liberum, 5- (vel abortu 4- vel 3-) loculare. Ovula in loculis 4-5, angulo centrali 
alternatim inserta, pendula. Styli 5, abortu 4 vel 3, liberi vel plus minusve connati ; stigmata 
capitellata. Capsula 5- (abortu 4- vel 3-) locularis, dehiscens, loculicide 5-, vel in abnormis 3-4- 
valvis, valvis medio septiferis, axi centrali persistente, faciebus seminifero. Semina in loculis abortu 
solitaria, rarius gemina, inversa, festa nucamentacea, umbilico apicali impresso. Zmöryonis ex- 
albuminosz cotyledones crassz, inzequales, radicula brevissima, supera. ; 

Arbores vel frutices sempervirentes, Indiæ orientalis, Cochinchinæ, Chinæ et Japoniæ ; foliis. alternis, 
petiolatis, coriaceis, nitidis, serratis, gemmis magnis, perulis distiche imbricatis, floribus solitariis vel 
aggregatis, axillaribus, erectis, sæpe speciosissimis, albis, roseis vel purpureis. 

Camellia, Linn. Gen. Plant. n. 848; Endl. Gen. n. 5425 (excl. sp.); DeCand. Prodr. i. p. 529 (excl. sp.). 

Sasanqua, Nees in Sieb. Nipp. ii. p. 13 (excl. sp.). - 

Calpandria, Blum. Bijdr. bl. 178; Endl. Gen. n. 5546. | 

Obs. Nees von Esenbeck has proposed to divide Camellia into two genera (Camellia 
and Sasangua), adopted as subgenera (Kissi and Sasanqua) by Endlicher ; to the one is 

assigned a dehiscent, to the other an indehiscent fruit; but as all Camellias have a 

dehiscent fruit, though in Camellia Sasangua the dehiscence takes place rather late, they 

fall to the ground. 


foliis ovatis vel ovato-oblongis 
petalis (rubris, albis, flavidis 
staminibus, 


l. ©. Japonica; arborea; ramulis petiolisque glabris, 
acutis vel acuminatis, subtus subaveniis, floribus inodoris, ox 
variegatisve) rotundatis emarginatis (vel in var. hortens. fimbriatis), 
ovariis, stylis connatis capsulisque glabris. (v.s. sp. et V- e.) 

Camellia Japonica, Linn. Sp. Plant. p. 698 (1753), non Champ. ; Thunb. Fl. Jap. p. 272 (1784) å De 
Cand. Prodr, i. p. 529 (1824); Booth in Hort. Soc. Trans. vii. p. 529. t. 14 (1830) ; — så = 
F L Jap. p. 155. t. 82 (1835-44) ; Chois. in Mém. Soc. Gen. xiv. p. 147 (1855) ; Seem. in Bonplandia, 
Vi. p.278 (1858). 


à : : z , 
Chinensis pimente Jamaicensis folio, flore roseo, Petiver, Gazophylacium, t. 33. f. 4 (1702 


342 DR. SEEMANN’S SYNOPSIS OF THE 


San sa, vulgo Jamma Tsubakki, Kæmpf. Amoen. Exot. p. 850, cum ic. p. 851 (1712). 
Thea Camellia, Hoffm. ex Steudl. Nom. Bot. i. p. 265 (1841). 
Cameilia Kæmpferiana, Reboul, Atti della Tercia Riunione, p. 494, ex Wlprs. Ann. ii. p. 178 (1851-52), 
Nomina vernacul. In Japonia, “ Tsubaki, Jabu tsubaki” (i.e, Camellia sylvestris), a Chinensibus 
* San tsja” (i.e. Thea montana) vocatur. 
Geogr. Distr. Throughout Japan (Kempfer! Thunberg | Siebold !), forming dense woods, which, accord- 
ing to Siebold, look like those of our young Beeches ; cultivated in China and in European gardens. 
I have not seen wild specimens of this species from China, nor am I acquainted with 
any account of its having been found wild there. Champion thought he discovered it 
at Hongkong; but the species he took for Japonica turns out to be quite a distinct one 
(C. Hongkongensis, Seem.). The Chinese have from time immemorial cultivated C. 
Japonica in their gardens. In Europe it became known in the beginning of the eighteenth 
century, and the first figure of it was published in 1702 in Petiver's * Gazophylacium. 
Strange to add, though there are thousands of representations of the various varieties of 
this Camellia, yet we do not possess a single plate exhibiting the normal state of it. The 
form figured by Siebold and Zuccarini in their ‘Flora Japonica’ has semi-double 
flowers. 


2. C. Honekoncensts (Tab. LX.) ; arborea; ramulis petiolisque glabris, foliis ovato-lan- 
ceolatis vel lanceolatis acuminatis, subtus venis tenuibus distinctis, floribus inodoris, 
petalis (rubris) obovatis emarginatis, staminibus glabris, ovario stylisque liberis 
lanatis; capsula (glabra?). (v.v. Sp.) | 

Camellia Hongkongensis, Seem. MSS. 

C. Japonica, Champ. in Hooker’s Journ. of Bot. and Kew Misc. iii. p. 309 (1851), non Linn. !; Champ. 
in Trans. Linn. Soc. xxi. p. 112 (1853) ; Seem. Bot. Herald, p. 367. n. 68 (1857). 

Geogr. Distr. Cochinchina-Tourane (Gaudichaud, n. 271, in Herb. Par. !); Island of Hongkong (Eyre! 

Bowring! Champion! Hance! Seemann N. | 


This species was discovered in January 1837 by Gaudichaud in Cochinchina, and about 
1849 by Lieut.-Colonel Eyre, of the Royal Artillery, in the island of Hongkong, where it 
grows In company with Castanea concinna, Quercus bambusæfolia, Thea salicifolia, & ; 
it was afterwards collected by Bowring, Champion, Hance, and myself. Only three 
troos are known to exist in Hor kong. In a paper read November 5, 1850, before the 
Linnean Society, and published in 1853 in our Transactions, Capt. Champion took it for 
the true Camellia Japonica of Linnæus; and so did Mr. Bentham and myself in ow 
respective enumerations of the plants of Hongkong. A more recent examination and 
comparison with a large set of specimens of the genuine C. Japonica, Linn., has, bow- 
ever, led me to consider the Camellia found by Gaudichaud and Eyre as indeed allied to, 
but very distinct from, C. Japonica, Linn. It differs from C. Japonica in the shape ad 
dark-green colour of its leaves, in its free styles and woolly ovary; otherwise it has very — 
~~ habit of the common. single pink variety of C. Japonica, and is easily mistaken Ds 
or that species if the important differences pointed out be overlooked. Champion de 
scribes the capsule as * smooth,” thereby meaning most probably “ glabrous," as he his 


previously described that of C. ectabili xim r that, it is more 
than an inch in diameter. 7 is as * sericeous,” and he adds tha 


GENERA CAMELLIA AND THEA. 343 


3. C. RETICULATA; arborea; ramulis petiolisque sericeo-pubescentibus, foliis oblongis vel 
lanceolatis, subtus reticulatis, floribus inodoris, petalis (albis vel in var. fi. pl. varie- 
gatis) rotundato-obovatis emarginatis, staminibus glabris, ovario, stylis subliberis 
capsulaque sericeis (v. s. sp. et v. c.). | 

Camellia spectabilis, Champ. in Hook. Journ. of Bot. and Kew Misc. iii. p. 310 (1851); Champ. in Trans. 
Linn. Soc. xxi. p.111 (1853); Chois. in Mém. Soc. Genève, xiv. i. p. 148 (1855); Seem. Bot. 
Herald, p. 367. t. 78. p. 432 (1857) ; Seem. in Bonplandia, vi. p. 276 (1858). 

Var. flore pleno ; fl. pleno, Hook. Bot. Mag. t. 4976 (1857) ; Van Hout. Fl. des Ser. t. 1282-3 (1857). 

C. reticulata, Lindl. Bot. Reg. t. 1078 (1827) ; Booth in Hort. Soc. Trans. vii. p. 528 (1830); Chois. in 
Mém. Soc. Genéve, xiv. i. p. 147 (1855). 

Geogr. Distr. Island of Hongkong (Eyre! Champion!). Cultivated in European gardens. 


The double state of this Camellia was first figured and described by Dr. Lindley in the 
* Botanical Register,” t. 1078, from living specimens imported from Chinese gardens, 
while the normal state (with single flowers) was entirely unknown till discovered about 
twenty years afterwards in the woods of Hongkong by Capt. Champion. The discoverer 
did not, however, recognize it as such, but mistook it for a new species, to which he gave 
the name of C. spectabilis in a paper read November 1850 before the Linnean Society, and 
published in our Transactions. Bentham enumerated it under Champion's name in his 
‘Florula Hongkongensis.’ When I went over the same ground (Bot. Herald, p. 367), I 
was struck with the great resemblance existing between C. reticulata, Lindl., and C. spec- 
labilis, Champ. ; but not having at that time good specimens for comparison, I contented 
myself with remarking, in the Supplement to my Hongkong Flora (Bot. Herald, p. 432), 
of C. spectabilis— This species is closely allied to C. reticulata, Lindl.” Afterwards I 
was fortunate enough to obtain complete specimens of C. reticulata, Lindl., through the 
kindness of my friend Mr. Edward Otto of Hamburg, and was thus enabled to establish 
(Bonplandia, vi. p. 276) the identity of C. spectabilis and C. reticulata as species: as 
varieties they are distinct : the form described by Champion, having single white flowers, 
is the normal state, while that described by Lindley, having double red flowers variegated 
with white, is the abnormal state. In a horticultural point of view, this identification is 
of some importance. Although our florists have as yet taken little notice of c. reticulata, 
we have already the two principal tints displayed by its ally the C. Japonica, viz. the 
White and the red. It therefore now behoves them to take this species In hand, and 
endeavour to raise it in horticultural eyes to a standard of perfection; for although the 
flowers of C. reticulata are destitute of that compactness peculiar to C. Japonica, and 
rather remind us of a Pæony, yet they have this in their favour, that they are of gem 
arger size than those of C. Japonica—some specimens of the double variety observ 
by Sir W. J. Hooker actually measuring 20 inches in circumference, and being the larges 


floral development as yet recorded among the Camellias. 


e puberulis, foliis ellipticis 
odoris, petalis (albis) ob- 
lanato, stylis connatis, 


4. C. SASANQUA ; fruticosa vel arborescens ; ramulis petiolisqu 
vel ovato-lanceolatis acutis, subtus subaveniis, floribus in 
cordato-emarginatis vel bilobis, staminibus glabris, ovario 
capsula pubescente. (v. s. sp. et v. ©.) 


344 DR. SEEMANN’S SYNOPSIS OF THE 


Camellia Sasanqua, Thunb. Fl. Japon. p. 273. t. 30 (1784) ; Cav. Diss. vi. p. 306. t. 160 (1790) ; Staunt, 
Embas. to China, ii, p. 466 cum ic. (1797) ; Bot. Reg. t. 12 (1815), t. 1091 (1827) ; Lodd. Bot. Cab, 
t.1275 (1827); Booth in Hort. Soc. Trans. vii. p. 521 (1830) ; Sieb. et Zucc. Fl. Jap. p. 158. t. 83 
(1835-44) ; Chois. in Mém. Soc. Genéve, xiv. i. p. 147 (1855); Seem. in Bonpl. vi. p. 278 (1858). 

Sasangua, Kempf. Delineat. plant. Japon. (MSS. in Mus. Brit.) fig. 25; Amoen. Exot. p. 853, ex parte 

1712). 

len RER Abel, Journ. in China, p. 174 cum ic., App. p. 363 (1818); Lindl. in Bot. Reg. t. 942 
(1825); Lodd. Bot. Cab. t. 1065 (1825); Booth in Trans. Hort. Soc. vii. p.524 (1830); Chois. in 
Mém. Soc. Genève, xiv. i. p.147 (1855) ; Seem. in Bonpl. vi. p. 278 (1858). 

Thea oleosa, Lour. Fl. Cochinch. p. 339 (1790) ; Chois. in M&m. Soc. Genöve, xiv. i. p. 156 (1855). 

Thea longifolia, Nois., et Thea Sasanqua, Nois., teste Steudl. 

Nomina vernacul. Nomen Cochinchinense, teste Loureiro, “ Yeu-cha.” Nomen Chinense, teste Abel, 
“ Tscha-Yeoa,” teste Loureiro, * Che-deau,” et teste Staunton, * Cha-whaw.” Nomen Japonicum, 
teste Siebold, * Sasankwa.” Nomen Japon.-chinense, teste Siebold, * Tsja-bai.” 

B. var. flore semi-pleno, Lindl. Bot. Reg. t. 12 (1815), t. 1091 (1827); Lodd. Bot. Cab. t. 1275 (1827). 

Geogr. Distr. Japan (Thunberg in Mus. Brit.! Siebold). China, Prov. of Kiangsi (Sir George Staunton 
in Mus. Brit. !); Prov. of Kwangton (Sir G. Staunton in Mus. Brit.! Adel, Loureiro); Chusan 
(Dr. Cantor in Herb. Bth.!); Loo-choo (C. Wright, n. 28!). Cultivated in European gardens. 


Camellia Sasanqua, Thunb., and C. oleifera, Abel, are in most works regarded as distinct 
species. They are here united, because I could not find a single character by which they 
could be distinguished from each other. . I have also added as a synonym the doubtful 
Thea oleosa of Loureiro, which, from the description given by its author, agrees tolerably 
well with the old C. Sasanqua. I am the more convinced that this identification is 
correct, as there are only two Camelliaceous plants that yield seeds sufficiently oily to be 
used for economic purposes, viz. Camellia Sasanqua and C. drupifera; and Loureiro states 
emphatically that his Thea oleosa yields an oil and grows wild in the province of Canton, 
—a fact fully agreeing with what we know of C. Sasanqua. What is preserved in the 
Parisian Museum as the original specimen of Thea oleosa, Lour., is Thea Chinensis, Var. 
Bohea. The latter plant (I mean 7. Chinensis, var. Bohea) was well known to Loureiro, 
and named by him Thea Cantoniensis ; and, as it neither yields an oil nor grows wild in 
Canton, I do not think that we are far wrong in assuming that the specimen preserved at 
Paris as Thea oleosa has obtained that name by a misplacing of the label, and that the 
genuine specimens of Thea oleosa, Lour., like those of Camellia drupifera, Lour., and 
Thea Cochinchinensis, Lour., have been lost. The genuine C. Sasangua has always white 
flowers; and when Keempfer, who first brought it into notice, attributed pink ones to it, 
he undoubtedly confounded another species with it (viz. Thea maliflora= Camellia mali- 

flora, Lindl.), which, until Lindley pointed it out as distinct, was always regarded as à 
variety of C. Sasanqua. Siebold and Zuccarini, in their * Flora Japonica,’ still kept up 
this old error. There are no specimens of C. Sasanqua in Keempfer’s herbarium ; but in 4 
collection of manuscript drawings of that author, preserved in the British Museum, there 


is a figure of his * Sasanqua,’ which represents the genuine C. Sasanqua of Thunberg: 
(Conf. Thea maliflora, Seem.) 


5. en arborea ; ramulis petiolisque puberulis, foliis lanceolatis vel oyato- 
ceolatis longe acuminatis, subtus venis tenuibus distinctis, floribus odoratis, petalis 


GENERA CAMELLIA AND THEA. 345 


(albis) obovatis emarginatis, staminibus glabris, ovario lanato, stylis subliberis: cap- 
sula pubescente. (v.s. sp. et v. c.) 


Camellia drupifera, Lour. Fl. Cochinchinen. p. 411 (1790) ; DeCand. Prodr. i. p. 529 (1824). 

Camellia Kissi, Wall. in Asiat. Res. xiii. p. 429 (1820) ; DeCand. Prodr. i. p. 529 (1824); Wall. Cat. 
n.977 (1828); Booth in Trans. Hort. Soc. vii. p. 525 (1830) ; Wall. Plant. Asiat. Rar. iii. p. 36. 
t. 256 (1832); Lodd. Bot. Cab. t. 1815 (1832); Chois. in Mém. Soc. Genève, xiv. i. p. 147 (1855) ; 
Seem. in Bonplandia, vi. p. 278 (1858). 

Camellia Chamgota, Ham. MSS. (1814), teste Wall. in Asiat. Res. xiii. p. 429 (1820) ; Chois. in Mém. 
Soc. Genéve, xiv. i. p. 147 (1855). 

Camellia Keinia, Hamilt. MSS. in Don Fl. Nepal. p. 224 (1825). 

Camellia? oleifera, Wall. Cat. n. 976 (1828). — 

Camellia symplocifolia, Griff. Itin. Notes, p. 40, n. 652 (1848), et Notule ad Plant. Asiat. iv. p. 560. 
t. 604. f. 2 (1854). 

Camellia (Thea) Mastersiana, Griff. Notulae ad Plant. Asiat. iv. p. 559 (1854). 

Mesua bracteata, Sprengl. Syst. Veg. iii. p. 127 (1826). 

Nomina vernacula. In Cochinchina, teste Loureiro, * Cay Deau so ;” in Sylhetia, teste Hamilton, * Cham- 
gota ;” in Nepalia, teste Wallich, * Kissi” seu * Kissi- Soah," et, teste Don, * Keng-na" vocatur. 
Geogr. Distr. Nepal (Wallich! E. Gardner!) ; Khasia Mountains, 3-5000 feet (Hooker and Thomson |, 
Griffith), Th.Lobb!); Bootan (E. Ind. Comp. Herb.!); Sikkim (Hooker and Thomson !); Assam 
(Griffith!, Jenkins!); Cochinchina (teste Loureiro). Cultivated at the Botanic Garden, Ham- 

burgh. 


This species was discovered by Loureiro in Cochinchina, and has since been found in 
different parts of the East Indies; as it has an extensive geographical range, and grows 
both in the high and low lands, it looks so different in different localities, that we can 
scarcely be surprised if one and the same author has not recognized it again when he met 
with it in a new place. Its nearest ally is Camellia Sasanqua, Thunb., and, like that 
species, it yields an oil. There are no original specimens of Camellia drupifera, Lour., 
either in London or Paris,—a misfortune the more to be regretted, as, owing to the very 
imperfect description given by Loureiro, this species has always been regarded as a doubt- 
ful one. I endeavoured to show (Bonplandia, vii. p. 48) that the characters furnished 
might be reconciled with those of Pyrenaria serrata, Blum.; but I am now convinced 
that the balance of evidence inclines more to Camellia Kissi, Wall. ; Pyrenaria serrata, 
Blum., possessing no oleiferous properties, as I have since learned. That Camellia dru- 
pifera, Lour., has nothing whatever to do with Mesua, to which it has been referred by 
Sprengel, it is hardly necessary to demonstrate. 


6. C. LANCEOLATA ; arborescens; ramulis petiolisque puberulis, foliis lanceolatis vel ovato- 
lanceolatis acuminatis, subtus discoloribus, venis obscuris, floribus inodoris, petalis 
(albis) obovatis obtusis, staminibus glabris, ovario stylisque connatis lanato-pubes- 

_centibus, capsula . . ... (v. s. sp.) 


Camellia lanceolata, Seem. MSS. 
Calpandria lanceolata, Blum. Bijdr. 178; Korth. Verhandel. p. 148. t. 31. | 
Geogr. Distr. Borneo (teste Korth.) ; Sumatra (Marsden, in Herb. Bth. et Hook. !). 


, foliis oblongo-ovatis acutis, 


7. C. QUINOSAURA ; arborescens; ramulis petiolisque . . . - - = 


VOL. XXII, 


346 DR. SEEMANN’S SYNOPSIS OF THE 


subtus..... , floribus inodoris, petalis (albis) ..... , staminibus...., ovario stylisque 


connatis glabris, capsula . . . . 


Camellia quinosaura, Seem, MSS. | 
Calpandria quinosaura, Korth. Verhandel. p. 149. 
Geogr. Distr. Java (teste Korth.). 


I have not seen any specimens of this plant. 


Species excluse : 


C. assimilis, Champ.= Thea assimilis, Seem. C. Sasanqua fl. incarnato multiplici, Sims= Thea 
C. axillaris, Roxb. = Polyspora axillaris, Sweet. maliflora, Seem., fl. pl. 
C. Banksiana, Lindl.*. C. Sasanqua fl. rubro simplici, Sims= Thea mali- 
C. Bohea, Griff.= Thea Chinensis, Linn. flora, Seem. 
C. caudata, Wall.= Thea caudata, Seem. C. Sasanqua rosea, Hort.= Thea maliflora, Seem., 
C. euryoides, Hort.= Thea maliflora, Seem. fi. pl. | 
C. euryoides, Lindl. — Thea euryoides, Booth. C. Sasanqua stricta, Edw. = Thea maliflora, Seem., 
C. integrifolia, Chois. = Actinodaphne Chinensis, fl. pl. l 

Nees, C. ? Scottiana, Wall. = Andinandra dumosa, Jack. 
C. maliflora, Lindl. = Thea maliflora, Seem., fl. pl. C. Thea, Link= Thea Chinensis, Linn. 
C. roseflora, Hook. = Thea maliflora, Seem. C. theifera, Griff. = Thea Chinensis, Linn. 
C. salicifolia, Champ. = Thea salicifolia, Seem. C. viridis, Link = Thea Chinensis, Linn. 


THEA, Kæmpf. 


CHAR. GEN. EMEND.—Calyæ bracteatus, 5-phyllus, foliolis imbricatis, interioribus majoribus, persisten- 
tibus. Corolle petala 5, 7 vel 8, hypogyna, imbricata, cohærentia, interiora majora. Stamina hypo- 
gyna, biserialia, exteriora numerosa petalis adhærentia et in tubo concreta, interiora petalórum 
numero æqualia (5-7-8) libera ; filamenta subulata, antheræ incumbentes; biloculares, oblongæ, con- 
nectivo crassiusculo, loculis longitudinaliter dehiscentibus. Pollen rotundato-oblongum lineatum vel 
globoso-subtrigonum. Ovarium liberum 3-loculare. Ovula in loculis 4-5, angulo centrali alterna- 
tim inserta. Styli 3, connati vel subliberi, stigmata capitellata. Capsula 3-locularis, dehiscens, 
loculicide 3-valvis, valvis medio septiferis, axi centrali persistente, faciebus seminifero. Semina in 
loculis abortu solitaria, rarius gemina, testa nucamentacea, umbilico apicali impresso. Embryonis 
exalbuminosæ cotyledones crassæ, carnosæ, inæquales, radicula brevissima, supera. . 

Frutices vel arbusculæ sempervirentes, Indiæ orientalis, Chinæ et J aponiæ; foliis alternis, petiolatis, 


coriaceis, nitidis, serratis, floribus pedunculatis, pedunculis bracteatis, axillaribus, solitariis vel aggre - in 


gatis, nutantibus, albis vel roseis. 


Thea, Kempf. Linn. Gen. (edit. vi. 1764) p. 269. n.668; DeCand. Prodr. i. p. 530 (excl. sp); End 
Gen. Plant. n. 5426. 


Camellia, sp. auct. 


l. Tura MALIFLORA; ramulis petiolisque puberulis; foliis ovatis vel ovato-lanceolatis 
acuminatis, utrinque glabris, pedunculis imbricato-bracteatis, sepalis ovatis obtusis, 
dorso pubescentibus, petalis 7-8 obcordato-emarginatis erectis glabris (roseis), stami- 
nibus, stylis ovariis capsulisque glabris. (v. v. eult.) 

* “Camellia Banksiana, 


as neither Lindley nor any 
p. 278. 


Lindl. (mentioned by Champion in Hook. Journ. and Kew Misc. iii. p. 310), is a nonentity, 
other botanist has ever published such a name."— Seem. Bot. Herald, p. 367 ; Bonpl. Y! 


GENERA CAMELLIA AND THEA. 347 


Thea maliflora, Seem. MSS. 

Camellia roseflora, Hook, Bot. Mag. t. 5044 (1858). 

Sasanqua, Kempf. Ameen, Exot. p. 853! ex parte (1712). 

Camellia Sasanqua a, flore rubro simplici, Sims, Bot. Mag. sub t. 2080! (1819). 

Camellia euryoides, Hort. Germ. et Angl. ! (non Lindl. !). 

Var. flore pleno, Seem. 

Camellia Sasanqua, B. stricta, fl. pl. carneo, Edwards, Bot. Reg. t. 547 ! (1821) ; Lodd. Bot. Cab. t. 1134 
(1826). 

Camellia Sasanqua, B. flore incarnato multiplici Palmer's Double Sasanqua, Sims, Bot. Mag. t. 2080! 
(1819). 

Camellia Sasanqua rosea, Hort. ! 

Camellia maliflora, Lindl, Bot. Reg. sub t. 1078 ! (1827); Booth in Hort. Soc, Trans. vii. p. 526! (1830) ; 
Chois, in Mém. Soc. Phys. de Genève, t. xiv. pt. i. p. 147 ! (1855). 

Geogr. Distr. Japan. Cultivated in European gardens. 

Kæmpfer (Amcæn. Exot. p. 853) was the first who noticed the normal state of this 
species under the name of * Sasanqua," though he does not seem to have preserved any 
specimens of it, for the only species of Camellia to be found in his herbarium at the 
British Museum is C. Japonica, Linn. (Herb. Keempf. fol. 23. n. 2, et fol. 32. n. 2). He 
distinctly states that it has single red flowers. Thunberg (Flora Japon. p. 273), although 
he ascribed to his Camellia Sasanqua white flowers, quoted Kæmpfer's “ Sasanqua ” as 
an entire synonym of it (though it is only so in part), and thus, by confounding two 
very distinct species, laid the foundation of an endless series of mistakes committed by 
subsequent authors. DeCandolle, too (Prodr. i. p. 529), quoted Kæmpfer’s ** Sasanqua ” 
as a synonym of C. Sasangua, Thunb., further remarking that the latter species varied 
with white, flesh-coloured, pink and red flowers,—a remark not borne out by facts, as the 
true C. Sasanqua, Th., never bears any but white petals. Sims (Bot. Mag. t. 2080) 
figured, from garden specimens introduced by Capt. Rawes in 1816, the double state of 
the species under consideration, which he mistook for a variety of C. Sasanqua, Thunb. ; 
but he felt the necessity of bringing the various synonyms, supposed to belong to C. Sa- 
sanqua, Thunb., into some kind of order, by arranging them as follows :— | 


Camellia Sasanqua. 


Var. a. flore rubro simplici =“ Sasanqua;” Kempf. Ameen. Exot. p. 853. 

Var. 8. flore incarnato multiplici= Palmer’s Double Sasanqua, Bot. Mag. t. 2080. 

Var. y. flore albo simplici= C. Sasanqua, Thunb. Fl. Jap. p. 273. t. 30 [excl. syn. Kempf. ex parte! he 
ought to have added.—B. S.] ; Bot. Reg. t. 12; Staunton. Emb. to China, ii. p. 466, cum icon. 


Of these three varieties only g. and ß. belong to Thea maliflora, y. appertaining to, or 
rather being, the veritable C. Sasanqua, Thunb. The double state of our plant was again 
figured from the same garden plant in the Bot. Reg. t. 547, under the name of C. Sasanqua 
B. stricta, ji. pl. carneo. At last (1827) Lindley (Bot. Reg. sub tab. 1078) pointed out 
that this so-called double pink variety of C. Sasanqua was in reality a distinct species, to 
Which he gave the name of C. maliflora. Booth (Hort. Soc. Trans. vii. p. 526) adopted 
this view, and at the same time drew attention to the very different habits of C. maliflora 


and C. Sasanqua. Unfortunately, Siebold and Zuccarini (Fl. Jap. p. 158) overlooked the 
; 2.22 


348 DR. SEEMANN’S SYNOPSIS OF THE 


result of Lindley’s investigation, and probably misled by the name which Kæmpfer had 
adopted for two plants, they quoted, like Thunberg, DeCandolle, Sims, and others, Kæmp- 
fer's * Sasanqua ” as quite synonymous with C. Sasanqua, Thunb., and in their detailed 
description ascribed to C. Sasanqua “ petalis puniceis vel albis." Meanwhile the normal 
state of this species, i. e. the single-flowered variety, had found its way into our gardens, 
and was cultivated for many years, both in England and on the Continent, under the 
name of C, euryoides, Hort. (non Lindl. !), until (1858) Sir W. J. Hooker described it (Bot. 
Mag. t. 5044) as C. roseflora. 

There is undoubtedly a certain resemblance in the foliage of Camellia Sasanqua, Thunb,, 
and Thea maliflora, Seem., that of both being coriaceous and almost glabrous, but on 
closer inspection the difference between the leaves becomes as evident as is that in their 
habit. The characters by which the two species may instantly be distinguished are, that 
Camellia Sasanqua has white petals and a woolly ovary, while Thea maliflora has pink 
petals and a glabrous ovary. When some authors ascribe flesh-coloured and red flowers 
to the latter species, they mean pink, the pink being more or less intense. 

I have not seen any wild specimens of Thea maliflora. Whence and how the single 
state came into our gardens, is a matter of uncertainty; but as Keempfer met with it in 
Japan, it is probably indigenous to that empire. The double variety was introduced (1816) 
from Chinese gardens, and the Chinese may perhaps have obtained it from J apan. 


2. THEA EURYOIDES; ramulis petiolisque pilosis; foliis ovato-lanceolatis acuminatis, 
subtus sericeis, pedunculis imbricato-bracteatis, sepalis ovatis obtusis, dorso pilosis, 
petalis 7-8 obovatis obtusis erectis glabris (albis), staminibus stylis ovariis capsulisque 
glabris. (v. s. sp.) 

Thea euryoides, Booth in Hort. Soc. Trans. vii. p- 560 (1830) ; Chois. in Mém. Soc. Phys. de Genève, 
xiv. i. pp. 149, 156 (1855). 

Camellia euryoides, Lindl. Bot. Reg. t. 983 (1826); Lodd. Bot. Cab. t. 1493 (1828) ; Seem. Bonpl. vi. 
p. 278 (1858). 

Geogr. Distr. Chusan (Dr. Cantor, in Herb. Benth. N. 

I have only seen a single specimen of this species, from China, preserved in Bentham's 
herbarium. It was first imported in a living state by the Horticultural Society of 
London in 1822, and again in 1824, and is used by the Chinese for grafting upon it their 
varieties of Camellia Japonica. Itseems to have entirely disappeared from our gardens, 
what goes by the name of Camellia euryoides amongst horticulturists being the normal 
state of Thea maliflora. | 


* Des CAUDATA ; ramulis petiolisque puberulis; foliis lanceolatis longe acuminatis - 
utrinque glabris, pedunculis imbricato-bracteatis, sepalis ovatis obtusis, petalis 5 
obovatis obtusis, dorso villosis, patentibus (albis), staminibus stylis ovariisque villosis 
capsula glabra. (V. S. sp.) 

Thea caudata, Seem. MSS. | 

Camellia caudata, Wall. Cat. n. 978 (1828); Wall, Plant. Asiat. Rar. iii. p. 36 (1832); Wiprs. Rep. på 
ve t P-375 (1842) ; Chois. in Mém. Soc. Phys, Genève, xiv. i. p. 148 (1855); Griff. Itinerary 

ove’, p. 40. n.651 (1848) ; Griff. Notulæ ad Plant. Asiat. iv. p. 559. t. 601. fig. 2 (1854). 


GENERA CAMELLIA AND THEA. 349 


Geogr. Distr. Khasia Mountains and Sylhet from the sea-level to 4000 feet (Wallich! Griffith! Hooker 
and Thomson !). | 

4. THEA SALICIFOLIA ; ramulis petiolisque pubescenti-villosis; foliis ovato-lanceolatis 

~ longe acuminatis, supra (nervo medio except.) glabris, subtus pubescenti-villosis 
pallidioribus, pedunculis imbricato-bracteatis, sepalis ovato-lanceolatis longissime 
acuminatis, petalis 5 obovatis obtusis vel acuminatis, dorso villosis, patentibus (albis), 
staminibus stylis ovariisque villosis, capsula rostrata glabra. (v. v. sp. et cult.) 

Thea salicifolia, Seem. MSS. 

Camellia salicifolia, Champ. in Hook. Journ. of Bot. and Kew Misc. iii. p. 309 (1851) ; Champ. in Trans. 
Linn. Soc. xxi. p. 112 (1853); Chois. in Mém. Soc. Genève, xiv. i. p. 148 (1855); Seem. Bot. 
Herald, p. 367. t. 76 (1857) ; Seem. in Bonplandia, vi. p. 278 (1858). 

Geogr. Distr. Island of Hongkong (Champion! Hance! Seemann! Wilford! C. Wright, n. 63). Culti- 
vated at Kew. 


9. THEA ASSIMILIS; ramulis petiolisque adpresso-pilosis; foliis lanceolatis acuminatis, 
supra glabris, subtus ad nervos adpresso-pilosis, pedunculis imbricato-bracteatis, 
sepalis rotundatis obtusis, petalis 7 obovatis emarginatis patentibus (albis), dorso 
sericeis, staminibus stylis ovariisque villosis, capsula rostrata glabra. (v.s. sp.) 

Thea assimilis, Seem. MSS. ; 

Camellia assimilis, Champ. in Hook. Journ. and Kew Misc. iii. p. 309 (1851) ; Champ. in Trans. Linn. 
Soc. xxi. p. 112 (1853); Chois. in Mém. Soc. Genève, xiv. i. p. 148 (1855); Seem. Bot. Herald, 
p. 367. t. 77 (1857); Seem. in Bonpl. vi. p. 278 (1858). 

Geogr. Distr. Island of Hongkong (Champion! C. Wright, n. 61 !). 


6. Tura CHineNsis (Tas. LXI.) ; ramulis petiolisque glabris vel sericeo-puberulis ; foliis 
lanceolatis, obovato-lanceolatis vel ovato-oblongis acuminatis, utrinque glabris vel ad 
nervos plus minusve puberulis, pedunculis 2-3 bracteatis, sepalis rotundatis obtu- 
sissimis, glabris vel sericeis, petalis 5 obovatis obtusis patentibus (albis) dorso glabris 
vel pubescentibus, staminibus stylisque glabris, ovario villoso, capsula glabra. 
B B Sp ot Ys v). 

Thea Chinensis, Linn. Spec. Plant. edit. i. p.515 (1753); Sims, Bot. Mag. t. 998; DeCand. Prodr. i. 
p. 530 (1824); Chois. in Mém. Soc. Genève, xiv. i. p. 155 (1855). 

Thea Bohea, Linn. Sp. Plant. edit. ii. p. 735 (1762). 

Thea viridis, Linn. Sp. Plant. edit. ii. p. 735 (1762). 

Thea Cantoniensis, Lour. Coch. i. p. 339 (1790). 

Thea Cochinchinensis, Lour. Coch. i. p. 338 (1790). 

Thea Japonica, Kempf. Amen. Exot. p. 605 (1712). 

Thea Assamica, Masters. Thea stricta, Heyne. 

Camellia Bohea, Griff. Notul. ad Plant. Asiat. iv. p. 553. t. DCII. fig. 1 (1854). 

Camellia theifera, Griff. Not. ad Plant. Asiat. iv. p. 558 (1854). 

Camellia Thea, Link., ex Steudl. nomen. bot. 

Camellia viridis, Link., ex Steudl. nomen. bot. “Ta.” « Sa” vel * Cha” 

Nomina vernacula. In Japonia, teste Kæmpfer, * Tja,” in China, teste DM MT = 
et in Cochinchina, teste Loureiro, * Che dn näm” vocatur. TOT d^ 

Geogr. Distr. Upper Assam (Griffith | Masters!) ; cultivated in the East Indies, China, and Japan; also 


in European gardens. 


350 DR. SEEMANN’S SYNOPSIS OF THE 


I agree with Choisy, that there is only one species of the plant yielding the tea of com- 
merce, for which I adopt the name of Thea Chinensis given to it by Linnæus in the first 
edition of his *Species Plantarum, and afterwards sanctioned by Sims and others asa 
collective one for Thea Bohea, viridis and Assamica. The propriety of taking the name 
“ Ohinensis” may be open to discussion, since we have no wild specimens of Tea from 
China, but only from Upper Assam, and a Chinese tradition of great antiquity states 
that the Tea-plant was introduced from India by the Buddist priests, so that it is possible 
that we may term a species “ Chinese ” that is in reality ** East-Indian" in its origin, and 
thus furnish another instance of * lucus a non lucendo." Dr. Hooker, to whom I com- 
municated these doubts, thinks that the plant may yet be found wild in North-western 
China, and his intimate knowledge of the Flora of India renders that opinion of the 
utmost value. 

Respecting the varieties of Thea Chinensis, I think that they are supposed to be much 
more marked than they really are. There is indeed a vast difference between the narrow- 
leaved forms of what is called Thea Bohea in our gardens, and the broad-leaved specimens 
of the wild Tea of Assam (figured in our Plate LXI.) ; but the transition from Thea Bohea - 
to Thea viridis, and thence to Thea Assamica, is so gradual, that it is almost impossible to 
draw up any precise definition of these three great varieties. Besides, it must not be sup- 
posed that when we have furnished characters for these three principal varieties, our 
labours have terminated. As in all cultivated plants, there are minor varieties, and forms 
of the highest agricultural and commercial value, that claim our attention, and for the 
working up of which but limited materials are at present at hand. The belief so long 
entertained that Thea Bohea yielded the black, Thea viridis the green tea of commerce, has 
long been exploded; but a slight doubt seems still to linger in some quarters whether the 

fact that Thea Bohea suffers less from frost, and begins to put forth its young leaves later 
than Thea viridis, does not prove a specific difference between them. The reply to this 
is, that Thea viridis of the gardens does certainly begin to grow earlier than Thea Bohea, 
and therefore is apt to suffer more from night frost than Thea Bohea; but it does not 
follow that it must on that account be specifically distinct, as, according to the same rule, | 
we should have to make species of all the early and late sorts of our kitchen vegetables 
and of our fruit-trees *. | 
Species excluse : å 
Thea Assamica, affin. Chois. = P. yrenaria attenuata, | Thea imperialis, Hort.= Micromeria obovata, Bth. 


Seem. : Thea longifolia, Nois.= Camellia Sasanqua, Thunb. 
den amellia, Hoffmsg.— Camellia J aponica, Thea oleosa, Lour.— Camellia Sasanqua, Thunb. 
nn. 


Thea Sasanqua, Nois. = Camellia Sasanqua, Thunb. 


* For an interesting paper on the tea of co he Dirai 
xiv. p. 451 (1858). mmerce, by Charles Wach, see Ed. Otto's * Hamburger | 


GENERA CAMELLIA AND THEA. 301 


ADDENDA. (October 1859.) 


After the foregoing paper had been printed off, a fine specimen of Fortune's * Yellow 
Camellia ” flowered at Kew, affording me an opportunity for examination. It proves to 
be a variety of Camellia Sasanqua, Thunb., of the Warratah or Anemone Class, and may 
be termed 


CAMELLIA SASANQUA, var. y. ANEMONÆFLORA; foliis ovato-lanceolatis longe acuminatis, 
floribus plenis, petalis exterioribus (albis) obovato-oblongis bilobis, staminibus fere 
omnibus in petalos spathulatos (flavos) exterioribus multo breviores mutatis, stylis 5 
(rarissime abortu 4) liberis vel connatis, fructu ..... (v. v. cult.). 

Camellia Sasanqua, Thunb., var. y. anemoneflora, Seem. MSS.; Bot. Mag. t. . (ined.). 

Yellow Camellia, R. Fortune, Journ. to Tea Countr. p. 339 (1852); Gard. Chron. for 1852, p. 259; Seem. 
in Gard. Chron. for 1859, p. 807. 

Fortune’s Gelbe Camellia, Bonplandia, vii. n. 19 (1859). 

«Those who have read my * Wanderings in China, " says Mr. Fortune (Journey to the 
Tea Countries, p.339. London, 1852), * may remember a story I told of my endeavours to 
find a Yellow Camellia,—how I offered five dollars for one,—how a Chinaman soon found 
two instead of one,—and how he got the money and I got taken in! In one of those 
nurseries (about ten or twelve miles eastward of Shanghae), however, I found a Yellow 
Camellia, and it was in full bloom when I bought it. It is certainly a most curious plant, 
although not very handsome. The flowers belong to the Anemone or Warratah Class; 
the outer petals are of a French white, and the inner ones of a primrose-yellow. It 
appears to be a very distinct species in foliage, and may probably turn out more hardy 
than any of its race.” 

Fortune’s supposition that his * Yellow Camellia ” might be a distinct species has not 
been verified, nor is the plant in question to be regarded as å variety of Camellia Japonica. 

The latter flowers towards the spring, and has quite glabrous leaf-buds, petioles, leaves, 

and ovaries; whilst the Yellow Camellia flowers in the autumn, and has pubescent leaf- 

buds, petioles, and veins of the leaves, and a woolly ovary, characters agreeing with 

.. C. Sasanqua, and it may therefore be safely referred to that species as a Warratah or 

Ånemone-flowered variety, the stamens of which are nearly all transformed into short 

spathulate petals of a primrose colour. Hitherto the Warratah state was only known = 

occur in C. Japonica; and the yellow colour is certainly quite a new feature x this 

genus, deserving the greatest attention of horticulturists. That C. mee ur i 

tendency to assume a yellow tinge is evident even from the single flowering state, as 

be seen from the figure in the * Botanical Register, t. 942, where the outer series of sta- 

mens displays the primrose colour peculiar to the Yellow Camellia. I may add that m 

plate quoted represents the form nearest approaching the plant under consideration, the 


leaves of the Yellow Camellia always being more acuminate than those of the ordinary 


form usually found in the gardens, and the styles generally (not always) entirely free. This 


latter particular Fortune's novelty shares with C. Hongkongensis, Seem., and C. drupifera, 


352 DR. SEEMANN’S SYNOPSIS OF THE GENERA CAMELLIA AND THEA. 


Lour. (C. Kissi, Wall.). As a cause of the sickly appearance generally presented by the — 
Yellow Camellia in our gardens, I have assigned (Gardeners’ Chronicle for 1859, p.807) — 
its being grafted upon C. Japonica, and, as a cure, recommended its being grafted upor 


its nearest ally, C. Sasanqua. 


EXPLANATION OF THE PLATES. 
TAB. LX. 


Camellia Hongkongensis, Seem. 


Fig. 1. Petal. Fig. 2. Stamens of the monadelphous series. Fig. 3. Ovary and styles. Fig. 4. Ovary 
cut across. All, with the exception of Fig. 1, magnified. 


Tap. LXI. 


Thea Chinensis, Linn. + 
Fig. 1. Petal. Fig. 2. Stamens of the monadelphous series. Fig. 3. Ovary and styles. Fig. 4. Ovary = 
cut across. Fig. 5. Ripe fruit. Fig. 6. A seed. Figs. 1, 2, 3, and 4 magnified. T 


Vincent Brooks. Imp 


Camellia Hongkongensis. Seem 


fn 331. 


Trans. Linn. Soc Vel. ML tah6! 


Thea Chinensis, Linn: 


ee 


å 


- 


[ 858 J 


XXVI. On Tomopteris onisciformis, Eschscholtz. By Wirra B. CARPENTER, Esq., | 
| M.D., F.R.S., FLS., F. G.S. | 


Read Jan. 20th, 1859. 

THE animal which forms the subjeet of the present communication is one which has 
not very frequently presented itself to the notice of zoological inquirers; and it is not a 
little singular that almost every one under whose observation it has fallen has given 
an account of its structure differing in some essential particulars from that of other 
observers.. This circumstance may of course be readily accounted for on the hypothesis - 

_ that the several specimens examined have belonged not to the same, but to different - 
specific types. I cannot but believe, however, that a careful comparison of the pub- 
lished accounts will lead others, as it has led myself, to the conclusion that all the speci- 
mens described are referable to one and the same type, and that the differences are 
chiefly those of phase or stage of development. For it is remarkable that the accordance 
should be closest in those details of structure which might be expected, on the hypothesis 
of specific difference, to be most liable to vary; whilst the diversities are greatest in those 
features which seem most liable to undergo modification in the progress of development. 

I shall first give a detailed account of my own observations, and shall then compare 
them with the descriptions which I have since found to have been given of this creature 
by those to whom it had already. presented itself. My observations were made in the. 
month of September last, on specimens captured in Lamlash Bay, Arran. I was for- 
tunate in being able at the time to obtain the assistance of Mr. George West, whose 
intelligence and scrupulous accuracy as a microscopic draughtsman are well known to all 
Who have employed him; and the drawings which accompany this communication having 
. been all executed by him with the objects immediately before him, and under my own - 

direction as to the points to which his attention should be specially given, I cannot but 
consider that they have a value much greater than can be attached to many of the — 

_ Sentations heretofore published, most of which seem to me to be little better than ideal 

| diagrams. ; | j z . . e 

The study of this animal, I may say i» limine, presents peculiar difficulties, from its 
incessant restlessness, and from the circumstance that its delicacy is such, that confinement 
is Speedily fatal to it, its tissues and organs exhibiting a manifest tendency to disintegra- 
tion some time before it ceases to move. We had, however, the great advantage of being 
able to make our obsérvations on several specimens in the younger and simpler stage which 
I shall first describe ; and I feel sure that we had accurately mastered all the most important 
features of its structure, before I met with the remarkable, and not a little perplexing form, 
Which I think I cannot be wrong in regarding as a more advanced stage of the same. 

VOL. XXII. : 9A 


354 DR. W. B. CARPENTER ON TOMOPTERIS ONISCIFORMIS. 


The first phase under which the Tomopteris presented itself to me is delineated in ~ 
Tas. LXII. fig. 1, under a power of 10 diameters. Its entire length was about *20 of an inch. 
Its body was elongated, not quite cylindrical, but somewhat flattened, and extremely trans- 
parent; there was no other segmentation than that which was marked by the projection 
of the lateral appendages; but a delicate transverse wrinkling became apparent when a 
sufficient magnifying power was employed, as is indicated in figs. 2, 3, the former showing 
the dorsal, and the latter the ventral surface, under a power of 38 diameters. The part 
which may be designated as the head, from its containing the mouth, the nervous ganglia, 
and the eye-spots, and from its bearing what seem to be the principal organs of feeling, 
is not separated from the body by any constriction ; and the perivisceral cavity is not only - 
continued into it, but extends also into its appendages. These are of a very remarkable 
nature. In front of the head there projects (figs. 2, 3) what might be regarded as a short 
continuation of the body, narrowed into a kind of peduncle ; and this terminates anteriorly 

in a pair of large horn-like appendages that project transversely on either side from their 
common base, somewhat after the fashion of the horns of certain Ruminants. Between 
the foregoing and the head, there arise from the under side of the peduncle a second pair 
of these horn-like appendages, which are smaller than the first; the inferior position of 
these is seen in fig. 4, which gives a front view of the head and its appendages. The 
perivisceral cavity is distinctly continued into the neck and the first pair of horns; but I 
could not satisfy myself that it penetrated into the second. From the sides of the head 
itself there projected a far more remarkable pair of appendages, nearly half the length of 
the body (fig.1), which may for convenience be distinguished as the styliform. These 
were supported at the base by a lateral projection of the head on either side (figs. 2, 3), into 
which the perivisceral cavity extended, and which also contained the muscular appa- 
ratus that acted on the styliform appendage itself. This consisted of a long slender rod, 
which, though obviously very firm in texture, was not inflexible, and which, therefore, I 
should judge to be horny rather than calcareous. This rod was invested by a prolonged 
membranous sac, tolerably wide at the base, but gradually narrowing towards the apex, 
which appeared to close-in around the base of the rod, so that its cavity seemed com- 
pletely isolated from the perivisceral cavity in which it lay; and it was to each side of 
this membranous envelope that the muscular bands were attached, which gave free 
motion to the styliform appendage. ila 
The body bore {ex pairs of lateral appendages, of which the third was the largest; —— 
the size of the 2nd being about the same as that of the 5th and 6th; that of the Ist 
about the same as that of the 7th; and that of the 8th somewhat smaller; whilst that — 
of the 9th was not above half that of the rest; and the 10th (of whose character I was 
not at first by any means certain) was merely rudimentary. Each of these appendages 
consisted atits base of a nearly cylindrical prolongation of the body, transversely wrinkled 
like it, and containing an extension of the perivisceral cavity ; this basal portion gradu- 
ally narrowed itself, and then underwent bifurcation into two lobes, each of which sup- 
ported a flattened fin-like expansion of somewhat oval form. In this expansion å sort of 
A a could be seen with a sufficient magnifying power (figs. 2 4, 9 a). = : 
position these two fins seemed to be nearly at right angles to each other; v 


DR. W. B. CARPENTER ON TOMOPTERIS ONISCIFORMIS. 355 


and by their incessant action the animal kept up a rapid swimming movement, varying 
continually in its direction, and very much resembling that of the Branchiopod Ento- 
mostraca, the strokes of the several appendages succeeding each other from before 
backwards. | | | 

Owing to the glass-like pellucidity of. this interesting little creature, its interior struc- 
ture can be made out as well as its continual restlessness allows; a further allowance 
having to be made for the circumstance, that owing to the difficulty occasioned by this 
very transparency in judging as to what is near and what remote, whilst its restlessness 
necessitates a continual shifting of the focal adjustment, the relative position of its in- 
ternal parts cannot always be determined without some liability to error. 

The mouth ordinarily opens by an elongated slit (fig. 3) on the underside of the head, 
into a thick-walled and apparently muscular pharyngeal cavity. This is shown in fig. 3 to 
be connected with the wall of the body by two bands on either side, one passing anteriorly 

- and the other posteriorly ; and it can scarcely be doubted that these are muscles for its 
protrusion and retraction, since I occasionally saw the pharynx protruded as a proboscis, 
after the fashion of many Annelids. About half-way between the styliform appendages 
and the first pair of fin-bearing appendages, the pharynx terminated by a well-marked 
constriction in the intestinal tube, which was a straight and simple canal whose diameter 
was usually no more than about a quarter of that of the body itself, and passed direct to 
the posterior extremity of the body, where it terminated with a pouch-like dilatation in 
the anal orifice (fig. 5). I never saw any solid matter in this canal; but it was frequently : 
distended in parts by water, a wave of which would occasionally pass peristaltically from 
one extremity to the other. When thus distended, its wall could plainly be seen to be 
chiefly composed of ovoidal cells very compactly arranged, as shown in figs. 8,5. No 
indications whatever of a dorsal vessel could be distinguished ; but the fluid which occupied 
the large perivisceral space of the body, head, and appendages, could be seen to contain 
within it minute semipellucid granules of irregular form and size, by whose movements it 
was made obvious that this fluid was continually shifting its place,—rather in conse- 
quence, however (as it appeared to me), of the general movements of the body, than of any 
more special provision for its circulation. . 

I could not detect any organs of respiration, 
any part of the surface. For the reasons already 
upon myself to affirm its non-existence. | | 

In the head there could be plainly distinguished a bilobed mass having all the appear- 
ance of a nervous ganglion; and upon this lay two little masses of pigmentary matter ‘ 
each of which bore a small pellucid lens-like body (fig. 2). There can scarcely, I think, 
be a reasonable doubt of these being ocelli. From the central portion of the ganglionic 
mass, I thought that I could distinguish something very like the axis-band ofa nerve-fibre 
without its tubular sheath, passing backwards along the dorsal surface of the body, keep- 
ing near to the median line, but not exactly upon it, and passing at intervals not very 
regular through red spots, which seemed like aggregations of granules, or very minute 
cells, and of which there were commonly six or seven in each of the divisions of the body. 
From each side of the bilobed ganglion I thought that I could trace a 


and I did not observe ciliary motion on 
mentioned, however, I would not take 


similar fibre pass- 
3A2' 


356 DR. W. B. CARPENTER ON TOMOPTERIS ONISCIFORMIS. 


ing to the styliform appendage; and the membranous sheath of its stylet was studded at 
intervals with isolated red granules or minute cells, which appeared to me to be connected 
by delicate fibres having the same general resemblance to the axis-band of ordinary nerve- 
fibres. I do not speak with any confidence on this point; more especially on account of 
the å priori improbability of the longitudinal nerve-cord being on the dorsal aspect of the 
body. But I think it right to record the fact of the constant presence of these red spots 
(of which very little notice has been previously taken), and of the apparent existence of 
fibres passing between them. 

The impression at first left on my mind by the study of this interesting creature, of 
which I had never before seen or heard any account, was that it was a larval form, per- 
haps of an Annelid, but not impossibly of some Isopod Crustacean ; the latter notion . 
being suggested by its strong resemblance in habit to the Branchiopod forms of that 
class. I tried every expedient I could think of, for preserving specimens for subsequent 
examination and exhibition; but I could not succeed. Any medium in the slightest 
degree different in density from sea-water caused either an immediate endosmose, by 
which the body was made to burst, generally at the ends of the members; or an exosmose, 
by which it was at once shrivelled-up; and the tendency to disintegration which I have 
already noticed as showing itself even during life, operated still more strongly so soon as 
life became extinct. vo 

Not many days after I had made the observations already detailed, I captured a speci- 

men of larger size, having twelve pairs of fin-like appendages, and a caudal prolongation. 
destitute of appendages,—its structure being in every other respect the same. And after _ 
a few days more I was fortunate enough to obtain (along with several specimens of the 
younger form) the specimen represented in fig. 6, which can scarcely be considered as 
anything else than another phase of the same type. Its entire length was about 24 times 
as great; but of this only about the anterior three-fifths corresponded in structure to the 
original, the posterior two-fifths being obviously an addition formed upon a very different 
plan. Looking first to the anterior portion, we observe that it differs from fig. 1 in the 
following particulars. The head appears to carry on its front only one pair of horn-like 
appendages; a careful examination, however, shows that the second or smaller pair is 
really present, though, from its arising from the under side of the peduncle (which is 
here relatively shorter), it is concealed by the anterior part of the head when viewed from 
the dorsal aspect. The styliform appendages are greatly increased in relative length, soas _ 
even to pass considerably beyond the anterior three-fifths of the body. | | 


The number of fin-bearing ppendages is now siæteen; of these the first eight are nearly 


of equal size; and there is then a gradual diminution down to the 16th, which, though 
comparatively small, present 


+ Presents the structure characteristic of the rest. The alimentary 
canal, instead of being limited to a portion of the cavity of the body, now fills it entirely, 
except in the pharyngeal region; and it' seems to be in a state of constant distension, 
except where narrowed at 


ally passes from the mouth to the anus. 
13th pairs of fin-like appendages, and is s 
structure of the canal remains quite 


Such a narrowing is seen between the 12th and 
hown on a larger scale at the top of fig. 7. The 
simple as far as the 16th pair of appendages, and 


any part by a wave of peristaltic contraction which occasion 


DR. W. B. CARPENTER ON TOMOPTERIS ONISCIFORMIS. | 357 


elosely resembles that which is seen in the narrow canal of the earlier phase when partially 
. distended with fluid. | * 
The posterior portion of the body, however, constituting about two-fifths of its entire 
length, presents a number of very marked differences in conformation. Its aspect is more 
eylindrical and worm-like; its parietes are relatively thicker and stronger, and their 
. transverse wrinklings far more decided. It bears eight pairs of slightly-developed ap- 
pendages, which might at first sight be supposed (especially in regard to their terminal 
bifurcation) to be rudimentary forms of the fin-feet of the anterior part of the body, but 
which a more careful examination will show to be formed upon a different plan; for 
whilst the appendages that bear the fin-feet, as already described, contain extensions of 
the general cavity of the body, the parietes of which correspond with those of the body 
itself both in their tenuity and in their delicate transverse wrinkling, those of this vermi- 
form continuation are solid tubercles of ovoidal shape, put forth from the comparatively 
thick ‘integument, and which do not receive any prolongation of the perivisceral cavity. 
On reverting to figs. 1 and 5, it will be seen that in that period of the animal's life at which 
itis most rapidly undergoing increase upon the ordinary type, the last and consequently 
least developed pair of appendages is still formed upon the same plan with those anterior 
to it; and the same is the case with the 16th pair in the more advanced form we are now 
considering. If the 17th and seven succeeding pairs appertaining to this vermiform con- 
tinuation had any relation to the preceding, we should expect that tliere would be some 
gradation from one type to the other; so far is this, however, from being the case, that 
the 17th pair (the first of the vermiform continuation) shows the most decided unlikeness 
fo the 16th (the last of the anterior series) ; the 18th, 19th, 20th, and 21st are of nearly 
the same size and aspect; whilst the 22nd, 23rd, and 24th become progressively smaller, 
the last pair being scarcely distinguishable. But further, at the very same point at which 
this change manifests itself in the condition of the body and of the appendages, there is a 
very marked change in the condition of the intestinal canal; for this, instead of being | 
continued upon the straight and simple plan which so remarkably characterizes it in the 
anterior portion of the body, abruptly begins to present the appearance represented in 
figs. 6,7. Whether this appearance is due to convolution or to sacculation, I am unable 
to state with certainty. I was unwilling to submit my only specimen to the compression 
Which would have been requisite to ascertain this point, until I had secured a good | 
` delineation of it; and as it died and began rapidly to disintegrate before I could subject — 
it to further investigation, I was deprived of the opportunity of subsequently doing 80. 
The extreme delicacy of structure and the low grade of development of the earlier 
phase of this interesting creature having strongly suggested to me that it wasa larval 
form, the resemblance presented by the vermiform continuation, in the more advanced 
specimen, to the ordinary Annelidan type, together with its higher development and its 
appearance of greater permanence, led me to conjecture that this posterior portion would 
become the true type of the species, the anterior probably separating itself from Jt So 
soon as it should have acquired the power of sustaining itself independently, after the 
fashion of the Bipinnaria-zooid of the Star-fish. And notwithstanding that the informa- 
tion I have subsequently received as to the phenomena witnessed by other observers 


ES. 
e 


358 ' DR. W. B. CARPENTER ON TOMOPTERIS ONISCIFORMIS. 


has thrown much doubt on this notion, yet I think it right to record it, for the sake of 


the suggestion it may afford to those who may have the opportunity of following up the _ 


inquiry. Certain it is, that if all the accounts of it be correct records of actual phe- 
nomena, and if they all refer (which, for the reasons I have already stated, I can scarcely 


doubt) to the same specifie type, the creature's life-history must vary considerably in 


different circumstances. | : | 28 
Before referring to the published observations, I shall take advantage of the kindness 


of Prof. Huxley, who has given me permission to append to my own account of Tomo- 
pteris his descriptive notes and figures of two specimens captured by him in Torres Strait, 
during the voyage of the * Rattlesnake,’ Aug. 1, 1849 :— | 

* Body elongated, perfectly colourless and transparent, with thirty-one pairs of lateral 
appendages, exclusive of what may be termed the head (fig. 8). The body is transversely 


wrinkled, but not, properly speaking, annulated, there being a great many rings between 


every two pairs of appendages, especially anteriorly. Posteriorly the wrinklings become 
coarser and wider, but they are still numerous in proportion to the appendages, and are 
not true annulations. | 

* What may be considered to be the head, on account of its containing the mouth and 
the organs of sense, is provided with two pairs of appendages. The first pair terminate 
the anterior extremity of the body, the two appendages being united by their base, and 
disposed transversely. "They give the animal the appearance of a hammer-headed shark. 
Their anterior edge is thin and delicate, the posterior rounded ; and this part contains a 
cavity continuous with that of the body. | 

“ Behind the first pair of appendages the body suddenly narrows into a sort of neck, 


which expands again into two other transverse processes stouter than the former. Each | 


of these is prolonged at its extremity into a long, slender, curved process nearly equal to 
half the length of the body, and this process contains a still more slender cylindrical 
spine, apparently of a horny nature. The end of the spine, covered by a membrane, pro- 


jects into the cavity of the wide base of the appendage, and its extremity there gives - 


attachment to several bundles of muscular fibres, which go to be attached to the sides of 
the cavity, and move the spine on occasion. | 


“ In the cavity of the body, situated just where the narrow neck expands into the space — 


which lies between the second pair of appendages, is a transversely elongated mass, which 
appears to be divided into two parts by a central line. Towards the outer side of each of 


these parts, there is, upon the dorsal surface, a rounded mass of black pigment, and, im- å 


bedded in the outer edge of this, a spherical (?), transparent, crystalline body ; these are 


doubtless eye-spots, and the mass a double ganglion. Two delicate cords connect this 


double ganglion with the parietes, and I think I could trace a bundle of fibres running 
down on each side of the mouth ; but I am not certain about this, and I am quite sure 
there was no chain of ganglia along the ventral surface of the body. x 

* In the cavity of the body likewise, and extending between the second pair of append- 


ages of the head and the first pair of the body, is a narrow, thick-coated, pyriform phe 


ys It commences in a rounded oval aperture on the ventral surface, and terminates 
’y Opening into the wide stomach. This has much thinner walls, and continues a8 # 


DR. W. B. CARPENTER ON TOMOPTERIS ONISCIFORMIS. 359 


plain tube as far as the first fourteen or fifteen pairs of appendages. It then becomes 
transversely folded and wrinkled, and thus passes, preserving a very even diameter, to 
the end of the body. | | | 
« The cavity of the body contains, floating about between the intestine and the parietes, 
certain rounded free masses. These are made up of other more or less rounded masses, 
flattened when their sides are applied to one another (fig. 9), which have perfectly the 
structure of ova. These ova had a diameter of z37th of an inch, and less. In those of 
the former size the germinal vesicle measured 35th of an inch, and was clear and delicate. 
The germinal spot, vesicular and thick-walled, measured ygs5th of an inch. 
“The appendages are hollow processes of the body, and their cavities are continuous 
with that of the body, as was evident from the passage of the ova from the one into the 
other. They increased in size to the fourth or fifth pair, and diminished again from the 
seventh or eighth. The first sixteen or seventeen pairs consisted of a stout basal portion 
terminated by two divisions, each of which was provided with a flat, vertical oar-like 
expansion. The remaining appendages became smaller and smaller and more rudi- 
mentary. The anterior ones were provided with two conical processes merely, while the 
posterior ones were themselves nothing but short simple processes. There was a short 
space between the last pair of processes, which were mere buds, and the truncated anal 
extremity. | | 
“At the re-entering angle between the second pair of cephalic appendages and the 
narrow neck, there is, on each side, a rounded elevation, from which a sort of band or 
ridge runs back upon the dorsal surface. i 

“On the ventral surface, close to the two rounded elevations, a long, curved, spine-like 
process arises upon each side. In their natural position these two processes lie parallel 
with one another, one on each side of the mouth. i 

“ Aug. 28th.—A small specimen obtained to-day appears to be a male, for it contained 
masses of round cells, each rather more than s¢yoth of an inch in diameter, in the place 
of the masses of ova of the previous specimen; these were perhaps young spermatozoa. 
In other particulars its structure agreed with the for ing.” i 2, 

It is obvious that Professor Huxley’s speeimen must have been in a more advanced | 
stage of development than mine, since it had no fewer than thirty-three pairs of append- 
ages, and was also maturing its sexual organs. His figure and deseription indicate that 
the conformation both of the body and extremities underwent à change at about the six- 
teenth or seventeenth pair; but the transition seems to have been far less abrupt than it 
Was in my specimen; so that he does not appear to have been struck with any very 
decided difference in the conformation of the anterior and posterior parts of the As 
Ånd his figure of one of the appendages of the latter would seem to indicate "ue "— 
more advanced condition these approach more nearly to the type of those of the an , 


than they seemed to me likely to do. 
| 12:34 aeiicunpiotd 


chscholtz, who briefly characterized and 


T. . Se, . 1 
omopteris onisciformis was first described by Es assigning to it a place 


rudely delineated it in the ‘Isis’ for 1825 (p. 735, pl. 9 fig. 9), 


-860 DR. W. B. CARPENTER ON TOMOPTERIS ONISCIFORMIS. 


among Heteropodous Mollusks. He would appear to have met with it only in its earlier 


phase; for his figure shows no more than twelve pairs of lateral appendages, without any 
caudal prolongation. | A | di 
It seems, however, to have been previously (?) observed by Quoy and Gaimard, who 
met with itin the Bay of Gibraltar during the voyage of the * Astrolabe;' but as their 
account of it was first published in 1827 (Annales des Sciences Naturelles, tom. x. p. 235), 
the name Briarea scolopendra, by which they designated it, must yield in priority to that 
of Eschscholtz. The specimen which they describe as having been taken in the Bay of 
Gibraltar, is stated to have attained a length of 4inches. If there be no error in this - 
description, it must have been far larger than any that has been elsewhere met with. 
They speak of it as possessing 24 or 25 pairs of lateral appendages or fin-feet ; of these, - 
however, their figure shows only 21 pairs, of which 17 belong to the body, and 4 to the 
caudal prolongation. If this figure (which strikes me as in many respects rather an 
ideal diagram than a true representation) approaches in any degree to the real propor- 
tions of their specimen, its lateral appendages must have been very much longer than 
those figured by any of the other naturalists who have described it, their maximum 
length being in the anterior portion of the body, and a progressive diminution taking 
place as far as the commencement of the caudal prolongation, behind which they are 
merely rudimentary. Thus the outside contour of the entire animal is not very dissimilar 
to that of a boy's kite, the caudal prolongation representing the tail. It is somewhat 
singular, that, notwithstanding the extreme transparency of this animal, MM. Quoy and 
Gaimard were unable to make out its alimentary canal, although they described and 


figured what they believed to be ova. They considered Briarea to be a Mollusk, nearly 
allied to Glaucus: 

In Müller’s Archiv for 1847, there are a description and a figure of Tomopteris onisci- 
Jormis by Busch, who met with it in the North Sea. This figure nearly corresponds, 


_ except in the proportional shortness of the lateral appendages, with that of MM. Quoy — 


and Gaimard. The number of these appendages which Busch represents is 18 pairs, the 
form of all being the same, and their size diminishing gradually from the 1st pair to 


the 18th, which is close to the posterior extremity of the body, there being no distinct — 
caudal prolongation. Busch described and figured the ova in the perivisceral cavity, 3$ —— 


Professor Huxley has done subsequently ; and he also noticed other bodies which seem = 
to correspond with what Professor Huxley regarded as bundles of spermatozoa. We 

In the sueceeding volume of the same Journal (1848) is a very elaborate memoir by 
Grube on Tomopteris, based on specimens collected by Krohn (probably in the Medi- 
terranean), and preserved in the Museum of St. Petersburg. This memoir is specially 
devoted to the description of some of the minutest details of the structure of the animal, 
and to the inquiry into its place in the zoological scale. Grube’s figure of the entireanimal — 
IS not only small in scale, but is somewhat rudely sketched ; it represents twenty pairs of ; e 


lateral appendages, and a caudal prolongation of cylindrical form, apparently without * 


any appendages at all. In his description, however, the author speaks of this caudal pro- — 


longation as bearing rudimentary fins, in the form of whitish protuberances; these, Él 


remarks, are as yet undivided ; whilst the fully-developed appendages of the anterior part — 


DR. W. B. CARPENTER ON TOMOPTERIS ONISCIFORMIS. 861 


. of the body are bifid, and in large specimens the lobes of the posterior not-fully-developed 
appendages are relatively larger than the former. One of the most important parts 
of Grube's memoir relates to the nervous system. He asserts that he has distinctly seen, 
in many specimens, a double nervous cord passing backwards along the ventral surface of 
the body from an cesophageal ring, of which the upper part is formed by the cephalic 
ganglia. This double cord, he says, does not anywhere present ganglionic enlargements ; 
but, when examined under a high magnifying power, it presents a series of transverse 
striæ at short intervals. I cannot but express some doubt as to the existence of this 
ventral cord; for Grube’s description of it is entirely based upon preserved specimens * ; 
and although I looked for it carefully and repeatedly in living specimens, I was not able 
to detect it. Grube's description of the muscular structure also, especially in the fins, is 
very minute; but I can state confidently that this presents itself under a very different 
aspect during life; indeed, as to the general histology of the creature, I was struck with 
the remarkable change which it seemed to undergo when the animal appeared to be 
suffering from confinement, but while as yet far from having lost its vital activity. 

The most recent notice of Tomopteris is that given of it by Mr. Gosse in his * Natu- 
_ralist’s Rambles on the Devonshire Coast’ (p.356), under the name of Johnstonella 
Catharina; it adds nothing, however, to what had been previously recorded, excepting as 
to the occasional eversion of the pharynx so as to form a protruded proboscis, which 
Mr. Gosse seems to have been the first to observe. His specimens appear to have been 
in the same stage of development as my most advanced specimen, having 16 pairs of 
fin-like appendages, and a caudal prolongation without any obvious appendage. He - 
seems to have overlooked the posterior pair of frontal * horns," which, for the reason I 
have stated, are not to be seen in the larger specimens unless carefully looked for. In 
his ‘Marine Zoology ” (vol. i. p. 106), Mr. Gosse has introduced Tomopteris (by its proper 
designation) amongst the * animals of doubtful position, probably belonging to the Anne- 
lida ;” and he has given a figure of it, which, like the figure in his * Devonshire Coast, 
wants the second pair of frontal * horns," and which is further erroneous as giving not 
_ the least indication of the bifid character of the fin-like appendages. 


There cannot, I think, be any longer a reasonable doubt as to the Annelidan character 
of Tomopteris; and the principal point which now requires investigation seems to me to 
be the question of the real nature of the caudal prolongation, —that is, whether it is 
simply an extension of the body, or is a zooid destined to be cast off and to maintain an 
independent existence. The former view seems to be that of all previous observers ; no 
such marked peculiarity having been observed by them, either in the conformation of its 
appendages, or in the structure of its integument and of the portion of the alimentary 
canal included in it, as that which seemed to me to differentiate it completely from the 
anterior part of the body, and to bring it nearer to the ordinary Annelidan type. I speak 
With diffidence, however, on these points, since I find myself at issue In regard to them 


* It would be interesting to know in what fluid these specimens were mounted ; as I tried all the ordinary media 
without succeeding in preserving in any specimens an approach to their life-like appearance. : 
VOL. XXII. B 


.. Fig. 4. 


362 DR. W. B. CARPENTER ON TOMOPTERIS ONISCIFORMIS. » 


with observers of unquestionable competency. And if, as would seem, scarcely to admit . 
of doubt, the process of sexual generation is carried on in the portion which I imagined 

to be larval, such a notion of its character becomes obviously untenable. I have deemed 

it right, however; to record my first impression, that any observers to whom the animal: : 
may present itself may have their attention more strongly directed to the peculiarities of. 
structure by which it was suggested. And as the animal does not seem to be very uncom- 
mon on our coasts (having been taken by the late Dr. Robert Ball in the Bay of Dublin, 
and by Dr. 8. J. Salter in Poole Harbour, as well as by Mr. Gosse and myself), it may be 
hoped that further light may ere long be thrown upon this question, as well as upon the - 
nature and distribution of the nervous system of this beautiful and interesting creature. - 


[SUPPLEMENTAL Nore.—Subsequently to the reading of the foregoing paper, another 
memoir on Tomopteris has been published in Miller's Archiv, 1858, p. 588, by Drs. 
Leuckart and Pagenstecher. These observers describe, under the name of Tomopteris 
onisciforinis, a. specimen with twelve pairs of fin feet and a caudal prolongation destitute 
of appendages, corresponding closely in its grade of development with a specimen I have 
mentioned in p.354. In the perivisceral cavity of the anterior part of -its body, they 
distinguished sexual products, as Busch and Huxley had done. They also describe, under 
the name of Tomopteris quadricornis, a smaller specimen, having only ten pairs of fin feet, 
and obviously identical with those first seen by me; this they consider to be specifically 


distinguished from the preceding by the presence of two pairs of horn-like appendages, 


the T. onisciformis seeming to have but one. I have already mentioned, however, 
that the second pair really exists in the larger specimens as in the smaller (p: 354); 80 _ 
that there is no reason for regarding the two as otherwise than specifically identical, the ' 
difference in the number of fin feet being obviously a character of age merely.] : 


EXPLANATION OF THE PLATE. 
fas. LXE 


Fig. 1. Young specimen of Tomopteris with ten pairs of fin feet; enlarged 10 diameters; — ^ d 
Fig. 2. Head ‘and i anterior part of the body, as seen upon the dorsal aspect, enlarged 38 diameters + s 
: 4, bifid extremity of one of the fin feet. TAR of: ng GE M 
Fig. 3. The same, as seen upon the ventral surface. . d. SE . 
Front view of the head, showing the position of the two anterior pairs of cephalic appendages, and bs 
TE the opening of the mouth upon the ventral surface. . EG UT 
Fig. 5. Caudal ‘extremity, showing the last two pairs of least-developed fin feet, and the termination of - = 
the intestine in an anal orifice; io 


Fig. 6. 


ed Advanced specimen of Tomopteris with sixteen pairs of fin feet; enlarged 10 diameters. 


sn extremity of the body of the specimen represented in fig: 6, with its last four pairs. å 1 
ordinary fin feet, giving origin to à caudal prolongation of very: différent conformation, furnis THE 
with eight pairs of rudimentary appendages. ipto dg 
Fig 8. Specimen of Tomopteris with thirty-three pairs of ebene XERaréy). 
Fig. 9. Masses of ova in the perivisceral cavity of the preceding (Huzley). 


Frans Linn Soo Vol II vad 68 7.808. 


ie ve ue ET rr n 


iren. 


Jarman, sc. 


fr 


ferro 


Vrve West del. 


[ 363 ] 


. XXVII. On some new forms of Entozoa. By T. SPENCER COBBOLD, Es9,, MD., F.L.S. 
Read January 20th, 1859, 


Br permission of the authorities of the Zoological Society, I have enjoyed the opportunity 
of examining a large proportion of such animals as have died at the Society’s Gardens, 
Regent’s Park, during the spring and winter months of last season. 

The materials thus afforded for a search after Entozoa consisted of six species of Reptiles, 
thirty-one of Birds, and the same number of Mammals. With three duplicate exceptions 
in the case of birds, and one in mammals, only a single example of each species came 
under notice, thus giving a total of seventy-two animals subjected to dissection. Of these, | 
but three reptiles, nine birds, and ten mammals were found infested,—a result of con- 
siderable interest to.the Zoological Society, as it proves these animals to enjoy greater 
freedom from internal parasites in their condition of. domesticity than in the wild state. 
This indeed is precisely what we should expect, as it is well known that the larvæ of Entozoa 
occur.only in a limited number of hosts; and it is equally evident, in the case of Trematoda 
for example, that the uninfested foreign animals can have had no opportunity of devour- 
ing the various forms of molluscan hosts which under ordinary circumstances supply the 
cercariæ destined to become adult flukes in their viscera. 


1. DistomA comPACTUM (mihi). Body smooth, ovate, oblong, not compressed ; oral sucker 
terminal, orbieular; ventral sucker subcentral, aperture triangular ; reproductive 
pore immediately below, a little to the left. Length j-j in.; breadth 3-3 in. 
(Pl. LXIII. figs. i, 3, 3.) HÆ oun gat 
—. Remarks.—On the 19th of February I removed five specimens. of this trematode from 
pustular cavities in the left lung of an Indian Ichneumon (Viverra mungos, L.). All con- 
.. Jecture as to the source of its larvae must be useless; but the adult form is well marked, 
and easily recognized by the compact condition and arrangement of the internal organs, It 
is provided with a short æsophageal bulb, giving off two simple, tortuous and unusually 
broad digestive canals. The twisting is so marked and regular as to approach the peculiar 
zigzag condition of Campula—a genus which I have recently established (Linn. Trans. 
Vol. xxii, part 3. p.168). The vitelline cæca almost entirely cover the lateral and dorsal - 
surfaces, and are connected in the middle line by two primary transverse branches ; these 
Again unite to form à trunk which opens into the base of the short and folded uterine 
tube. The testes lie directly below. hk Wl 
The nearest resemblance to this species is a fluke discovered by N atterer at Matogrosso, 
in cavities of the lungs of the American Otter. Under the title of Distomum rude, Diesing 


has described and figured it in his ‘ Neunzehn Arten von Trematoden : p. o ge. 


364 - DR. T. S. COBBOLD ON SOME NEW FORMS OF ENTOZOA. 


2. Distoma MINUTUM (mihi). Body flat, smooth, oblong, narrowed in front; oral sucker 


oval, subterminal ; ventral sucker orbicular, central. Length 135 1% in.; breadth 
147-135 in. (Figs. 4, 5.) | ( 
Remarks.—This exceedingly small trematode was found in great abundance in the 
duodenum of an Oyster-cateher (Hæmatopus ostrealegus, L.), and could scarcely be dis- 
cerned with the naked eye. . Its form is entirely different from D. brevicolle described by 
Creplin as infesting this bird; moreover the latter is a much larger species. The pre- — 
sence of a long uterine canal crowded with ova showed its mature character, while few 
traces of other organs were to be seen. I did not succeed in detecting with certainty the 
position of the genital pore, but appearances seemed to indicate its presence directly above 
and to the right of the ventral sucker. : 


3. Distoma Boscı (mihi) Body subclavate, covered with minute spines, slightly eom- — 
pressed, thickened below; oral sucker orbicular, subterminal; genital pore above the 
ventral acetabulum. Length 4-4 in.; breadth 73—% in. (Figs. 6, 7.) 

Fasciola colubri, Bosc, Hist. Nat. des Vers, i. p. 271, with figs. : 

Distoma colubri Americani, Rudolphi, Entoz. Hist. vol. ii. p. 434; and Synops. p.121; Diesing, Syst. 
Helminth. vol. i. p. 398. e | : | 

Remarks.—This species has hitherto been only very briefly noticed, and is placed by 

Diesing among doubtful and imperfectly-described forms. Like Bose, I found several — 

specimens in the buccal cavity of an American serpent of the genus Coluber, the specific 

name of which I am unable to give. The worm is provided with a muscular oesophageal — 
bulb, two simple digestive cæca, and largely-developed reproductive organs. 


4. BILHARZIA MAGNA (mihi) Body smooth, linear, gradually narrowed anteriorly ; oral | 
sucker oval, subterminal; ventral acetabulum round, very prominent. . Length — 
upwards of 1 in. (Figs. 8, 9.) ! EG E 

| Remarks.—Up to the time of Bilharz’s discovery of Distoma hematobium, all the flukes — 
were considered hermaphrodite, and I am not aware that any other species excepting the Jå 

above has been since found to present a deviation from the general type. Trematodes 2 

regn: ic marked a structural peculiarity as D. hæmatobium, associated as it is with — 

the presence of a gyncecophoric canal in the male, deserve, I think, to be generically sepa- = 
rated from Distomata properly so called, and I have therefore employed the indefatigable — 
discoverer’s name for this purpose. Only a single male example of the species here recorded 
jin observ ed by me, while engaged in the dissection of a Sooty Monkey (Cer copithecus — 

P uliginosus), in some blood which had escaped from the divided portal veins. It isa mue 

larger species than Bilharzia (Distoma) hæmatobia ; Sut us precise length I am unable — 

to record, as å portion of the caudal extremity had been aceidentally removed. Unfortu- — 
nately it was at the time regarded as a common nematode; but hud its true nature vr 


earlier detected, a more careful examination of the blood-vessels would probably d 
brought other individuals to light. | 


DR. T. S. COBBOLD ON SOME NEW FORMS OF ENTOZOA. 365 


. 5. ASCARIS TRIBOTHRIOIDES (mihi). Head truncate, with three sucker-like processes ; 
neck constricted; body uniformly linear; tail conical, with a short blunt extremity. 
Length about lin. (Figs. 10,11.) 


Remarks.—1 am not certain that this eccentric-looking nematode should be considered 

a true Ascaris. Two examples only were detected in the small intestine of a Dusky 
Duck (Anas obscwrus), and neither of these was sufficiently fresh to exhibit its internal 
structure to advantage. ; 


6. Canurus. From a specimen of the Ring-tailed Lemur of Madagascar (Lemur maco), 
which died at the Zoological Gardens on the 30th December 1857, and which had been in 
this country only four months, I procured a remarkable series of hydatid-like cænuri 
(fig..12). They existed in such abundance in the liver, and on both sides of the thorax, 
as to become the immediate cause of the animal’s death. Those in the chest were con- 
nected to the pleura, and occurred in semitransparent pedunculated masses, split up, as it 
were, into numerous lobules, the entire parts of each separate bunch being connected 
together and to the surface of the extremely atrophied lungs by short pedicles. Here 

-and there small colonies, consisting of only one or two lobules, were in process of develop- 
ment. Each lobe presented a variable number of small, round, papillary elevations, which 
in some places assumed a more or less regular linear arrangement. Under a low magnify- 
ing power, the surfaces of the imperfectly-formed papillæ exhibited a central oval depres- 
sion (fig. 13), while each of the more completely developed eminences was found, on dissec- 
tion, to contain a single well-formed tape-worm head. Enlarged forty diameters, every 
head displayed four suckers and a short proboscis armed with thirty-two hooks disposed 
in two rows (figs. 14, 15). No loose scolices occupied the interior of the lobules, which 
were filled, however, with a pale-yellow serous fluid. 


Appendix.—Among known forms of Entozoa, I may mention the occurrence of Tenia 
paradoxa in the Oyster-catcher (figs. 16-19 inclusive); and in the Dusky Duck (Anas 
obscurus) of numerous partially decomposed Tenia, referable, I think, to T. lanceolata 
(fig. 20). From the duodenum of a Night Heron (Ardea nyctocorax) I obtained ten or | 
twelve examples of Tænia multiformis (fig. 23) ; also three fine specimens of Hustrongylus 
- papillosus (fig. 24) from the mouth of a Crane (Grus antigone). From the small intestine 
of an American Barn Owl (Strix perlata) were procured nine individuals of Distoma equale, 
and from a Horned Pheasant several examples of Tenia infundibuliformis (fig. 25). The 
cæca of a Ring-necked Pheasant (Phasianus torquatus) were crowded with Ascaris vesicu- 
laris (fig. 21); and the intestines of a Sandwich Island Goose contained several worms very 
like Heterakis dispar (figs. 26 and 27), besides other nematodes requiring further investi- 
gation. The liver of an Axis Deer (Cervus axis) contained a few degenerated Cercarie, 
While a large aborted Cenurus or acéphalocyst occupied the left lung of a Goat (Aries 
 fragelaphus). Several nematodes (fig. 28) were obtained from the lungs of a Peccary 
(Dycoteles torquatus), while the rectum of a Weasel-headed Armadillo (Dasypus sea- 
, Cinctus) yielded several specimens of Ascaris retusa (fig. 22). From the lungs of pi 
Four-horned Antelope (Antilope quadricornis) I procured several very large opm o 


366 DR. T: a. COBBOLD ON SOME NEW FORMS OF ENTOZOA. 


Echinococcus veterinorum (altricipariens of Kiichenmeister), precisely like those described 
by Professor Huxley from the Zebra. I also procured some nematodes (figs. 29 and 30) = 
and cestodes (fig. 31) from an Indian Rat Snake (Coluber Blumenbachii) measuring six 
feet long, as well as a single round worm from an Indian Lizard (Calotes versicolor), 
which, together with another large Ascaris (figs. 32 and 33) from the intestine of the 
American Snake formerly alluded to, I have not at present been able to identify. 


— DESCRIPTION OF THE PLATE. 
| Tas. LXIII. 


Fig. 1. Distoma compactum. Enlarged ith. 
Fig. 2. Another example. x 5 diameters. 
Fig. 3. Egg of the same. x 360 diameters. TIU 
Fig. 4. A few drops of mucus, &c., taken from the duodenum of the Oyster-catcher, and spread on a. 
thin square of glass. It shows several small flukes. Nat. size. 
Fig. 5. Distoma minutum, from the same. x 180 diameters. 
Fig. 6. Distoma Bosci. x 8 diameters. 
Fig. 7. Egg of the same. x 200 diameters. | 
Fig. 8. Bilharzia magna. Nat. size. Assumed proportion of the lost end of the tail in outline. 
Fig. 9. Upper two-thirds of the same. x 10 diameters. — 
Fig. 10. Head of Ascaris tribothrioides. x 160 diameters. 
Fig. 11. Tail of the same. x 160 diameters. 
Fig. 12. Colony of Cenuri from Lemur maco. Nat. size, 
Fig. 13. Part of an immature lobule. x 20 diameters. 
Fig. 14. Tape-worm head from ‘one of the mature papillæ. x 40 diameters. 
Fig. 15. Hooks from the same. x 260 diameters. 
Fig. 16. Tenia paradoxa. Nat. size. * > 
_ Fig. 17. Head of the same, x 260 diameters. _ : 
Fig. 18. Everted extremity of the rostrum.. x 260 diameters. 
Fig. 19. Upper joints of the same. x 150 diameters. | 
Fig. 20. Tape-worm head. T. lanceolata? x 250 diameters. : EXC 6 
Fig. 21. Tail of Ascaris vesicularis. Male. x 30 diameters. | Be 
Fig. 22. Tail of Ascaris retusa. Male, x 95 diameters. ye 
Fig. 23. Head of Tenia multiformis.. x 900 diameters, 
. Fig. 24. Head of Eustrongylus papillosus. Enlarged. 
Fig. 25. Head of Tenia infundibulum. x 80 diameters. 
Fig. 26. Tail of Heterakis dispar? Male. About 60 diameters. 
Fig. 27. Tail of Heterakis dispar? Female. About 60 diameters. 
Fig. 28. Tail of a female Ascaris from the Peccary. x 100 diameters. 
Fig. 29. Head of an Ascaris from Coluber Blumenbachii. x 60 diameters. _ | 
Fig. 30. Larval nematode from å cyst in the heart of the same Serpent. x 60 diameters. 
Fig. 31. Head of a cestode from the muscles of the same. Enlarged. -— 
Fig.32. Head of an Ascaris from the intestines of an American Snake. - 
Fig. 33, Tail of the same. x about 20 diameters. | | 


Brans. Linn. Soo Vel ET, tab.63 fr. 866. 


y————— 


ri re 


XXVIII. On the Anatomical Characters of three Australian species of Tunicata referable 
to Savigny’s subgenus Cæsira. By Jonn DENIS MACDONALD, F.R.S., Assistant Sur- 
geon of H.M.S. * Herald, commanded by Captain H.M. Dunnam, R.N, FRS. 

Communicated by GrorcE Busk, Esq., F.R.S., FLS. ER, 


Read January 20th, 1859. 


1. IN ‚another paper, on the anatomical characters of Perophora Hutchisoni, I had occa- 
sion to notice the frequent occurrence of a small sessile and solitary Ascidian, attached to 
the same branches of Amphibolis, covered with a similar granular coating, and so. far 
- exhibiting a corresponding habit. As this little tunicary forms the type of a well-marked 
genus, of which we have discovered several species, a short description of it may not 
be uninteresting. 

The body is of a depressed pyriform shape, and from one-half to three-fourths of an - 
inch in length. It generally rests a little on one side, as well as on its base, being thus 
in part sessile and in part recumbent. | 

The two external openings lie nearly on the same plane, and in general appearance, 
more especially in the contracted state, resemble those of Boltenia, a resemblance which 
is heightened by an infolding of the test extending between them. The branchial aper- 
ture, however, is obscurely divisible into six rays instead of four, — å | 

- The test is exceedingly thin, and so densely studded with fine grit, that it is rather 
difficult to investigate its structure satisfactorily ; an internal glistening coat of a fibrous 
texture is nevertheless distinctly traceable. | HE, 

The pallio-vaseular system, which is so highly developed in other cases, is scarcely at 
all visible in the present species, a circumstance most probably to be accounted for by 
the thinness of the test, the greater part of whose apparent thickness is due to extraneous 
matters. en i | : 

On removing the test, an elaborate system of reticulated vascular canals, invested with 
à greenish-yellow pigment, presents itself beneath the epithelium of the. ausnile and the 
‚More superficial fibres of the muscular coat. Piece Oe oP Sag | 

The outer part of the branchial opening is armed with a circlet of simple, pointed or 
bifid tentacula, the equivalents of which are also present in Boléenta ; and, pe the latter 
genus, the inner rim of the same opening is surrounded by compound tentacula, divided 
Into pretty equal branchlets and pinnæ. | | | 

The branchial network is composed of rather stout transverse poe 
longitndinal nervures, strengthened at intervals by stronger ones assuming the c er 
of folds. - Tu | = 

The mouth is situated at that part of the respiratory | hamber which 18 — to v = 
eloacal cavity, and leads by a very short æsophagus almost directly into an elonga 


bars and very delicate 


368 MR. J. D. MACDONALD ON THE ANATOMICAL CHARACTERS 


stomach which curves gently downwards and backwards, and gradually diminishes in size - 

to the point where it becomes continuous with the intestine. The latter courses back- 
wards and then upwards on the right side of the body, until, having arrived opposite the 
branchial opening, it bends suddenly forwards, forming a simple loop, and again passes 
towards the stomach, above but parallel with its former course, to terminate in the cloacal : 
chamber close to the position of the mouth. ^ 

The liver is of a rudimentary character in this species, consisting of minute sacculations, 
which emboss the surface-of the stomach, and impart to it a rich amber or brown hue. 

The ovaria are two in number, one lying on either side of the body, between the 
branchial membrane and the muscular coat. The right ovarium does not occupy the lop | 
of intestine, as it does in Boltenia and most other Tunicata, but is placed in a recess in 
front. of it. 'These organs are pyriform in shape and laterally compressed, with the ` 
smaller end and the short duct issuing from it directed towards the cloaca. The testes | 
are also two in number, consisting of numerous . elongated and finely-divided lobules 
radiating round the base of the ovaria. | i : 


II. In Hamelin’s Harbour, Shark Bay, a second species of this genus, but of much larger 
size, is rather plentiful. Unlike that just described, it appears to lie loosely, or witha . 
very slight attachment; on the sandy bottom, being too massive to be supported on the 
delicate stem of a Zostera or an Amphibolis. - ; | | 

The body of the animal is somewhat rounded, compressed on the- sides, and averages - 
one inch and six-tenths in antero-posterior measurement, and one inch and three-tenths in 
the vertical direction. | | | | $ 

The test itself is very thin ; but numerous branched and interlacing fibres, often tubular, 
and containing prolongations from the mantle, arise from every part of its outer. surface, 
and form a matrix for the lodgement of shell and coral grit, foraminifera, and other extra- 
neous matters. -In this way the coat assumes a thickness of about one-fifth of an inch. 

The external openings lie nearly on the same plane upon the upper or neural surface; 
but they are usually so surrounded with small and irregular processes of the test, that it 
is difficult to determine their actual configuration; they are, however, connected by å 
straight fold of the test, as in the former species. 7 | ee 

The mantle presents an almost uniformly greenish-yellow tint, produced by minutely x 
divided and reticulated vessels. pn | | EE 

The fibres of the muscular coat are delicate, and rather loosely disposed over the general — 
surface of the body, though very strongly developed round the branchial and cloacal 
openings, in circular and radiating bundles. The lining membrane of the latter opening 
presents a rosy hue; and just within the margin of that which leads to the respiratory 
chamber spring several tentaculiform bodies, which appear to correspond with the dupli- 
catures of the contracted aperture. The true tentacula, however, always lie internal AE 
these or their modifications in the Tunicata generally. 

The tentacula in the present, as in the former species, are compound, consisting of a 
tapering central portion, with six or seven pairs of subramose lateral processes, openly set ` 
with minute papillæ. Both the axis and rami are crested or angulated in the middle; = 


OF THREE AUSTRALIAN SPECIES OF TUNICATA. 369 


gently curved, and ornamented with yellow vascular lines. The curvature of these beau- 
tiful organs is due to the presence of longitudinal muscular fibres on the concave border, 
having no others to antagonize them on the opposite side. 

The branchial network is minute and regular, and the membrane is thrown into about 
fourteen longitudinal folds; seven on either side of the body extending between the 
branchial and oral apertures, and increasing in length from before backwards, in which 
direction also they gradually become more curved. 

The mouth is small and surrounded by the converging ends of the longitudinal folds of 
the branchial membrane, leading into a short esophagus, which curves downwards and 
backwards to join an elongated stomach, from whose tapering pyloric end the intestine 
takes a very similar course to that described in the former case, and terminates in the 
cloacal chamber near the position of the mouth. 

The liver consists of numerous short and compressed glandular sacculi of a rich brownish- 
. red colour, lying on the left side and along the inferior border of the stomach, into which 
their contents are poured. 

The products of digestion are usually to be seen in the intestine, connected together by 
a plastic substance, and rolled into a filiform, continuous, and highly convoluted mass, 
extending from the stomach to the vent. Ps 

The testes and ovaria are identical in character, position and relations with those of the 
foregoing Ascidian. 

The heart lies on the left side of the body, just below and in front of the corresponding 
ovarium. It rests on a small brown cylindroid body with rounded extremities, and 
exhibiting a slight curvature, with its concavity looking upwards and forwards. This 
body appears to lie loosely in a blood-sinus adapted to receive it, and is chiefly composed 
of minute cells, cell-nuclei, and an amorphous matter enclosed in a membranous sace, 
but exhibiting no very definite arrangement. What its real nature is I have not been 
able to determine, though I am disposed to believe that it may be the representative of 


the * elæoblast” of the Salpian, 


III. In Shark Bay also, and in about three fathoms water, we obtained another small 
Ascidian, so nearly allied to those just described as to merit brief notice here. It is nearly 
of the same size as the King George’s Sound species, but rather more narrowed and pro- 
duced above, and more rounded at the base. | 

The branchial and cloacal openings lie on nearly the same level, at the extremes of the 
upper border, and are tubular and prominent, though capable of considerable retraction. 
— The test is thin, smooth, colourless, and beautifully transparent, s0 far calling to mind 

the pelagic Tunicata generally. It is, however, loosely covered over with the fine sandy 
Particles of the sea-bottom on which it rests, being apparently quite unattached, though 
Perhaps unable to change its place at will. 

- The muscular coat id its epithelial covering are also quite transparent and free trom 
Pigment. : «ie : 
The external openings, particularly at their base, are surrounded with pue: an 

VOL. xxıı, i 


370 MR. J. D. MACDONALD ON THE ANATOMICAL CHARACTERS . 


circular muscular fibres, while otters are scantily distributed in- Various directions over 
the body. 
The branchial network is very delinate and rests upon a stout transparent membrane, 
which is saceulated or embossed towards the respiratory cavity. | 
The internal tentacula are compound, as in the former species; and the mouth, stomach, 
_ intestine, testes, and ovaria exhibit the same general disposition, so that the description of 
those organs above given will equally apply to the present case, with the following excep- 
- tions, namely, that the stomach in the latter -is covered with a very dark pigmentary 
matter, probably including biliary follicles, and the intestinal loop is more > Opens so that 
the right testis and ovarium lie within it. 
- Finally, the heart holds a slightly oblique position below the left ovarium, and rests on 
a curved cylindroid body tinted with a rich madder-brown pigment, unquestionably the 
equivalent of the organ which I have assumed to be an “ elæoblast ” in the last-described 
species. | 


June 1858. 


EXPLANATION OF THE PLATE. 
Tas. LXIV. 


[The same general references apply to all.] 


L Cæsira ‘parasitica. King George's Sound species, 


Fig. 1. Animal in situ, with a cluster of zooids of Perophora Hutchisoni. 

a. Branchial, and 

5. Cloacal opening (both being contracted and radiately puckered). E 
Fig. 2. The animal removed from the test, to show its internal parts. 

€. Nervous ganglion. 

d. Position of the mouth. 

e. Stomach, and e. Liver, E Ep oc 

f. Intestinal canal and loop. i | | xs 

9. Rectum, | 

h. Right testis. - 

i. Right ovarium. 
Fig. 3. Simple external tentacula, magnified. 
Fig. 4. One of the compound internal tentacula. 
Fig. 5. Immature ova. 


Fig. 6. Perfect ovum with additional yelk-granules and outer covering. 


IL C. ficus. Shark Bay species (No. 1). 


Fig. 1. The animal as it appears in its natural State, with one or two Sertularians grow 
granular coat. ; 


ing on its 


: QU. 3-3 
UV DDE. VOL. 


r 
/ 


» 
N 
S 
e 


+ 


W West imp 


- 
d 
å 
3 
p 
2 

ive 
td 

m 

& 

z 
A. 

G 

F5 


Fig. 2. 
3. Left view of ditto. 
Fig. 4. 


Fig. 


Fig. 5. 
. Fig. 6. 


Fig. 1. 
Fig. 2. 


E Kur. 3. 


Fig. 4, 


OF THREE AUSTRALIAN SPECIES OF TUNICATA. 371 


Right view of the animal removed from its test, 


Portion of the test, under a low power. 

Portion of the test, highly magnified. 

Part of the internal fibrous and granular brit of the test, seen in face, and showing the aper- 
tures of nutritive canals. 


III. C. pellucida. Shark Bay species (No. 2). 


The animal in its natural state. 
Right view of the animal removed from its test. 
Dorsal view of the same. 
k. Sacculations beneath the branchial ie 
i. Endostyle. 


Left lateral view. 


m. Left testis and ovarium. 
n. The héart. ; à 
9. Elongated body on which the heart rests, as alluded to in the text. 


3c2 


[ 373 ] 


XXIX. On the Anatomical Characters of a remarkable form of Compound Tunicata. 
` By Joux Denis MACDONALD, F.R.S., Assistant-Surgeon of H.M.S. * Herald,’ com- 
manded by Captain H. M. DEXHAM, R.N., F.R.S. Communicated by GEORGE Busk, 
Bigy FRS. BLS ot 2 ! 
. Read February 17th, 1859. . ` 


ON examining an encrusting form of Compound Tunicata, taken from the ship’s copper 
while refitting in Sydney Harbour, I was surprised to find that each little cluster of 
viscera was surmounted by two similarly-constructed branchial chambers or thoraces, as 
though two zooids had been combined together. | 

‘ Each branchial chamber was supported on a narrow pedicle, and both pedicles arose 
from one short trunk, which suddenly expanded into the abdomen, while little gemma- 
tions were frequently seen clustering near its base. 

About four delicate and unbranched tubules, with a dilated and glandular extremity 
(derived from that part of the mantle which invests the viscera), extended themselves into 
the connecting substance. | 

A stout endostyle oceupied the dorsal region, and the branchial network exhibited 
three or four principal transverse bars, intersected at right angles by the more numerous 
longitudinal nervures. The orifice of entry was guarded by a cirelet of six broad and 
short tentaculiform processes—organs which are so often mistaken, in other cases, for 
the true tentacula; and there being no proper atrium, the anus opened directly upon the 
surface near the middle of the ventral aspect. The existence of a superficial common 
cloacal system was clearly indicated by the low position of the rectum, and the absence of 
pigment-cells within a circumscribed space on the fore-part of the body. The actual 
disposition of this system, however, I have not succeeded in determining, on account of 
the peculiar delicacy of the connecting substance. ser 

A small spur:like appendage or caudex was sometimes distinctly observed, projecting 
from the dorsal surface of the pedicles, just below the branchial chamber. i 

. The viscera formed a large subglobular mass, in which a voluminous stomach, testis, 
and ovarium were plainly discernible. In the specimens examined, the diameter of the 
ova, visible in the ovarium, far exceeded that of the pedicles, through which, according 
to the present view of the subject, they were destined to pass; moreover, numerous ovo, 
scarcely further advanced than those within the ovarium, were scattered through the 
connecting substance, in which they were perfectly enclosed. The ova of this genus, like 
those of the larger solitary Ascidians, were invested with a stout chorion, supporting a 
beautiful epithelial pavement, and containing å dull amber-coloured or reddish yelk. 4 
The process of yelk-cleavage was easily traceable in a selec series of ova ; and where 


that of differentiation had commenced, the vitelline mass appeared to be encircled almost 


completely by a long and gradually tapering tail, while three short — — 
m an opposite point. In more advanced examples, the transparent polygo a " : ^ 
cluding the true embryonic structures, formed an oval tadpole-like body, from the delica 


374 MR. J. D. MACDONALD ON THE ANATOMICAL CHARACTERS 


investing membrane of which the tail received a covering. These cells no doubt form 
the proper test of the embryo, and as such must also be the basis of the connecting sub. 
stance or common test of the mass. 
. . The part destined to become the future abdomen was divided into several rounded - 
lobes, from the confluence of which the tail emerged posteriorly, and two distinct bran- i 
chial chambers arose in front. : = | e 
. A nervous ganglion with an otolithic sac, and what may probably be a rudimentary 
visual organ, were connected with one of the thoraces; while the three sucker-tubes and 
four glandular stolons; above referred to, originated near the pedicle of the other. 

The incipient branchial network was marked by several transverse rows of short slits, 
which appeared to have thickened or everted edges, on account of the highly-refracting 
property of the epithelium at this stage of development. | 

There are many interesting questions connected with this double form of existence, to 
be answered perhaps by further observation of its structure, evolution, and history. 

The unusually large ova, which undergo the ordinary round of changes subsequently to 
their liberation from the body of the parent, and are perfectly enclosed in the connect- 
ing substance, seem to negative the idea of their being discharged in the usual way, and 
rather suggest their escape by rupture of the abdominal walls, or by pedicellation and — 
ultimate separation from the abdomen. With this consideration before us, the query 
naturally arises, Does the embryo at any stage become free and locomotive, as in other 
cases ? and if not, which I am much disposed to believe is the fact, what purposes can the 
transitory suckers, organs of sense, and tail subserve ? >H 

There being two respiratory chambers, with their proper openings, two mouths and . 
two vents, with a visceral mass common to both, forming perhaps the most perfect 
diplozoon in creation, —now that the originally simple nature of Nordmann's celebrated 
example has been satisfactorily explained,—are we to recognize here two distinct entities ? 
~ mia the sexes and generative function common to both, divided between them, or 
dapes to either? Some species of Polyzoa clearly exhibit the union of two zooids 

ra Skre of the so-called ds ovicell” and its contents, which are very pro- - å 
ea du ne : c im of a definite cycle of gemmations. These ideas may be pe 

so long as our jud and; i rpm en mn ae, font Pe = 
ereire =. ER = = tion to Immaterial and abstract things is i 
litio aha Men. material things afford. Be it as it may, however, its 
Ltd: botluti demise Seu, Just as in the case of simple Ascidians, propagei® à 
FE ND. " sg = oviparous development ; and the offspring A gr 
union pervades a rs = ei a E MER mon Khaipstenti Dole ME ue | 
ing substance, it would ln i zu _ Een For pe: of 
et : mine cult to regard it as any more than a simple aggregation of 
simple Tunicata blended Sets zn m — bec gir 

BERTE ass mdi e een of ‚their test or outer cuvena a de^ 
ae considered a valid reason for classifying such with ©” 


What then are the most important distinguishing features of the latter? I think it 


OF A REMARKABLE FORM OF COMPOUND TUNICATA. 375 


may be affirmed that, when the zooids of a compound genus are pedunculated on a central 
axis, vascular continuity is the most striking characteristic; but, where they are immersed 
in the connecting substance, the importance of vascular communication as a character 
yields to the existence of a common cloacal system. The curiously modified forms of the 
latter system, taken as one of the grounds of classification, I shall have to notice in a 
summary of Australian genera, now in progress. 

It is now full time to give a name to the little subject of this paper; an appropriate 
one would be Diplosoma, as at once sufficiently expressing the peculiar nature of the 
animal. I therefore propose to call the species Diplosoma Rayneri, after Mr. F. M. Ray- 
ner, Surgeon of H.M.S. * Herald, who investigated its anatomy with me, and satisfied 
himself of the truth of the descriptive particulars above given. 

June 1858. ! 


EXPLANATION OF THE PLATE. 


Tas. LXV. Div. I. 


- Fig. 1. A diplozooid, separated from the mass, and highly magnified, exhibiting the more important 
points of its structure. 
a, a. The two distinct thoraces. 
b. The pedicles, bearing gemmations near their confluence, and a caudal appendage at the upper 
part of their dorsal surface. 
c. Branchial opening. 
d. Anal aperture. 
e. Visceral mass. ; 
J, f. Pallio-vascular tubules, with their dilated glandular extremities. 
g. Stomach, showing a sort of valvular cardiac orifice. : 
h. Ovarium, containing one principal ovum, and several others in an earlier stage of develop- 
. ment. 
i. The large sacculated testis. 
Fig. 2. Ovum in which the process of cleavage is going forward. 
Fig. 3. Ovum further advanced. 
Fig. 4. Primary state of the embryo, showing the central vitelline mass, the three sucker-tubes, and 
caudal process. i 
Fig. 5. Embryo i the ovum more fully developed, presenting, besides the frontal suckers and tail- 
| process, nearly all the parts discernible in the adult state. 
a, a. The two distinct thoraces. 
b. Otolithic sac, occurring only on one of the thoraces. 
c. The frontal suckers arising in common with 
d. The pallio-vascular processes from the pedicle of the thorax to the left. 
e. Vitelline mass, in which the viscera are faintly marked off. * 
f. The caudex. i 
g. Polygonal cells of the primordial test. 
Fig. 6. Cells and intercellular corpuscles of the connecting substance. 
Fig. 7. Marginal coronet of the branchial opening. tels 
Fig. 8. Caudal process of the adult, springing just below the root of the endostyle. 
Fig. 9. Cæcal extremity of one of the stolons. 
Fig. 10. Nervous ganglion with otolithie sac and eye-speck (?). 


[ 377 ] 


XXX. On the Anatomical Characters of an Australian species of Perophora. By Jonn 
Denis MACDONALD, F.R.S., Assistant-Surgeon of H.M.S. * Herald, commanded by 
Captain H. M. DENHAM, R.N., F.R.S. Communicated by GEORGE BUSK, Esq., F.R.S., 
PLS. 


Read February 17th, 1859. 


DURING our stay at King George’s Sound, the dredge obtained several specimens 
of a very interesting compound Tunicary, belonging to the “social” division so called, 
and which I believe to be a new species of the genus Perophora. The points in which it 
differs from the known species appear to me to be too trifling to warrant the fabrication 
of a new genus for its reception. I shall therefore name it Perophora Hutchisoni, after its 
first discoverer, Lieut. John Hutchison, R.N. 

The zooids are about one-fifth of an inch in length, pyriform in shape, and supported on 
alternate tubular foot-stalks of a corneous texture, with one or more transverse articula- 
tions. These pedicels are continuous at their base with a tubular axis, from which they 
appear to be derived by simple extension of its walls without articulation. 

The principal trunk in the specimen before me is firmly fixed upon a sprig of Amphi- 
bolis antarctica, over which its rather irregularly-divided branches extend themselves, 
occasionally drooping freely, like the tender sprays of a climbing plant. 

No articulations are anywhere to be found except in the pedicels of the zooids, whose 
test commences at a definite line, where the corneous tissue of the pedicels terminates. 

The test is finely coated over with a minute siliceous grit interspersed with Forami- 
nifera and calcareous atoms. 

When all these adhering particles are removed by cautious immersion in dilute nitric 
acid, which destroys all the carbonate of lime and loosens the attachment of the siliceous 
elements, their impressions are left on the surface of the test, and here and there on that 
of the axis, which thence presents a deeply pitted appearance. 

The internal surface of the tubular axis is lined throughout with a very distinct epi- 
thelial membrane, obviously connected with its nutrition ; and in the little buds, which 
spring, botanically speaking, in the indefinite mode, from the growing branches, the 
cecal dilatation of the rudimentary test is lined with an extension of the same membrane. 
This lining, however, is not to be confounded with the true pallio-vascular system which 
is contained within it, and consists of a simple branched tube exactly corresponding with 
the trunks and ramifications of the corneous axis, while the coats of the tubules which 


enter the pedicels of the zooids are continuous with the mantle. Thence results a sort of 


“ cænosare,” exhibiting a remarkable analogy to that of the Sertularians, an analogy which 
us axis just described. This, however, 


is still further sustained by the nature of the sclero a 
may be regarded as perfectly homologous with the repent tubes and free pedicels of 
Laguncula, Pedicellina, and other Polyzoa. 

The branchial and cloacal openings present little or no external 
VOL. XXII. 


prominence, and are 
8 D 


378 ANATOMICAL CHARACTERS OF PEROPHORA HUTCHISONI. 


still further concealed by the gritty coating. The former is situated on the upper surface 
near the free extremity, holding a subterminal position, so that in the mutual relation- 
ship of these openings, Perophora, and indeed also other pedunculate forms, may be 
said to exhibit an approach to Boltenia. 

The surface of the mantle is beautifully marked with yellow branched and reticulated 
lines, similar, no doubt, to those which have been noticed in Clavelina, indicating the 
distribution of vessels. | | 

The endostyle occupies a portion of the dorsal surface extending from near the 
branchial opening to the inner extremity of the sac, where it presents a slight curvature 
towards the oral orifice. 

The transverse bars of the respiratory network are about five in number, and join the 
endostyle in a direction nearly parallel with a line drawn through the branchial and 
eloacal openings, while the much smaller and more numerous longitudinal nervures lie 
parallel with the endostyle. The beautiful network so formed is invested with a stout 
epithelium, and its meshes are fringed with vibratile cilia. 

The mouth is situated at the anterior and inferior part of the respiratory cavity, and 
opens into a short cylindrical æsophagus which passes downwards and backwards to join 
a comparatively small and subglobular stomach. The intestine arises from the dorsal 
surface of the latter organ, and after having performed a graceful flexure by turning 
forwards and outwards, terminates in a very ample atrium or open space between the 
exterior of the branchial network and the muscular coat. The lining of this space is very 
obviously continuous through the branchial slits with that of the respiratory chamber. 

The generative organs, as in most compound Tunicata, are included within the intes- 
tinal loop, and, on account of the close proximity of the cloaca, the ducts are necessarily 
very short. dr 

Two distinct canals, alternately interchanging the office of artery and vein, are eircum- 
stantially described. as occupying the pedicels of the British species of Perophora, but I | 
cannot say that I have found such to be the case in the Australian one. There is, how- 
ever, much difficulty in determining the question, on account of the obstacles which the 
superficial gritty particles and the pigmentary matter of the mantle and tubules oppose 
to the observation of the vital phenomena of the animals. I think I may safely affirm - 
that no such arrangement is traceable in Didemnium, Botryllus, and numerous other 
Tunicata, both simple and compound, that might be adduced ; and if it be actually present 


in the British Perophora, I cannot help regarding the fact as something quite exceptional 
in the group. + 


May 1858. 


ANATOMICAL CHARACTERS OF PEROPHORA HUTCHISONI. 379 


EXPLANATION OF THE PLATE. 
Tas. LXV. Div. II. 


Fig. 1. A sprig of Amphibolis, with Perophora Hutchisoni elimbing over the stem and leaves. 

a, a. Drooping branches. 

b. Solitary Ascidian, usually occurring with Perophora, and, like it, having a thin test closely 
covered with sandy particles. [I have reserved the more particular description of this species 
for a separate communication. ] 

c. Encrusting Polyzoon. 

Fig. 2. À growing branch, magnified to show the disposition and mode of gemmation of the zooids. 

a. Growing point, always slightly in advance of 

b, b. The gemmations. 

c, d, e. Zooids further advanced, with the gritty coating removed. 

J. Corneous tubular axis. 

g. Epithelial lining of f. 

h. Pallio-vascular membrane. 

Fig. 3. One of the zooids, more highly magnified. 

a. Corneous axis. 

b. Lining membrane of a. 

c. Pallio-vascular canal. 

d, d. Articulations of the pedicel. 

e. The test, showing the pitted surface remaining after the removal of the gritty matter. 

J. Branchial orifice, and 

9. The cloacal orifice (both somewhat contracted). 

h. Endostyle. 

i. Branchial sac. 

Jj. Cloacal chamber. 

k. CEsophagus. 

l. Stomach. 

m. Intestine, embracing the generative organs in its flexure. 


3 D 2 


Trans Jinn.S0c Vol XATI. tab öp 979 
+ x 


JD Macdonald R N.ad nat del. Tuffen West lith. n "s 4 


[ 381 ] 


XXXI. Observations on the “ Bitentaculate Slug” of New Zealand (Limax bitentacu- 
latus, Quoy and Gaimard; Janella antipodarum, Gray; * Aneiteum Slug” ?, Mac- 
donald). By Cuarrzs Knient, Esq., F.L.S. 


Read June 2nd, 1859. 
Auckland, New Zealand, 


27th December, 1858. 
DEAR Str, 
I Have lately compared Mr. J. D. Macdonald’s observations on the Bitentaculate Slug 
found at Aneiteum (Annals and Mag. Nat. Hist. 2 ser. xviii. p. 38, 1856) and Dr. Gray's 
description of Janella antipodarum (Annals and Mag. Nat. Hist. 2 ser. xii. p. 414, 1853) 
with the living Gasteropod found in New Zealand, and trust that the following notes 
and the accompanying drawing will interest the members of the Society. 

The New Zealand mollusk is little more than an inch in length. The mantle (?) covers 
the whole of the upper surface. The space which Dr. Gray describes as existing between 
the edge of the mantle and the foot, is on the under surface of the animal; so that the 
edge of the mantle forms the outline of the body as seen from above. The mantle (?) is 
mottled with light-coloured spots, the larger of which are at regular intervals, and are at 
times elevated above the surface in sharp conical points (fig. 14). The “ raised margin ” 
of Dr. Gray’s description is due to the effects of aleohol. 

In alcohol the animal becomes subcylindrical, and answers to the description of 
Dr. Gray and to that of Lamarck (Anim. sans Vert, 2nd ed. vol. vii. p. 723, Limax biten- 
taculatus, Quoy). 

The aperture of respiration (4, fig. 3) is on the right side of the slightly depressed roof 
of the pulmonary sac. There are four small, semitransparent, calcareous (?) granules in 
the anterior wall of the pulmonary sac (fig. 13); and immediately anterior to the sac is a 
small foramen (a, fig. 3), affording passage to a fluid which flows in pulsatory waves over 
its roof. Generally this pulsatory movement occurs at regular and frequent intervals ; 
and the wave extends around the pulmonary aperture, as if a limpid fluid were pro- 
pelled under the glutinous secretion which covers and protects the body of the mollusk. 

The buccal mass is armed above with a single horny plate or tooth (figs. 7 , 8). In feed- 
ing, the animal forces forward the buccal mass, and scrapes or hoes up with the horny 
plate the surface of the substance it is feeding on. 

The lingual sae, instead of being furnished with the single, symmetrical, ume 
appendage of the common Slug of these Islands, has posteriorly two c P 7 ; 
convolute appendages armed with transverse rows of dental processes (figs.11,12). The 
sac itself is furnished with a complete armature of rasps (figs. 9, 10) placed in conma 
transverse parallel rows, which meet in the median line at a very obtuse angle. eg jte 
has several minute teeth, and is set diagonally in the row, as are also the spine-like plat 


from which the minute rasps project (fig. 10). 


382 ON THE BITENTACULATE SLUG OF NEW ZEALAND. 


There is no such mantle as that described by Mr. Macdonald in the Aneiteum Slug as 
covering the roof of the pulmonary sac; but there is a depression instead of it, and the 
roof of the sac is extremely thin. 

In deference to the views of Dr. Gray, I have termed the covering of the whole of the 
upper surface of the animal “the mantle;" but, taking into consideration that it would 
be extraordinary that the eyes of the animal should be protruded through the secretory 
organ of the shell, this interesting Gasteropod may more probably be regarded as quite 
naked. 

Believe, dear Sir, yours very truly, 


E ER CHARLES KNIGHT. | 
J.J. Bennett, Esq., F.R.S., 


Secretary of the Linnean Society. 


EXPLANATION OF THE PLATE. 
Tas. LXVI. 


Figs. 1, 2. Dorsal aspect of Janella antipodarum. 
Fig. 3. Lateral view. 
Fig. 4. Portion of dorsum, enlarged, and showing— 
a, Small foramen anterior to pulmonary sac. 
.. "' d. Respiratory orifice. 
Fig. 5. Lingual sac laid open. 
| a. Plicated folds. 
b. Transverse rasp-like processes. 
Figs. 6, 7. Horny plate of buccal mass. 
Fig. 8. Lingual sac and its tubular appendage. 
a. Tubular appendage. Mu : 
b. Sac. sort Op HE 
Fig. 9. Armature of lingual sac magnified. Eg || 
Fig. 10. Magnified view of rasps. 
g. Rasp. 
J. Plate. | 
Figs. 11, 12. Dental processes and plates on plicated appendages. 
Fig. 13. Calcareous particles in wall of respiratory cavity. 
Fig. 14. Conical cutaneous papilla. 


LIT tab. OG p.382 


fa 
O, 


Bans Zmmn.smo, 


[ 383 ] 


ol 


XXXII. Further Remarks on the Organs of the Antenne of Insects, described in « 
Paper published in the ‘ Transactions of the Linnean Society,’ vol. xxii. p.155. By 
J. Braxton Hicks, Esq., M.D. Lond., F.L.S., &e. 


Read May 5th, 1859. 


ON the 2nd of June, 1857, I read a paper before this Society, describing some * peculiar 
structures in the Antennæ of Insects,” which was published in the * Transactions,” vol. xxii. 
p.155; since which time I have extended my inquiries among the different tribes, the 
results of which I have now the pleasure of laying before you. But before doing so, it 
seems to me that it would be well to give an outline of what had been previously accom- 
plished towards the determination of the structure and uses of the antennæ, and of the 
various opinions founded upon those researches. 

I shall first of all quote Burmeister, who, in his ‘Manual of Entomology,’ translated 
by Shuckard, 1836, p. 295, $ 195, says, * But as insects doubtlessly hear—as some, for 
example the Cicada, Grasshopper and May Beetle, &c., produce a peculiar sound, which 


serves to attract the attention of the female—they must evidently be provided with an i 


organ of hearing, which is either very recondite, or referred to organs whose form does 
not evince their function. The antennæ are doubtlessly of this class; and, indeed, Sulzer, 
Scarpa, Schneider, Bockhausen, Reaumur, and Bonsdorff considered them organs of 
hearing. That they are not organs of touch is proved, anatomically, by their hard, horny 
upper surface, and physiologically by the observation that they never use them as such, 
this function being exercised by other organs, viz. the palpi. Besides, the analogy of the 
Crabs, in which it is well known that the organ of hearing lies at the base of the larger 
antennæ, speaks in favour of the opinion of their being in general organs of hearing. If 
after this hint we look to the insertion of the antennæ, we likewise detect aero N "- 
articulating membrane, which lies exposed, and which is rendered mn by He eon T" 
the antennæ. This membrane, beneath which the nerve of the antenna runs, might, 
Without much inconsistency, be explained as the drum of the ear, and thus would the 
antennæ be transformed helices, which, as very moveable parts, would receive the ae 
tions of the air caused by sound, and act as conductors to it. | Whoever has — 
tranquilly-proceeding Capricorn Beetle which is suddenly surprised by a loud A 2 5 
have seen how immoveably outward it spreads its antennæ, and holds them qutm E 
were with great attention, as long as it listens, and how carelessly the insect p I 
its course when it conceives that no danger threatens it from the unusual noise. = for 
Straus-Diirckheim, and Oken are of the same opinion, a m nn 
years, and ende to confirm myself in by numerous experiments. : 

$ 196. « vr doubt and NG attend the observations Få — 
"pon the organ of smell of insects. Reaumur, Lyonet, and several m ? "Th hard 
anatomists consider the antennæ as such; but I would ask, with what right e , 


id 


E 


384 DR. J. B. HICKS ON THE ORGANS 


horny organ, displaying no nerve upon its surface, cannot possibly be the instrument of 
smell, for we always find in the olfactory organs a soft, moist mucous membrane fur- 
nished with numerous nerves. No such tunic is to be found in insects, at least in their 
head, or upon the surface of their bodies.” 

Robineau Desvoidy (see Ray Society vol. for 1845) considers there is still doubt on the 
the point. * He has already shown, in year 1827, that in the Crabs, as the outer antennæ 
are evidently the seat of hearing, so the inner ones are the seat of smelling ; and afterwards 
proved, in his * Recherches sur l’organisation vertébrale des Crustacés, des Arachnides, 
et des Insectes, 1828, that in the Isopodes the sense of hearing is no longer doubtful; 
in the Arachnides it is wanting, while, on the other hand, the parts pointed out as the 
mandibles are here organs of smelling, and the poison-canal in them corresponds to the 
lacrymal passage of the higher animals. In the Insects the antennæ are organs of 
smelling, and usually also of touch. They have no organs of hearing at all.” 

Kister (see Ray Society vol. for 1847, p. 306) assigns to the feelers (antennæ) of 
insects the function of smelling. In some experiments with turpentine in glass tubes 
the reporter says, * If these experiments show that the feelers betray a sensibility to the 
effect of powerful odours, it is not yet proved that these act on them directly; and so the 
evidence that the sense of smell has its seat in the feelers is defective.” 

Newport has given us a valuable paper on the use of the antennæ of insects, in the 
‘Transactions of the Entomological Society,” vol. ii. p. 229 (read in 1838, but prepared in 
1831). He examined the antennæ of Ichneumon Atropos ; he found all the joints, except 
the second, perforated all around by very minute holes. This, he says, is the structure of 
nearly all the setaceous antennæ. He observed two tracheæ passing up its whole length, 
which gave off branches at every joint, and which seemed to him—but of that he was not 
quite certain—to communicate with the holes in the antenna-wall. He also noticed å 
nerve passing throughout its length. 

From this, and from analogy with other animals, he judged it probable that the antennæ 
were not organs of smell. He instituted a series of interesting experiments which led 
to the same conclusion, and mentions that * Copris Molossus,” when in motion, ex- 
tends the plates on the end of its antenna, as if to direct the insect in its course; but 
that on the occurrence of any loud and sudden sound, it instantly closes the plates and 
retracts the antennæ, as if injured by the percussion, while the insect stops and assumes 
esi, al ht hr nn hetere in El maner. es D 
QE ind ies P ; as been observed of the antennæ, it seems probable that in s 
funetion of hearing are nid Es A — the means by which they are fitted m i 
minene : es ried in different insects, to adapt them to the perception 0 

, 3 to the habits of the species: that in some species they are eem 

; ; hey are of great, although not of vital importance 
the insect; and that the loss of both of them, more attire when as also with 
; will clearly explain in every instance the agitation, delirium, or stupor 


. OF THE ANTENNÆ OF INSECTS. 385 


the occasion of reading my former paper, were those by E. F. Erichson, published at Berlin 
in 1847, and contained in his * Dissertatio de fabricå et usu antennarum in Insectis, in 
which he states that he has arrived at the following laws :— j 

*1. Antennarum testam in insectis nequaquam solidam, sed numerosis poris perfora- 
tam esse. 

* 2. Poros hosce ab interiore parte membraná tenui clausos esse. 

. “8. Poros in variorum insectorum antennis variis modis dispositos esse.” 

He then proceeds to show the position of these closed pores in the various forms of 
antennæ, and that they are never found on the basal joints. He describes the dilated third 
joint in Musca as alone possessing these pores, and considers the seta to be the true con- 
tinuation of the antenna, which view had already been promulgated by Mr. Curtis. He 
also notices the numerous hairs on the antenna, between the pores, which he considers 
to form a protection to them from extraneous bodies. Now these pores he regards as 
organs of smell; because, he remarks, the olfactory organs of the higher animals are moist 
membranes in order that the odorous particles may be dissolved by the humour secreted, 
therefore he doubts not that these membranes perform the same office, protected by the 
downy hairs, and retained moist by them. Another reason for which he considers them 
organs of smell, is that they are more numerous in those tribes whose scent is acute. 

Of this work I was not aware at the time of reading my former paper, it being very 
scarce. The plates accompanying it simply show holes or pores, closed in by a delicate 
membrane. : 

Vogt (Zoologische Briefe, vol.i. p. 516-17, Frankfurt a. M., 1851) says, * If these uniform 
feelers are examined with a sufficiently high magnifying power, the outer surface of all 
the divisions, except the articulating joint, is found to be covered with minute ne 
Which are closed in at the bottom by a thin membrane that appears to be clothed wit 
numerous hairs. In the antennæ that are not of a uniform shape throughout, — 
Shaft or style, and these pores or indentations are then found only upon the too 


Processes, branches, and feathers of the feelers, whilst the integument of the shaft is con- 


i . iar structure of the 
Stituted lik ini rtions of the body. The peculiar ; 
e that of the remaining po one: but it would;be go ing far to 


ante TA ith their func 
nn is, no doubt, closely connected with the able proofs of the correctness of the 


say, wi 1 ds undenia V o 
y» With some recent observers, that it affor " He then says, “We are of opinion 


oft-contested theory that they are organs of smell only. 
ed thi : bining those of smell 
that these minute pores, filled with fine hairs, perform a function com — cef gia 


and touch; for it is an undoubted fact that many insects, such as 


: : ize objects, and that nocturnal 
others, constantly employ their antennæ to touch and "— of touch in the organs ; 


creatures of this class, especially, possess a remarkably acute se å 
Whilst others, again, as re Sr in which these organs are ———Q 
in size and form, make no such use of their feelers. On the bes = em 
ee organs of smell in insects ; and all attempts to find the seat o 

itherto ineffectual.” ; i ctions 

Thus we see how many different opinions have been entertained = rs 
of the antenne. Sulzer, Scarpa, Schneider, Ie cnn a are all in favour 
Straus-Dürckheim, Oken, Burmeister, Kirby and Spence, and Bor 3r 

VOL. XXII. 


386 DR. J. B. HICKS ON THE ORGANS 


of their being auditory organs; whilst, on the other hand, we have Lyonet, Robineau- 
Desvoidy, Küster, Erichson, and Vogt in favour of their being olfactory organs. In point 
of number, the majority are evidently in favour of their being organs of hearing, though 
it appears to me that some on either side base their opinions on but slender foundations, 
and that the most important advocate on the olfactory side, Erichson, has formed his 
opinion upon an imperfect knowledge of the true nature of the organs: he has thought 
them to be closed perforations opening internally ; whilst I have, in the paper mentioned, 
shown them to be closed sacs, sometimes considerably convoluted, to the inner aspect of 
which the nerve passes. I shall, in the course of the present paper, show that by no means 
in all cases is the closing-in membrane thin and delicate, but that in some insects it is 
thickened and even raised into a conical and hair-like eminence; his arguments, therefore, 
regarding its analogy to the pituary membrane clearly cannot hold. None of these writers, 
and, so far as I am aware, no one else, had at the time when my former paper appeared 
pointed out these closed, chambered sacs ; and it is evident that the opinions concerning 
the functions of the antennæ, hitherto entertained upon an imperfect knowledge of their 
structure, will undergo considerable modification when the structures I have already, and 
those which I am now about to describe, are duly considered and re-investigated. 

I shall now proceed to the detailed description of the other antennæ which I have 
examined since my last paper. At that time I had not investigated the antennæ of the 
Coleoptera to any extent; but I find they form no exception to the rest of the Insecta. 

lst. In Necrophora Vespillo the organs are only found on the last three joints, two of 
which are foliaceous, while the terminal one is rounder and pointed at the apex, forming 
the club or clava; and a fourth, foliaceous joint does not possess these organs. 

The sacs are found on both sides of the leafy joints, and all round the terminal one, 
though they are larger on the back. At A. fig. 1 (Pl. LXVII.), I have shown the surface 
enlarged, with the covering-in membranes and position of the hairs between them. At 
Å. fig. 2. is the section of the same. The average diameter of the sacs is about 3560 inch. 

The distribution of the nerve, spreading by many branches throughout the breadth of 
the joint, can be seen by focusing for the centre of the lamella. This antenna is men- 
tioned by Erichson, who, after describing its form, says of the last three joints, “ Testa ei 
est poris subtilissimis creberrimisque perforata, relictis nonnullis areis minutis im- 
pressisque, e quibus setula nascitur ” (op. cit. $ 10). It is these pores I have drawn at 
A. fig. 2, showing a sac which can be clearly made out in the bleached antenna. 

In the genus Silpha there are eleven joints: the last three are dilated and covered with 
fine hairs ; and scattered equally over the surface is to be found a number of sacs, one- 
fifth less in quantity than the hairs, B. fig. 1 bb. Besides these, there are larger sacs; 
B. fig. 1 aa, principally found on the terminal joint, about twenty-four in number. These 
show clearly the form of the organs from above, and at B. fig. 2 their nature can be 
further understood in section. The covering-in membrane is delicate, and rises as a low 
cone from the general surface-level; between the organs the hairs arise and overhang 
pasa zn - the larger ones at the surface is i7sg inch. This is the character 

ghout all the genus, so far as I have ascertained. 


In Creophilus, and probably in kindred genera, the organs are found only on the 


OF THE ANTENNÆ OF INSECTS. 387 


terminal joint, sen is more elongated than the rest: they consist of the smaller and 
larger sort, as in Silpha; but arranged in a manner very peculiar, and confined to the 
lower half of the terminal joint. The smaller forms are disposed in groups of from six to 
sixteen in each. The groups are diamond-shaped, with a row of hairs between them: at 
the angle are occasionally found the larger organs, as in Silpha, as also a few scattered on 
the upper half of the joint. Their covering-in membrane, however, is not so conical. 
The diameter of each of the small organs is 5-55 inch. (See C. fig. 1.) 

In Goerius olens the organs are distributed thickly over the surface, rather more fre- 
quently than the hairs, and show their sacculated nature very well (D. aa). 

In Carabus there are eleven joints in the antenna ; only the last five possess the organs. 
They are not so numerous as those I have described before, and are principally placed on 
one side of the antenna, as shown in E. 

In all the species examined of this genus, I have found the closing-in membrane raised 
considerably above the surface-level, more so than in Silpha, and having six or eight ridges 
running from apex to base, as is shown in E. figs. 2, 3. The greatest diameter of the sac 
is +550 inch, while that of the closing-in membrane at the surface is 3557 inch. The 
Section, fig. 1, shows the form of the sacs and closing-in membrane. 

In six species of Ælater which I have examined, I find the same conditions, namely : 
chiefly on the serre of the joints, generally in front of the base of the hairs, are small 
sacs about 3357 inch in diameter, as is shown in F. figs. 1, 20 &b. The closing-in mem- 
brane is slightly raised above the surface; and on the apex are from three to six, generally 
four, small papillæ or tubercles, as is shown in F. fig. 2 0, & fig. 3. 

In Strangelia elongata these sacs are very numerous, disposed over eight of the twelve 
joints, and becoming larger and more frequent towards the apex. Theyare generally situ- 
ated in front, or near the roots of the hairs; and their closing-in membrane is much smaller 
than the diameter of the sacs beneath. These sacs vary much in size, being from 37; to 
sovo inch in diameter. The larger sort seem convoluted, or two- or three-chambered. 

The antennæ of the Musk Beetle (Aromia meer v = similar ; and the same | 
description will apply, except that the sacs are larger t wem 

In iie iab: the didi lamellæ are furnished with sacs, which are placed close 
together, and are found on both sides of the middle one only, and on that side of the first 
and third which is in contact with the middle one. ng ie 

In the antenna of Dynastes Hercules the sacs are very — P and the un 
membrane well marked, their diameter being about 3760 inch. They are = seco 
same parts of the lamellæ as in Cetonia aurata. In consequence of the sacs no 

| from the inner aspect of 

through the whole thickness of the antenna-wall, a tube passes "rom Chus dho 

each to connect it with the interior of the lamella: towards the margin of : : 

‘thin the thickness of the integument ; 

form is modified, the sac being placed halfway wit : smaller one extend- 

and there is not only a tube connecting it with the interior, but also penes csse 
ing to the outer surface (H. figs. 1,2). Numerous depressions, in . 


: i I have hitherto 
cavities, ate to be found in the antenna; and in all the Lam ET gre thus common 
examined, the sacs are situated on all portions of these depressions, 


to them. | 3E2 


388 DR. J. B. HICKS ON THE ORGANS 


In some of the Lamellicorns the closing-in membrane is prolonged to a point, sometimes 
rigid, so as to be easily mistaken for a hair; but closer inspection and comparison, I am 
sure, will satisfy any one as to its true nature. I can best explain what I mean, by the 
antenna of Geotrupes stercorarius. The surface of the lamellæ is patterned all over in 
the same parts as in the former Lamellicorns, and in a very similar manner to those of 
Andrena fulva, shown in fig. 8 of my former paper. The surface is occupied with about an 
equal number of true hairs, and articulated projections very like hairs (I. fig. 20), beneath 
each of which is a sac, as in the former kind of antennze; and as the sac does not extend 
through the whole thickness of the antenna-wall, there is, as in Dynastes, a tubular com- 
munication with the interior of the lamella. (See I. figs. 1, 2.) 

This structure will help to explain other forms in the Lamellicorn groups, as for 
instance in the Stag Beetle (Lucanus Cervus). Over the same parts as in the others 
are a number of apparent hairs, slightly curved, projecting from the antenna-wall. On 
looking more closely, it will be observed that they are not all true hairs; on the contrary, 
only a few of the larger ones are hairs, while beneath the rest an elongated sac is placed, 
as is shown at K. figs. 1, 2. In this insect the terminal lamella has a large cavity— 
sometimes there are two—extending deeply into its interior (fig.3). The diameter of the 
sac is 5555 inch. 

A similar condition exists in Hydrophilus piceus ; but the nature of the sacs is well shown 
by the fact that some have the closing-in membrane level, and others slightly raised, whilst 
some have it prolonged into a spine (G. fig. 2). The diameters range from teso to 
3760 inch. This peculiar-shaped antenna has nine joints; the last three are liberally 
furnished with these sacs, and of a larger size than any I have hitherto found: possibly 
the size of the insect will account for it, though it may generally be said that the size of 
the insect has but slight relation to the size of the sacs, as will be seen by comparing 
the various measurements I have given. | | 

That these sacs with hair-like apices are the same as those which are found in Melolontha, 
Dynastes, &c. is evident from the facts—1st, that they are situated in the same position; 
2nd, that some have level, others spinous apices on the same antenna; 8rd, so far as I have 
found on examination of a large number of insects, I think it may be laid down as a rule, 
that no cuticular projection or spine has a cavity or sac beneath it in any other part of the 
insect, except in the antennæ. In some Hemiptera I have found beneath the true hairs a 
cavity, but not like these sacs, inasmuch as the whole antenna-wall was protruded, with a 
true hair at the apex. m 
= are ee of the covering-in membrane is an important point, because : 
© - : tese with what is generally understood to be the structure z 
od us when we consider the habit of Hydrophilus and compare : 
may obtain a clue as to the ete — sa stp a 
Hint aril) ba: soen-thot = wa of these organs in the antennæ. If watched a er 
moment he is placed in wat ë h Rae es Je release t 

VEN mar = : — brings them beneath the joint of thé head = 
the under surface a the tho ges maner: by the silvery film of air which we 
rax, and which is collected by the numerous downy hairs 0? 


OF THE ANTENNÆ OF INSECTS. 389 


that part of the body. If this insect employed its antennæ to scent its food, which is 
aquatic, does it not seem very improbable that the olfactory organs should be carefully 
shut off by a peculiar provision, from the medium in which both the inseet and the food 
are, and which medium is known to be well fitted for the diffusion of odorous particles ? 
Now in Dyticus marginalis we shall see the reason that its antennæ are bathed with the 
water, —not because it uses its antennal organs for scent, but because it uses them as in- 
struments of touch, and has them provided with special organs for that purpose. 

The antenna of Dyticus marginalis (L.), very similar in general appearance to the 
palpi of Hydrophilus piceus, consists of nine joints, all nearly alike, except the terminal 
one, which is pointed. The antenna is almost universally smooth: the only representa- 
tives of the hairs are on the prominent parts of the joints and apex, which I will describe 
shortly. But on one side of the upper part of each of the joints are a number of 
circular depressions (L. fig. 1552, fig. 3.), of the diameter of 3455 inch, more numerous 
towards the apex of the antenna; in the centre of each circular depression is a short 
canal, which enters a sac having its interior aspect in communication with the interior of 
the antenna; whether closed or not it is difficult to determine, but it appears to be so by 
a firm substance frequently found on the inner surface of the antenna where these organs 
are present. The nerve is to be seen very plainly giving off branches as it passes up the 
antenna to each of these groups, and finally expends itself in those of the apex, sending 
off a branch to the roots of the hairs. 

To these hairs, of which I have given a drawing (L. figs. 2, 4), I would also request atten- 
. tion, because of their peculiar form, and because such a form is not uncommon in insects, 
both on the antennæ and palpi, when those parts are used as organs of touch. That all the 
hairs, long as well as short, possess to some extent the faculty called in the higher animals 
“ general feeling,” I think cannot be doubted for a moment; but hairs of the form I am 
about to describe seem, from their position and shape, to be specially adapted as instruments 
of that more refined tactile sense called “touch.” Perhaps the most marked example 
is to be found in the antenna of Dyticus marginalis. The organ in question consists of 
a widened follicle (L. fig. 1 a a a a, figs. 2, 4), becoming gradually narrowed to the centre, 
from which a delicate membrane, in the form of a depressed cone, stretches across; from 
the centre of this membrane springs the modified hair, something like a ninepin, its apex - 
reaching a little above the general surface of the antenna. There are three on the apex 
of the antenna, three on each side of the widest part, and two or three on cach — 
the upper end of each joint at its widest part. Their situation on the prominent parts 
will show their use for touch; and the habits of the insect also show how valuable as a 
taetile organ is its antenna. : pens 

The whole of the antenna of this insect is wetted by the water, in which it passes the 
. greater part of its life, and where it finds its sustenance. In form the organs on p 
are not essentially different from those before described in other insects; but it is diffi- 


cult to suppose that odorous particles could pass through a membrane, then through a 


i i Josed, before they could affect the nerve placed 
very small tube into a sac, itself probably closed, be y eg IRON 


at the inner surface; while it is easy to understand how suc 
i ; al them 
for hearing in water. In the larval antennæ no such organs are found: the hairs on 


390 DR. J. B. HICKS ON THE ORGANS 


are few and long; whilst the apex, which has three spinous processes which are supplied 
with a nerve, doubtless possesses the sensation of touch (see M. figs. 1, 2, 3). 

In Meloe the antennæ consist of eleven joints, of which the 1st, 2nd, 3rd, and 9th are 
not furnished with the organs, as all the rest are, especially the 6th, 7th, 10th and 11th: 
those in the last, or 11th, are the largest. They have the same form as those of the Neero- 
phora Vi espillo, but rather smaller. 

In Clytus arcuatus the sacs are found interspersed between the hairs, of the diameter 
of 305 inch, in considerable number on the last five joints, and of the same form as in 
Necrophora Vespillo. 

I shall next describe the antennæ of the Hymenoptera. 

In the genus Vespa, whose antenna I have described in my former paper, I have since 
had an opportunity of examining the antenna of the male, female, and neuter, both of 

V. vulgaris and also V. Crabro. In all these, as there described, the last ten joints are 
covered on all sides from base to apex by organs having the canoe-shaped closing-in mem- 
brane. The number of these on each antenna of F. vulgaris is prodigious; on each seg- 
ment there are rather more than 2000. The 3rd segment possesses rather fewer, and the 
12th rather more, making altogether on the ten segments 20,000 for each antenna. There 
are also on this antenna many dwarfed hairs, as on that of Dyticus marginalis. There is 
no particular difference between the different sexes and neuters; and I may add that å 
distinet sac is visible behind the oval opening, which is seen behind the canoe-shaped 
closing-in membrane. | 

In Apis mellifica, however, the antenna of the worker or neuter (described in my 
former paper) possesses the organs only on one side; whilst in the drone or male the 
antenna is entirely covered with them, similar to those of the male Æucerus longicornis 
(see former paper, Pl. 30. fig. 7). I have, unfortunately, not succeeded in obtaining a 


female or queen, to enable me to institute a comparison: still one would argue that if 


these organs were olfactory, we ought to find them in a peculiar degree developed in the 


worker, and scanty in the male, who seldom leaves the hive, and then only for a short 
distance; but the contrary is the case. | 


Odynerus murarius possesses organs very like those of the neuter Apis mellifica, situated 


, on one side; and as the other side is free from any markings, the nerve and its distribu- 
tion and the accompanying tracheæ can be beautifully seen. The nerve, accompanied by 
two tracheæ, passes from base to apex, through the centre, giving off a branch from 
either side, about the centre of each segment, which branch passes through the next joint, 
to be distributed to the organs in the succeeding segment, and again giving off numerous 
fibres to supply the organs with sensation. i 


A remarkable antenna among the Hymenoptera is that of the Red Ant, Myrmica rubra. 
. It consists of twelve segments 


right angle; the segments after the second gradually increase in width to the last, which 


is longer than any, except the first, and tapers gradually to a point, towards which the 


antenna-wall gradually becomes thinner, so that at the extreme apex it is very delicate, 


the hairs also abbreviated. The surface is furnished tolerably thick with hairs, as in most 


antennæ. It also possesses the sacs very well marked, as may be seen at N. fig. 1 4 34,4 


: the first is very elongated, the rest being joined to it ata — 


OF THE ANTENNÆ OF INSECTS. 391 


They exist on the last nine segments more towards the distal end of each. The different 
forms are shown at N. fig. 3 a a, fig. 4. : the diameter of the closing-in membrane is about 
5700 inch; the length of the sac is from qayo to tyy inch. 

But, besides, there is another form of what seems to be the same structure, and which 
has a rather less-marked parallel in the antenna of Pronœus inetabilis (to be described 
next). There will be observed at N. fig. 1 bbb, a number of small closing-in membranes, 
of a diameter of 3550 inch; behind each is a very small sac, leading to a long delicate 
tube, which, bending towards the base, dilates into an elongated sac having its end in- 
verted, as may be also seen in the larger sacs (see N. figs. 2&3). What their specific 
use may be, it is at present impossible to say ; but, supposing these organs to be auditory, 
we may easily conjecture that they would be able to appreciate notes in a higher key. 
The nerve in this antenna is well seen, throwing off branches to the organs in its course 
upwards. 

The antenna of Pronœus inetabilis consists of thirteen segments, and possesses three, 
if not four forms of these organs, disposed in groups on one side of each segment :— 

First. In the middle of the segment are a number of small closing-in membranes, 
of the diameter of zzo inch; they are raised above the level of the surface, and have a 
small elongated cavity behind them, which is shown at O. fig. 1 dd, fig. 3 d. 

Secondly. On the side are level, circular or slightly oval closing-in membranes, with 
an oval opening just beneath, leading into sacs; in the centre of each inner surface is 
a papilla pointing inwards. These are shown at O. fig. 100, fig.35. The diameter is 
3300 inch. 

Thirdly. There is a scanty group of organs towards the middle of the segment, whose 
closing-in membrane is also level and quite circular ; and beneath it is a shallow cavity, 
having a small opening in the centre leading into a tube, which soon dilates, and is con- 
nected with a curved tube, which expands in the form of a trumpet. The exact method by 
which these trumpet-shaped expansions end internally is difficult to decide, owing to the 
delicacy of the parts and the thickness of the antenna-wall. It is impossible to make out 
the exact nature of the junction at the curve. This form I have shown at o. fig.1ec, 
fig.3cc. There is a strong resemblance between these and the long expanding tubules 
in Myrmica; and they probably have the same function. 

Bourthly. On shay sede portion of the half-antennal surface occupied by ur 
structures, are a number of low projections, at first sight like dwarf hairs ; but uec 
inspection they will be found to be conical projections of a delicate pe > 
from the centre of a depression of the cuticle, and having a sac behin ss 
nerve can be plainly seen passing, as in the other forms; m i rn ri real 
matter may be observed at the junction of the nerve with the sacs. å Fu m 
analogy between these organs and true hairs. Their diameter is 5560 inch, an pw 
shown at O. fig. 1 a a, fig. 2. å | 3 

That these different å are capable of imparting to im PER se Å EE 

iati : : - ied, if we admit that they are auditory 
ciating notes of different pitch, will scarcely be denied, «må mlt to explain the reason 
Organs; while, if considered as olfactory, it would be ——— | 
of such a variety of forms. 


392 DR. J. B. HICKS ON THE ORGANS 


Sirex gigas possesses two forms of these organs, as is shown at P. They are found 
freely scattered over the segments on one side, the other being free from them. The 
consist of— | l 

lst. A number of depressions, diameter 315 inch, in the centre of which is a closing-in 
membrane; beneath is a tube passing through the antenna-wall, and leading to å mem- 
brane-sac, as is shown at fig. 14, fig. 2, fig. 35. | 

2nd. A number of closing-in membranes, having a distinct cavity behind them, in the 
antenna-wall. They are shown in section at fig. 4, and from above at fig. 3 a. Diameter 
at surface 3965 inch. These appear to be nearly the same as at O. fig. 2, and similar to 
those in Carabus. | 

We come now to the Lepidoptera. In the Diurna I have examined 

Gonepteryx rhamni, 

Pieris, or Pontia brassice, 

Pontia rape, 

Vanessa urtice, 

Hipparchia Janeira, 

Lycæna Phleas ; E 
and I find the same description will apply to them as to the antenna of Argynnis Paphia 
(of which I have given a description and figure in the paper before referred to), with the 
exception of a variation in the comparative number of organs, and of the number of 
cavities. 

In the Moths I have as yet only found the smaller organs described in Argynmis 
Paphia : their position on the pectinated forms in the Bombicide and Geometride seems 
to be universally towards the apex of the division; and in the forms less deeply pecti- 
nated they are found on those sides where the deepest indentation exists. They are more 
numerous towards the apex of the antenne. 

lg Oi ti 
E g. 2. At fig. 3 the same is magnified, showing 
in the dotted lines the existence of a cavity in the interior. | 

Q. shows them on the dilated end of the combs in Odonestes potatoria (Drinker 
Moth) : fig. 5 shows one enlarged. | 

This condition I have found to exist in | 


På 


| d of each segment, as in the Buff Tip (Pygæra buce- 
phala), and Yellow Underwing Moth (Tryphæna Bomba). The latter I have figured at 


OF THE ANTENNÆ OF INSECTS. | 393 


them. In the interior of the antennæ, underlying the organs, is found a lobulated 
membranaceous mass (fig. 2 b), in which the extremities of the nerve lose themselves. 
It is difficult to make out the precise structure of the mass; but it seems composed of 
a number of membranous sacculi. It occurs in a great number of insects, and princi- 
pally, if not always, in those whose organs are not dilated into a distinct sac in the 
antenna-wall. 

In Pygera bucephala I have noticed a depression om each segment of the antenna, 
similar to those observed in some Coleoptera. 
.. ln the Ghost Moth (Hepialus humuli) the organs are in appearance more like those of 

the simple sort in the Diptera, and are dispersed over the whole antenna. The male and 
female antennæ and antennal organs are precisely alike; and it may be stated, as a rule, 
that those Moths which possess such an exquisite sense of smell as Orgyria antiqua have 
the organs more developed in the male than in the female, but they do not seem to be 
more numerous than in those Moths which are reputed to possess a more obtuse olfactory 
power. 

In Anthocera loti (Burnet Moth) the organs may be easily detected, being very 
distinctly placed in a slight depression between the cuticular projections covering that 
part of the antenna in which they are situated. The closing-in membrane is irregularly 
circular, of a diameter of 445,5 inch, and has a small papilla in its centre (S. fig. 1 #, 
fig. 2 b). Behind this is a bowl-shaped sac, which has projecting into it a small papilla 
in the centre of its inner wall, probably where the nerve impinges. This is shown in 
the section at S. fig. 2 a, while at c the hair-like cuticular projections are seen partly over- 
hanging it. : : 

The antenna of Macroglossa stellatorum is peculiar in having one side of each segment 
furnished with numerous rows of hairs. In the centre of each facet may be seen sacs 
(fig. 194). These hairs give a peculiar appearance to the antenna. They gradually 
enlarge towards the exterior of each facet, and possess large roots or follicles. 

In Sphinæ ligustri (Privet Hawkmoth), the organs are to be found from base to apex 

Pheng ligustri (Priv å ede | bou 
on every segment, on one side only, and in considerable profusion ; and, so far as 
‚make out, are on the same plan as in the Burnet Moth. 

I have examined many specimens of foreign species of 

organs are on precisely the same plan, and as numerous, 


my former paper. | | ig). F 
| | : : g) Fourteen 
At U, I have drawn the antenna of Forficula auricularis en — me res 
segments are found to possess the sacs ; from one to — - 9 a: their diameter at 
end, as in fig. 1 aaa. Their shape may be seen : section at fig. å 4; 
the surface is about 5455 inch | 
2300 . bout three or 
I have also examined the antennæ of an Agrion å they as in Libellula, 
four) of the simple cells I mentioned as occurring in Suc be d but a very rudimentary 
While in the common May-fly (Ephemera diurna) they exist m 
condition, | | tV 
: uc s, as shown at V., 
In the parasites also, the antennæ are furnished with similar organ 3F 
VOL. XXII. 


Tetrix of all sizes, and find the 
as in the Zetrix described in 


394 DR. J. B. HICKS ON THE ORGANS 


where the antennæ of the Pediculus of the Crow are represented. Two organs are found 
on each joint, except the last two. In the Pediculus of the Dog the organs are very 
large; they are shown in profile at V. fig. 1 b. 

Pulex also has them on each joint, as seen at V.fig.2. An enlarged view of those of 
the Pulex of the Mouse is shown in fig. 3. 

Having now detailed all the principal facts I have been able to secure since my last 
paper, I shall again endeavour to point out the great advantage to be derived from the 
process of bleaching which I described at that time, without which it would have been 
impossible, by reason of the depth of colour of the insect-integument, to have made such 
advances upon our previous knowledge. It is for this reason that I have no doubt the 
opinions of the naturalists I mentioned at the beginning of this communication would 
have been different, had they known the precise nature of the organs behind the closed 
pores; for it does seem to me impossible that the essential nature of an olfactory organ 
should be included in the structures just mentioned, and that odorous particles could 
pass, first through a membrane, sometimes even spinous, then through a cavity filled 
with fluid, and thirdly through another membrane to reach the extremity of a nerve, 
On the other hand, it is not difficult to conceive that such a structure would be well 
suited to the transmission of sound, upon the principles pointed out by Müller; and the 
numerous modifications of these antennal organs especially to be found in the Hymeno- 
ptera, seem to form an additional reason for supposing them to be auditory—namely, 
that they may give the insects a power of appreciating sounds of various pitch. 

Amongst entomologists, some misconception as to the nature of the antennæ has 
arisen from their not duly considering, in their observations on the habits of insects, that 
the antenna has (whatever other function it possesses), in a great number of insects, å 
faculty of feeling superadded, at least in those insects whose antennæ are tolerably mobile. | 
There can be no doubt that most of the Hymenoptera use them as tactile organs; and I 
am sure that the Honey-bee, Andrena, and the Ichneumonide do so continually, though - 
most writers have considered that the numerous movements of the antennæ of the Ich- 
neumons are for the purpose of smelling; but I am confident that strict observation 
would confirm my opinion that these motions are for feeling—and thus to enable the 
Insects to detect the opening of the holes in which the larvæ they are seeking reside ; 
and that the antenna could be used as well for hearing any noise the larvæ made, as for 
smelling out their position. The Ant has been quoted as an instance strongly showing 
E pa — pe ee for smelling, as is manifested in the power of mutual dec 
vit agus are 3 "eps Last summer I had opportunities of oe 
duals met one another " ; ee ege pe brom oe bt 
their antennæ into conta s be er | 
palpi together, and we u es ERDE Fe er ad pec 
never touched: each other à ES ee = — = re it is 
SM to my; bat I à Ep ve What means they communicated their thoug 

dd = uspect the palpi played the most important part. + 

arks made in this paper, as well as from the observations 


OF THE ANTENNÆ OF INSECTS. 395 


Newport on the habits of insects, I think it may be concluded that the antennal organs 
are formed upon a plan in accordance with our present ideas of an auditory apparatus, 
and are therefore capable of hearing, and that :—1st, they consist of a cell, sac, or cavity 
filled with fluid, closed in from the air by a membrane analogous to that which closes 
the foramen ovale in the higher animals; 2nd, that this membrane is for the most part 
thin and delicate, but often projects above the surface, in either a hemispherical, conical, 
or canoe-shaped, or even hair-like form, or variously marked; 3rdly, that the antennal 
nerve gives off branches which come in contact with the inner wall of the sacs; but whether 
the nerve enters, or, as is most probable, ends in the small internally projecting papilla 
which I have shown to exist in many of these sacs, it is very difficult to say. The principal 
part of the nerve proceeds to these organs, the remaining portion passing to the muscles, 
and to the roots of the hairs, at least to those of the larger sort. The distribution of the 
nerve can be very beautifully seen in the antennæ of the Pronæus before mentioned, as 
also in Odynerus. 

Another point, which might be mentioned as rather tending by inference to the con- 
firmation of this opinion respecting the antennal organs, is that in the Shrimp and Craw- 
fish among the Crustaceans (which have a sac at the base of the antenna, commonly 
regarded as the auditory organ) there is no trace of any organs similar to those of the 
Insecta; the nerve simply supplies the hairs and muscles, 

More than a year after the reading of my former paper on this subject, one was read on 
August 30, 1858, by M. Lespés, before the Academy of Sciences, Paris, reported upon satis" 
factorily by a Committee, and subsequently published in the ‘ Annales des Sciences Na- 
turelles :’ Paris, 1858. The title of this paper was * The Auditory Apparatus of Insects." 
Both the author and the reporters seem ignorant of what I had already done on this 
Subject, as, by not being aware of the value of bleaching the integument, M. Lespés 
had very great difficulties to contend with, which would have been avoided had he 
used that process, and moreover he would have escaped a great error. He had in con- 
sequence to employ the most colourless species; and the most minute of his researches 
were upon the antenna of the Lamellicorns, Melolontha, Polyphyllus, &c., and he was 
obliged to view the organs perpendicularly only. He certainly asserts the enm of 
sacs behind the membranes, which he calls “éympanules,” and ge drawing and 
description of the ultimate branches of the antennal nerve pinsding to them. po 
states that the sac or cell (which he terms * cellule,” or “poche ") contains a ds ee 
transparent, solid body attached to the inside of the membrane; this body he " 
“otolithe.” Now, in all the numerous antennæ w ich I have observed with en à 
high powers, I have never seen this otolithe, the nearest approach to that La 2. | ? 
small chain of solid granules passing, in Tetris, from the centre of the o— = 

; Sr P a -o his directions for the observation of these 
bråne to the back of the sac. But by following his directions m de 
Structures in Melolontha, by splitting open one lamella of the antenna and viewing 


D. à 
wall from within, I have seen the very same appearance he describes, when pl 
arent otolithe is very soon resolved into its | 


With low powers. However, the app posed to be 
nature: for by using an }-inch objective and high eye-piece, the ... ee otolithe 
the otolithe is seen long before the rest of the sac; and by adjusting x s " 


396 DR. J. B. HICKS ON THE ORGANS 


melts away ; so that by the time the sac is in the proper focus, the appearance of the otolithe 
has vanished ; and on further examination it will be plainly seen, especially where the 
sac is viewed obliquely, that the appearance of a solid body was an optical deception, pro- 
duced by the looking perpendicularly at the tube which passes from the ‘sac behind the 
tympanum to communicate with the interior of the antenna: as the tube is seldom in the 
centre, the appearance of the **otolithe " is usually more or less on one side. Another 
great proof against the existence of an otolithe is the fact, that when looked at sideways 
the sac is, so far as I can see, perfectly empty. 

M. Lespés, with myself, considers these organs to be none other than auditory; and 
from the numerous researches which have now been instituted by Erichson, Lespés, and 
myself, I think we may safely say that their existence is very general if not universal 
throughout the insect tribes. 

M. Lespés has also pointed out the peculiar hairs which I had noticed some time 


since, and which I have figured in the Plate as they exist on the antenna of Dyticus 
marginalis. 


Note.—The bleaching process which I now adopt is a slight modification of that formerly 
recommended. I take chlorate of potash, say one drachm ; water one drachm and a half; 
mix these in å small bottle with a wide mouth, holding about an ounce; wait five minutes, 
and then add about 12 drachm of strong hydrochlorie acid: chlorine is thus slowly de- 
veloped ; and the mixture remains in action from one to two weeks. 


EXPLANATION OF THE PLATE. 
Ta». LXVII. 


À. Organs on the antenna of Microphora Vespillo. 
Fig. 1. Enlarged view of the organs, 


between the hairs. Diameter 3357 Of an inch. 
Fig. 2. Section of antenna wall, sho 


wing the sacs behind the membrane. 


B. Organs on the antenna of Silpha 
Fig. 1. Enlarged view of organs, 
organs. 


Fig. 2. Section of larger organs. 


À 
as seen from above, on the terminal joint : a a. large sort; 55. smaller 


Diameter at surface 1750 Of an inch. 


C. Organs on the terminal Joint of a Creophilus. 
Fig. 1. Shows the lozenge-shaped groups. 


a a. Organs from 6 to 16 in each group, 


divided by rows of hairs. Diameter 54; of an inch. 
b b. Large organs. 


OF THE ANTENNÆ OF INSECTS. 397 


D. Organs on antenna of Goerius olens. 


a a a 4. Organs rather more frequent about the hairs. 
b b. Hairs. 


E. Organs of Carabus. 


C. violaceus. Section of antenna-wall, showing true hairs, a a. 
b b. Sacs with covering-in membranes raised above the general level. 


. Enlarged view of section of antenna-wall, showing the peculiar markings of the covering-in mem- 


branes. Greatest diameter of the sacs „„45- Diameter at surface yoyoth of an inch. 
C. arvensis. The covering-in membrane seen from above, ribbed more than in C. violaceus. 


F. Segment of the antenna of Elater. 


Shows the sacs generally in front of the hairs. 
Enlarged view of portion of same; 4. still more magnified, showing four or five papillæ on each. 
Section of the antenna-wall: a a. organs; à. true hairs. 


G. Organs of antenna of Hydrophilus piceus. 


Enlarged view of a portion of surface. 
Section of antenna-wall, showing the various conditions of the covering-in membrane. 


H. Portion of antenna of Dynastes Hercules. 


Part of one lamella. aa. Normal sacs. 
b b. Sacs distorted at margin. 


. Section of antenna-wall. Diameter 7275 of an inch. 


I. Antenna of Geotrupes stercorarius. 


Enlarged view of upper surface of lamella. 
a a. True hairs. 
b b. Organs. 

Section of antenna-wall. 
a a. True hairs. 


b b. Organs. 
K. Antenna of Lucanus Cervus. 


Surface of lamella, showing the organs and their spines. 
Section of antenna-wall. 

a. True hair-follicles. 

b b. Organs with spinous terminations. 
Terminal lamella with cavity. There are sometimes two. 


L. Antenna of Dyticus marginalis. 


Last two segments of antenna, showing— 
aaaa. Dwarf hairs (tactile). 
bb b b. Peculiar structure on one aspect only (see Fig. 3). 
c. Nerve passing to the structures and dwarf hairs. 
Enlarged view of both tactile hairs and organs. 
Enlarged view of organs. Diameter 5000 of an inch. 


Enlarged view of tactile hairs. 


DR. J. B. HICKS ON THE ORGANS 


M. Antenna of larva of Colymbetes striatus. 


Terminal joints. 
a, Spine or hair. 
b. Apex with three hairs. 


2. Enlarged view of 6 (fig. 1), showing the nerve. 
. Enlarged view of a (fig. 1). 


N. Antenna of Myrmica rubra. 


. The terminal and next segment, showing— 


aaaa. Organs as in other insects ; 
b b b. Smaller, communicating with the tubes, as in fig. 2. 


. Shows the elongated tubes with expanded extremities, proceeding from 55 b (fig. 1). 
. Section of antenna-wall. 


aa. The larger and more usual form. Diameter 375, Of an inch; length from 957 to 4; o 
an inch. 
bb. Diameter 4... of an inch. 


. Another form of a a (fig. 3). 


O. Antenna of Pronceus inetabilis. 


. Portion of antenna. 
. Section of antenna-wall, showing structure of a a a (fig. 1). Diameter -4y of an inch. 
. Section of antenna-wall. — 


c c. Shows section of fig. 1 c c. 
b b. Shows section of fig. 1 55. 
d d. Shows section of fig. 1 d d. Diameter qyp Of an inch. 


| Diameter yay of an inch. 


P. Antenna of Sirex gigas. 


. One segment, showing two kinds of organs :— 


a. With elevated closing-in membrane (see P. fig. 4). Diameter 44... of an inch. 
5. Cup-shaped membrane (see fig. 2 and fig. 37). Diameter ,1, of an inch. 


. Section showing the cup-shaped organs. 
. Superficial view. 


. Section of antenna-wall through fig. 1 a d, fig. 3 a. 


Q. Antenna of Arctia Caja, 


: One segment of male antenna, The organs are chiefly on the extremity of the lateral peeten, 


scales being on the opposite aspect. 


- Segments of female antenna. Organs on serrated aspect. 
. Magnified apex of serra, with organs. The dotted lines represent the interior condition. 
+ Apex of club-headed pecten of Odonestis potatoria. 


aa. Organs. 55. Hairs, 


. Enlarged view of an organ (a) and hair (5), with the cuticular lines, 


R. Antenna of Tryphena Bomba. 


. External aspect of a segment. (All are alike.) 


a. Scales on one side. 


b b. Organs on opposite side, with small hairs between. 


- Longitudinal section of ditto, showing nerve: a and 5. lobulated matter beneath organs. 


aged view of organs. aa. Organs, Between are hairs and cuticular markings. 


Fig. 


Fig. 


Fig. 


Fig. 
Fig. 


Fig. 


Fig. 
Fig. 


2 
3 


OF THE ANTENNÆ OF INSECTS. 


S. Organs on antenna of Anthocera loti. 


. Enlarged view of surface. 


a. True hair. 
b bb. Organs. Diameter 44... of an inch. 
c c. Cuticular projections. 
Section of antenna-wall aud sacs. 
a. Section with papilla at back. 
b. Covering-in membrane, with papilla in centre. 
c. Cuticular projections. 


T. Antenna of Macroglossa stellatorum. 


. One segment of antenna. 


a. Organs, large and small. 
b b b. Rows of hairs with large follicles. 


U. 


Last two segments of antenna, showing organs, a a a d. 


at b b. 
y: 


. Antenna of Pediculus of Crow. 


aa. Organs as in other antennæ. 
bb. Hairs (tactile) at apex. 
Section of organs. 
Antenna of Pulex of Mouse, showing an organ on each segment. 


Fig. 4. Organs on antenna of Pulex of Mouse. 


399 


2. Section of antenna-wall, showing the organs at aa.; diameter sgygth of an inch; and true hairs 


Frans. Linn .Soo Vol II tab 67 p493. 


D E 


nt) 
tofu, 


7» 49% 
N fl 
a} dtf dii 


rim 


NE 


NE RTE 
- 
à x 


ie titt Y 


EAR å 


P 
XXXIII. On the Embryos of Ende and their Germination. 
By BENJAMIN CLARKE, Esq., F.L.S. 


A^ 


Read June 16th, 1859. 


IT is now some years since L. C. Richard, in his original and very accurate work, the 
‘Analyse du Fruit *,' divided phanerogamous plants, with reference to their germina- 
tion, into Endorhizæ and Exorhize; but notwithstanding that exceptions to the exo- 
rhizal character as regards the secondary radicles have been repeatedly observed in 
exogenous plants, and its universality in the primary radicle doubted, yet in general 
works in which germination is referred to, this division is considered practicable, and the 
Endogens also are still regarded as being constantly endorhizal. 

In examining the germination of the Endogens, however, for the purpose of ascertain- 
ing the position of the first leaf of the plumule relatively to the cotyledon, the primary 
radicle proved, contrary to my expectation, to be perfectly exorhizal in the greater number 
of instances, and so obviously without any trace of a sheath (coleorhiza) in Zris, Alströ- 
meria, Smilacina, Butomus, Tamus, and Arum, as to place the fact beyond question. 
The secondary radicles in the Endogens, it is true, are usually, if not always, endorhizal, 
inasmuch as they protrude from beneath the cellular integument common to the primary 
radicle and young stem, although no sheath may be present; but this is a character also 
common to certain Exogens, as Menyanthes trifoliata, in which the secondary radicles 
and their branches not only protrude from beneath the surface, but have small sheaths at 
their bases; and some of the more remarkable instances of this kind have been described 
as occurring in the germination of Ranunculaceæ, Composite, and other families. The 
germination of Nuphar lutea also presents another remarkable instance in which the 
primary radicle is perfectly exorhizalt, but the secondary radicles above it are ms 
rhizal, the elongated sheaths remaining attached during the early stages of growth. 
(Pl. LXVIII. figs. 1, 2, & 3.) EE BER ! | 
_ Finding, therefore, the Endogens to be frequently exorhizal, as many of om og oppor- 
tunity afforded were germinated, and the probability appears to be that the primary or 


- true radicle is known to be endorhizal (with two additions afterwards noticed) only in 
and Graminaceæ. 'To these families should 


figured Zannichellia palustris as having 


& 


Marantacee, Commelynaceæ, Cyperaceæ, 
perhaps be added Naiadeæ, since Richard has ; 

all the appearance of an endorhizal germination ; bat whether they ge ji Pet 
be a question, because the embryos of some genera in their radicular po | e E 
materially from those in which the germination is exorhizal ; and al = e ti 
Zannichellia no trace of a sheath is perceptible in a longitudinal section, = 


* Démonstrations, ou Analyse du Fruit, 1808; H. A. Duval, Ed. ` SR 
en ing endorhizal, but as I have not had an opport ty 
+ The germination of Nymphæa alba has been referred to as being ESO d nie give it in 


of seeing it, this observation on Nuphar lutea was made with the greatest care ; ! 
x : AR wer s to be entirely an appearance. 
some degree the appearance of being endorhizal, but on examination it proves to be entirely TÅ 


VOL. XXII. 


+ 


402 MR. B. CLARKE ON THE EMBRYOS OF ENDOGENS 


marina the radicle is readily discernible lying in a canal directed obliquely towards the 
apex of the seed. 

Although the germination of the Endogens is as yet known only in part, the subjoined 
instances of the exorhizal character occurring among them will, I believe, lead to the 
anticipation that the endorhizal is the exception rather than the rule, as most of those 
families in which the germination is unknown scarcely differ from one or other of the 
following, in which the primary radicle, as far as my observations have extended, is 
strictly exorhizal :—Liliacee, Dioscoreaceæ, Juncagineæ, Alismacee, Butomacee, Iridee, 
Amaryllideæ, Orchidee, Palmacee, Aracee, Typhacee, and Briocauloneæ *. 

The inquiry relating to the position of the first leaf of the plumule originated in 
having observed that in the Graminacee the first leaf of the plumule is next the coty- 
ledon, whilst in all other Endogens I had seen germinating it was directly away from the 
cotyledon, so as to be alternate with it, as in the ordinary genera of Liliaceous plants. 
For some years this phenomenon had appeared inexplicable, but further comparisons 
have led to the conclusion that the suggestion of L. C. Richard, as regards the embryo of 
the Graminacee, viz. that the apparent cotyledon is in reality an enlarged radicle, is the 
only practicable explanation of this departure from the regularly alternate position of the 
leaves of the embryos of the Endogens. But as objections exist to regarding the apparent 
cotyledon of the Graminacee as the radicle or a part of it, it may with equal propriety be 
compared to the enlarged cauliculus occurring in exogenous plants, or to the cormus as 
occurring in endogenous plants, and then the first leaf of the plumule will be the coty- 
ledon, containing within it the leaves of the plumule occupying their usual alternate 
position. For the purpose, however, of being more readily understood, I shall, with 
Richard, still call it a radicle, though at first sight this view does not appear at all pro- 
bable ; but yet the position of the first leaf of the plumule next the apparent cotyledon in 
Graminacee and Oyperacee (fig. 20 c) appears otherwise an anomaly not to be accounted 
for. But admitting that the cotyledon, as at present generally understood, is either an 
enlarged radicle or a cormus, the steps by which such an inference may be arrived at 
form an almost unbroken chain. ~ 

Beginning, for example, with Caulinia, Naias, and Potamogeton, we find that the first 
leaf of the plumule alternates with the cotyledon, being directly away from it; and the 
embryo does not otherwise differ materially from those of Orontiacee, Aracee, and 
Liliacee, excepting that the plumule is situated higher up, in consequence of its radi- 
— P" n having become elongated. But in Zunnichellia we find a commencing 
cipe x structure; the radicle has become more enlarged, and a partial ridge is 
gn Br: part round the base of the cotyledon (figs. 4 & 5); but no further | 

: s ervable, as the first leaf of the plumule is directly away from the coty- 
ledon, as in Naias and Potamogeton. | 
m drake ee we find the cotyledon depressed and lying almost horizontally Ja 
té Ei radiele, which forms a shallow ridge surrounding it, so that it appear 

a furrow on its upper surface. This ridge is found to be divided into two parts; 


* The primary radicl : å 
in aa Å cle, as afterwards described, becomes suddenly arrested in its growth, but, as far as it developes, 


AND THEIR GERMINATION. 403 


and this, I believe, offers a practical explanation of the nature of the two portions of the 
cotyledon (as at present understood) of Triticum, Oryza (fig. 20 a, b), and other Grami- 
naceæ, in which traces of a stipule-like process exist, regarded by some botanists as a 
second cotyledon. The two portions of the ridge of the embryo of Ruppia consist of, 
1, the larger portion, which forms nearly the whole of the ridge; and 2, a small process, 
which fills up a notch in it, and which is always in apposition with the membrane which 
conceals the plumule, and is therefore always next the fissured or marginal side of the 
cotyledon * (fig. 6). There is, however, no further difference between Zannichellia and 
Ruppia, the first leaf of the plumule being directly away from the cotyledon, as in that 
genus f. 

Again, passing on to Zostera, we find in Z, marina (figs. 7, 8, 9, & 10) all the parts of the 
embryo of Ruppia much enlarged; thus the ridge, which is there but feebly marked, has 
become an enlarged two-lobed body, appearing when artificially opened almost like a two- 
lobed cotyledon, and the second portion of the ridge before alluded to, which in Ruppia 
is only a minute body, has become sufficiently prominent to look like a second cotyledon 
(figs. 8 & 9a). The cotyledon has become elevated on a cauliculus, which is curved (fig. 8); 
but this is the only difference between it and the embryo of Ruppia, as the smaller lobe or 
process of the radicle is opposite the membrane concealing the plumule, as in that genna 
(fig.9a). The cotyledon becoming elevated on a cauliculus is not uncommon, as it occurs 
in the Zrideæ, Juncacee, Alismaceæ, &e., and is afterwards further noticed. 

If this is admitted as the only practical explanation of the structure of the embryo of 
Zostera marina, it appears to follow that the same explanation is the only mode of 


accounting for the relative position of the parts of the stis à V igit = 
th : : Zostera, being next the larger lobe oi the vidi 
e cotyledon, both in Ruppia and Zostera g cotyledon is in reality a 


should expect to find it so in the Graminacee, if the apparent 20 

radicle, and this I have found to be the case in both British and exotic pne å > 
All the parts of the embryo have precisely the same relation » på pénis ael 
Mays the cotyledon (plumule as at present understood) in germination un nn 
on a cauliculus (fig. 21 a), making the embryo in all respects like that o m. 
except that it wants the minute secondary lobe, which is, mad being 
Avena, and Oryza (fig. 20 b). Zostera also further agrees — ft might almost be 


RE, | afterwards noticed ; from 
endorhizal in a remarkable degree, as ve to be the same. 


anticipated that the structure of their embryos would pro Ruppia and- Qymodocea in the 
Most of the Graminaceæ I have seen also ag - b Ge medic but lying in à 

cotyledon (plumule as now understood) not being enclosed by si 

a: : ly show its 

* "This small process is distinctly figured by Richard in zu. are mg EEE xvii. pl. 9. 

Position relatively with the marginal side of the cotyledon. (Vide Ann. du 

figs. 43, 58. i connecting link between 
t E Webbiana and Posidonia Caulini, as figured by M —Ó part of the radiele, and 

Ruppia and Zostera: in Posidonia the cotyledon is more A agri gin and they cannot be 

the position of the cotyledon is distinctly shown in both of them by the p meras ces Nat. 2™ série, xi. Bot. pl. 17. 

Supposed to differ in this character from Ruppia and Zostera. Aisi ds 5 

figs. 15, 16.) da 


404 MR. B. CLARKE ON THE EMBRYOS OF ENDOGENS 


furrow on its surface, as in Avena and Hordeum. 'The embryo of Hordeum vulgare is in 
this character much like that of Cymodocea Webbiana, as figured by Ad. de Jussieu*, the 
position of its leaves being precisely the same; so that the evidence of the identity of the 
embryo of Zostera with those of the Graminaceæ appears to be complete, and the com- 
paratively large radicle of this genus and its allies may be regarded as supplying the place 
of the deficient albumen. 

As a further argument that the apparent cotyledon of the Graminacee is a radicle, it 
deserves remark, that in germination it scarcely enlarges, and its smaller lobe, by some 
botanists regarded as a second cotyledon +, also remains almost stationary; whereas the 
stipulary process of the cotyledon, which occupies the same position in Smilacina, Tamus, 
Tris, and Canna, grows in favourable circumstances so as to be as long or longer than the 
cotyledon, or even as long as the seed, and sometimes forms a sheath for the young stem. 

In this view of the structure of the embryo of the Graminaceæ, the second or rudiment- 
ary cotyledon, which is a minute body, usually referred to as being alternate with, and 
rather lower down than the larger (fig. 20 5), is the smaller lobe or process of a two-lobed 
radicle, as in Zostera (fig. 8 a) ; so that the division of flowering plants by Ray into Mono- 
cotyledonous and Dicotyledonous is quite correct, and the Endogens are separated from 
the Exogens more completely by this character than by any other. | 

For the purpose of further explanation as regards the germination, the position of the 
first leaf of the plumule, and a proposed division of the Endogens, particular notices of the 
embryos of the families examined are added. | 

Cyperacee.—I have seen germinating the seeds of Cladium Mariscus, of a Cyperus, 
and of a Carex from Japan, and find that they agree with the Graminaceæ in having the 
first leaf of the plumule always next the cotyledon, as at present understood, from which, 
as this family is so nearly allied, it may be inferred that the embryo has the same struc- 
ture, and therefore consists of a radicle partially enclosing a cotyledon. There is nothing 
in the germination to contradict this supposition, unless it is that the radicle is endo- 
rhizal in the lowest degree, as no coleorhiza forms, and the nascent spongiole emerges 
from beneath the surface without a distinct appearance of a rupture of tissue taking 
place. This is the only family I have observed to agree with the Graminacee in the 
altered position of the first leaf of the plumule, but it not improbably occurs in other - 
glumaceous Endogens, as Restiaceæ and Desvauxiacee. 

Zosteracee.—The embryo of Zostera marina has been accurately figured by Richard }: 
the cotyledon is seen elevated on a cauliculus, and the position of the plumule is distinctly 
shown, and even the smaller lobe or process of the radicle is faintly indicated, but it 
is imperfectly seen in consequence of the embryo being closed; but its endorhizal cha- 
racter does not appear to have attracted notice. This, however, is quite obvious, as the 

* Ann. des Sciences Nat. 2"* série, xi. Bot. pl. 17. fig. 16. 


s : T ieri nun that this is not a second cotyledon, but a part of the cotyledon itself, which beoe pi 
à T and Mate It as being of the same nature as stipules, to which it certainly has a close resemblance ; but, as 
oes not enlarge in germination, it wants one of the characters of stipulary processes, as far as my observation has. 


race. (Vide Sehleiden's Principles of Scientific Botany, translated by Dr. E. Lankester, p. 272.) 
f Ann. du Muséum d' Hist. Nat. tom. xvii. pl. 9. figs. 47,48. | 


ÅND THEIR GERMINATION. 405 


radicle lies in a canal directed obliquely towards the base of the embryo (fig. 9), and is so 
unattached that it may easily be taken out entire. But in Z. nana the radicle is short, 
less oblique in its direction, and the orifice of the canal is not closed, as in Z. marina, 
being covered only by a delicate semitransparent membrane (fig.22). In Triticum, 
Avena, and Hordeum, there is a tubercle immediately under the radicle, which looks not 
unlike a closed orifice. 

As thus described, therefore, the embryo of Zostera marina differs from those of Oryza 
sativa, Zea Mays, &c., only in the cotyledon and its cauliculus being curved by becoming 
bent-downwards, and in the smaller lobe of the radicle being removed from the base of 
the cauliculus (fig. 8 a), which may be the effect of the pressure of the bent cauliculus, as 
it makes a depression in the substance of the radicle. 

Eriocaulone@.—The germination of Eriocaulon septangulare is very singular, differing 
from that of any other plant I have seen. The embryo first protrudes a horizontal 
process, having a small speck on its outer part, consisting of a circular portion of the 
testa, in the same way apparently as Mr. Wilson has described to occur in the germi- 
nation of Lemna gibba (Bot. Misc. i. 145. t. 42) ; but in its after stages it differs in sending 
up a leaf, which is directly away from the cotyledon, as in Juncacee and Ljliacee, and it 
_ cannot, therefore, be compared in this character to either Graminaceæ or Cyperacee. 

Subsequently to the appearance of the first leaf, the horizontal process protrudes a root 
from its under surface, which has no coleorhiza, although it most probably escapes through 
a fissure in the tissue, as the subsequent radicles do. As growth proceeds, the horizontal 
process becomes fissured by the enlargement of the root, which soon breaks through it, 
and its remains are finally left as a root-like process on the root itself. (Figs. 11, 12, 18, 
14, & 15.) så i» : 

On referring to Klotzsch's figures of the germination of Pistia lexensis , I was ned 
ably surprised to find a satisfactory explanation of the germination of Eriocaulon : Fe 
no doubt it will also serve to explain that of Lemna, as figured by Mr. Wilson (loc. > » 
Which is admitted to be very obscure. In Klotzsch's figures the horizontal cap. > D 
doubt the primary radicle, as it has so entirely the appearance of rs Amen > 
Araceæ and their allies, and the same therefore must be the conclusion wit regard E r 
the horizontal process of Briocaulon septangulare (fig. 11) and the un De mai 
 Lemna gibba. But whether this kind of germination is eser rå eene 
a question, because the growth of the radicle is arrested about AER endorhizal and 
- first appears; and possibly such a germination may be eo rd a i 
exorhizal plants, so that Pistiacee and Lemnaceæ may 96% of the loiuale of Pistic 
am. obliged, however, to differ from Klotzsch in his deserip a er u process, 
terensis: what he calls the first leaf, I should undoubtedly Gea å ide calls the 
or perhaps only as the lips of the fissure in the cotyledon; and w3 


ich i sd as being directly away from 
second leaf must, of course, be the first leaf, which is figured as being directly 


the cotyledon, as in Arum. med ER 
nn TIG endorhizal character is here well € rå E e «i 
ever, very thin, and soon disappears. The first leaf of the plum ) 


in, 1853. Taf. ii. figs. Q,R, S. 
* Ueber Pistia ( Abhandl, der K. Akademie zu Berlin, aus dem J. 1852). Berlin 


406 MR. B. CLARKE ON THE EMBRYOS OF ENDOGENS 


the cotyledon. On the face of the germinated cotyledon in Commelyna tuberosa, and 
another species, there is always present à small process looking like a bud, which is not, 
however, present on any of the succeeding leaves; possibly it may be analogous to the 
buds that occur on the ribs of the inferior paleæ of the Grasses. 

Juncaceæ.—As this family is intermediate between Liliacee and Cyperacee and their 
allies, and its limits are not well defined, its germination presented a more than usually 
interesting subject of inquiry, and was rather expected to prove like that of the Liliacee. 
Luzula campestris is, however, distinctly endorhizal, being more so than the Cyperacee, 
and, I believe, strictly agrees with the Commelynacee in this character; it may therefore 
serve to show that Juncacee have no near affinity with either Ziliacee or Palmacee, 
from which they further differ in the cotyledon, as growth advances, becoming elevated 
on a cauliculus. The coleorhiza is of delicate texture, and disappears within a day or two 
from the enlargement of the root*. The cotyledon has a well-defined opening for the 
plumule, the first leaf of which is directly away from it. (Fig. 23.) 

Hydrocharidee.—This family has the embryo (the plumule only being rather larger) 
as well as the habit of Naiadee and their allies, and so may, perhaps, supply a connecting 
link between the hypogynous and epigynous Endorhizæ. It also agrees with them in 
having the first leaf of the plumule directly away from the cotyledon. 

Marantacee.—The germination of Canna indica has been figured by Richard with _ 
his usual accuracy +, with the exception of the primary radicle, which, from his figures, 
might be supposed to possess a coleorhiza of the same kind as the secondary, which is not 
at all the case. The coleorhiza of the primary radicle is of comparatively delicate texture, 
and unless the germination is daily examined from its commencement, may escape 
observation, as it very soon disappears entirely; whereas the secondary radicles break 
through the epidermis from a considerable depth, and the torn fissures through which 
they escape remain for a much longer time. At the last stage figured, the coleorhiza of 
the primary radicle has almost or entirely disappeared, so that the appearance of it there 
represented is, I believe, only that of the lateral radicle on either side reftected down 
upon it. The first leaf of the plumule alternates with the cotyledon, and as germination 
advances, the cotyledon sends up a stipulary process, forming a short sheath for the 
growing plumule. 

Alismacee and Butomacee.—In the germination of Actinocarpus Damasonium the 
plumule becomes elevated on a cauliculus, which elongates, while the radicle has scarcely 
rg the first two leaves of the plumule are lateral, i. e. neither directly 
dote. hes E = nor directly away from it, but yet, as growth advances, the external 

; à tendency to turn more directly away from it. Butomus umbellatus 
sr ss Ese way, the cotyledon having a very distinct fissure for the emussio® 
€: iem e, > its first leaf is directly away from the cotyledon. The germination 

y observed, and no trace of a coleorhiza was perceptible. (Figs. 24 & 25.) 


* $ vir 
The coleorhiza cannot be seen unless the seed is first completely deprived of its mucilaginous coat by placing d : 
e 


water fi i : : : 
DE or two or Mme days, when it swells, and is easily removed. If this is not done, the radicle becomes cov 
with mucus, so as entirely to prevent observation. | 


t Ann. du Muséum d'Hist, Nat. tom. xvü, pl. 5. figs. 3, 4, 5, 6. 


AND THEIR GERMINATION. 407 


Juncagine@.—The germination of Triglochin palustre agrees with that of Arum, show- 
ing no trace of the endorhizal character; but after the radicle has protruded, a slight 
ridge forms, which marks its junction with the cotyledon, and is produced by the base of 
the cotyledon ; and this gives the radicle in some degree the appearance of having been 

endorhizal. 

-~ Typhacee.—Repeated examination failed in showing any traces of the endorhizal 
character in Typha latifolia, the germination of which is much like that of Butomus, in 
the cotyledon becoming elevated on a short cauliculus, which is distinctly separated from 
the radicle by a slight ridge, and it agrees with it also in the position of the first leaf of 
the plumule. A fringe of minute radicles gives it the appearance of being endorhizal, 
and renders the examination difficult unless made in water. Richard's figures do not 
represent it as endorhizal *. | 

Aracee.—It was after repeated unsuccessful attempts that the seeds of Arum macu- 
. latum and Dracunculus germinated, and neither of them proved endorhizal At first 
there is no distinction between cotyledon and radicle (fig. 16), and subsequently only a 
faintly marked ridge shows the junction between them (figs.17 & 18); this consists of 
the base of the cotyledon, and is seen at the bases of the succeeding leaves (fig. 19). In 
4. Dracunculus the first leaf of the plumule is alternate with the cotyledon, being directly 
away from it; but A. maculatum does not produce its first leaf till after it has formed a 
small cormus, and in the meantime its cotyledon has withered. Supposing therefore 
the cormus in the latter to develope rather earlier than the cotyledon, so as partially fo 
envelope it on its dorsal side, or only to form a concave tuberosity beneath it, then it 
might be compared with the embryo of Zosteracee. 

Palmaceæ.—The germination-of Phenix dactylifera agrees with that of the a 
and Liliacee, except that it protrudes its radicle like a Commelyna, forcing = € ies 
tega away in much the same manner +; a shallow furrow, extending the lg å 
the cotyledon, shows its marginal side, and at its base, close to the radicle, prs hes 
for the plumule, looking like a small scar. It is, however, if pe a. vas 
_ the plumule escapes by an artificial opening, sometimes through the FEE Taking 
. ledon, but often through the furrow, in consequence of the tissue being tt i v 
this furrow as a guide, the first leaf of the plumule is directly away 
ledon. .— Å t rimary 

Liliace@.—Smilaeina, Funkia t, Asparagus, and Allium, all agree in having the p Te 

e A 7 E ule directly away from the coty A 
radicle exorhizal, and the first leaf of the plum | of the cotyledon, as in Iris 
Asparagus has the plumule sheathed by a stipulary accade bs oibie ds. 
and Canna; it is found on the inner side of the plumule, an i 


* Ann. du Muséum d' Hist. Nat. tom. xvii. pl. 5. figs. 8, 9. ; | : liel times exfoliates in 
+ About ten days after germination has commenced, the cuticle at the apex a bii: Merc same kind of ex- 
minute irregular scales, having in some degree the appearance s sen becomes an inch or more in length, 
foliation soon takes place from the whole surface of the Be cotyledon, w 
and therefore, I believe, is not at all analogous to a coleorhiza. hich, growing during germina- 
T One of the species of this genus is remarkable for a triple plumule, the go E Before germination the 
tion, produce a singular appearance. It is in cultivation in the Botanic Gardens, 
$ int, 
Plumules are enclosed within the fissure, and all spring from one poin 


408 MR. B. CLARKE ON THE EMBRYOS OF ENDOGENS 


cumstances, as long as the cotyledon, but no trace of it is present before germination. It 
cannot be a rudimentary leaf, because it has no connexion with the base of the plumule, 
but only with the margins of the opening in the cotyledon. 
` Iridee.—The embryos of several species of Zris agree in their germination with those 

of Argcee and Liliaceæ; they are, however, remarkable for the cotyledon sending up a 
sheath surrounding the plumule, which sometimes becomes longer than the cotyledon — 
itself, and terminates by elongating internally between the plumule and cotyledon (fig. 26). — 
In other species of the same genus, this sheath exists only as membranous margins of the 
elongated opening in the cotyledon, which proves beyond doubt it is of the nature of 
stipules. | 

Amaryllidee.—The germination of Alströmeria aurantiaca agrees precisely with the 
species of Iris in the primary radicle being exorhizal, and the secondary endorhizal, or at 
least breaking through the cellular surface; but the stipulary process, as I have termed 
it, is wanting, which is a further proof of its real nature in Tris. i 

Digscoreacee.—The germination of Tamus communis agrees most with that of the 
Liliaceæ. The first leaf of the plumule alternates with the cotyledon, and not only is it 
exorhizal, but it is some time before any distinetion between the cotyledon and radicle is 
apparent. The opening for the emission of the plumule is horizontal, and a sheath, like 
that which forms in Asparagus, rises round the nascent plumule, and, as growth advances, 
becomes external to it instead of next the cotyledon, which gives it in some degree the 
appearance of à second cotyledon; it is, however, often bifid, and the young leaves, as well 
as those which are mature, have lateral stipules, which alone might be sufficient to explain 
its true character *. | 

With reference to the value of the characters, it may be observed that they will probably 
prove of some practical use in subdividing the Endogens; Graminaceæ, Zosterace@, ` 
Naiadeæ, and their allies forming a natural section, and containing, it may be antici- 
pated, all the truly endorhizal and macropodal hypogynous families; and, on the other 
hand, Aracee, Liliacee, Dioscoreaceæ, Trideæ, and their allies, in which these characters 
are wanting, are all otherwise in near affinity to each other. | 


Such a subdivision, however, would involve the placing of the Marantaceæ, Musaceé .— | 


and Amomee, and also the Orchideæ and Burmanniaceæ on account of their large 
radicle, m the endorhizal section, with which they have scarcely any connexion except 
the affinity between Orchidee and Triuridee ; but yet their epigynous character should 


not perhaps be regarded as excluding them, because of the truly epigynous character e 


the Hydrocharideæ. If, however, Von Martius is right in his comparison of Burmanni- 
ace@ with Hydrocharideæ, to which he refers them, another link of connexion between 
Orchidee and the hypogynous endorhizal families may be traced; and it may also deserve 
notice, that the embryo of Orchis Morio in germination becomes in shape much like that 
of a Ruppia, the young plumule also springing from the same part. The ovules of Bur- 
manmaceæ are orthotropal, and the distichous inflorescence of Burmannia dasyantha t 


* i 3 = ; ; 

å This stipulary DEM NE the ungerminated embryo of Dioscorea cordifolia, as figured by Ad. de Josse 
and 15 represented as bifid. (Ann. des Sciences Nat, gme Série, xi. Bot. pl. 17. fig. 13.) 

t Nov. Gen. et Sp. Plant. Brasil. tom. i. tab. 5. fig." 


AND THEIR GERMINATION. 409 


occurs in Alismacee, a near ally of the Hydrocharidee ; besides which, they may not im- 
probably prove to have the embryo of Triurideæ, which would lend some further support 
to such a comparison. And if the Bromeliacee should prove endorhizal, such a subdi- 
vision of the epigynous families would not be unnatural, as they connect Hydrocharideæ 
and Marantacee. | 

The following is added as a provisional arrangement of the Endorhizæ and their allies, 
including especially all those families in which the radicle is enlarged; and the remainder 
of the class Endogens thus separated will, I believe, be found to form by themselves a 
very natural section. But it must be confessed that no entirely natural arrangement 
. ean be expected to be made until the structure of the embryo and its germination in all 
the families is definitely ascertained. 


ALL, 3 
Musacem. 
å Amomeæ. 
S Marantaceæ 
å ALL. 1. ^ op 
åg : LL. 4. 
& Mane Aun. 2. ? Bromeliaceæ 
~ Apostasiaceæ. : 2 . 
urmanniaceæ. Hydrocharideæ. ? Taccaceæ. 
ALL, 9, dax. d 
Commelynaceæ. ad 
å Mayaceæ. ALL 4. Triurideæ. 
S Xyrideæ. Philydraceæ. me 
£;| Eriocauloneæ. Juncaceæ. — 
EN ABL. 9. 
ar en Potamogetoneæ 
= | Restiaceæ. Nai xd ; 
Desvauxiaceæ. E T? : 
Cyperaceæ. 
Unner 
SE Zosteraceæ. 
RE 
BS 
~ 
= 


EXPLANATION OF THE PLATE. 


Tas. LXVIII. 


first protrusion, before the first leaf appears. 


Fig. 1. Nuphar lutea. The radicle at the time of its 4. his leaf is rudimentary, having no 


Fig. 2. The seed of the same after the first leaf has appeare 


DER secondary radicle protruded, which has a coleo- 


Fig. 3. The same, after the second leaf has formed, and a adicles, which succeed. 
| rhiza, The same coleorhiza appears on the MT N N Que plumule having been laid 
Fig. 4. An embryo of a Zannichellia, the membranous sheath con g 
open to show the position of its first and second €— is perfectly adherent to the surround- 
Fig. 5. A longitudinal section of the same, showing that the penes of a canal is formed for it. 
; “ | 
ing tissue, and that in the ungerminated embryo no 58 


VOL. XXII. 


410 MR. B. CLARKE ON THE EMBRYOS OF ENDOGENS. 


Fig. 6. The embryo of Ruppia maritima, the membranous sheath containing the plumule having been 
laid open to show the position of its first and second leaves: a. the smaller lobe of the radicle. 

. Fig. 7. The embryo of Zostera marina, showing the smaller lobe of the radicle coiled over the cotyledon 
in its cavity. 

Fig. 8. The same, opened, and the cotyledon raised out of its cavity: a. the smaller lobe of the radicle. 

Fig. 9. The same, in longitudinal section, showing the radicle lying in a narrow canal at its base. The 
membranous sheath containing the plumule is opened to show that it is situated on that side of 
the cotyledon which is next the smaller lobe of the radicle, a. 

Fig. 10. The plumule more magnified, showing that it consists of two leaves: the first leaf, a, is directly 
away from the cotyledon. 

Fig. 11. A germinating seed of Eriocaulon septangulare, showing the primary radicle protruding, with a 
small circular portion of testa on its apex. 

Figs. 12 & 13. The same more advanced, fig. 13 showing the position of the first leaf of the plumule. 

Fig. 14. The same, showing the commencement of the formation of the first secondary radicle. 

Fig. 15. The same more advanced, showing that the secondary radicle has broken through the primary, 
and the first leaf of the plumule become elevated on a cauliculus. From the cauliculus another 
secondary radicle has protruded; it has no coleorhiza, but the surface of the cellular tissue 
divides to allow its escape. 

Fig. 16. A seed of Arum Dracunculus commencing to germinate, showing that it has no trace of a 
coleorhiza. 

Fig. 17. The same more advanced, showing a slightly elevated line marking the base of the cotyledon. 

Fig. 18. The same more advanced, showing the position of the first leaf of the plumule. 

Fig. 19. The same more advanced, the cotyledon having been removed to show a circular line at the base 
of the first leaf, similar to that at the base of the cotyledon in fig. 17. r 

Fig. 20. A germinating embryo of Oryza sativa: a. the upper part of the radicle as understood by 
Richard ; 4. the smaller lobe of the radicle (the epiblastus of Richard), which has been regarded 
as a second cotyledon ; c. the cotyledon, showing its relation to the upper part of the radicle. 

Fig. 21. A germinating embryo of Zea Mays: a. the cauliculus; à. the cotyledon as understood by 
Richard, the plumule having been partly drawn out of the fissure to show its position. 

Fig. 22. The embryo of Zostera nana, showing the aperture for the emission of the radicle, covered by a 
semitransparent membrane. 

Fig. 23. A germinating embryo of Luzula campestris extracted from the albumen. The stricture in the 
cotyledon is produced by the pressure of the testa after the radicle has protruded. 

Fig. 24. A germinating embryo of Butomus umbellatus. ’ 

Fig. 25. The same more advanced: a. part of the cauliculus (not cotyledon) is seen above the radicle. 

Fig. 26. A germinating embryo of an Iris, extracted from the albumen : a. the sheath of the. plumule in 
its earliest stage, : ; 


[ 4E ] 


XXXIV. Notes on the Structure and Affinities of Batidex, Callitrichaceæ, Vorne. 
| and Cassytheæ. By B. CLARKE, Esq., F.L.S. 


Read June 16th, 1859. 


1. On the Structure and Affinities of Batideæ and Callitrichaceæ. 


THE principal points in the structure of the genus Batis, P. Br., have become well known, 
but its affinities have not been decided, while those which have been suggested for it are 
proved to be, for the most part, if not entirely, impracticable; further analysis therefore 
seems required before its affinities can be brought to light; and the following additional 
particulars, as regards its structure, and especially the relative position of the ovules to 
the axis of the ovary, will, I believe, make it evident that it must be a near ally of the 
Verbenacee. 

The male flowers are arranged in 4-rowed spikes, giving them a habit like that of the 
Verbenacee with 4-rowed spikes of flowers; but in one species the inflorescence becomes 
compound, which gives it in some degree the habit of a Globularia, as of G. orientalis. 
_ The calyx is reduced to one sepal, posterior ; but, since in the Acanthaceæ also the calyx is 
sometimes very nearly obsolete, this negative character is of little value. The corolla is 
attached by a very slender basis to the calyx, and consists of four petals with long claws 
. and oval or almost circular laminæ, and of four stamens alternate with them, two of 
which are placed anterior and posterior, and two right and left of the axis *,—the two 
lateral stamens having the rudiment of an ovary slightly adhering to them, which is 
sometimes bifid, the segments being lateral. 
The female flowers gti siterip which are 4-rowed, as in the male. The floral 
envelopes are deficient; but each ovary has at its base a quickly deciduous bractea rar 
LXVIII. fig. 27), so that in the early stage the spike is completely enveloped in — 
and is also terminated with three or four pairs of minute flowerless scales. | 'The rare 
are two, right and left of the axis, and the cells of the ovary four, so placed aon idi 
them stand right, and two left of the axis, so as to correspond in their deniqne T ot 
two stigmas, each pair having no bundle of vessels between them, but m vhi 
by very distinct bundles from the opposite pair (fig. 27 a). Each pek LÉ kii 
anatropal ovule, the raphe and foramen of which are very distinct. : e wn ER 
in relation to the inner angle of each cell; but in each pair of cells the rap 


: Labia ving beyond 
turned towards each other (fig. 27 a), as in the Ver nn rar ped ae ls y 
i , consis is sona MD 
doubt that the structure of the ovary is the same, co cin Vostoitües vily tà being 


Accordi is view of its structure, Batis differs from | 
di rense tole unless it is in the position of the stamens; for in the female 


* 7 h l 1 s LPG $ og> Le 3 - ' 
3 > 1r posi 10n in B. mar ı > y y q * 


diclinous and polypetalous, 


412 MR. B. CLARKE ON THE STRUCTURE AND AFFINITIES 


flower no positive difference is apparent except in the carpels being right and left of the 
axis, instead of anterior and posterior: and in its deficiency of floral envelopes, it may, as 
far as relates to-the corolla, be compared to the Stilbaceæ, near allies of 7 erbenacee, the 
female flowers of which are apetalous. 

The position of the Batideæ, therefore, appears undoubtedly to be with the mono- 
petalous families; and however contrary to the rules of affinity it would have appeared 
only a few years since, this makes it in some degree probable that Callitrichacee should 
also be stationed there, and near Batideæ, as the ovary has precisely the same structure, 
being 4-celled, and consisting of two carpels, right and left of the axis. In Callitriche, 
however, the ovules are suspended, amphitropal, with the raphe next the placenta, in which 
it corresponds with Boraginee, or very nearly so*; the seed is albuminous, and the 
cotyledons are very short, as in Stilbaceæ, with which it further agrees in its anther, which 
closely resembles that of Campylostachys in shape, and in the peculiarity of the two cells 
being confluent at the apex, so that at the time of dehiscence it appears as if it had but 
one cell. 

Callitriche may be compared with Elatineæ, especially if the structure of the ovary of 
Tetradiclis can be considered as analogous to that of Callitriche ; but Batis does not 
appear to have any affinity with the polypetalous families, except that the long claws 
of its petals are like those which occur in Caryophyllacee. Callitriche differs from Eu- 


phorbiaceæ in its amphitropal ovules; besides which, hermaphrodite flowers are not 
uncommon, 


2. Note on the Structure and Affinities of Vochysiaceæ. 
One of the most remarkable characters in this family is that the carpel when single is 


with the single carpel posterior; and these characters occur in Daphnaceæ, of which the 
Vochysiaceæ may be a polypetalous form +, standing in the same relation to the Chrys0- 
balanee and Leguminose, and especially to the former, as Daphnace to Proteacee. 
The relation, however, between 7. ochysiaceæ and Daphnacee is rather one of analogy 
than affinity, although perhaps there is no character to separate them except the absence 


* + . 
ad ^is Boragineæ the Sier IS scarcely suspended, being attached near or at the base of the cell, at the inner angle ; 
ut the raphe of the amphitropal ovule is on its under surface—which is, doubtless, equivalent to the ovule being 


OF BATIDEÆ, CALLITRICHACEÆ, VOCHYSIACEÆ, AND CASSYTHEÆ. 413 


of petals in Daphnacee; and it is doubtless among the polypetalous families that the 
more immediate relations of Vochysiaceæ are to be sought for,—the vicinity of Rosacee, 
Leguminosæ, and their allies (which are regarded as polypetalous forms of Daphnaceæ 
and Proteaceæ) appearing to be the true station of this family. 

The relation of the Vochysiacee to Ohrysobalanee confirms this view of their aflinities, 
as they may be regarded as differing from Ohrysobalaneæ only in the carpel when single 
being posterior instead of anterior ; for it is on the posterior side of the flower that the 
stamens are deficient in both families, and the calyx in Vochysiaceæ is sometimes so 
nearly regular that the spur is scarcely apparent. 

To the Amygdaleæ they are doubtless equally allied, agreeing with them in the ovules 
when suspended having the raphe next the placenta * ; but as the flowers in this family 
are always regular, the approach between them is less obvious. 

To the Rosaceæ they may be compared in the remarkably convolute æstivation of the 
corolla; to Pomaceæ in the occasionally inferior ovary, the ovules when erect having the 
raphe next the placenta, and in the cotyledons being either flat or convolute; and to 
Calycanthaceæ in the imbrication of the calyx, which occasions the flower-buds to look 


like leaf-buds; and in their large anthers. 


3. Note on Cassytheæ. 


The affinity of this family having been for some time well known to be with the 
Lauracee, I have only to add that it agrees with them in the ovary consisting of a single 
carpel +, as is shown by a furrow on one side of the style, which is also of a paler colour 
(fig. 29), and by the ovule being attached to the same side of the ovary, so as to be in rela- 
tion with the furrow in the style, the ovule being pendulous, not from the apex of the cell, 
but from one side near the apex, as in Lauraceæ (vide Ann. Nat. Hist. ser. 2. vol. xi.). 

The carpel is also variable in its position in nearly the same degree as in the Lauracee, 
being occasionally posterior, and otherwise variable from anterior to lateral; and it agrees 
also with them in the ovule having the raphe dorsal (fig. 29), so that there does not appear 


to be any distinction between them except in parasitism. 


Spee i ith | next the 
* In those genera where the ovules are numerous, they are suspended, amphitropal, with the raphe 


placenta, and therefore the foramen is uppermost. : à 

f The ovary of Sassafras officinale consists of a single carpel (vide Am. Nat. er 2nd I, Eh De de 
sionally, although very rarely, it becomes dicarpous, with an imperfectly bifid et gemis of : TON 
of two ribs on the opposite sides of the ovary, which are alternate with the styles. ; Vat Saanyhes being 
momum, and Tetranthera have also decidedly the appearance of — of Gå carpel ; d aat p 
occasionally dicarpous, makes it probable that Nees v. Esenbeck is right in descri ren — 
when the ovules are two, and attached separately, the ovary would — NM n npe 


* 


414 > MR. B. CLARKE ON THE STRUCTURE AND AFFINITIES OF BATIDEÆ, ETC. 


EXPLANATION OF THE PLATE. 
Tas. LXVIII. 


Fig. 27. A portion of a spike of female flowers of Batis maritima, showing the scars left by the hade 
a. a transverse section of the front ovary, showing the relation of the cells to its two stigmas, and 
bundles of vessels interposed between them, anteriorly and posteriorly, so that they form pairs 
right and left of the axis. 'The ovules are also seen to be in pairs right and left of the axis, 
indicated by the position of their raphes. 

a 28. A longitudinal section of the ovary of Erisma violacea, showing the relative position of the 
spurred sepal and stamen, and that the ovules are attached on the anterior side of the ovary. 

Fig. 29. A longitudinal section of the ovary of Cassytha filiformis, showing the attachment of the ovule, 


and that the raphe is dorsal; also a furrow in the style on the same side as the attachment of 
the ovule. 


[ 415 ] 


XXXV. On the Origin and Development of the Pitchers of Nepenthes, wii an 
Account of some new Bornean Plants of that Genus. By J.D. es Pa 
F.R.S. & LS. $c. 


Read June 16th, 1859, 


Ina paper read before the Linnean Society on the 16th June, 1857, and published in 
this volume of the * Transactions” (p. 137), I mentioned, in a note, that I had examined 
and prepared drawings of the development of the pitchers of Nepenthes, from plants in 
the Royal Gardens at Kew, and that these confirmed Griffith’s observation * that the 
pitchers are modifications of the excurrent midrib of the leaf. I have now the honour 
of laying before the Society the principal facts observed during the examination in 
question, which have further led me to the conclusion that the pitchers are modifications 
of a gland situated at the apex of the midrib of the leaf; and I have added the necessary 
illustrations. I shall also append to this paper an account of the most singular and 
gigantic plants of the Order, which have lately been discovered by H. Low, Esq., on the 


lofty mountain of Kina Balou in Borneo. 


Part I.—On the Development of the Pitchers of Nepenthes. 

At the earliest period at which I have examined the undeveloped leaf of the full-grown 
plant of Nepenthes levis of our gardens t, I find it to consist of p — ue ou 
body, about 45th of an inch long (Tas. LXXIV. fig. 12), with a slight longitu " 
depression on the anterior surface, leading to a shallow oval cavity, which is cedet 
diately below the apex. At this period the cuticle is scarcely sri rper ort 
depression has a more shining appearance and less defined rer t ^g] + thie | ve 
any other part of the cone. On a vertical section carried tak; i f ves ol å ee 
and oval depression (Ag. ) the whole substanco is found to be formal oa mer 
dense parenchyma, becoming looser, and as it were deliquescent : e bb À id 
oval cavity is all that represents the future pitcher, and it is simp mi; d that at which 

_ The next period (as defined by a marked progress oe specializa Dis ol Dé teak død à 
there is an evident differentiation of the conical body into wis rolonged upwards, and 
superimposed body. At this stage (fig. 3) the nn — | eom as defined margins. 
about 35th of an inch long. The frontal groove 1s joa it most marked 
There is an evident contraction about halfway between ne tester à ær deb cn 
on the edges of the groove. The apex of tb ec Pen 2 d downwards. There is a 
the oval cavity, which has deepened, and is directed inwards and do | 


* In “Calcutta Journal of Natural History,’ vol. iv. no. xiv. July one 

t It is the N. gracilis, Korth. I have to record my obligations to Tt © 
Messrs. Veitch and Son, of the Exotic Nurseries, Chelsea and verte 
aided me by specimens of rare species in their establishments, for examination. 


1843, p. 231. | : 
í Messrs. Low, of Clapton Nurseries, and 


liberal manner in which they have 


416 -. DR. J. D. HOOKER ON THE ORIGIN AND DEVELOPMENT 


more evident cuticle to the whole surface of the cone, except over the gland, which now 
appears to be a secreting surface, and is rather pulpy or viscid. 

At the next succeeding period, when the whole body is about 2;-æ5th of an inch long, — 
a considerable further change has taken place (fig. 4). The contraction towards the middle - 
has greatly lengthened and forms a neck, which separates an elongated conical base (the 
future lamina) from an oblong terminal body (the future pitcher). The groove is still 
shallow on the intermediate portion (the excurrent midrib) and on the pitcher above it, 
but is deepened into a canal with incurved edges on the lamina below it. The gland is 
now å deep cavity, with a large quadrangular orifice, over which hangs the incurved apex 
(the lid of the future pitcher) of the original conical body. A vertical section (fig. 5) 
through the centre shows a narrow pale line ascending close to the dorsal margin, curving 
over the apex and terminating in the incurved apex; this line indicates the position of 
the future vascular bundles. At fig. 4 a minute conical body is seen at the base of the 
groove, and is a nascent second leaf. 

At the next following marked stage, the lamina, excurrent midrib, and pitcher are 
externally very well defined, and the lid of the pitcher differentiated. The lamina is still 
a cone, but much elongated, and presents a cylindrical groove with the edges slightly 
turned in, representing the earliest appearance of that organ in its involute condition of 
vernation. The contracted portion above it (the excurrent midrib) has elongated more in 
proportion than the lamina or pitcher, and its frontal groove is less marked. The apex of 
the pitcher has grown upwards and backwards ; the incurved portion, again, has so grown 
downwards as to close the orifice of the cavity. On a vertical section (fig. 7) the cavity 
is seen to be prolonged downwards as a sac parallel to the walls of the pitcher, whilst the 
incurved portion or lid has both grown downwards over the mouth of the cavity, and 
inwards towards its dorsal face; its protuberant inner surface is more papillose than any 
part of the sac, As the pitcher enlarges, the apex, which points upwards and backwards, 
elongates, at first considerably more than the lid itself, and becomes the styliform process 
at the junction of the lid and pitcher. On a vertical section (fig. 8) the vascular bundle is 


seen fo run up to the apex of this process, which thus continues to be the true organic z | 2 


apex of the whole foliar organ. E 
The succeeding stages of growth were examined in N. Rafflesiana and phyllamphora M 


and present few external characters but what have been previously observed. In ie 
former species the terminal process often spreads and divides, and, together with the Å = 
whole pitcher, is covered with appressed hairs. These hairs point upwards everywhere mes 
except on the young lid, where they either project or point in the opposite direction. On 


à vertical section at this time, the lid is found to have reached the lower portion of the 
orifice, and it afterwards closes the mouth of the pitcher (fig. 10) ; and one or two recur- 
rent fascicles of vessels are found to originate in the principal vascular bundle in its 
course towards the apex of the young styliform process, which bundles enter the sub- 
stance of the lid, and ramify in it. ES 
The vernation of the leaf in Nepenthes is well known to be involute, the opposite 
he. dm being ‚each rolled in towards the midrib. This curious arrangement, whichis ` 
no% peculiar to the genus, appears to me to be a secondary one, or rather an induced con- 


OF THE PITCHERS OF NEPENTHES. 417 


dition; for in some very young leaves of N. Rafflesiana I observed, that as the frontal 
groove which represents the anterior (or upper) surface of the leaf deepened and its 
margins expanded, these margins decidedly overlapped (fig. 11), thus forming a convolute 
vernation ; but that, owing to a more rapid growth of the overlapping hemiphyll, which 
still retained its initial tendency to curve inwards towards the midrib, it eventually 
formed a Cylindrical roll parallel to, and in contiguity with, the originally overlapped 
hemiphyll. 

If these observations should receive confirmation, they will tend to prove that the 
involute vernation of leaves may be, in some cases at any rate, a modification of the more 
frequent convolute condition of these organs. | 

In the above description I have used the term adopted by Griffith, of * excurrent 
midrib ;” but I need hardly say that the intermediate organ, or stalk of the pitcher, is no 
more an excurrent portion of the lamina, than is the petiole in those species where that 
organ is developed: it is a body more or less strictly analogous to the terminal cirrhus 
of the leaf of Gloriosa, Flagellaria, various species of Convallaria, &e.; and though the 
development of a gland at its apex into a pitcher is very anomalous, and possibly unique*, 
the existence of terminal glands in the leaves of plants, which is physiologically the most 
important feature, is by no means so unusual a fact as may at first sight appear. ‘They 
are very conspicuous at the apex of the leaf of Limnocharis, Caladium, and other genera, 
both of monocotyledonous and dicotyledonous plants. In young Limnocharis Plumieri, 
especially, a gland is placed at the apex of the midrib, and being hollow, resembles the 
early condition of the Nepenthes pitcher. 

Since the above observations were made, I have had the opportunity of examining some 
seedling Nepenthes, for which I am indebted to Hugh Low, Esq., of the Nursery, Clapton, 
and which are both extremely singular in themselves, and throw much light on the whole 
subject. One of these (species unknown) is figured of the natural size in Tax. LXXIV. 
fig. 19, and magnified at fig. 20. The first pair of leaves (fig. 20 a & 21), the cotyledonary, 
are opposite, lanceolate and acute; the following at once bear ascidia at their apices, of a 
size, form, and in a position that cause the whole leaf strikingly to resemble that Z 
cenia. The youngest leaves I have examined, namely those immediately me a; 
cotyledonary, have a more or less dilated lamina, and a hollowed-out —— dp 
midrib, partially closed with a small ciliated lid; as the leaf grows, 2 c = 
upwards on each side, so as to become cuneate, and finally obobidete A Kost La inch 
apex, with the pitcher in the sinus. When fully developed, these — € u F if Pa siå 
long ; they present on the upper surface a shallow mesial groove, Er re — ; 
slightly tumid area towards the apex, which area answers to the anterior face p ; 


beyond the apex the neck of the pitcher protrudes, with its lid; the margins of the leaf are 


carried up as wings along the sides of the neck of the pitcher to near its orifice, where they 


meet, and form a transverse, reflexed, ciliate membrane. 
Lookine at the under surface of the leaf, the midrib appears very år = je m 
dually Baines as it advances towards the apex of the leaf, forming the belly o p 


s to the origin of the pitcher in Cephalotus and nm 
31 


* I have not yet satisfied myself a 
VOL. XXII. 


418 DR. J. D. HOOKER ON THE ORIGIN AND DEVELOPMENT 


The examination of these remarkable seedling plants suggests the following obserya- 
tions :— 

1. The difference in the development of the leaf and pitcher in these seedling plants, 
and in the full-grown ones previously described, is very great, and at first sight anomalous. 
In the full-grown plant, the lamina, petiole, excurrent midrib, and pitcher are very inde- 
pendently differentiated, and the pitcher itself is first developed in the most rudimentary 
conceivable condition, that of a simple naked gland. In the seedling plant, on the other 
hand, the pitcher and lid appear to be developed in the earliest discernible condition of 
the leaf, which is that of a hollow midrib open at the apex and there closed with a lid, along 
each side of which midrib the lamina becomes developed in one plane (not with a convo- 
lute or involute vernation). As the plant grows, the upper part of the hollowed midrib 
of each succeeding leaf becomes more and more inflated, its apex protrudes beyond the 
lamina, as the neck of the pitcher, and the orifice and lid of the latter assume the usual 
highly organized condition of these parts in the genus. 

2. The position of the pitcher, occupying chiefly the underside of the leaf, is very 
remarkable, the appearance of the whole being not that of a pitcher with foliaceous 

margins, but of a leaf with a piteher partly adnate to its under surface; and tbe larger 
the leaf is, the more independent does the pitcher appear to be, and the more confined to 
the apex of the leaf; so that I expect that in more advanced states of the seedlings of 
this species, the pitcher will be found to be wholly free from the lamina of the leaf, 
though continuous at its base with the midrib *. When the plant arrives at such a stage _ 
of growth that the lamina of the leaf becomes a larger and more important organ than the 
pitcher, then the vernation of the leaf will assume the normal condition which obtains in 
the old plant. 

3. The horizontal development of the lamina on the sides of the pitcher, and the pro- 
longation of the margins of the lamina on the neck of the pitcher, at first sight seem 
to suggest the view that in the old plant the lamina of the leaf is represented by the 
wings of the pitcher, and that the apparent lamina is only a winged petiole. But in the 
seedlings the produced margins of the lamina do not reach the mouth of the pitcher; on 
the contrary, they converge, and form a transverse membranous wing below its orifice ; 
and the older the leaf is, the longer is the neck of the pitcher produced beyond this trans- 
verse lamina: and if the oldest of these seedling leaves be compared with that of a full- 
grown Nepenthes, it would appear possible that the transverse lamina is the true apex of 
the leaf, which in the old plant forms an elevated ridge on the anterior face of the base of 
the stalk of the pitcher+. This ridge, though generally small in most full-grown Jeaves, 
is often very prominent, so much so in N. Rajah (Tas. LXXIL) that the stalk of the 
pitcher is there peltately attached to the back of the leaf. , 


` Since the une observations were made, I have received from Messrs. Veitch more advanced seedlings, which 
ren this—the pitchers being wholly free from the lamina, but continuous at their base with the midrib. (October, 
se rette do not confirm this idea : it appears that the anterior wings of the old ye 
E: P ds of - lamina in the seedling pitcher, and that the transverse process uer 
üs ge an ors ^ the young pitchers is not developed in the youngest leaves of the seedling pps = 
need it occupies the whole space between the mouth of the pitcher and the transverse lamina. 


OF THE PITCHERS OF NEPENTHES. 419 


4. In my description of the development of the leaves in full-grown plants of Nepen- 
thes, I have stated that the gland which is developed into a pitcher occupies a position 
towards the apex of the nascent leaf; the examination of the seedling plants would tend 
to show that the position of the gland indicates the organic apex of the future midrib, and 
no doubt it does so (as may be seen in Limnocharis Plumieri). Such glands often occur 
on the margins, midribs, veins or petioles of leaves, and are most frequent at the anasto- 
moses of the veins, as at the base of the lamina (Cucurbitaceæ, Leguminose, &c.), or at the 
serratures where the lateral nerves meet the marginal (Aurantiaceæ, Myrsineæ, &e.), or 
where the lateral veins finally converge at the apex of the midrib (as in Nepenthes, Limno- 
charis, &e.). 

5. The sudden transition from the simple cotyledonary leaves of a seedling Nepenthes, 
to the pitcher-bearing leaves immediately contiguous to them, is extremely remarkable. 
There is, in the species I have examined, no transitionary stage of development whatever. 
This renders the formation of the pitchers of Sarracenia and Cephalotus (which, so far as 
I have observed, never present the appearance of ordinary leaves) less anomalous, than if 
a seedling Nepenthes presented a graduated series of more and more highly organized 
leaves connecting the simple cotyledonary with the fully developed pitcher-bearing ones. 

The resemblance between the pitcher of a seedling Nepenthes and that of Sarracenia 
purpurea is very close, and leaves little doubt in my mind that that organ is strictly 
homologous in the two genera. I have never seen seedlings of Sarracenia, nor of Cepha- 
lotus, but a comparison of young leaves of the latter with those of Nepenthes presents 
several curious similarities. In Cephalotus the ordinary leaves are perfectly simple, and 
similar to the cotyledonary leaves of Nepenthes ; and the pitcher-bearing leaves are at 
once developed as such, having the cavity and ciliated lid in their earliest discernible con- 
dition: though these occur both above and below the ordinary leaves, and in immediate con- 
tiguity with them, there are no intermediate stages whatever, the transition from cauline 
leaf to pitcher being as sudden and abrupt as from cotyledonary leaf to piteher ce 
Nepenthes. The appearance, too, of the young Cephalotus pitcher and stalk is p 
stout petiole, with a hollowed-out terminal head obliquely adnate to its lower surface. Pu 
the analogy with Nepenthes holds good, the stipes of the Cephalotus pitcher represents 
midrib of a leaf on whose sides no lamina is developed. 


Parr IL— Notes on the Bornean Species of Nepenthes, with descriptions of the new | 

The want of any important characters in the flowers and fruit of Nepenthes is : de re- | 
markable feature of these plants. The leaves differ considerably kr ge and ut 
more or less petioled. The pitchers of most, when young, are shorter, rovided 


420 DR. J. D. HOOKER ON THE BORNEAN SPECIES OF NEPENTHES. 


NEPENTHES. | 
$ I. Ascidia magna, ore lamellis latis disciformibus annularibus remotis instructo. 


1. NEPENTHES VILLOSA, H. f. (Hook. Ic. Pl. t. 888). Ascidia magna turgida late pyri- 
formia coriacea 5" longa 32” lata, alis anticis mediocribus grosse dentatis, ore aperto 
annulo maximo! lamellis annularibus distantibus disciformibus rigidis 1" diam, 
cristatis posticis in spinas rigidas 3" longas fundum ascidii spectantibus productis, 
collo elongato erecto, operculo orbiculato intus densissime glanduloso dorso basi 
longe cornuto. (Tas. LXIX.) 

Hab. Borneo (Lobb). Kini Balou, alt. 8000-9000 feet (Low). 


This most remarkable pitcher resembles that of N. Edwardsiana in so many respects, 
especially in the size, form, and disposition of the distant lamellze of the mouth, that I am 
inclined to suspect that it may be produced by young plants of that species, before it 
arrives at a stage when the pitchers have elongated necks. 

The whole inner surface of the pitcher is glandular, except a very narrow area beneath 
the mouth at the back. 


2. NEPENTHES EDWARDSIANA, Low, MSS. Foliis (6" longis) crasse coriaceis longe petio- 
latis ellipticis, ascidiis magnis crasse pedunculatis cylindraceis basi ventricosis 8-18" 
longis, ore lamellis annularibus distantibus rigidis magnis cristato, collo elongato 
erecto, operculo cordato-rotundato, racemo simplici, rachi pedicellisque ferrugineo- 
tomentosis. (Tas. LXX.) 


Hab. Kina Balou, north side, alt. 6000-8000 feet (Low). 


Mr. Low desires that this magnificent plant should bear the name of the Honourable 
George Edwardes, Governor of Labuan, who has materially assisted him in his expeditions. 
Under N. villosa, I have stated my suspicions that this may be the more mature form of 
that plant with elongated pitchers. The leaves, ascidia, and pitchers sent by Mr. Low 
are all old, and nearly glabrous; but the young parts—rachis, peduncles of the panicle, 
and the calyx—are covered with ferruginous tomentum. One of the pitchers sent is 
18 inches long from the base to the apex of the erect operculum; it is 21 inches in dia- 
meter below the mouth, 14 at the narrowest part (about one-third distant from the base), 


and the swollen part above the base is about 2 inches in diameter. The beautiful annular 
dises which surround the mouth are 4 inch in diameter. 


$ IL. Ascidia magna, curva, basi inflata, medio constricta, dein ampliata, infundibuliformia ; ore maximo, 
latissimo, annulo 0. 


3. NEPENTHES Lown, H. f. Caule robusto tereti, foliis crasse coriaceis longe crasse petio- 
latis lineari-oblongis, ascidiis magnis curvis basi ventricosis medio valde constrictis, 
ore maximo ampliato, annulo 0, opereulo oblongo intus dense longe setoso. (TAB. 
LXXI.) 

Hab. Kina Balou, alt. 6000-8000 feet (Low). 


Å noble species with very remarkable pitchers, quite unlike those of any other species. 
They are curved, 4-10 inches long, swollen at the base, then much constricted, and suddenly 


DR. J. D. HOOKER ON THE BORNEAN SPECIES OF NEPENTHES. 421 


dilating to a broad, wide, open mouth with glossy shelving inner walls, and a minute row 
of low tubercles round the circumference ; they are of a bright pea-green, mottled inside 
with purple. The leaves closely resemble those of Edwardsiana and Boschiana in size 
form, and texture, but are more linear-oblong. p 

I have specimens of what are sent as the male flower and fruit, but, not being attached, 
I have not ventured to describe them as such. The male raceme is 8 inches long, dense- 
flowered. Peduncles simple. Perianth with depressed glands on the inner surface, ex- 
ternally rufous and pubescent. Column long and slender. Female inflorescence : a very 


. dense oblong panicle; rachis, peduncles, perianth, and fruit covered with rusty tomentum. 
Capsules 3 inch long, å broad. 


$ III. Ascidia magna, ore mediocri, annulo latissime explanato, dense lamellato v. costato. 


4. NEPENTHES VEITCHII, H. f. 
N. villosa, Bot. Mag. t. 5080; non Ic. Plant. t. 888. 
Hab. Borneo (Lobb), alt. 1000 feet. Gunoong Mooloo, alt. 3000 feet (Low). 


5. Nepentues Rasam, H. f. (Frutex 4-pedalis, Low.) Foliis maximis 2 pedalibus 
oblongo-lanceolatis petiolo costaque crassissimis, ascidiis giganteis (cum operculo 
1-2 pedalibus !) ampullaceis ore contracto, stipite folio peltatim aflixo, annulo 
maximo lato everso crebre lamellato, operculo amplissimo ovato-cordato ascidium 
totum æquante! (Tas. LXXII.) 

Hab. Borneo, north coast, on Kina Balou, alt. 500 feet (Low). 


This wonderful plant is certainly one of the most striking vegetable productions 
hitherto discovered, and in this respect is worthy of taking place side by side with the 
Rafflesia Arnoldii; it hence bears the title of my friend Rajah Brooke, of whose services 
in its native place it may be commemorative amongst botanists. Mr. Low describes it as 
a shrub 4 feet high; and the pitchers being of that short aniipallacdons form which nen 
times denotes that the plant producing them is ee it = possible that, at a differen 
season, it may attain a greater stature and bear different pitchers. . 

I have En two er of leaves and pitchers, both quite similar, but one twice as 


large as the other. Of these the leaf of the larger is 18 inches long, exclusive of the 


i ichi i ; broad, very coriaceous and glabrous, with 
petiole, which is as thick as the thumb; and 7-8 ry ee t 


indisti T i is gi ff below 

. indistinct nerves. The stipes of the pitcher 1s given 0 iore 
: : ampullaceous pitcher is 6 inches 1n 
20 inches long, and as thick as the finger. The broad amp ao Mifruty 


i i i ings in front, is cov 
diameter and 12 long; it has two fimbri ted wings in , nd 
hairs above, is wholly studded with glands within, and the broad annulus is eve 


: TET ipi 10 inches long and 8 broad. 
1-11 inch in diameter. Operculum shortly stipitate, 30 inches long, of which 20 are 


: : i ion. Male raceme 
The inflorescence is hardly in proper rs clothed with short rusty pubescence. 


occupied by the flowers; upper part and flowe 13 inch long, 
Ponidi ide simple or pifid. Fruiting raceme renere rd 
often bifid. Capsule $ inch long, $ broad, rather turgid, ; 

tomentum. . 


422 DR. J. D. HOOKER ON THE BORNEAN SPECIES OF NEPENTHES. 


6. NEPENTHES BoscHiANA, Korth. ; Miquel, Flora Ned. Ind. i. 1074, cum syn. 
Hab. Borneo (Korthals). Mount Mooloo, alt. 3000 feet (Low). Also (var. 8) found in Sumatra. 
Pitchers 15 inches long. Plant 15-20 feet high. 


7. NEPENTHES RAFFLESIANA, Jack; Miquel, Flora, i. 1070, cum syn. &e. 


Hab. Labuan (Mottley). Kina Balou, alt, 3500 feet (Low). Also found in Sumatra, Malacca, and Sin- 
capore. 


8. NEPENTHES PHYLLAMPHORA, Willd.; Miquel, Flora, i. 1069, cum syn. 
Hab. Labuan, alt. 2500 feet; very rare (found by Mr. T. Lobb) (Low). Also found in Malacca, Sinca- 
pore, Java, New Guinea, and China. 
I have no Bornean flowering or fruiting specimens of this species, which may be 
known by its stout terete stem, long petioled ciliated leaves (when young), and cylindrical 
pitchers, which are swollen at the base, and have a contracted mouth surrounded with a 
broad flat annulus. Its foliage resembles that of N. Jimbriata, from which it differs chiefly 
in the broad annulus. 


§ IV. Ascidia cylindracea, ore mediocri, annulo angusto creberrime costato v. striato v. sublevi. 


9. NEPENTHES REINWARDTII, Miq. Flora, Z. c. 
Hab. Mount Mooloo, alt. 3000 feet (Low). 


Apparently chiefly differs from N. gracilis in the pitcher bulging at the base. 


10. NEPENTHES GRACILIS, Korth.; Miquel, Flora, i. 1071, cum syn. &c. 
N. levis, Hort. 

Hab. Borneo (Korthals). Common along the N.W. coast (Low). | 

8. elongata. Borneo (Low). Also Malacca and Sincapore, 


Varies much in size, and in the length of the decurrent part of the leaf. 


11. NEPENTHES FIMBRIATA, Blume, Mus. Lugd. Bat. ii. p.7; Miquel, Flora, i. 1072. 
Hab, Borneo (KortAals). Also found in New Guinea and the Louisiade Archipelago. 


~ Apparently differs from N. gracilis chiefly in the narrower annulus. 


12. NEPENTHES ALBO-MARGINATA, Hook. MSS. Glaucescens, caule tereti apice foliis 
subtus et inflorescentia rufo-tomentosis, foliis anguste lanceolatis basi longe angus- 
tatis vix petiolatis coriaceis acuminatis, ascidits cylindrieis, annulo angusto minute 
creberrime striato margine lato albo-tomentoso cireumdato, racemo tenui, pedicellis 
subternis filiformibus, floribus parvis. (Tap. LXXIII.) 

Hab. Borneo (Lobb). Maritime rocks near the mouth of the Lokotan and Tanjong-poe Rivers (Low). 
Also found at Sincapore, Lobb, 

Species pulcherrima, distinctissima, colore pallido, foliis angustis non decurrentibus, ascidiis plerumque 


tomentellis, annulo angusto late albo-marginato. 
This very beautiful species is sent by Lobb both from Borneo and Sincapore. The 


DR. J. D. HOOKER ON THE BORNEAN SPECIES OF NEPENTHES 423 


pitchers are usually covered with a short white fömentum, and the broad white velvet 
. band surrounding the narrow polished brown annulus affords a very marked vist 


The pitchers of the Sincapore specimen are (when dry) of a beautiful rose-colour blotched 
with purple. 


18. NEPENTHES MELAMPHORA, Reinwardt ; Miquel, Flor. i. 1072, cum syn. &c. 


B. lucida, Blume, Mus. Lugd. Bat. ii. p. 8. 
Hab. Southern Borneo (Korthals, Mottley). Also found in Eastern Bengal and Java. 


l have received Bornean specimens of this plant, collected by Mottley at Banjarmassing, 
on the south coast of Borneo, where it was first detected by Korthals. The originally 
described form grows at an elevation of 3000—5000 feet in Java. 

I take the East Bengal Nepenthes to be the same species; it was sent from that country * 
by Carey to Europe, and published by Graham in the * Botanical Magazine ' (t. 2798) as 
the Ceylon N. distillatoria, L. (from which it is very different). It has also been gathered 
by Griffith in the same place, and by Dr. Thomson and myself near Amwee, on the 
Jyntea Hills—the eastern continuation of the Khasia Mountains and north of Silhet, 
whence probably all the specimens were sent, as the plant and that particular habitat are 
well known to the natives of those districts. The figure in the * Bot. Mag.’ well repre- 
sents the Bengal plant in every respect ; but the dried raceme in Herb. Hook., from which 
the drawing was made, is much more tomentose than any specimens gathered by Thomson, 
Griffith, or myself, or by Korthals, Blume, or Reinwardt in Java. One of Griffith's 
Bengal specimens is in this respect intermediate between ours and those of the Edinburgh 
Gardens. Korthals’ figure is not characteristic, except of a small half-starved-looking 


_ form. 
I have a pitcher of what I take to be this species upwards of a foot long ; it was grown 


in Mr. Rollisson's Nursery. 


F 


$ V. Ascidia brevia, ore contracto, annulo lato creberrime sulcato, operculo parvo. 


14. NEPENTHES AMPULLARIA, Jack ; Hook. Comp. Bot. Mag. i. 271; Miq. Flora, i. am 


cum syn. &c. 


. Hab. Borneo (Lobb, Low, Mottley). Also found in Malacca and Sumatra (Jack, Cuming, Korthals) 


EXPLANATION OF THE PLATES. 


Tag. LXIX. 


Nepenthes villosa, HE a... 
i i . Fruit. ; 5. Fo f 

Fig. 1. d raceme; nat. size. Fig. 2. d flower. Fig. 3. 9 flower. Fig. 4. Fruit. Fig 
under surface of lid :—all magnified. 


å : here—the 
å Khasia Mountains are those meant 
* From the Circar Mountains in N.E. Bengal, Bot. Mag. Le. The % 


Circars being in the Peninsula. 


424 DR. J. D. HOOKER ON THE BORNEAN SPECIES OF NEPENTHES. 


Tas. LXX. 


Nepenthes Edwardsiana, Low. 
Fig. 1. Fruit. Fig. 2. Portion of under surface of pitcher :—both magnified. 


Tap. LXXI. 


Nepenthes Lowii, H. f. 


Fig. 1. Portion of 3 raceme, and Fig. 2. of 9; both nat. size. Fig. 3. g flower. Fig. 4. Young fruit. 
Fig. 5. Portion of under surface of pitcher :—all magnified. 


Tas. LXXII. 


Nepenthes Rajah, H. f. 


Fig. 1. d flower. Fig. 2. Portion of fruiting raceme. Fig. 3. Ripe fruit. Fig. 4. Portion of under 
surface of pitcher :—Figs. 1, 3 and 4 magnified. | 


Tas. LX XIII. 


Nepenthes albo-marginata, Hook. 
Fig. 1. d flower. Fig. 2. 9 flower. Fig. 3. Portion of under surface of pitcher :—all magnified. 


| Tas. LXXIV. 


Illustrations of the Development of Nepenthes pitchers. 


Fig. 1. Conical terminal bud of N. gracilis, Korth. (N. lævis, Hort.) (a, nat. size). 
Fig. 1 b. Apex of the same rather more advanced, representing a leaf in its earliest state. 
Fig. 2. Longitudinal section of the same, | 
Fig. 3. More advanced condition of ditto, showing the differentiation of the lamina below, and exeurrent 
midrib and pitcher above. e | 
Fig. 4. Still further advanced state of terminal bud and leaf. a. is a second leaf. 
Fig. 5. Vertical section of apex of the same. ` 
Fig. 6. Apex of leaf after the lamina, excurrent midrib, and pitcher are all differentiated. 
Fig. 7. Apex of pitcher from the same, cut vertically. 
Fig. 8. Vertical section of young pitcher of N, Rafflesiana. 
_ Fig. 9. Further developed Pitcher of ditto. 
. Fig. 10. Vertical section of pitcher of ditto. 
Fig. 11. Convolute vernation of rudimentary lamina of leaf of N. Rafflesiana. 
‚Fig. 12. More highly developed pitcher of ditto, cut vertically. 
Fig. 13. Very young pitcher of N. phyllamphora ? 
Fig. 14. Vertical section of ditto. 
. Fig. 15. Further advanced pitcher of ditto. 
Figs. 16 and 17. Vertical sections of still further developed pitchers of ditto. 
Fig. 18. Vernation of very young leaf of N. Rafflesiana. 
Fig. 19. Seedling Nepenthes from Borneo, from Mr. Low’s Nursery :—nat. size. 
Fig. 20. Ditto magnified, a, cotyledonary leaves. 
Fig. 21. Cotyledonary leaf of ditto. 
Figs. 22-95. Various views of leaves of ditto. 


All the above figures but fig. 19 are more or less highly magnified. 


SENS ST 


Lg 2 u 


» 
~ 


^ ANN tiis 


* Å t 


te 


mul 


+ Kat PEUT bal 


FRE, 


fød 


x 


M À MMÀ— € 


jui t P t, 


+ rei a m_ TY, * T m T T 1 
LINN. Soc. TRANS. Vor. XXII Tas LXXI ` 


ine 
{R= SF 
I 


2277771 LÀ M 


NEPENTHES LOWII, 


€ 


a t Vea" 
quem omen: 7 = 


1 
t 
e 
1 


. 


Lin. Soc. Trans. Vor. XXII. TAB. 


ee nn game 


3 RO E MÀ 


Sp 


SS Lc 


INIT 
P aae 


x ad 


Fitch, del et lith. NEPENTHES ALBO -MARGINATA, Hook 


Days Son, Litto the Que 


Liny Soc Vor, XXILzas 1 XXIV » 424 


RANS 


T 


mnt 
Mert strate ne PT 


IT? 


Der, 


Ap 


Development. of Nepenthes leaves. 


TDF ae [itch Tah. 


[ 425 ] 


XXXVI. On a New Genus of Balanophoreæ from New Zealand, and Two New Species of 
Balanophora. By J. D. HooKERy M.D., F.R.S. $ L.S. $c. 


. Read June 16th, 1859. 


SINCE laying before the Linnean Society my observations on the structure and affi- 
nities of Balanophoreæ *, I have received two additional species of Balanophora, and a 
remarkable new genus of the Order, which I shall now proceed to describe under the 
name of Dactylanthus. 


DACTYLANTHUS, n. g. 


Dioicum. Rhizoma tuberosum, lobatum. Pedunculi plurimi, squamati, apice spadices plurimos gerentes. 
Flores in spadices dense conferti, inferiores dissiti. Fr. masc. Stamen solitarium sessile, filamento 
brevissimo, anthera 2-loculari. Fr. raw. Perianthium ovario globoso subsessili adnatum, lobis 2- 
subulatis, stylo filiformi, stigmate simplici. 


1. DACTYLANTHUS TAYLORI, H.f. (Tas. LXXV. A.) 


Hab. Nova Zelandia ad radices Fagi et Pittospori montibus prope Wanganui, alt. 4000 ped. (Rev. R. 
Taylor, M.A.). Nom. vern. Pua reinga (Flos Hades). 

Rhizoma magnitudine pugilli et ultra, tuberculatum, superficie celluloso, pustulis papillisve nullis, intus 
sicco rubrum. Pedunculi numerosi, 4-6 unc. alti, sicco fragiles, rugulosi, opaci, ferruginei, intus 
rufescentes, squamis imbricatis laxe tecti, apice in receptaculum floriferum dilatati. Squamæ ob- 
longæ, obtusze, lata basi sessiles, inferiores dissitæ, 1 unc. longæ, brunneæ, superiores sensim majores, 
supremæ inflorescentiam involucrantes, 1-2 unc. longæ, lineari-oblongæ, membranaceæ, albæ medio 
sordide brunnez v. rubræ.  Spadices (v. pedicelli floriferi) 10-30, conferti, erecti, 1-2 unc. longi, 
cylindracei v. fusiformes, crassitie pennæ corvinæ, stipitati, basi nudi v. floribus deformatis bracteolati. 
Flores dense conferti. Masc. Stamen solitarium nudum, sessile, filamento brevissimo crasso; an- 
thera adnata, 2-locularis, transverse oblonga, rimis subverticalibus dehiscens, loculis crassiusculis, 
inferiores in spica imperfectæ dissitze stipitatæ. Pollen pallidum, globosum. Fr. raw. sicco nigri, 
sub i unc. longi. Ovarium stipitatum, subglobosum. Perianthium supra ovarium paulo angusta- 
tum, dein bi-, rarius 3-partitum, a dorso compressum, cum stylo connatum ; segmentis respectu 
axeos lateralibus, subulatis, inzequalibus. Stylus cylindricus, teres, superne paulo nr ey apice 
obtusus.—“ Planta viva odorem subgratum Melonis, sed demum ingratum humi spirat.”— Taylor. 


For a specimen of this singular plant I am indebted to my friend the Rev. R. Taylor, of 
New Zealand, who brought a fragment of it to England in 1856, and on my pointing out 
its probable interest, promised to procure more on his return to New Zealand. This he 

did; and early in the present year I had the pleasure of receiving from him a dried 
specimen of the female plant, a perfect male inflorescence in a letter, and a pen and ink 
sketch of the peduncle and flowers, with notes on the same. Owing to the great con- 
traction of the cellular tissue, and blackening and fragility of the only entire specimen, 
which was unfortunately loosely packed in a box and much damaged, I have had con- 
siderable difficulty in so restoring the parts as to enable Mr. Fitch to complete the 
accompanying Plate, and myself to make the necessary analyses. 
| * Page 1 of this volume. 


VOL. XXII. Ex 


426 : — DR. J. D. HOOKER ON A NEW GENUS OF BALANOPHOREÆ 


The most remarkable character of Dactylanthus is its inflorescence, which, instead of 
presenting the solitary capitulum or spadix of most of its allies, or the branched cha- 
racter of Sarcophyte, consists of numerous erect spadices, densely covered with flowers. 
Of these the males consist of a solitary stamen, not articulate with the spadix, and without 
any trace of perianth, and hence present by far the most reduced form of male flower in 
the Order. The female spadices present a crinite appearance from the long flexuose styles 
of the flowers. The ovary appears to be compressed, furnished with a short thick stipes, 
and its upper part is contracted into a short thick neck, apparently consisting of the tube 
of the perianth connate with the base of the style: this contracted part varies much in 
width and breadth, and sometimes appears as if reduced to the adnate bases of the subu- 
late lobes of the perianth. 

I have not been able to ascertain the nature either of the contents of the ovary, which 
is extremely minute, and crumbles into black powder on being touched, or of the tissues 
of the rhizome and peduncle, which do not seem to differ materially from those of other 
Balanophoree. 

The position of Dactylanthus is amongst my first group of Balanophoree, the Mono- 
styli; but in many respects it partakes of the structure of various genera of Distyli, and 
is not very nearly related to any described genus. In habit it presents the rhizome of 
Balanophora. In the male flowers reduced to a solitary stamen, it agrees with Lopho- 
phytum and Thonningia, and in the subulate perianth-lobes of the female flower with 
Cynomorium. Upon the whole it may perhaps rank nearest to Thonningia, with which it 
agrees in habit, in some points of structure of the female flower, especially in the tube of 
the perianth being connate with the style above the ovary, in the absence of imperfect 
flowers amongst the females, of bracts, and of a perianth to the male flower. 

The natives of New Zealand call this plant * Flower of Hades,” because it grows almost 
buried in the soil; they further consider it allied to Freycinetia, which it strikingly | 
resembles in the inflorescence. 

Mr. Taylor informs me that he has heard vaguely of the existence of another species 
with blue flowers in the forests of Mount Egmont. 


2. BALANOPHORA HARLANDI, H. f. Dioica, rhizomate parvo lobato epustulato, pedunculo 
ima basi bracteis subfoliaceis involuerato, capitulo globoso, floribus foemineis capitulo 
sessilibus. (Tas. LXXV. B.) 

- Hab. Sylvis insule Hongkong (Harland) ; in convalle Mont. Gough (C. Wilford, Dec. 1858). 


Species pusilla, 1-3 pollicaris, ab omnibus differt pedunculo basi tantum squamato; et preterea a 


B. divica, capitulo globoso, et rhizomate epustulato ; a B. fungosa capitulo unisexuali globoso, floribus 
fæmineis capitulo (non pedicello bracteolæ) sessilibus. 


3. BALANOPHORA Lown, H. f. Dioica, rhizomate parvo papilloso et pustulato, pedun- 
culo brevi, squamis inferioribus parvis, superioribus magnis numerosis imbricatis 
concavis capitulum velantibus, capitulo fl. 4 oblongo, receptaculo columnari, peri- 
anthio 4-fido, lobis lateralibus angustioribus. (Tas. LXXV. C.) 


Hab. Sylvis umbrosis montis Kina Balou, insulæ Borneo (H. Low). 


FROM NEW ZEALAND, ETC. 427 


Rhizoma pallidum, intus laxe cellulosum, in exemplaribus meis parvum, simplex, magnitudine nucis 
Avellane, superficie cellulis hyalinis papillosum, pustulisque conglobatis conspersum. Pedunculus 
solitarius, 2-3 unc. longus, ob squamas superiores amplas obpyriformis, basi volva brevi rhizo- 
matis 3-4-lobi cinctus. Squamæ sicco atro-rubrz v. piceæ, nervis subparallelis, inferiores laxe im- 
bricatæ, 1-1 unc. longæ, oblongæ, obtuse; superiores sensim majores; suprema maxime, 1} unc. 
longæ, valde concavæ, subeymbiformes, vix coriacez, imbricate. Capitulum d pollicare. Flores 
erecto-patentes, pedicellis receptaculo non immersis? cum pedicello sub 3 unc. longi. Perianthii 
lobi antici et postici oblongi, obtusi, laterales lineari-oblongi. Capitulum antheriferum oblate sphæ- 
ricum, apice depressum, loculis polliniferis sub 20-30, parallelis, supra verticem capituli continuis. 
Pollen trilobum, minutissimum. Capitulum 9 valde immaturum globosum, bracteole clavate ; 
floribus stipitibus bracteolarum sitis. 

The exogenous root on which this species grows appears soft, and swells out into de- 
formed, spongy, lobed branches when attacked by the parasite. In allmy three specimens 
the rhizome is simple and small, but probably only small specimens were gathered; 
their surface is covered with a layer of lax, transparent, empty, prominení cells, giving the 
whole a minutely papillose appearance. Prominent pustules or lenticels also occur in the 
rhizome, formed of 4-6 rounded radiating lobes. There is a considerable quantity of 
balanophorine in the cells of the rhizome; not enough, however, to give it a very waxy 
texture. 

The most remarkable character of the species are the very large concave scales of the 
pedunele, which greatly exceed the capitulum, and, imbricating, arch over and entirely 
conceal the flowers. 


Balanophora globosa has also been found on Kina Balou in Borneo by Mr. Low; his 
specimens, which are male, have the flowers almost an inch long and half an inch broad. 
The lateral lobes of the perianth are much narrower than the others, and the anthers are 
many-celled. 


EXPLANATION OF THE PLATE. 


Tap. LX XV. 


A. Dactylanthus Taylori, H. f. 


Fig. 1. g spike, nat. size. Figs.2 and 3. Stamens. Fig. 4. Pollen :—all magnified. Fig. 5. 9 spike, 
nat. size. Figs. 6, 7 and 8. 9 flowers, magnified. 


B. Balanophora Harlandi, H. f. 


Fig. 1. 3,and Fig. 2. 9 plant :—both nat. size. Figs.3 and 4. d flower. Fig.5. Pollen. Fig.6. Young 
9 flowers and bracts :—all magnified. 


- 


C. Balanophora Lowii, H. f. 


Fig. 1. Young 9 plant. Fig. 2. Full-grown d :—both of nat. size. Fig.3. d flower. Fig.4. Pollen. 


Fig. 5. Very young 9 flower and bracts :—all magnified. 
3K 2 


CX 
29 "raha 


PARK 


Irans Linn. Soc. Vol. AX rab. 75 p427 7 


JDE dd J Fitch ith 


Bal:Lowiiz 


Bal Harlandi zz 


[ 489 ] 


DIRECTIONS 
FOR 
PLACING THE PLATES 


OF 


THE TWENTY-SECOND VOLUME. 


Tap. A 
to | Genera and Species of Balanophoree . . . . . . . . . . . . . to face page 68 
16. 
x } Development of the Ovule of Santahim album + ~ v e n u mn Pe 78 
19. Anomalous development of the Raphe in Seeds . . . . . . , . . . . s . . 112 
EE NTN 
JE SETT EE NN as NE 
3$. eR AY RON rasfaren 5 s = GIN GSE PSA EE A A. 5 
23. 
"A | Phozantius heterophylla > s - 2. REN RENN "12 198 
.95. Skepperia convoluta, Mitremyces Ravenelii, and Mesophellia arenaria . . . . . . .132 
20. OANOUUNS CORP ONE T à ee eO +. 4 «.. oo , 149 
27. 
se | Organs found on the Halteres and Wings of Insects . . . . . . + . . . . . 144 
29. 
"} Antennae of le NE MONT VU 154 
30. 2 | 
31. | 
ee RE qi 172 
33. 
34. 191 
ss | Cutaneous Muscles of the Larva of Pygæra bucephala . . 
36. l 
to ; Agamic Reproduction and Morphology of Aphis . 236 
1o. | 
4l. Palb cidis > 4... : 4 233 


42. Metamorphosis of Pedicularia, &c. . 243 


DIRECTIONS FOR PLACING THE PLATES. 


; Tas. 43. Eurybia Gaudichaudi . Rue Wo. EG 


44. Phoronis hippocrepia . . ++ . + + . . + + + . . 

45. l 
Ei Fructification of Compound Sphæriæ . . . . . . . . . 

M 

50. 


| Structure of the Stem in certain Caryophyllee and Plumbaginee 


* 


Henriquezia verticillata, H. obovata and H. nitida 


aa eo 


58. | Fructification of Simple Spherie . . . . 2 . . 2 . . . . 


sd 
E 
z | Ovules, &c. of different species of Gnetum 
E 
T 
Dann 
61. Thea Chinensis . Fa 
62. Tomopteris onisciformis . . . . . 
63. New forms of Entozoa 
64. Australian species of Tunicata . 
65. Compound Tunicata . . . Ke 
66 Janella antipodarum . |. Kcd ee os 
67. Organs of the Antennz of Insects . a, 
68. Embryos of Endogens, Structure of Batideæ, &c. 
69. 
ER species of Nepenthes . . . . .. 
73. J | 
74. Development of the Pitchers of Nepenthes . " Ew. 
25. Daciylontons Taylori, Balanophora Lowii and B. Hei à 


v ” 


END OF THE TWENTY-SECOND VOLUME. 


PRINTED BY TAYLOR AND FRANCIS, RED LION COURT, FLEET STREET. 


to face page 249 
x 4 


2 287 
. 294 
. 298 


. 312