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THE ANNALS
AND
MAGAZINE OF NATURAL HISTORY,
INCLUDING
ZOOLOGY, BOTANY, ann GEOLOGY.
(BEING A CONTINUATION OF TITE ‘ANNALS’ COMBINED WITIT LOUDON AND CHARLESWORTI’S ‘ MAGAZINE OF NATURAL IH{STORY.’)
CONDUCTED BY
ALBERT C, L. G. GUNTHER, M.A., M.D., Ph.D., F.R.S., WILLIAM CARRUTHERS, F.R.S., F.LS., F.G.S.,
AND
WILLIAM FRANCIS, F.1.S.
VOL. XIII.—SEVENTH SERIES.
———eeee - -
LONDON: PRINTED AND PUBLISHED BY TAYLOR AND FRANCIS.
SOLD BY SIMPKIN, MARSHALL, HAMILTON, KENT, AND CO., LD. ; BAILLIERE, PARIS: HODGES, FIGGIS, AND CO., DUBLIN ;: AND ASHER, BERLIN,
1904,
“Omnes res creat sunt divine sapientie et potenti testes, divitiz felicitatis humanz :—ex harum usu donitas Creatoris; ex pulchritudine sapientia Domini ; ex ceconomid in conservatione, proportione, renovatione, potentia majestatis elucet. Earum itaque indagatio ab hominibus sibi relictis semper zestimata ; & yeré eruditis et sapientibus semper exculta; malé doctis et barbaris semper inimica fuit.”—Linna£vs.
“Quel que soit le principe de la vie animale, il ne faut qu’ouvrir les yeux pour voir qu’elle est le chef-d’ceuvre de la Toute-puissance, et le but auquel se rappor- tent toutes ses opérations.”—Brucxner, Théorie du Systéme Animal, Leyden, 1767.
Seis Wun» ene ae He svi van inp wens Obey our summons; from their deepest dells The Dryads come, and throw their garlands wild And odorous branches at our feet; the Nymphs That press with nimble step the mountain-thyme And purple heath-flower come not empty-handed, But scatter round ten thousand forms minute
Of velvet moss or lichen, torn from rock
Or rifted oak or cavern deep: the Naiads too Quit their loved native stream, from whose smooth face They crop the lily, and each sedge and rush
That drinks the rippling tide: the frozen poles, Where peril waits the bold adyenturer’s tread, The burning sands of Borneo and Cayenne,
All, all to us unlock their secret stores
And pay their cheerful tribute.
J. Taytor, Norwich, 1818,
ALERE § YFLAMMAM.
2! Museum.
S65, 42
-_
CONTENTS OF VOL, XIII.
[SEVENTH SERIES. ]
NUMBER LXXIIlI. Page I. The Prototheca of the Madreporaria, with Special Reference to the Genera Calostylis, Li ds., and Moseleya,Quelch. By Henry M. Beensnp, MA; Cantabi, B.S: *(Plate Ty) ee Sie all
II. Some Parasitic Bees. By T. D. A. CocKERELL..........05 33
II. Description of a new Genus of Frogs of the Family Dysco- phide, and List of the Genera and Species of that Family. By G) AShbOvEENGEE, H.i.oo "(Plate FR yo. 2.05 Oe. ee eee oes aig 42
IV. The Collections of William John Burchell, D.C.L., in the Hope Department, Oxford University Museum :—
I. Introduction. By Epwarp B. Povutton, D.Sc., M.A., F.RS., ee CE les BEBO rs wintry he Alas rare ire y aia c Viste 45
II. On a new Stridulating-organ in Scorpions discovered by W. J. Burchell in Brazil in 1828. By R. I. Pocock, F.Z.S.
GE Tn Gee EW ofa te aerators ssineievintas a «.dciine a anateees . 56 V. Notes on Depastrum cyathiforme, Gosse. By E.S. Russet, EA ey Wal tain Sather ee cin a Sania. Wile g's = ode ae ovs.si nish 62
VI. On a new Genus of Spiders from Bounty Island, with Remarks on a Species from New Zealand. By H. R. Hoge, M.A., F.ZS. .. 65
VI. On new Forms of Anomalurus and Sciurus from Tropical Aiea (Ey ELAR Ory SCHWANN Fe nayecins cgiideis ote encedds'coess 70
VIII. On new Species of Lycenide from Sierra Leone. By D. GATOR oye s acs ar esse cereeerteseterseciccnencccstecseetes 73
New Books :—Catalogue of the Collection of Birds’ Eggs in» the British Museum (Natural History). Vol. III. By KuGene W. Oates and Capt. SaviLE G. Rer.—The Geological Structure of Monzoni and Fassa. By Marte M, Oeitvie Gorpon, D.Sc., | 100 eee cr OUR LT MORO ORE rect oee Cof mmr 76,77
A Correction to “Notes on some Meduse from Japan,” by R. Kipkpatwicks FZ.505 5 v.0c. vcedr thes Wea A er Seer ere 80
1V CONTENTS.
NUMBER LXXIV.
Page TX. Notes on Mantide in the Collection of the British Museum (Natural History), South Kensington, with Descriptions of new Species. By Wim. Kiney WL Si BS... 2.0. uos ence ee cere toll
X. The Collections of William John Burchell, D.C.L., in the Hope Department, Oxford University Museum :—
III. Rhipidoeérides et Malacodermes recueilles par W. J. Burchell dans ses voyages en Afrique australe (1810-1815) et au Brésil (1825-1830) ; avec la description de quatre espéces
nouvelles’ Mo ara). <DOURGHOIS aie <> gst hee seen 89
XI. Rhynchotal Notes—XX. By W. L. Disrant............ 108 XII. A Contribution to the Characteristic of Corals of the Group
Rugosa. dy. Prof. N. YaAKovaaer yds aac terpiee ss sess sven ee 114
XIII. On the Distribution of Marine Animals. By Prof. M‘Inrosu, ANISD) |, MMO eS, Whores ERE SEAR UR ecco lok Le LOS bs eee ke 17
XIV. Descriptions of new Frogs and Snakes from Yunnan. By Sorel ESOP RINGO ME TLS In sorte AG ac) a wae Gus SEE Caen Peouea een 130
XV. On some Fishes from the Lakes of the Cameroon Mountain. By Dr tawan Lownpaec, CMU.) de. 6). 9 8. 0. RR 185
XVI. Descriptions of new Species of Lycenide from Borneo and New Guinea. By Hamitton H. Drucs, F.Z.S., F.E.S. ........ 140
XVII. Two new Mammals from South America, By OLpFIELD BIO MIGAS 2ohac, Joes cols mo ie eft tahs cede laa efayayaheate Mepe cies ees 142
XVIII. On the Classification of the Crustacea Malacostraca. By Ke CMAN, DISC: 5 cee eens ss cena s Se cla oot e teem nee eee 144
New Books :—Memoirs of the Geological Survey of the United Kingdom. The Cretaceous Rocks of Britam. Vol. II. The Lower and Middle Chalk of England. By A. J. Juxxs- Browne, B.A., F.G.8. With Contributions by Witi1aMm Hitt, F.G.8.—A Treatise on Zoology. Edited by E. Ray LankesTerR, M.A., LL.D., F.R.S., &c. Part I. Introduction Gnd PrOpezod, Ge sic sjcwioe 4 y «cewek Nee «Wj ee cutee Bs 158, 159
NUMBER LXXV,
XIX. A Synopsis of the Suborders and Families of Teleostean Fishes, Wy G, A. BouLENGEn, MSs, 2... .5 162s ato ke ween 161
XX. On a Collection of Fishes made by Mr. John Graham at Yunnan Fu. By C. Tarr Ruean, B.A.
XXI. Rhynchotal Notes —XXI, By W. L. Disranr
XXII. New Bats from British East Africa collected by Mrs. Hinde, and from the Cameroons by Mr. G. L. Bates. By OLDFIELD Tuomas. 206
CONTENTS. Vv
Page XXIII. Descriptions of new Species of Aculeate and Parasitic Hymenoptera from Northern India. By P. CamERon 211
oree eee eee
XXIV. Preliminary Note on certain Points in the Anatomy of Eryx and other Borde, partly indicative of their Basal Position among the Ophidia. By Frank KE. Bepparp, M.A., F.RS. .... 238
XXYV. Description of a new Genus of Spatangoids. By F. Aborgoriped aril & gh: Of (paws: Reis: aera Tors 3 eee ek See a
XXVI. Description of a new Barbus from Cameroon. By G. A. Boupiaasany WS Ws 29 nou tire ae cect. Peed ele clon alas Blok orale 237
XXVII. Notes on the Structure of the Teeth of some Poisonous Snakes found in Travancore. By R. SaunKARA NARAYANA PILLAY. 238
Onituary Nobies s Wr WIELEAne BRAN CIS, shen ss, 5 caretip crates: om = os 239
NUMBER LXXVI,
XXVIII. Descriptions of some new Species of Lepidoptera Hete- rocera from Tropical South America, By HERBERT Druce, ee On Ceee osha ag hal oft Ppetmaehs a) mor diated cr viay er seo Ciacehas Pell ang ey ola.) ¥ 0s 241
XXIX. New Forms of Saimiri, Saccopteryx, Balantiopteryx, and Thrichomys from the Neotropical Region. By OLprirtp THomas. 250
XXX. Descriptions of new or little-known Fishes from Mexico and British Honduras. By C. Tats Rea@an, B.A. .............. 255
XXXI. Descriptions of Holocentrum osculum, Poey, and of a new Fish of the Genus Centropomus. By C. Tare REGAN, B.A. ...... 259
XXXII. Descriptions of Two new Genera of Frogs of the Family Ranide trom Cameroon. By G. A. BouLENGER, F.R.S. ........ 261
XXXII. Rhynchotal Notes—-XXII. By W.L. Distant ....
XXXIV. Description of a new Fish of the Genus Chetodon from the New Hebrides. By C. Tats Reean, B.A.
efia! pues ja ie ¥l4 (se ov, ple
XXXYV. On some new Species of Hymenoptera from Northern Beidteia, Es yg re 3 WMEGRON onc ta aon sah inode nin Ao sig entddepataales coals 277
XXXVI. An undescribed Genus of Coretde from Borneo. By eye PS ARNTIT ot cnc ee itaet eet Aaa eke ea, RAT Ae Cae f 303
XXXVII. The Collections of William John Burchell, D.C.L., in the Hope Department, Oxford University Museum :— IV. On the Lepidoptera Rhopalocera collected by W. J, Bur- chell in Brazil, 1825-1880. By Cora B. Sanprrs, of Lady Margaret Hall, Oxford. (Plate VI.)............ 305
XXXVI. Note on an undescribed Weasel from the Atlas Mountains, and on the Occurrence of a Weasel in the Azores. By Gis Hed Baran ERM ROY 505 a waves ajers at m,s) ein heiesivnich a olaes wide SR 323
New Bovk :—Report on the Sea Fisheries and Fishing Industries of the Thames Estuary. Prepared by Dr. JAMES MvRIE .... 3265
Proceedings of the Geological Society .......... : 326—828
@ er ee orate (0.8
vi CONTENTS.
NUMBER LXXVII.
Page
XXXIX. The Phylogeny of the Teleostomi. By C. Tarr Reean, BA. (Plate VIED” testes Abony ee ae santo icles OW Shae 329 XL. Rhynchotal Notes—XXIII. By W. L. Distanr ........ 349
XLJ. The Collections of William John Burchell, D.C.L., in the Hope Department, Oxford University Museum :— IV. On the Lepidoptera Rhopalocera collected by W. J. Bur- chell in Brazil, 1825-1850. By Cora B. SanpErs, of (lady Marcoretstlall “Oxtord:: 0, ss petits nee uia oicatens 506
XLII. Notes on Phasmide in the Collection of the British Museum (Natural History), South Kensington, with Descriptions of new Species —No.d. Sy WW. Karey, WS. BORIS... 2)-.e sae eee 372
XLII. On the Genus Ortmannia, Rathb., and the Mutations of Pertam wAbyids;: “yeh, Ms, SsOUVAER. @ 0)... S dis «<bean gieie be Patter @ ea
XLIV. Notes on a new Species of Acts. By W. D. HeNpDERSON, M.A., B.Sc., Zoological Laboratory, the University, Aberdeen .... 381
XLY. A new Bat from the United States, representing the Euro- pean Myotis (Leuconoe) Daubentonit. By OLprintp THomas .... 382
XLVI. Three new Bats, African and Asiatic. By OLpFIELD WISETO NUAS! © sais seve ales EEG da TR rR tat one ag Ee ree tats tele Mee 384
XLVII. Notes and Descriptions of some new Species and Sub- species of Mustelide. By G. EK. H. BAnrrrt-HaMILTon ........ 388
New Books :—Mostly Mammals. By R, LypEKKER.—Catalogue of the Lepidoptera Phalzenze in the British Museum. Volume LY. Catalogue of the Noctuide in the Collection of the British Museum. By Sir Grorex F. Hampson, Bart—The Fauna of British India, including Ceylon and Burma. Published under the authority of the Secretary of State for India in Council. Edited by W. T. Biranrorp. Rhynchota: Vol. II. Part 1 (Heteroptera), By W. L. Distanr.—Memoirs of the Geo- logical Survey. Paleeontologia Indica, Series IX. The Jurassic Fauna of Cutch. Vol. Ill. Part 2. The Lamelli- branchiata. No. I. Genus Trigonia, By F. L. Kircutn, M.A., Ph.D., Geol. Survey England.—Cireulars on Agricultural Economic Entomology. Issued by the ‘Trustees, Indian MSSM Geen k ys ke cei cbse a bmke ob pep etacts 395—399
Proceedings of the Geological Society ....ceeeessesseuvees 399, 400
The Action of Human Serum on certain Pathogenic Trypanosomes, Action of Arsenious Acid upon Trypanosoma gambiense, by A. Laveran; Relations between the Development of the Tracheal Apparatus and the Metamorphoses of Insects, by Piles AMZ1AS’ .\, isles itwye oc0ee eiio e sopee whe ees pre o pee 40R——40g
CONTENTS. Vil
NUMBER LXXVIII.
Page XLVIII. On Mammals from Northern Angola collected by Deo W. J, Ansorgen By Onprtern THOMAS ..cu. esc ce se cases 405 XLIX. On Felis ocreata, better known as Felis caligata, and its Subspecies) by HAROLD SCHWANN) fo 000. cee- 5-0 reese sna 421 L. On certain African Butterflies of the Subfamily Pierine. By riven Ga Durem ADs PVE Se. SCs... .en gece ds mane «ane 426
LI. Notes on Phasmide in the Collection of the British Museum (Natural History), South Kensington, with Descriptions of new Genera and Species.—No. II. By W. F. Krrsy, F.LS., F.E.S. ., 429
LII. Diagnoses of Three new Species of Barbus from Lake Wicora,- by Gy A. BOUmnNGWR, HOR Sts vane Magi ones ceil aes . 449
LIII. Descriptions of Three new Snakes. By G. A. BoULENGER, eR am oa iste alg eter a oles oe RNS RES AeA MND ie cots af do ktaps 450
LIV. Descriptions of some new Species and Varieties of Cataulus from the Collection of the late Hugh Nevill, Esq. By Huau FULTON
LV. Natural History Notes from H.M. Indian Marine Survey Steamer ‘Investigator,’ Commander T. H. Heming, R.N.—Series III., No. 1. On Mollusca from the Bay of Bengal and the Arabian Sea. By PGA Ap OMrney TOs wees wake oa ig cig aedentweee ns 453
New Books:—Memoirs of the Geological Survey of the United Kingdom. The Cretaceous Rocks of Britain. Vol. III. The Upper Chalk of England. By A. J. Juxkxes-Brownr, With Contributions by WintaM Hit, F.G.S.—Pictures of Bird- Life. By R. B. “LopGE
Teleostome Phylogeny; a Correction ........cccsscscecenens Br {i>
Index 6 6 0 0 88 b 8 6. 6 Oe © 60) 86 6 B60 6 6 6 «e606 0 0 0) O 6 60) 6) 00 0) 986 0 6 Bee e 2 6.6 476
PALE le II.
III.
lV.
WV:
VI
VII.
PLATES IN VOL. XIIL
Prototheca of the Madreporaria. Colpoglossus Brooksii.
Map: W. J. Burchell’s Travels in Brazil. Stridulating-organ in Scorpions. Depastrum cyathiforme.
Lepidoptera Rhopalocera from Brazil. Phylogeny of the Teleostomi,
THE ANNALS
AND
MAGAZINE OF NATURAL HISTORY.
[SEVENTH SERIES.]
A oii cetedoccuce er litora spargite muscum, Naiades, et circiim yitreos considite fontes: Pollice virgineo teneros hic carpite flores: Floribus et pictum, dive, replete canistrum. At vos, o Nymphe Craterides, ite sub undas ; Ite, recurvato variata corallia trunco Vellite muscosis e rupibus, ef mihi conchas Ferte, Dez pelagi, et pingui conchylia suceo.” N. Parthenii Giannettusi, Bel, 1,
No. 73. JANUARY 1904.
I.— The Prototheca of the Madreporaria, with Special Reference to the Geneva Calostylis, Linds., and Moseleya, Quelch. By Henry M. Bernarp, M.A. Cantab., F.L.S.
[Plate I.]
THE task I have set myself is to sketch what appears to have been the leading features in the evolution of the Madrepo- rarian skeleton. ‘The researches on which the arguments are based have been almost entirely limited to the skeleton, not because the importance of a close study of the soft parts is not recognized, but because, for the attainment of accurate results, the widest possible survey of homologous structures is indispensable. ‘This condition can never be supplied by the soft parts. ‘They can at the most be studied in a few recent specimens, whereas the vast majority of the forms presented by the Madreporarian system are fossil. Further, let me add in passing that I do not believe that the study of the individual development of a few living forms can by itself establish anything certain about the past history of the group, for the simple reason that we cannot tell whether any special
Ann. & Mag, N. Hist. Ser. 7. Vol. xiii. 1
2 Mr. H. M. Bernard on the
developmental feature is a repetition of some ancient con- dition or a recent adaptation*. As I have already often maintained, lines of phylogenetic growth can only be satis- factorily established by the discovery of connected series of variations, morphologically and chronologically arranged. The skeleton alone can supply us with such series, and that of the corals probably with a more complete series of forms, extending from the Paleozoic era to the present day, than will ever be obtained of any other animal group. Whether, there- fore, the skeleton be of great or of little importance in itself in the morphology of the corals, it alone supplies us with what we want—a continuous series of homologous structures. On this account alone, then, when our aim is taken into account, we are obliged to confine our attention to the skeleton. As a matter of fact, the skeleton is of paramount importance in the coral organism. There is a sameness in all the soft parts which limits their morphological importance in any comparative study. Their chief variations may, for practical purposes, be said to be repetitions of the variations of the skeleton which they secrete. ‘The skeleton is, par excellence, the chief structural feature of the coral, its relation to the soft parts being extremely simple. It is, as we now know, thanks to the researches of von Koch, Heider, Fowler, Bourne, Ortmann, and Miss Ogilvie, an excretion of the basal parts of the outer wall of the body, and hence morphologically it is external to the organism. At times very complicated, it is an organ of protection and support for the body of the polyp, or, in colony formation, for the colonies of polyps, the polyps themselves, thus protected, having, as arule, remained simple and primitive. ‘The corals, indeed, present us with a group of organisms still primitive enough to illustrate the fact that, of the earliest morphological modifications of the living matter, skeletal formations were the most pronounced. This is strikingly exemplified by the Foraminifera and Radiolaria, in which there is a wealth of skeletal formations with little or no visible variation of the soft matter. Again, in the sponges the skeletal variations far outrun those of the soft parts. ‘The same is true of the stony corals. In what follows, therefore, I shall make no detailed refer- ence to the soft parts or to the excellent work which is being done with their help by Dr. Duerden towards the elucidation
* N. Guldberg and Nansen, “On the Development and Structure of the Whale,” Bergens Museum, 1894, p. 89; also Sedgwick, Proc. Fourth International Congress of Zoology, 1898, p. 74.
Prototheca of the Madreporaria. 3
of the same problem as that which here interests us. I shall confine myself solely to showing how some of the chief transformations of the skeleton can be linked into series and how, in a few cases, the causes which led to those transforma- tions are apparent. We are justified in hoping that the conclusions obtained from the continued studies of the soft parts on the one hand and of the skeleton on the other will ultimately coincide.
I wish to make it specially clear that only a few of the lines of modification can be dealt with, but those few, being some of the earliest, are, I believe, the most fundamental and important for the elucidation of all the later transformations of the coral skeleton. To deal with the whole of these latter would be to write a complete systematic account of the stony corals. This is the aim of the great catalogue now being prepared and published by order of the Trustees of the British Museum, and must be a work of years *.
The researches of the writer in reference to this work so far hardly entitle him to speak with confidence on any other of the larger divisions than the Perforata; no other has as yet been systematically dealt with by him, at least in the thorough manner required for a British Museum Catalogue. It would not, however, have been possible to discover the morphology of these highly specialized Perforate forms without a study of and constant reference backwards to earlier and simpler types. In this way certain lines along which the stony corals have travelled, viz., those leading from the most primitive to the most specialized, have been growing clearer.
* The last attempt to deal with the whole of the coral system in the ‘ Hist. Nat. des Coralliaires’ of Milne-Kdwards and Haime, completed in 1860 by Milne-Edwards alone, was founded on comparatively small collections and written at a time when the relations between the skeleton and the polyps were not understood. The excellence of the results which were nevertheless obtained is, on the one hand, a tribute to the genius of the great French naturalists, and, on the other, a witness to the compa- rative unimportance of the polyp, morphologically, as compared with the skeleton. aa
The new catalogue projected by the authorities of the British Museum, and rendered necessary by the immense increase in the collections due especially to the sending out of scientific expeditions, was started in 1876, but was interrupted by the death of Dr, Briiggemann, who was engaged for the purpose. After fourteen years Mr. George Brook undertook the work, but again death intervened soon after the first volume was published in 1893. Two years were again lost, when the present writer was appointed to continue the work. There are now four volumes published, and the fifth is rapidly approaching completion. Each volume is practi- cally a monograph of one, or at the most two, genera, and, like the earlier attempt of Milne-Edwards and Haime, it now describes the fossil as well as the recent forms.
4 Mr. H. M. Bernard on the
The following pages sum up the principal conclusions he has arrived at.
The most important stage to establish in an evolutionary history is the first, or that which we may consider as the first, inasmuch as from it all the modifications we wish to compare can be deduced. ‘The first stage in the evolution of the coral skeleton was first dimly recognized by me in the minute saucer-shaped cups of young Madreporidan colonies— so young as to consist only of a parent calicle and one or two daughters. In none of the Madreporids have I yet found the earliest stage in which the cup containing the parent alone was cup-shaped. Such astage, however, may be legitimately assumed.
The discovery of such colonies made three points clear to me :—
1. The parent calicle of a colony rises out of a basal cap— the PROTOTHECA *.
2. This prototheca is not a composite structure, but a morphological unit, the rim of which can be bent up, flattened completely down, and indefinitely expanded in any direction as a film, from the upper surface of which, as originally from within the cup, the coral skeleton arises.
3. This film is the EPITHEGA fF.
These conclusions received complete confirmation from a study of the Paleozoic form, Kavosites and of its modern descendant Alveopora. I have already described and figured the prototheca of the latter genus}. Its rim, as shown in the figures referred to, does not usually flatten down, but grows upwards and outwards to form the irregular film-like invest- ments characteristic of the colonies of this genus.
In both cases—that is, in Madreporide and Favositide alike—it was easy to see the bars of the intracalicular skeleton rising directly out of the wall of the cup as internal projections from its surface; this point is of importance, because von Koch, whose developmental researches also revealed to him the prototheca, was led by what he saw to regard it as a composite structure consisting of a basal portion (‘‘ Basal-
* Lindstrom suggested the word “ initium” for the earliest cup-like skeleton ; the term “‘ prototheca” was suggested to me in conversation by my friend Prof. Jetfrey Bell.
t The fact that the skeletal elements rise from the surface of the pee ‘was pointed out by Martin Duncan in 1884 (Journ. Linn. Soc.,
ool. xvii. p. 861) as indicating the importance of that element of the coral skeleton.
{ Journ. Linn. Soe., Zool. xxyi. 1898, p. 495, pl. xxxiii.
Prototheca of the Madreporaria. 5
platte,” sole) and of a peripheral portion (epitheca). This appears, however, now to have been a too literal rendering of the facts of his observations, for no one who had seen several of these epithecal saucers of different sizes and with edges turned up to different heights at different curves, and the skeletal bars springing indifferently from the sides and the base, could possibly divide it into a basal and a peripheral portion.
Besides, in a young saucer-shaped colony it is obvious that the turned-down side (the ‘ epitheca”’) of the parent becomes the “‘basal plate” of the daughter, and in this successive flattening down of the rim we can see the explanation of the characteristic wrinkled appearance of the supporting epitheca of so many horizontally expanding corals, whether single or compound. Each furrow represents a pause in the outward growth long enough to allow the rim of the widening saucer- shaped epitheca to grow upwards a short distance. The next period of growth carries it downwards and outwards again. ‘This process has been actually seen by Lacaze- Duthiers * in the development of Balanophyllia regia. This writer observed three attempts of the basal secretion of the Jarva to turn up to form a cup or “ envelope calicinale,” but they were always futile; the septa overran them and the edge was flattened down again and continued as a_ basal secretion icjo Fl. . fre, 10):
Before continuing with the history of this prototheea—that is, with our account of some of its earliest modifications—it will strengthen our argument to mention a few instances in which earlier writers have come near to recognizing this identity of the prototheca with the epitheca. As we might expect, such an identification would be more probable in relation to Paleozoic forms, in which the primitive cup remained longest in evidence and had not become so distorted and masked as it is in the majority of the modern forms. Milne-Edwards f, in describing the Paleozoic genus Za- phrentis, which, from its appearance in time, might have been expected to have retained the prototheca, says that it is completely surrounded by an epitheca. Nicholson could not distinguish the epitheca of these same corals from the wall. Miss Ogilvie ${ declared that in Zaphrentis the epitheca “supplied the primitive base and periphery in one,” and again that the primitive wall of corals was epithecate ; and
* Arch. Zool. expérimentale, (3) vol. v. 1897, pp. 179-183 & 280, pl. x. figs. 19-24. ; tt
+ ‘Les Coralliaires,’ iii. p. 385 (1860).
{ Phil. Trans. 1896, p. 820 &e.
6 Mr. H. M. Bernard on the
again the same writer recognized the wall of Zaphrentis as “euthecate,” which means that the persistent prototheca in these early corals is the eutheca or true primitive wall of Heider and Ortmann, as compared with which all other thecze are secondary. ‘To this last opinion we shall return.
Mention should also surely be made of Ludwig *, who, so long ago as 1866, attempted to found a classification upon his recognition of the prototheca as the primitive shell (“ Gehiiuse ”’) of the coral polyp. But beyond the interest attaching to the fact that he thus emphasized the importance of the prototheca in Madreporarian morphology his work has no value, for he was led astray in his further analysis by a fancied analogy with the shell of the mollusk.
In the present paper, then, we start again from the recog- nition of the prototheca, but this time, avoiding Ludwig’s mistake, we shall try to analyze some of the actual modifica- tions which this primitive coral skeleton has undergone in the progress of its evolution. So far from being as simple as Ludwig appears to have assumed it to be, it isa task of considerable complexity to follow and of no small difficulty to describe. This paper, indeed, was begun five years ago, and has been frequently rewritten.
As I have shown, those parts of the coral skeleton called epitheca must for the future be referred to the rim of the prototheca. This seems simple and clear now, but in the past the epitheca has been the stumbling-block of coral morphology. It has been this for the very reason that it waited for the discovery of the prototheca betore there was any possibility of its elucidation, The fact of the confusion in the prevailing views as to what the epitheca is is familiar to every coral student. For instance, Prof. Gregory, of Melbourne, after all his years of work at corals, characteristically summed up his despair of ever making anything out of it by declaring that “there was no part of the coral skeleton over which more time had been wasted” ft. This attitude and that which is taken in this paper are poles asunder. Between these two, authors and text-books hover. None are so bold as Prof. Gregory, yet none have succeeded in formulating an intelligible doctrine.
We may here state that there is ample excuse for this confusion, for even now that we know that the epitheca, as it occurs in the majority of specimens, is only an extension of the rim of the original cup, still in each case the problem as
* ‘Paleeontographica,’ vol. xiv. } Paleontol. Indica, ser. 1x. vol. ii. p. 11 (1900).
Prototheca of the Madrepora rid. 7
to how this can be requires unravelling. It may, for in- stance, be the rim extended indefinitely and continuously as a chalky film round a colony (e. g. Alveop ora), or, again, it may be discontinuous and represent the separate rims of an aggregation of corals, each with its own cup, as in so many Paleozoic forms. In this case it depends upon the way in which the corals are aggregated whether the rims are easy or difficult to recognize. Add to these difficulties the fact that apparently any part of the surface of a polyp may dis down and secrete a calcareous film* which is purely adventitious and has no morphological significance, and it is obvious that until we had a key to its elucidation the epitheca could not fail to be a source of bewilderment.
Diagram 1 (PI, I.) shows the three earliest growth periods of a primitive Madreporarian skeleton. All that we see is a deep cup with three tabular floors. ‘The process is explained in diagram 2, in which we see three cups progressively modifying their shapes. The lowest of these is the prototheca in the strict sense of the word}, bat it is advisable to apply the term to all simple repetitions with free edges. Fig. 2 is so far diagrammatic, inasmuch as with cups of this shape it is impossible to say how far the rim of each cup extended before the soft parts of the base of the polyp became detached from the base of its prototheca, Cases, however, do occur in which the change in the shape of the new thece was rapid, and for this and also for other reasons the rims of the separate
* The formation of calcareous films somewhat irregularly over the skin of corals is hardly to be wondered at. The prototheca was but the primitive secretion of the basal portion of the polyp, forming a protective cup into which the animal could retract the oral and exposed end of its body. Above the rim of this cup calcareous secretions were not usual, otherwise they would have interfered with the process of retraction, but the power of secreting them was not lost. Indeed, some forms actually secreted lids, which, when the polyps retracted, closed down over the prototheca (Calceola, Goniophyllum). A histological difference between these secondary films and true epitheca may sometimes be noticed. The former may be built up of separate plates, each of which starts round some point of the skeleton and grows by concentric increments.
+ The prototheca is here drawn quite diagrammatically. Figure 8, after Lacaze-Duthiers, is one of the best figures from life. My own figures, already referred to, of a young <Alveopora are of a prototheca somewhat distorted. Theoretically we might expect a slight constriction above the flattened sac, for as the soft larva settled down we might expect its aboral end to flatten out somewhat wider than the neck carrying the oral disk and tentacles. The base of the second prototheca might easily be rounded or pointed, for it would hang down in the hollow of the prototheca proper, The later development of conve tabule and vesicular dissepiments may have been due to the pulls of mesenterial muscles,
8 Mr. H. M. Bernard on the
cups may be distinguishable. For instance, the development of exsert laminate septa may lift the cups above one another (see Pl. I. figs. 3, 11, 12).
Fig. 3 refers to Monélivaliia, of especial interest because it was the irregular bands of epitheca round specimens of this genus which induced Dr. Gregory to give up this element of the coral skeleton in despair. We shall now show that an understanding of these bands is essential to a true insight into the morphology of the skeleton.
It is frequently stated* that in Montlivaltia there is epitheca, but no theca. There was, however, certainly a prototheca, and examination of the coral shows that the successive protothece gradually flattened out until, after reaching a certain size, they formed a series of flat saucers (tig. 3, e, e, e...) of nearly uniform size, and piled up one above the other as tabule with edges which may either only just reach the surface or be bent sharply upwards to varying heights according to the accidents of secretion. On the left of the figure a few of the septa are shown supporting and raising the successive saucers above one another. ‘The septa of each polyp continue those of that which went before it, so that these radial structures naturally run up continuously through the whole skeleton. On the left of the diagram the saucers alone are shown in optical section as a series of flat or wavy floors with turned-up rims.
Here, then, we have the three facts necessary for the understanding of the case in hand :—
1. A series of shallow thecz or protothecal saucers ending abruptly at the surface or with edges bent up externally.
2. The septa which, being exsert, support and lift these saucers above one another, so that, while the septa are con- tinuous, the rims of the cups may be free and separate, or, when bent up, may run together as irregular epithecal bands.
3. ‘Lhe extreme irregularity of the bands is due to the want of uniformity in the height to which the secretion of the rims of the saucers, if bent up, extends.
These three factors fully explain the puzzle presented by the epitheca of Montlivaltia.
It is obvious that in diagram fig. 3 the saucers might contain not single polyps, but gradually expanding colonies
_* £. g., by Miss Ogilvie (J. e. p. 158), who, however, followed Milne-
Edwards and Haime, who wrote with reference to A mplecus, in which the succession of saucer-shaped protothece is very pronounced :— “ Quelquefois méme la muraille puiait manquer et ‘le polypier n'est constitué que par une série de cornets trés évasés et naissant les uns au-dessus des autres” (Ann. Sci. nat. (3°) ix. p. 84, 1848).
Prototheca of the Madreporaria. 9
(cf. the minute colonies of Madreporids already mentioned), Such series of gradually expanding colonies might grow into columnar or massive stocks widening at the top. In all such stocks the tabule which run through them must be regarded as the floors of successive saucers. This is well exemplified in the genus Goniopora, as I have already explained *. In this genus too we have, as we have in Montlivaltia, irregular bands of epitheca running round the stocks. These are the rims of the protothecal saucers showing irregularly at the surface. In Alveopora the rims all run together to form continuous epithecal investments, except, perhaps, in their branching forms, in which the protothecee may be lifted up above one another by the growth of the spiny septal skeleton.
For an understanding of the morphology of the coral skeleton we must bear in mind that essentially the same process, viz. a succession of epithecal cups or saucers, occurs throughout the whole of the Madreporaria. They may be simple conical cups fitted one into the other (Zaphrentis) or flat plates piled up (Montlivaltia, Goniopora), or their epithecal floors may be thrown into complicated folds and both the cup and its repetitions may be difficult to unravel, but the fundamental principle is the same throughout. There is only one group I| can think of in which the epitheca is not nor- mally repeated, namely in the highest Madreporids—Madre- pora, Turbinaria, Montipora, Astreopora, and their simpler ancestors the Hupsammiids. In these the purely septal skeleton rises rapidly above the original flattened prototheca which is then left behind. This is the reason of that well- known characteristic of these forms that the calicle cavities run continuously for long distances through the skeleton.
We repeat, then, for the sake of emphasis that wherever the epitheca occurs it represents the rim or the coalesced rims of one or more protothecal cups or saucers, the floors of which are represented by the tabulee. In any individual case the tabula below the living layer is the zth repetition of the original prototheca of the parent polyp.
‘The main problem, then, of the student of coral morphology, that is, taking the skeleton alone into account, is to trace the various moditications of the prototheca from its earliest simple cup stage to the many different shapes and positions it now assumes and occupies as part of the coral skeleton.
Roughly speaking, we may say that there are two periods in the evolution of the Madreporaria—that in which the prototheca, though modified, remains in evidence, and that in
* Cf. vol. iy. Brit. Mus. Madreporaria, p. 24, diagram A,
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which it has disappeared from view or is difficult to unravel. Only in the few Madreporids (the chief families of the Perforata) above mentioned can it be said to have been aborted, and then only in a limited sense, for the whole coral skeleton is its product. If the original rim of the cup is replaced as the edge of the theca by new thece formed either by the rising up of concentric folds from its floor or of radial plates from its sides, or by complicated combinations of these two, these new thece are strictly infoldings of the prototheca. The prototheca, then, however obscured its early cup shape, being replaced by secondary cups produced by its own infoldings, remains throughout the fundamental element in the Madreporarian skeleton.
I propose here to trace some of the more obvious trans- formations of the prototheca, treating them entirely morpho- logically—that is, simply as forms which admit of explanation and deduction from simpler forms, and without regard to their real phylogenetic sequences.
The working out of these latter—that is, the attempt to discover the real places of these transformatory processes in the genealogy of the Madreporaria—must be a work of time. I am convinced, however, that it will at once give a new and much needed interest to the student of the stony corals.
We return, then, to our simplest form (diagram fig. 1). It shows us a conical cup standing on a flattened slightly ex- panded base and gradually thickening upwards. The problem of increasing instability must obviously have been one of the first which the polyp inhabiting such a skeleton had to solve. I shall endeavour to show that the earliest divisions of the Madreporaria were due to the different ways in which this problem was solved.
I. Falling over and recovery of the upright position.—The simplest of all methods was to fall over so that the flesh of the polyp could come once more into contact with the sub- stratum and secrete a new cementing layer where it touched. From this new base the polyp could bend upwards once more securely attached. The following is some of the evidence which shows that this actually took place :—
(a) The earliest period is specially characterized by the great number of single corals which are conical but curved. ‘he curve is exactly what is required ; that is, it is most pro- nounced at the tip, e. g. Zuphrentis, Menophyllum, &c.
(b) All these curved corals have what is known asa fossula, that is a deep depression within the calicle and most frequently on the convex or what is called the “dorsal” side. The fossula has a very simple explanation, if the assumption of
Prototheca of the Madreporaria. 11
the falling over is correct (see diagram fig. 4). As the soft parts detach themselves from the base of the prototheca they might be expected to bag down, and they will continue to be acted upon by gravitation and drawn over towards the convex side of the coral until the vertical position has been regained. It is possible that this bearing over to the side may be due to the efforts of the polyp itself to bend up, but gravitation is a causa efficiens.
In some forms, however, the fossula is not on the dorsal, but on the ventral side. ‘There is abundance of scope for variations of all kinds: a deep cup (that is, the cup of a polyp which grew very slowly in width, for instance) would lie very prone and its fossula would fall over to the dorsal side (diagram fig. 4); but a shallower more open proto- theca (that is, one in which the polyp grew very rapidly in width) would, in the prone position, have one (the “ventral””) wall nearer the vertical, and this would keep the skin of the point while it hung loose for the while near the ventral side, and the fossula would consequently also appear on this side (diagram fig. 5) *.
(c) The falling over of the prototheca will explain the departure from a ‘strictly radial symmetry of the septa seen in these curved Paleozoic corals. It is obvious that, as the coral is bending to the vertical, seen from above, the septa would have the arrangement shown in diagram fig. 6, which is after the classical figure of Kunth showing the septal formula typical of the group called Rugosa. ‘The position of the fossula with relation to this modification of the septal arrange- ment shows that this is the true explanation. Further, it has long been known that, as such corals gradually reacquire a vertical position, the septal arrangement slowly gives up the bilateral and returns to the radial symmetry. Thus the character on which it was proposed to found a great division of the stony corals was nothing but a slight mechanical
* This is not the first time that this origin of the fossula as a repetition of the tip of the prototheca has been recognized. Ludwig’s figures made it quite clear in 1866 (‘‘Corallen aus “paliolithischen °Formationen,” Palontographica, xiv. 1866). But, regarding the skeletons as analogous to the shells of mollusks, to whose shapes he thought they were adapted, he failed entirely to understand the true character of the coral skeleton or
of the causes of its changes.
The use assigned in text-books to the fossula, viz. as a sort of erypt for the sexual products, is probable enough, but need not have been the cause of its origin. I fail to see the evidences for the existence of more than one true fossula in any coral I have examined. Superficial irregularities in the septa, due perhaps to the presence of sexual products, may y be quite distinct from the true fossula, A longitudinal section or a fracture showing a complete tabula is the only ey idence which can be relied on,
12 Mr. H. M. Bernard on the
adaptation to a passing phase in the life of each individual coral. But it is only fair to say that the whole tendency of recent works on corals has been to discover the invalidity of the supposed division Tetracorallia.
Into the interesting questions which this suggests as to the value of the existing divisions of the corals, we cannot here enter, but content ourselves with merely pointing out that while probably all the very earliest corals fell over and, if they bent up again, became Tetracorallia during the process, it is possible that many, which later had learnt a different method of acquiring stability, might easily be knocked over and in their efforts to become vertical again might become Tetracorallia by accident.
(dq) The falling over of the prototheca enables us to find an origin for several groups which are usually regarded as corals, but whose position is still a matter of uncertainty. It is quite within the limits of probability that a certain number of these overturned polyps in their small protothece should remain prone and bud in this position. One such case we know of for certain (see p. 28, on Heliolites). We ask whether the creeping branching stocks of Aulopora might not also have been formed by the early budding of a parent whose proto- theca had fallen over.
From Aulopora the genus Syringcpora might be deduced. Syringopora is said to begin with the same horizontal creeping stock as Aulopora, and then to bend up and form its tufts of wavy tubes freely communicating with and supporting one another. In these erect tubes very irregular tabule are formed by the constant rising of the polyp in the tube as the latter lengthens. The very presence of tabule and of the rudi- mentary septa, consisting of rows of points, clearly indicates an affinity with early Madreporaria. Add to these the proto- thecal outer covering, and we have the same three structures which make up an Alveopora or a Favosites. It is only their dispositions and the relative developments of the parts which differ *,
Halysites could also be deduced from such a prone theca by ra}id continuous budding, in such a way that the parent and its buds bent up in rapid succession into the vertical, as shown in the diagram fig. 7, each then continuing to grow as a thin flattened tube. ‘These in contact and mutually sup- porting one another would supply the typical skeleton of this
* The apparent affinity between Syringopora and Favosites has been pointed out by Mr. Bourne (Phil. Trans. vol. 186 B, p. 474). But Favosites is structurally indistinguishable from Alveopora, and was not therefore an Aleyonarian (Proc. Linn. Soc., Zool. vol. xxvi. (1898) p. 495.
Prototheca of the Madreporaria. 13
remarkable genus. We have the same three elements, proto- thecal tubes, tabule, and spiny septa *.
(e) The habit of falling over is known still to occur in the genus Flabellum.
(7) Lastly, I appeal to the modifications of the prototheca which will be described in the following pages, every one of which may be regarded as an adaptation for the purpose of solving the problem of vertical stability, that is, how to avoid the natural consequence of having to stand on a point while continuing to grow in height and bulk. For we are surely justified in assuming that the falling over at the very outset of life of an organism intended, if we may say so, to stand upright, would mean considerable loss of time and energy during the reattainment of the upright position. Such a loss might be expected to delay budding, and it is probable that we may have to take this into account in our ultimate classi- fication. We may have to form a group which arose from the early budding of parents still in their protothece (sens. str.), and this would include such forms as Aulopora, Syringopora, and Halysites, in all of which the protothece fell over, and to these we might add Chetetes arising probably by fission. Whether the prototheca also fell over in this last case I have not ascertained. Such a group arising from parents still in their protothecz proper, would stand in contrast to another group in which the budding was delayed until the polyp had grown considerably larger and had again assumed the upright position, and our divisions of these latter would have, in the first instance, to be based upon the methods adopted to attain this end.
I]. Radicle-formation—This process has been carefully studied and described in F’labellum by Lacaze-Duthiers f. A small portion of the lip of a prototheca bends over until it adheres to the ground (see diagram fig. 8a,). I have myself seen a similar process as an occasional thing in young colonies of Alveopora. It is difficult to see how the great pear-shaped colonies in this latter genus could possibly stand upon the tip of the original prototheca without gaining support on this principle. Extensive droopings of the rim till it touched the ground with subsequent bends up again are probably more common in this genus than the formation of thin radicles.
Root-processes may come from the rims of different proto- thecz in those cases in which the corallum is built up of a
* Cf. Fischer-Benzon, Abhandl. wissench. Ver. Hamburg, Bd. y. 2
(1871), pp. 1-23. ¢ + Arch. Zool, expér, (3) i. 1894, p, 445, pl. xviii.
14 Mr. H. M. Bernard on the
series, like those shown, for instance, in fig. 3. Omphyma is a typical case.
But this whole process need not detain us ; it has no serious morphological value, being obviously a device for a certain end. When that is attained, it has no further influence on the shape of the skeleton *.
Ill. Early flattening out of the Prototheca.—It is obvious that if, by any means, the early prototheca could be trans- formed rapidly into a disk, a broad base could be acquired by the skeleton which would keep it upright. It seems to me clear that the morphology of many of the Paleozoic corals can be explained on this hypothesis. But the different ways adopted of so changing the primitive conical prototheca seem to have been very numerous, and a review of the forms from this point of view is a desideratum. It is, I believe, along this line that we shall find a more natural set of characters for the revision of such groups as those row included, e. g., in the Cyathophyllide, than any now adopted.
In the present place I can only give a few samples, an, to avoid doubt as to the forms meant, I propose to take _ as examples certain well-known figures accessible to every
student.
“ Zaphrentis gigantea,’ pl. iv. of Milne-Edwards and Haimes’s Pol. toss. d. Terr. paléozoiques. I give this in passing because it is interesting as a very irregular method of acquiring a broad flat base. Diagram fig. 9 (Pl. I.) shows my interpretation of the process. It may be that the coral did not actually become detached and fall over, but that the method may be compared with radicle-formation, only, instead of a narrow lip, the whole side of the prototheca bent out- wards and apparently became cemented to the substratum.
It will be seen from a comparison with Milne-Edwards and Haimes’s figures that in this diagram I am assuming what the early transformation of the prototheca was from the shape of the tabulze in the adult stages; and this is, I believe, perfectly justifiable. Unfortunately not sufficient attention has yet been paid to the variations of the prototheca, which are still to be discovered. In certain types of modifications, e. g. those shown in diagrams figs. 11 and 12, the very earliest
* Miss Ogilvie’ssuggested origin of the Perforata froma great elaboration of root-processes so as to form the reticular ceenenchyma is very ingenious. But it is hardly borne out by the development of young Madreporidan corals in which the cup- or saucer-shaped prototheca persists as a basal epitheca (see p. 4), and being flattened out from the first has no oppor- tunity to form radicles.
Prototheca of the Madreporaria. Iie
modifications can still be easily seen; but in others they are at once obscured, incorporated perhaps in the subsequent stock, or, again, in others worn or dissolved off.
The tendency has been to regard the variations at the extreme bases of these Paleeozoic corals as accidental, and hence of no real value in classification. This view will, [ hope, for the future be abandoned and special attention be paid to any traces which can be seen of the different ways in which the early prototheca was modified. It is quite possible, indeed, that many will be found to have been largely accidental. For instance, such a bend over as that shown in diagram fig. 9 may have been pure accident. ‘The same may be said of radicle-formation. More extensive comparisons, especially from this point of view, are necessary before we can say whether such a method of forming a broad base as that shown in fig. 9 became habitual in any group of early corals or not. It is worth noting that other corals are known which adopted it, as, for instance, the Dipterophyllum glans of Roemer (‘ Lethea Geognostica,’ i. p. 371).
More interesting, however, than these irregular, one-sided bendings over are those which took place more or less symme- trically all round. The most perfect of these methods, and, I believe, one of the most recent, is certainly that in which the edge of the prototheca is very early bent down, that is before the cup has any real depth, as already described above (see p. 5) as being the case in the Perforata. ‘The successive bendings down and attempts to bend up again of the edge of this proto- theca will, as we have seen, account for the successive wrinkling of the flattened epitheca (see diagram fig. 10). The Perforata owe their leading characteristics to this fact, that upon their flattened prototheca or epitheca a purely septal theca arises, and as the polyps bud the new thecz are also septal and may mount upwards to form enormous stocks built entirely out of radial septa mutually supported by con- ceutric synaptacule, leaving the epitheca, as in Turbinaria, as a film beneath the base of the stalk.
On the solution of the question as to when this very early flattening out of the prototheca arose depends that of the first appearance of the Perforata in the coral system. We get what appears to be a flat, very wrinkled epitheca in Cyclolites of the Secondary epoch, and again still earlier in the Paleozoic Paleocyclus. But an examination of specimens of these at once shows that their flattened epithecze were not continnous as in fig. 10. In Palewocyclus the conditions may be represented by the diagram fig. 11, and for Cyelolites by diagram fiz, 12
16 Mr. Hl. M. Bernard on the
the tabule in this latter case being represented internally by vesicular dissepiments*. In these cases, then, instead of there being one continuously expanding prototheca, there was the usual repetition of protothece which is so patent in the Paleozoic forms and still persisting, though disguised, in all corals. Even in the Perforata with tall conical septal calicles it must occasionally reappear, while in forms like Porites and Goniopora it is very marked (see above, p. 9).
These diagrams (11 & 12) are instructive because we see in the Silurian Paleocyclus that the original conical shape of the prototheca was not yet quite got rid of but persisted as a kind of stalk, whereas in Cyclolites it was quite flattened out. The process of flattening was apparently a slow one, and we may assume that the earlier forms always started from a deep prototheca, however rapidly (as in the case of Paleocyclus, tor instance) the following protothecze may have flattened out. Only in time was the flattening-out process so antedated that the very first larval prototheca appeared as a flattened saucer. And then, again, it was necessary to wait for the development of a septal theca to take the place of the flattened prototheca, before the latter could be left to grow outwards continuously as a mere basal support. One factor in bringing about this eradual flattening of the prototheca, as seen, for instance, in Cyclolites, might perhaps be seen in the delaying of the secretion of the rigid walls of the cup, which was probably rendered possible in the case of those forms which produced well-developed radial or septal thecee, the formation of which might, in the early stages, use up the available material f.
There was, therefore, apparently a long period during which the rim of the prototheca was undergoing modification in the direction of bending outwards and, if one may so describe it, a period of uncertainty and hesitation. I am convinced that the gradual steps by which the various flattened
* Tabule are secreted when the whole basal skin becomes detached at once and secretes a new continuous floor. Dissepiments are the secretions of portions of the skin coming loose at different times. We may see two reasons for this partial detachment, and, these if correct, would throw some light on the distribution of vesicular dissepiments :—(1) the mus- cular attachments of the mesenteries buried in the skeleton may hold the skin down at definite spots ; (2) the original floor becomes divided up by ery septa, and thus the skin could not come off in one continuous sheet.
In Cyclolites the rims of the tabule, the internal parts of which are broken up into vesicular dissepiments, can be traced round the corallum as sharp lines (see fig. 12).
+ Lacaze-Duthiers, /. c., found that the septa could be the first skeletal elements produced in developing Perforates, whereas phylogenetically the
prototheca came first. .
Prototheca of the Madreporaria. 1
protothece were brought about deserve much more attention than has ever yet been bestowed upon them. While I would not deny that the rise of the radial ingrowths from the inner (or upper) face of the prototheca, that is the septa, on which Milne Edwards’s classification is mainly based, may not supply during this period the best taxonomic characters, I still do not think that the variations in the curve of the protothecal rims, or, in other words, the shapes and dispositions of the tabulz, can be so completely ignored as has hitherto been done. A few examples will show what I mean.
Diagrams figs. 13 a-f show some of the forms assumed by the prototheca of adult single Paleozoic corals, They were built up of series of such protothece fitting into one another and usually raised above one another, sometimes by septal folds or ridges, sometimes by vesicular arrangements of the tabule of which only the edges showed clear and sharp, or sometimes the sloping sides being vesicular, while the more or less flattened bases are smooth.
It is impossible now to say how far these foldings outwards and downwards of the rims are of the nature of accidental variations. But until we know I can hardly think it right to ignore them so completely as has been done, say, in the genus Cyathophyllum as given by Milne-Edwards and Haime. For instance, we find specimens called Cyathophyllum which show the prototheca of the shape given in fig. 13a (e. g. C. turbinatum, Goldfuss*, said by Milne- Edwards and Haimet to be C. ceratites, although they themselves give a figure of it which appears to have the prototheca of the form 13 A). Again, Goldfuss (/.c. fig. 8d) gives other figures of his C. turbinatum with prototheca 13c, while his C. ceratites (pl. xvii. fig. 2) is shown with prototheca 13 d, with tabulate floor and vesicular sides. This latter M.-EXdwards and Haime called C. Dechent with the same form of prototheca as their C. Bouchardit. C. heterophyllum § appears to have a proto- theca of the form, 13g. Goldfuss again gives a Cyatho- phyllum helianthoides (in his pl. xx. fig. 2 e) with the same prototheca, 13e, as that given tor the genus Ptychophyllum.
It is quite true that considerable variation in the slopes of these flattening rims may be expected. For instance, in Goldfuss’s figures of C. helianthordes, just referred to, some have the protothcca 13c, others wih rims even more convex
* Petref. Germ. pl. xvi. fig. 8a.
+ Brit. Foss. Corals, pl. 50, fig. 2.
{ Pol. foss. Terr. paléozoiques, pl. x. fig. 2, § Ibid. pl fig. 1.
Ann. & Mag. N, Hist. Ser. 7. Vol. xiii. 2
18 Mr. H. M. Bernard on the |
than 13/. And again variations of curve are seen in the figured section of Chonophyllum perfoliatum* with proto- theca 13d.
Thus at the very outset we find ourselves face to face with the crux of all systematic work: What is the taxonomic value of these slopes and curves of the rim in any individual | case? We know from Mr. Pace’s observations 7 that great variability in the openness and flatness of the calicle can_be correlated with the degree of muddiness of the water. The sediment runs more easily off a coral with a flattened open theca than from one with a cup-shaped theca. Then, again, | we are justified in assuming that these forms were developed — in each case by slow modifications of an originally deep pro- totheca (age, therefore, may have something to do with the form) ; and, lastly, we can imagine many different accidents which would tilt or depress such rims.
Nevertheless we have a structure of fundamental importance in the coral skeleton, and the form-variations of this structure may justly claim to take the first taxonomic rank. But how are we to distinguish those of importance from those which are accidental in individual cases? The matter is further complicated in the case of these ancient fossils, because the transition-forms are preserved equally with those which have passed over finally to some well-defined type. It seems fairly clear that classification of such forms must be attempted on wholly different lines from that still in vogue. Before any form receives a name we should satisfy ourselves by a close study of series that it embodies some new principle of struc- ture. Three or four such distinct principles can be gathered from the forms of the prototheca given in PI. I. diagram 13a~g. In a the rim continues to show no sharp bend downwards, and is distinet from that in which the rim tends to bend out so as to form an open dish either as 13 or 137; and both these differ from the sharper curve of the edge all round (139). Fig. 13 A, in which the edge bends rapidly over and then either hangs straight down or shows a tendency to curve up again, seems to me to be very easily distinguishable from 13, for, even though the two might possibly pass into one another, a smooth curve and a sharp bend are very distinct. ,
I propose now to leave all but one of these early variations of the prototheca, hoping that I have said enough to claim greater consideration for them in all future work on Paleozoic
* Brit. Foss, Corals, pl. 50. fig. 5, The section perhaps does not run true.
+ Ann. & Mag, Nat. Hist. ser, 7, vol, vii. (1901) p. 385.
Prototheca of the Madreporar ia, 19
forms. Fig. 13, however, is of very great morphological interest and demands some further attention.
In the first place, it shows a simple and very efficient method of enabling the skeleton to stand upright. It differs from the radicle-formation in that the lip bends over all round.
The septa which come over the lip run down on the outside
just as we know that they run down inside the radicle (see fiz. 84). It is also obvious that the flesh of the polyp must have clothed the outer surface of such a theca, which is no longer the outer surface of the prototheca. ‘To the flesh thus hanging over Bourne’s term “ perisare”” may be applied, and lower down we will compare it with, and distinguish it from, another principle of structure which also involves the forma- tion of a perisare. A calicle built up of a succession of such -protothece as those now under discussion, one fitted inside the other, as in diagram fiz. 1, would have a rib-like arrange- ment of septa running down on the outside ; but in this case one would expect to find traces of the hanging rims appearing irregularly one above another as whole or portions of rings round the corallum. They would appear to be drooping or perhaps even show a tendency to curl up again. Iris because no such epithecal rims show in the fig. 2, pl. 50, ¢ British Fossil Corals,’ that I doubt whether its prototheca has this form (13 A) or belongs to the type I shall presently describe as also depending upon the formation of a perisare. This point, then, may be left for the present. It is clear at any rate that its true place is nowhere among the Cyatho- phyllidee.
This form 13h is of special importance, however, for the understanding it gives of the morphology of the Silurian Calostylis as developed by Lindstrém.
This coral has been announced as a Paleozoic Perforate, and this claim has had to be dealt with for the British Museum Catalogue, the first section of which, it is proposed, shall deal with the Perforata. As I have shown above, the true Perforates were only possible when the prototheca was flattened out as shown in diagram fig. 10. When thus flat- tened the septal ridges towered up above it and free of its rim, carrying on the skeleton by themselves alone. The thece being constructed solely of radial plates and their synapti- cule: were necessarily porous. In Calostylis the prototheca was not flattened out at all, but folded as shown in 13, and the septa were not laminate, but appear to have been repre- sented by a compact mass of large, irregular, rounded or subangular nodules, arranged roughly in radial rows. These come over the edge of the thecal fold and extend down to the
Q#
20 Mr. H. M. Bernard on the
rim of the prototheca. The compact layer of septal nodules on both the inner and outer surfaces of the calicles cause the walls to look as if they might be perforated—as if the deep depressions between adjacent nodules might run right through. » But this they do notdo. One of the chief puzzles of Calostylis | has been how to explain the pendent tongues of epitheca | which hang down irregularly and at intervals round the corallum and sometimes bend even slightly outwards. There | can be only one explanation of them, and that is supplied us as soon as we have unravelled the modifications of the proto- theca and recognized that its rim was bent in the way shown in this diagram. I repeat it was of importance to have this point settled, for a Silurian Perforate was a difficulty which| the British Museum Catalogue had now to dispose of one way or the other.
Turning from this to a somewhat kindred point which has too long been waiting for solution, and which may be partly dealt with in this connexion: Mr. Quelch* has raised the question as | to whether the Paleozoic Cyathophyllide are not still surviving | in the form which he has called Moseleya. Itis quite true that we have in both eases skeletons built up of the same elements, and at first sight similarly disposed. It has already been| pointed out by Mr. Pace that some of the suggested resem-; blances of Moseleya to a Cyathophyllum have no value, such’ as, for instance, the supposed tetrameral symmetry of Moseleya. But arguments based upon more or less similarity will not carry us far. The relationship can only be proved or disproved by an analysis of the principles on which the two corals are built. It is not merely the fact that both have similar elements somewhat similarly arranged, which is of importance, but the principles of their respective arrangements. Now whichever of the curves or series of curves of the rim of the prototheca shall afterwards be decided upon as that which shall charac- terize the genus Cyathophyllum, there is no question at all that the special forms which Mr. Quelch relied upon (e.g. C. Stutch- buryt and OC. regium, at least as figured by Milne-Edwards and Haime in the ¢ British Fossil Corals’) are of the pattern 13 d with the floors tabulate and the sloping sides vesicular. Hence unless Moseleya can show a somewhat similar arrange- ment of tabule or vesicles, the relationship between the two cannot be maintained. Now an examination of the available specimens of Moseleya shows a principle of protothecal modi- fication which, in some respects, resembles diagram 13’; but on closer analysis it appears to be nearer that other method
* Chall. Report, xvi, 1886, p. 110.
Prototheca of the Madreporarva. 21
of perisarcal formation referred to above, which will be de- scribed in detail in the next section. We shall have there- fore to postpone the further discussion of this point for a few pages, contenting ourselves with stating that a comparison of the protothecal specialization of Cyathophyllum with that of Moseleya shows them to have been well nigh as wide apart as they could possibly be.
One word before leaving these early flattenings of the prototheca as methods adopted by the early Madreporaria for the purpose of retaining the upright position. It is difficult to see how, as single corals, they would be efficient for the purpose unless the rims managed to touch the ground and re-cement a part of the animal to the solid substratum, and this, judging from some of the shapes assumed, does not appear to have taken place. But what is wanted is a closer study of the protothece and their earliest modifications. One advantage of early flattening out they would obtain, however ; they would grow more slowly in height, and the leverage would not be so great. Further, if this flattening out meant , ever so small an increase in the size of the base of the proto- theca, we can see that it might be of some value to the coral, even though the rims did not again touch the ground.
‘The moment these flat-calicled forms begin to bud and form colonies the advantages of the flattening become obvious, as will be seen in another section.
IV. The Perisarc.—One of the simplest of the really important methods of keeping the prototheca upright was for the soft parts to bag over all round the cup until they touched the ground, so as to form a secondary fleshy foot. This process differs from that shown in diagram fig. 13 A, for it in- volves no gradual bending over of the rim of the prototheca. I assume that the polyp simply overflowed the edge of the cup, that it reached the ground, and even expanded somewhat over the substratum all round the point of attachment of the skeleton. Since the under surface of this overhanging flesh is a continuation of that which secreted the prototheca, it might be expected not only to secrete a layer over the outer face of the cup, but also to deposit a continuation of that layer where it touches the ground. ‘This latter might be thickened to form a solid pedestal, in which the tip of the prototheca would be firmly fixed. ‘The fleshy foot secondarily formed in the way described may have taken almost any shape, even sending out radial prolongations or embracing the round stems of weeds, in which cases the solid pedestals which it secretes would encircle such stems, fixing the corals firmly.
When once fixed the coral may continue to grow in height
23 Mr. H. M. Bernard on the
and size without fear of falling over. If the rise in height 1s slow the soft parts hanging all round down to the ground may go on thickening the wall, and especially the base, almost indefinitely, so as to keep the corallite nearly cylin- drical. In such cases the septal ridges on the inner face of the cup may be continued over the edge as ridges (coste) or as rows of (costal) spines down the outside. On the other hand, as soon as the base of the prototheca is sufficiently firmly fixed the corallite may grow rapidly in height as well as in width, and in so doing may drag the soft parts away from contact with the ground. ‘The latter will then persist as the typical “ edge-zone” or “ Randplatte” vound the mouth of the corallite. The withdrawal of the parts that thickened the base while the coral grows in size leads to the latter being turbinate. Bin.
From this point of view the typical “edge-zone” is in reality a vestigial structure ; it is the remains of the perisare * which in the young stage formed the secondary fleshy foot. But even as such it may continue to fulfil some useful function. It will always continue to leave a layer of skeletal matter on the outer face of the prototheca, thus increasing the thickness and strength of the latter, and it will continue to form costal (=septal) ridges or spines. In Galawea advan- tage is taken of its gradual withdrawal from contact with the ground to secrete horizontal or arched films round the base of each ealicle. In this way the corallites of a Galaxea colony are embedded in and supported by an increasingly thick layer of irregular filmy vesicular tissue f.
We are now in a position to reconcile our statement that the epitheca, as usually seen in adult corals, is the rim of the protothecal cup perhaps indefinitely expanded, with the appearances which have led to the text-book statement
* I suggest this distinction between Bourne’s “ perisare” and the “ edge-zone” of Miss Ogilvie; the edge-zone is the vestigial perisare. It is important not to confuse the perisare which hangs over the solid edge of the prototheca with the sides of the polyp of a perforate coral in which the prototheca has been flattened down and the septa alone form a secondary internal theca, and no bagging over of soft parts ever took place.
f There is in the Natural History Museum a specimen showing a group of “ Caryophyllia clavus” growing on a piece of a telegraph-cable from the Caribbean Sea (700 fath.). The individuals are near together and their perisarcs have covered the intervening spaces with a chalky film. Here and there in the angles made by the corallites with the sub- stratum the film is raised and slopes outward and downward from the sides of the coral. It is this kind of free film formation which has been specialized in Galavea,
Prototheca of the Madreporarta. 23
that the epitheca is that part of the skeleton secreted by the edge-zone and left on the sides of the coral as it (the edge- zone) is drawn up with the growth of thecoral. This secretion may show periodical wrinkles or thickenings if the withdrawal is intermittent; and it is also clearly epithecal, inasmuch as, morphologically, it must be regarded as a doubling over of the rim of the prototheca, as can be gathered from the diagram (fig. 14). But this secretion is only one of many, and, moreover, one of the most highly specialized, modifications of the rim of the epithecal cup. Hence while it is quite correct to call it epitheca, itis quite incorrect to define epitheca in terms of this single specialization of it.
It is also clear that if the term“ eutheca” is applied to such cups as those shown in diagrams figs. 1, 2, 4, and 5, in which the lip of the prototheca grows straight on, we want some other term to designate acup in which the bagging over of the soft parts has practically doubled back the edge of the cup, so that the fold adheres to its sides (see fig. 14). But L would suggest that the simple unmoditied theca should be called prototheca, while the term eutheca would be more aptly applied to the theca which has been secondarily attached by a solid pedestal, thickened by the extra matter secreted on its outside, and strengthened and armed by ribs and spines. We might call the wall of Zaphrentis, Streptelasma, &c. (diagrams figs. 1-4) “continuously protothecate’’ and that of MJontli- valtia (diagram fig. 3), or at least of those specimens in which the septa can be seen between the edges of successive saucers, discontinuously prototkecate.
But although this eutheca, with the meaning just suggested, is due morphologically to a doubling of the wall of the proto- theca by the secretion of a layer on the outside of the cup, it can hardly be described as due to a bending over of its rim. I conceive of it rather as due to the rapid bagging over of the soft parts, without at the moment any actual continuous growth of the rim. A true bending over would have been a growth process of the rim itself (sée fig. 13g). I imagine that only when the soft parts had acquired their new position on the outside of the cup that they commenced secreting the external layer, which is nevertheless strictly a continuance of the rim down the outside and into the basal pedestal.
This explanation of the morphology and origin of the edge- zone throws an interesting light upon a very specialized and inorphologically puzzling group, viz. the small highly sculp- tured free Turbinolide. ‘Their origin can now be understood from diagram fig. 14, if we suppose that the powers of secreting
7
24 Mr. H. M. Bernard on the
carbonate of lime were for some reason restricted, perhaps locally *, In that case the outside fleshy foot might fail to secrete a solid pedestal, and then if, perhaps owing to the move- ments of the animal itself, the prototheca became detached from the substratum, it would be completely enveloped by the polyp and become a small internal cup-shaped skeleton. The ribs or spines coming over the edge of the cup could then run right down to the extreme tip of the original prototheca, as they do in typical members of the genus, If this origin is correct, the genus Turdinolva will have to be regarded as an extreme specialization of the “ Euthecate corals,” and can hardly, as it now does, give its name to a family.
It is evident then that a considerable reshuffling of the Milne-Edwards classification is required. For instance, as has already been pointed out by Bourne, the ‘ Turbinolide ” can no longer contain such purely protothecate forms as Flabellum and Rhizotrochus, while the Euthecate corals will have to include such forms as Galawea, Kuphyllia, and Mussa, which were placed among the Astraide by Milne-Hdwards and Haime. Yurbénolva itself will be a specialized offshoot of the Euthecate corals. It would, however, be premature to found such morphological divisionsas Protothecata, Euthecata, for it might be discovered, for instance, that the method of forming a perisarcal foot round the larval prototheca has been adopted more than once by different types of coral. Indeed, we seem already to have discovered two ways, viz. that shown in fig. 14 and that found in the Paleozoic Calostylis (fig. 13 h).
And this brings me back again to the much discussed genus Moseleya, already referred to as that which Mr. Quelch, working on a single specimen, took to be a Cyathophylilid. Fortunately Mr. Pace was able to bring more specimens of Moseleya, and I have found two others in the great collection made by Mr. W. Saville-Kent on the Great Barrier Reef. All these specimens are Lithophyllie. The only difference that I can detect between them and the ‘Challenger’ specimen lies in the fact that the latter has flatter and more open ealicles. ‘This, as Mr. Pace suggests f, may be merely an adaptation to the mud which we gather is present in the parts where the ‘ Challenger’ specimen was obtained, Examination of the specimens with a view to discover what was the principle of protothecal modification overlying them reveals ithe type of structure shown in the diagram fig. 15. It is
* They are plentiful in the Barton Clays, + Ann, & Mag. Nat. Hist. ser. 7, vol. vii. (1901) p. 385.
Prototheca of the Madreporaria. 25
essentially the same as that shown in fig. 14, but the proto- theca was shallow and open and the soft parts had bagged over the low walls on to the ground, doubling them as shown in the figure. Large wing-like septa come over the wall and also reach to the ground or to the rim of the epitheca all round outside. Between these flange-like septa, as they grow upward and outward, the polyps leave one basal secretion atter another, so that both inside the cup and outside it there is an increasing thickness of vesicular tissue. In the diagram (fig. 15) the lines are drawn as so many distinct tabule. But it would hardly be expected that the successive detach- ments of the polyp would take place simultaneously within each interseptal loculus, right from the centre of the calicle over the edge of the theca down to the ground. But as dissepiments are only portions of tabule, the diagram is the best way of illustrating the facts. ‘This type of structure, in which the vesicular tissue not only rises between the septa within the calicle, but also thickens the column between the coste outside it, is that which lies at the base of Lithophyllia. It is true that emphasis has not hitherto been laid upon this point, for the simple reason that the prototheca had first to be discovered. Milne-dwards and Haime merely remark that dissepimental tissue is very abundant, while their classing Mussa with Lithophyllia shows clearly indeed that the arrangement of the dissepimental tissue had not been analyzed. On the other hand Knorr, to whose figure among others Milne-Edwards and Haime refer as a type of L. lucera, mentioned the “stony films round the foot”? and described the impression made upon him by the words “new crowns continually covered up the old ones.” ‘lhe meaning of this otherwise enigmatical saying is quite clear when we glance at the diagram (fig. 15) here given. We conclude, then, that there is no generic difference between JJoseleya and Lithophyliia and that the genus A/oseleya is superfluous. At the same time it is due to Mr. Quelch to point out, (1) that the analysis of the essential structure of Moseleya was hardly to be discovered from the single specimen at his disposal at the time, and (2) if it had been, there was no existing descrip- tion of Lithophyllia which would greatly have helped him. ‘The calicle of which he made a section was old, very much flattened, and somewhat distorted, and with the tissue on its exposed side largely killed down. ‘This latter point is of gieat importance, for it is the structure of the sides of the column which is essential to a correct diagnosis. Once, however, the clue is given, which is supplied in abundance by the new specimens, the structure is easy to comprehend.
26 . Mr. H. M. Bernard on the
With the striking superficial resemblance to Cyathophyllidee to mislead him, it is no wonder that Mr. Quelch was misled. Nor do I see how his claim could have been disproved without a clear understanding of the position of the prototheca in coral morphology.
While on this subject I may point out that Mr. Quelch’s figure (/, ¢. pl. xii. no. 5) of a small calicle of Moseleya showing marked tetrameral symmetry is seen on the actual specimen to have been distorted by too close contact with the shell of a mollusk much larger than itself. Its internal arrangement is not quite normal. Mr. Pace has presented the Museum with over a dozen specimens, most of them single forms in all stages of growth, and not one shows any such striking tetrameral arrangement. On this subject of tetrameral symmetry in the so-called “ Rugose”’ division of the Madreporaria I would refer the reader to what is said above (p. 11).
Whether, after all, the subsequent classification of the Lithophyllide will ultimately admit of the existence of a genus Moseleya among them I cannot say. In this paper I am only concerned in showing that it has no place among the Cyathophyllide. The latter are characterized by extreme simplicity of protothecal modification, the Lithophyllide by great complexity ; they are at opposite ends of the evolution of the coral skeleton.
Betore closing this section I should like to refer once more to the difference between the principles of modifying the prototheca shown in diagram fig. 18 4 and diagram fig. 15. In both the soft parts bag over and reach the ground, but in the former the lip grows with the growing of the soft parts and its bend isa true bend. In fig. 15 the soft parts seem to overflow the edge of the cup too rapidly actually to bend the edge. Only aiter they have taken up their new position do they secrete a layer on the outer side of the cup, and this layer is practically the homologue ef the bent-down edge shown in fig. 13 2, ‘The two methods are thus clearly distinct, but it is not always easy to say whether a particular case belongs to the one or to the other. or instance, in those specimens of Lithophyllie in the Museum which have the corallites crowded together and forming pseudo-colonies, it is frequently noted that where the interseptal loculi of adjacent corals run into one another the dissepiments are every where arched, suggesting an open bend of the thecal lip, such as is shown in the diagram fig. 13 h, or even more resembling the bend of fig. 13 g, or even of 13f. But in the specimens with single corallites the actual lip of the theca is mostly a solid plate
|
Prototheca of the Madreporaria, yal
like those shown in diagrams figs. 14 or 15, and from it the dissepiments slope away on the one side into and across the calicle, and on the other down to the substratum. But it is doubtful whether an actual section of the wall would show that structure so straight and continuous as it is shown diagrammatically in the figure (15), and it is quite certain that the tabula would not be so regular and complete.
It was some such case as that just referred to (?a specimen of Acanthastrea), in which vesicular arched walls separated
calicle from calicle, that inspired the diagram given by me on
pl. xxxiii. fig. 10 in vol. xxvi. of the ‘Journal of the Linnean Society of London.” I am not yet, however, prepared to answer the question as to which of the two methods of edge- zone formation we have just been comparing—that of fig. 13 2 or of fig. 15—the actual case was due. For, as we have just seen, the Lethophyllie show that the smooth, arched, vesicular dissepimental wall might be a secondary modifica- tion, and due to colony formation, of the true edge-zone formation of fig. 14, which is the subject of this section.
V. Early Budding and Colony Formation.—In vol. iv. of the ‘ British Museum Madreporaria,’ Introduction, p. 23, I suggested a restricted use of the word “ astreeiform,” viz. to colonies of calicles all reaching to the same height and without any apparent tendency to grow and bud independently. The true astreiform colony is therefore that built up by a ealicle which is by habit low and whose buds spread laterally over the substratum all round the parent. The group Astreide as now understood consequently cannot be a natural one. It appears to me that we may have astreiform colonies of corals whose prototheces are modified upon very different plans. And it is on these modifications of this fundamental element that the ultimate classification will have to be based.
We might expect, then, a great development of astreiform colonies among the Paleozoic corals from the forms in which the prototheca was early flattened out in the ways described. We might also expect that it would be those methods of flattening out which were from the first symmetrical, because if the parent had acquired its flattening as a secondary matter, after having perhaps at one time fallen over, it could hardly be expected that the buds would appear with the necessary flattened symmetry straight away, although in some of the Astreid forms with very large calicles ‘this must apparently have taken place.
While I think these conclusions are perfectly justifiable, we learn from the researches of Lindstrém that one great group of Paleozoic astreiform corals with very small calicles,
28 Mr. H. M. Bernard on the
e. g. the genus Heliolites, developed from a prototheca which had fallen over. From Lindstrém’s figures * we gather that the lip which touched the ground expanded as a flattened epitheca over the substratum, and buds appeared at intervals upon it. Especially characteristic are the various wrinklings and ridges which appear on the upper face of the epitheca between the buds. As the living layers were periodically de- tached from and rose above this epitheca they secreted tabulate floors, which repeated its wrinkles and foldings. In this way the structure seen in the section typical of the Heliolitide was produced. Through the tabulate laminas which form the bulk of the coral the calicles run as tubes, while smaller tubes also appear in many cases in the intervening tabulate tissue. These smaller tubes receive their explanation as the continua- tion of the folds or wrinkles already mentioned through the whole series of tabule. Such folds or wrinkles would run as naturally through a series of tabule as the septa run appa- rently continuously through the tabule of Montlivaltia, as already explained in fig. 3 and p. 8.
If, however, we had had no knowledge of the origin of Fleliolites, we should have assumed that it had been built up of calicles with the form shown in fig. 137. And, indeed, this is the form which the calicles of the adult colony assume, but it is not arrived at by a symmetrical outward folding of the rim of the prototheca, but indirectly from a parent the unmodified prototheca of which fell over in the way already described. We owe the small size of the calicle of //eliolites to this fact.
The chief difference between the Paleozoic and Recent astraiform corals is due entirely to the more recent development of the radial or septal, as compared with the concentric, proto- thecal foldings. In Paleozoic times the former were not very pronounced, so that the flattened or curved sides of the proto- thecal cups with their tabulate floors formed the most charac- teristic portion of the skeleton. ‘lhe cup was, however, never quite flattened out, there is always the remains of the bend where the lip first turned over. These bends frequently form ring-folds (see fig. 137), which become the walls of the fosse, while tabule form not only the floors of these fosse, but also the areas which intervene between the fosse. These areas are variously sculptured with radial septa, and when the respective areas of the individual calicles are not marked off from one another, the septa of one may run into the septa
* See K. Sv. Vet.-Alkad. Handl. xxxii. (1899), pl. i. figs. 25-28. Com- pare the case of Paleocyclus referred to above.
Prototheca of the Madreporaria. 29
of those around it, as, for instance, in Darwinta and Phillipsastrea.
On turning to modern Astraide, we find that the tabulate character of the Paleozoic corals has become obscured, on the other hand the septa have become prominent. ‘These conspicuous radial folds of the prototheca make it difficult to discern the exact character of the concentric foldings of the protothecal wall.
I would suggest that, as a rule, the rising of the radial septal folds has also raised the concentric rim-folds. We might diagrammatically express it by imagining a calicle like that in figure 13 7 becoming changed into the form shown in fig. 16, which represents a calicle with high double walls, and on each side of it 'a smaller bud. We may assume that the tall ring-fold has been formed at the expense of the earlier horizontal tabulate area round the fossa. It is im- possible here to attempt any review of the many Astreid forms, but, speaking roughly, they are built of groups of low calicles with the protothecse modified in this way. The difference between this and that shown in diagram fig. 14 is that the calicle is shallower and more open.
Without professing any intimate knowledge, I am inclined to believe that most of the different forms now included among the modern Astraidae may be referred to variations :—
(1) In the distances of the corallites from one another: (a) they may be wide apart, as in Orbicella, Solenastrea, Echinopora, &c.; (6) they may be close together, Favia, Diploria, &c.; (c) they may be so close that the outer wall of the parent supplies the inner wall of the bud, Prionastrea, Goniastrea, Leptastrea*, &c.; (d) even these single division- walls may be incomplete, Hydnophora.
(2) In the ways the intercalicinal valleys are filled up.
(3) In the characters of the septa and in the way in which they come over the edges of the fosse and are distributed on the surface of the intervening tissue.
Concluding Notes on the Terminology of the Walls.
A wall built by a direct continuation of the edge of the prototheca should, I think, be called protothecay. These
* The Paleozoic Michelinia might be regarded as the morphological equivalent of these forms before the development of septa disguised the protothecal cups.
+ The term epitheca may be retained in its usual sense, and be under- stood to refer to all traces of the primitive undifferentiated protothecal wall and rim, even though they have lost all signs of haying been once parts or expansions of a cup.
30 Mr. H. M. Bernard on the
primitive protcthecal walls, recognized by Miss Ogilvie as equivalent to epitheca, have been hitherto called eutheca. But proto- is a more appropriate affix to express primitive simplicity than ew-, which better denotes some special excel- lence. Hence I propose, once more, that eutheca be applied to those walls which have been thickened, ornamented outside, and cemented firmly to the substratum in the way described above (p. 22) and illustrated in diagrams figs. 13 A, 14, and 15.
We now come to the term “ pseudotheca”’ of Heider. This is applied to cases in which the septa are so crowded together that they fuse along lines which together constitute a fairly symmetrical solid thecal ring. ‘The parts of the septa within this ring are septa proper and the parts without are coste. Now I cannot help doubting whether this differs in any respect from the eutheca, for it is obvious that a eutheca, as here understood, over which the septa ran close together, would give exactly the same result.
The suggestion that this wall is built wholly of fused septa does not take the possibility of a ring-fold into account. But from the review of coral morphology here set out it would appear that the ring-fold was a more primitive structure than the radial septa. Further, it is really impossible in a matter of such complicated folds to say how much at their points of crossing belongs to the radial and how much to the con- centric elements,
That the concentric element plays a part we gather from the fact that dissepiments frequently slope up the interseptal loculi just as if, had there been space enough, they would mount over the walls. This giving off of dissepiments means that the basal floor shares in the formation of the wall. What is usually called pseudotheca, then, is to my mind simply a modification of the eutheca as here understood, and the word, if retained at all, should have a new significance. My own proposal is to apply it in the sense of Ortmann’s “athecalia” *, ‘This term was suggested by that author for the Pertforata in which the protothecal cup, being entirely flattened out, a new secondary theca rises up formed entirely out of septa with their synapticular junctions. Now it is obvious that no part of the old protothecal rim is found in this new septate thecal wall, and to mark the total distinction between this and all the wall-formations made by folds of the true lip of the prototheca, it might well be called pseudo- theca.
* Zool, Jahrb. (Syst. Abth.) iv. 1889, p. 493.
Prototheca of the Madreporaria. 31
The term “ athecalia ” of Ortmann, it may be remarked, has not been very well received, for it is certainly not true that the corallites, say, of Madrepora have no thece. The very opposite is the case; the thecze are most pronounced. What we want to express is that these theca are morphologically distinct from the original thecze of the Madreporaria, and no better term could be employed than that here suggested— pseudotheca.
Tam aware that a long critique of the views and suggestions of other workers on the subject of the wall should be offered before proposing a revision. But I have the excuse that the revision of the terminology here suggested rests upon a somewhat far-reaching revision of the skeleton. A closer comparison of the terminologies would involve a closer com- parison and criticism of the views on the wall-structure of each different author, some of which are, I confess, not always clear to me. Indeed, we seem to have had enough of detailed and complicated discussion. The great want is some simple working hypothesis which will enable us to coordinate the facts.
My own work has convinced me that some order appears out of the chaos if we recognize the prototheca and give it the important place in the morphology of the coral skeleton here all too briefly sketched.
In conclusion, I should like to emphasize the fact that this paper is intentionally devoted to the prototheca and its con- centric modifications—that is, to those modifications which alter its cup-shape concentrically. Only occasionally and where necessary reference has been made to the great and complicated system of radial wall-folds which are the most characteristic structures of the stony corals. These have, however, claimed the attention of workers too exclusively in the past. We shall only be able to obtain a true insight into the evolution of the coral skeleton when we understand both systems of modification—the more primitive concentric and the later radial—and ean trace out their influences on one another. It is to me a matter of sincere regret that this paper was not published prior to Miss Ogilvie’s comprehensive and patient treatise on the septa, for her valuable observations would then, Iam convinced, have admitted of more precise and coherent treatment.
EXPLANATION OF PLATE I.
vg. 1. The three earliest growth-periods of a primitive Madreporarian. The thick basal part is the prototheca (sens. strict.), see fig. 2.
32
On the Prototheca of the Madreporaria.
The curved line a represents the secretion of the basal skin after it has been dragged (?) out of the prototheca by the growth of the walls in height; 6 represents the secretion © formed by the skin after it has become detached from a.
Fig. 2. The same regarded hypothetically as three separate cups, the
lowest thick-walled cup being the prototheca (sens. strict.) ; in it cup aaa is inserted, and cup 006 in aaa.
Fig. 8. A diagram to explain the morphology of Montlivaltia, The
Fig. 4. Fig. 5 Fig. 6 1G, 7 Fig. 8 Jif, Fig. 10.
early cups rapidly expand and eventually become a series of saucers, €e.., supported above one another by the septal folds which run continuously upwards. On the right half of the figure the upper part is in section, showing the tabulate floor and the irregularly bent-up rim. On the left half these rims are shown from the outside as irregular bands of epitheca running round the coral.
An early stage like that of fig. 1, but, having fallen over and resecreted itself at a, it bends upwards again. The bagging of the detached basal skins will take the shapes shown, and the fossula in the bases of the cups will be on the convex or dorsal side of the curved skeleton.
. A diagram to show how, if the prototheca proper was wide-
mouthed when it fell over, the fossula will come over to the ventral or concave side. q@is again the spot where the coral secretes a new attachment.
. The diagrammatic representation of the arrangement of the
septa in the so-called Tetracorallia, It receives a simple explanation as due to the necessary rearrangement of the septa in a coral which fell over and was bending up again. See text, Demale
. Diagram to illustrate the method of budding of a prone proto-
theca and the subsequent bending upwards of parent aud buds which might give rise to such a form as Halysvtes.
. Two figures of radicle-formation, after Lacaze-Duthiers.
Diagram to illustrate the one-sided bend-over of the prototheca such as it is suggested would give rise to the Zaphrentis gigantea of Milne-Edwards and Haime. See text, p. 14.
Diagram to explain the early flattening out of the prototheca in the Perforata. The rim of the cup creeps outwards all round, generally with successive slight bendings up and then down again.
Fig. 11. Diagram of the early stages in Paleocyclus. The prototheca
Fig. 12
proper seems to have fallen over and then suddenly to have widened out, the repetition of this is still more widened out, and so on. What appears to have been a wrinkled basal epitheca is not a continuous growth like that in fig. 10, but a repetition of so many separate protothecal rims.
. Diagram of the early stage of Cyclolites. The prototheca is nearly flattened out, but it is still repeated continually, only instead of the secretions of the successively detached skins forming continuous tabule, they are broken up into vesicular dissepiments. Here also what appears to be the wrinkled epithecal floor is in reality a concentric series of separate rims,
Figs. 18a-g. Various forms assumed by the protothecss in Paleozoic
corals, all in the direction of becoming flattened out. The developmental transitional stages between the deep proto- theca and these adult forms have still in many cases to be worked out.
On some Parasitic Bees. 33
Figs. 13h andj. Two types of foldings of the wall of the prototheea— h, seen in Calostylis; 7, common in early astreeiform colonies, e. g. Helivlites.
Fig. 14. Diagram to show the overflow of the prototheca by soft parts which bag all round down to the ground and form a new fleshy foot. This secretes a pedestal which can fix the proto- theca firmly to the substratum and doubles the thickness of the protothecal wall. This, it is suggested, should be called the eutheca.
Fig. 15. Diagrammatic section of a Lithophylha. Large wing-like septa radiate out over the wall, and dissepiments are formed on both sides of it; within the calicle they slope inwards, on its outer side they bend down and thicken the column with vesicular tissue between the coste. Mr. Quelch’s genus Moseleya is built on this plan, and cannot therefore be a Cyathophyllid with prototheca modified on one of the simpler plans shown in figs. 13 a-f.
Fig. 16. A diagram to illustrate the principle of structure characterizing the modern Astreeidz: 1 is the central parent calicle with the prototheca modified somewhat asin fig. 14; 2, 2 represent buds from the lateral edges, the budding thus resulting in the production of an astrzeiform colony.
IIl.—Some Parasitic Bees. By T. D. A. CocKERELL.
Celionys ribis, var. Kincaidi, n. var.
? .—Length 11-13 millim., the difference in size partly dependent on the extension or retraction of the apical part of the abdomen.
Similar in all structural characters to C. ribis, but the pubescence of the head and thorax is ochreous, the basal part of the third abdominal segment is more sparsely punctured, and the apical dorsal plate has the apex beyond the slight lateral constriction a little more produced. There are distinct and conspicuous transverse grooves across the middle of the second and third abdominal segments, but not on the fourth or fifth. Tibial spurs black.
Hab. Olympia, Washington State, June 9 to 24, 1895, June 26, 1896, five females (7. Kincaid).
This is the first Coeelioxys recorded from the north-west. It is quite different from ribis in appearance, but structurally it is almost the same, having the same sculpture on the penultimate ventral segment, &c. A male collected by Mr. Kincaid at Olympia, June 18, 1895, is presumed to belong to C. ribis Kineardt, though the pubescence (especially on the face) is white. ‘This male almost exactly agrees with
Ann. & Mag. N. Hist. Ser. 7. Vol. xiii. 3
34 Mr. T. D. A. Cockerell on
C. sodalis, Cresson, though the lateral teeth of the scutellum, while obtuse, are not short; the apical margins of the wings are only slightly dusky. ‘The tibial spurs are black, and the fifth abdominal segment has no lateral teeth, though there are minute nodules. The lateral teeth of the sixth segment are long. The upper apical teeth are flattened and rounded. The spines on the anterior coxe are large and blunt. It is to be remarked that while C. ribis was described from a locality in the upper austral zone, it is also an inhabitant of the boreal, and probably goes far north of New Mexico. On June 29, 1902, my wife took females of C. ribis and C. Portere at flowers of Frasera at Beulah, New Mexico, in the Canadian zone.
The exact relationship of C. sodalis to rib’s and Kincaidi will not be determined until the male of the first-mentioned is discovered. The localities given for sodalis are New York and Colorado; New York, being first mentioned, may be considered the type locality. I rather expect that rédis and sodalis will prove to be one species.
IPeoLvs, Latr. (sens. lat.).
Females.
Fifth ventral segment of abdomen strongly concave in lateral view ; fifth
dorsal segment truncate; size large.. 1. Fifth ventral segment not so........ 2. 1, Legs black.......... Seales ore ayoe een Triepeolus concavus (Cress.). Ces feMrUPINOUS 5 we See meee ce nes Triepeolus penicilliferus 2. Fifth dorsal segment with only a diffe- [( Brues). rentiated apical lunule; small species. (GEBCGIS:) W288 as shiek Pe ee 3.
Fifth dorsal segment with a large diffe- rentiated area; large species. (T'ri-
CREOIUS ates cceiajo'n s niete a mien ote 6. 3. Front with a tubercle on each side; Bembellam edn wes sm wwinkhe ede bifasciatus, Cress, Front simple; scutellum black or PHBE MCUGISD cics;0 elo cyn enemies vats 4,
4, First abdominal segment hairy all over; antenne red, suffused with blackish ; tibial spurs clear red............... erucis, Ckll. First abdominal segment with a black hairless area; antenne black, with little srf-anty, ed Wencewedie ee eet 5. 5, Hind tibial spurs black ; two submar- ginal cells; antennze entirely black ; lower half of pleura hairless ...... Phileremus americanus Hind tibial spurs clear red ; three sub- [Cresson marginal cells; second and third ;
some Parasitic Bees.
antennal joints largely red; lower
half of pleura covered with hair ,
G: Meersvblack, : 5 vsasdac eitachareen ae stae Legs TOU +. ssc: cata cay ate ees
7. Larger; black area on first segment
narrow, % e, not much produced laterally Sh a eee eee Rs ie 8
Smaller; black area on first segment SO ROR ee rea eh a EE a Se 0 3
10.
11,
bo
or
“I
. Scutellar
. Larger; labrum entirely black
. Legs black
. Lower part of pleura bare ...
. Larger; tegule clear red ....
teeth long and sharply pointed, at least partly red; dark area on disk of first segment very small.
Scutellar teeth shorter, entirely black .
. Larger; mesothorax mainly red
Smaller ; mesothorax black Area on first segment a broad trans- verse band Area on first segment small, not pro- duced laterally ; RIG PAPO OH ost onin aid « Tegule clear light red; ” mesothorax with an anterior patch, not two dis- tinct stripes Tegule dark reddish to dark fuscous ; : mesothorax with two distinct stripes.
Can Mah One) 0G ee) 88 Se Whew) Glee
Glse #6 0 6) 90 &) Be @ ee eel o 8m
Smaller; labrum with two red spots. . Males.
Abdomen with eight Sayan light MATIN ys: svalvhcia aise aaa Abdomen not so marked... Markings of abdomen orange, white on sixth seament Staion Asano Markings” of abdomen white or " pale cream-colour ..,...
se eete .
Oo) eens 9: 2. a
. Mark on first abdominal seoment semi-
lunar , Mark on first abdominal segment a transverse band
eH 6) 0 0m) 0) 8) @) 6) ev oy 6
Lees red Bands on second to fourth segments interrupted in middle line ; size small; femora black . Bands on third and fourth segments, at ner entire ; hind femora, at least, re
CO eC Ye I ee ar a Pa
«6 @\ s) 8.0 6 06 0.6
Lower part of pleura covered with hair.
Smaller; tegule darkened ....,
. Anterior femora red; abdominal mark-
ings white Anterior femora )black ;
markings cream - colour ; :
alae hears 9s oud ton ara Sy 5
abdominal antennee
Bi
beulahensts, Chl. The
8.
nevadensis, Cress.
donatus, Smith,
9.
10.
pimarum, Ckll, mesille, Chill.
TR
tevanus, Cress.
occidentalis, Cress.
12. heliantht, Rob., var. helianthi, Rob.
verbesine, Cll.
nautlanus, Ckll,
5)
ae
3.
concolor, Rob. lunatus, Say.
olympiellus, Ckll.
5.
6.
te
occidentalis, Cress, helianthi, vay. arizonensts,
(Ckll, isocome, Ckll.
Cressoni, var. frasere, Ckll, *
36 Mr. T. D. A. Cockerell on
Triepeolus nautlanus, sp. 0.
go .—Length 93-11 millim.
Agreeing with 7. lunatus and T. concolor except as follows :—Light markings of thoracic dorsum, and particu- larly of dorsal segments of abdomen, light orange; sixth dorsal segment with the pubescence entirely silvery white, in strong contrast with the orange of the other segments; the extreme sides of the second to fitth segments are touched with silvery white, which is most conspicuous on the fifth; the bands on the second and third ventral segments are silvery white, the erect curved hairs on the fourth and fifth being fuscous; the pubescence of the face is entirely silvery (not golden at the sides as in 7’. flavofasciatus) ; the lower part of the pleura is hairy, with an ill-defined bare central area ; mandibles with a dull red spot in middle of outer side; an- tenne black, first joint of flagellum red beneath ; tibiee and tarsi red, spurs on middle and hind tibia black; femora black, reddened at apex, and the middle femora sometimes red beneath ; the tibie vary to black, but the tarsi in such specimens remain red.
Hab. Vicinity of San Rafael, Rio Nautla, State of Vera Cruz, Mexico (C. Hl. 7. Townsend). The dates are March 13 and April 7; it occurs at flowers of plant no. 1 of ‘'ownsend’s collection, which is a species of Compositee.
The insect is a tropical representative of 7. dunatus, appa- rently constant in its bright colours. It is possible that T. nautlanus may prove to be the male of the species described by Cresson from the female as Hpeolus totonacus.
Triepeolus nevadensis (Cresson).
Albuquerque, New Mexico, Sept. 15.
Recorded erroneously in Bull. Denison Lab. as Epeolus remigatus (p.73) and LE. robustus (p. 61). It is easily known from robustus by the prominence between the antenne. E. robustus was described from New Mexico, but I have not met with it.
Triepeolus pimarum and T. mesille, spp. nn.
The females of these two species agree in the following characters :— Light markings of thorax and abdomen cream- colour; first abdominal segment with only a very small median black mark; second to fourth segments with broad even bands, that on second with no anterior lateral processes ; labrum, greater part of mandibles (at least), and first three
some Parasitic Bees, 37
joints of antenne and base of fourth red; considerable white hair about base of antenna (not soin 7’. bardus), but clypeus and adjacent sides of face hairless ; clypeus and face extremely closely but very distinctly punctured ; pleura very strongly punctured ; tubercles red; hind border of prothorax densely pubescent; mesothorax extremely densely punctured, not hairy, but having a sort of mealy appearance; two short anterior stripes of pubescence (slender and very weak in mesille) ; scutellum not or hardly at all bilobed, its lateral teeth very long and pointed; only the margins of pleura hairy ; tegule apricot-colour ; legs red, some blackish suffused markings on middle and hind femora; hind tibial spurs dark ; hair on inner side of basal joint of hind tarsi orange; abdo- men extremely closely punctured ; fifth segment without a band, convex, with fine silvery pubescence, and with a quadrate minutely roughened red area; apical plate red, punctured, sharply truncate ; ventral surface of abdomen not banded, but pruinose, with minute white pubescence. They differ as follows :—
T. pimarum. T. mesille, Larger, length about 125 millim. Smaller, length 11 millim. Clypeus red. Clypeus black, with anterior margin red, Mesothorax red, with a broad Mesothorax entirely black. median black band. Scutellum and pleura (except an Scutellum black, the ends of the oblique black band) red. teeth red; pleura black, with a faint reddish spot. Teeth of scutellum curved at Teeth of scutellum straight.
ends. Apical plate of abdomen not or Apical plate of abdomen keeled. hardly keeled. Punctures at sides of second and | = Punctures at sides of second and third ventral segments of abdomen | third ventral segments very diffe-
not conspicuously different, rent, those of second being much larger and less dense. Wings quite dark, nervures Wings not so dark, nervures piceous. | fuscous, Three submarginal cells. Nervure between second and third submarginal cells usually in- complete,
T. pimarum was found by myself at Alhambra, Salt River Valley, Arizona, in the autumn of 1899, at flowers of Verbe- sina encelioides. Of T. mesille 1 collected a number of specimens at Mesilla, New Mexico, Sept. 24. Fora long time I have had the latter species labelled with doubt T. bardus, Cresson, but I believe it to be distinct, though closely allied. According to Mr. Brues the scutellar teeth of bardus are incurved.
38 Mr. T. D. A. Cockerell on
Triepeolus donatus (Smith).
A female in the National Museum, from San Bernardino County, California, October (Coguillett), is referred here, as it agrees in every particular with the descriptions of donatus by Smith and Cresson, except that the pubescence of the abdomen is identical in colour with that of 7’. concolor. It is to be remarked that 7’, superbus (Provancher) has nearly the same characters ; but its pubescence is pale yellow and the markings of the abdomen appear to be different.
Triepeolus isocome, sp. n.
The male was taken at Albuquerque, New Mexico, Sept. 16, at flowers of Isocoma Wrightit, and was recorded in Bull. Denison Lab. xi. p. 73, as Epeolus occidentalis. It is certainly a distinet species, differing from ocetdentalis as
follows :—
T. isocomeé S.
Smaller, about 9 millim. long ; abdomen less tapering.
Markings pale cinereous.
Labrum with a little apical pit full of white pubescence, its sides projecting and subdentiform.
Labrum all black.
Stripes on mesothorax hardly separated, z. e, the area between them pubescent.
Scutellum strongly bilobed.
Lower part of pleura covered with hair.
Wings shorter, hyaline; venation more ferruginous, marginal cell more obtuse.
Hair on inner side of basal joint of hind tarsi black.
Second abdominal segment with large pyriform lateral hair-patches, pointed antero-mesad.
T. occidentalis $ (from Colorado).
Larger, about 11 millim. long; abdomen more tapering.
Markings cream-colour,
Labrum with two minute apical projections, but no pit.
Labrum with a red spot on each side.
Stripes on mesothorax well se- parated.
Scutellum feebly bilobed.
Lower part of pleura nude,except on anterior margin.
Wings longer, brownish; vena- tion more fuscous, marginal cell more acute.
Hair on inner side of basal joint of hind tarsi orange-ferruginous.
Second abdominal segment with rather small lateral patches anterior to the band.
The mandibles of 7’. isocome are perfectly simple, red in the middle ; the antenne are black, the flagellar joints with obscure reddish spots; the hind coxe are mainly red; all the trochanters, femora, tibia, and tarsi are red; the scutellar teeth are short and black; the hind tibial spurs are black. Eyes (at least when dry) light green.
LT’. segregatus (Epeolus occidentalis, var. segregatus) appears
some Parasitic Bees. 39
to be also a distinct species, allied by the punctuation of the pleura to 7. pectoralis (Rob.).
Triepeolus helianthi (Rob.).
I have confused this with 7’. Cressont, which it very closely resembles, I have a female from [llinois, sent by Robertson years agoas Hpeolus mercatus, Another female was collected by Mr. C. E. Mead, Sept. 19, 1898, at the Experiment Station near Aztec, New Mexico, at flowers of Verbesina en- celioides, A specimen from near San Ignacio, N. M., formerly recorded as Cressont, is nearly 13 millim, long, but appears to be the same species.
Triepeolus helianthi, var. arizonensis, var. nov.
3 .—Small, length about 8 millim.
Wings clearer, marginal cell considerably shorter and more rounded at end ; labrum red; pubescent margin of first abdo- minal segment not broken anteriorly or posteriorly ; fringe on fourth and fifth ventral segments fuscous.
flab. Phoenix, Arizona, at flowers of Helianthus annuus, October 9 (Cockerell).
Perhaps a distinct species. The legs are coloured as in helianth7, the anterior legs being very dark.
Triepeolus Cressoni (Rob.), var. frasere, var. nov.
3 .—Variable in size, from about 8 to nearly 11 millim.
Antenne and labrum entirely black ; mandibles black with a red spot ; hind femora red, middle femora with a black mark above ; tegule reddish to piceous; nervures black except towards base of wing, where they become reddish ; hair-stripes on mesothorax broad, flame-like, connected with a broad hairy anterior border.
Known from helianthi by the entirely hairy pleura, and from occidentalis by the black anterior femora &c.
Hab. Beulah, New Mexico, about 8000 ft., June 29, at flowers of Frasera (W. P. Cockerell), July 12 (W. P. Ckil.), July 12 (2. D. A. Ckll.) ; Las Vegas, N. M., at flowers of Spheralcea Fendleri lobata, July 24 (W. Porter). I think the insect recorded from Beulah by Mr. Viereck as Z. occidentalis must have belonged to the present form.
Epeolus crucis, sp. n,
? .—Length about 74 millim. This is the species, found at Las Cruces by Professor C. H. T,
4() Mr. T. D. A. Cockerell on-
Townsend, which has passed as /. compactus, Cresson, in New Mexico, having been so identified by Mr. Fox. It may have been included by Cresson among his specimens of compactus; but it surely is not the species described under that name. From the description (Tr. Am. Ent. Soc. vii. p. 89) it differs thus :—Not especially compact, the abdomen at least twice as long as broad; pleura with the upper part densely white- hairy, the lower densely and coarsely rugoso-punctate, nearly free from pubescence ; pale (hair) markings white, not buff; abdomen strongly pruinose all over with fine pubescence, so that the usual black markings, while indicated, are more or less obscured, the black surface being nowhere exposed ; the apical white bands on the first four segments are broad and entire, and are somewhat emphasized by the fact that the apical margins of the segments, beneath the pubescence, are white ; the transverse dark band (grey because pubescent) on the first segment is much produced and quite attenuated laterally. Labrum, mandibles, and first three joints of an- tenne ferruginous, the flagellum brownish grey with a sort of silvery sheen above, ferruginous beneath ; anterior middle of mesothorax with a white hair-patch, no separate stripes ; scutellum faintly bilobed, black with two reddish spots, lateral teeth red, quite sharply pointed, not extending so far as scutellum ; tegule bright orange-ferruginous, Wings rather short, faintly dusky, with an apical cloud ; stigma and nervures of basal half of wing ferruginous, nervures of apical half dark fuscous ; marginal cell rounded at end, appendiculate. Legs ferruginous, the femora strongly infuscated, spurs light ferru- ginous; silvery area on last dorsal segment of abdomen inconspicuous.
Epeolus beulahensis, sp. n.
¢ —Length 7 millim.
Black with yellowish-white markings due to pubescence ; face, including clypeus, covered with silvery-white appressed hair; mandibles and labrum dark ferruginous ; eyes strongly converging below; eyes (dry) grey ; antenne long, brown- black; end of scape and the two following joints ferrugi- nous beneath; tubercles and tegule ferruginous ; scutellum entirely black, faintly bilubed, the lateral teeth very short ; thorax, including pleura, covered with pubescence, except disk and anterior margin (except two broad short stripes) of mesothorax, anterior half of scutellum, metathoracic enclosure, and a spot on each side of metathorax, which are bare and consequently black ; legs clear red, including the spurs; an- terior femora except knees, and anterior tibiz except ends,
some Parasitic Bees. Al
black ; middle femora with a black stripe above. Wings quite clear, except the broad apical margin, which is faintly dusky ; nervures and stigma piceous; marginal cell obliquely sub- truncate, minutely appendiculate ; second submarginal cell nearly as broad above as third. Abdomen thick-fusiform, the black areas very distinct, that on first segment a broad trans- verse band, obliquely truncate laterally ; hair-bands on first and second segments narrowly interrupted medially ; second with large lateral oval spots touching the band ; light areas on fifth segment just meeting on disk ; pygidial plate ferru- ginous, broadly triangular, but subtruncate at tip; second ventral seement white with a large black (nude) patch on each side; third and fourth with white hair-bands.
Hab. Beulah, New Mexico, prox. 8000 ft., July 11 (Cockerell).
Allied to £. autumnalis, Rob., but differs by the clear wings, small spines of scutellum, &c.
Epeolus olymptellus, sp. n.
$ .—Length about 74 millim.
Stout, with an oval abdomen, black with the usual yellowish- white markings; labrum entirely black, with two prominent apical projections ; middle part of mandibles bright ferrugi- nous; lower part of face, down to middle of clypeus, covered with silvery hair; clypeus densely rugoso-punctate ; scape black ; tubercles ferruginous; tegule dark ferruginous, mi- nutely and densely punctulate. Wings nearly clear, apical margin broadly dusky, nervures and stigma piceous; marginal cell obliquely subtruncate, minutely appendiculate; second transverso-cubital nervure with its upper half wanting; if it were complete, the second submarginal cell would be fully twice as broad above as the third. Femora black, the knees red; tibie with the greater part black; tarsi ferruginous ; spurs light ferruginous ; lower part of pleura thinly pubescent, densely rugoso-punctate; mesothorax with the usual two stripes widely separated, and without erect pubescence ; scu- tellum subbilobate, wholly black, the lateral teeth short, but very distinct. Abdomen with the black areas well-defined ; apical bands on segments 1 to 4 interrupted in the middle, the approximating portions of the bands on 2 to 4 club-shaped ; black area on first segment a very broad band, obliquely truncate laterally, and not produced much more than halfway to the lateral margins; band on second segment broadened at the sides, but no oval patch ; apical plate broadly rounded, black ; ventral surface with two broad white hair-bands.
42 Mr. G. A. Boulenger on a
Hab. Olympia, Washington State, July 2, 1896 (Trevor Kincaid).
Allied to E. cnterruptus, Rob., but basal joints of antennee not red, legs with much more black, postscutellum without a tooth, &e.
Phileremus americanus, Cresson.
Hab. Beulah, N. M., at flowers of Apocynum androsemi- folium, July 8 (W. P. Cockerell).
New to New Mexico. Cresson’s description is not suffi- ciently detailed, but I think my identification is certainly correct. This species and P. mesdlle, Ckll., are to all intents and purposes Hpeolus with two submarginal cells. I am convinced that these insects stand nearer to Hpeolus as restricted by Rebertson than that genus does to T'riepeolus.
The black band on the first abdominal segment is much less produced laterally in P. americanus than in P. mesille. The fringes of erect hairs on the fourth and fifth ventral seg- ments of P. mesille are white. While P. americanus flies in summer in the Canadian zone, P. mesil/le is a spring insect of the Middle Sonoran; a male before me was collected at Mesilla Park, N. M., May 7, at flowers of Dithyrea Wislizenit. It has the face densely covered with white hair.
The female of P. mesille has not been described ; but I have a specimen (Ckll. 2810) collected at flowers of Sophia at Mesilla Park. The abdomen is longer than in the male, and the hind margins of the first four segments are broadly orange, with a coppery lustre, and practically hairless, though perhaps denuded. More than the apical half of the fifth segment is orange, and the very distinct white lunule is bordered behind by brown. The pygidial plate is truncate. ‘The knees, tibiz, and tarsi are all ferru- ginous. ‘The flagellum is ferruginous, darker above. The disk of the mesothorax is dark brown, and the two light stripes are very distinct ; in the male there are two very large light patches on the anterior part of the mesothorax.
I1I.—Deseription of a new Genus of Frogs of the Family Dyscophide, and List of the Genera and Species of that Family. By G. A. BOULENGER, F.R.S.
[Plate II.}
COLPOGLOSSUS.
_ Pupil vertically elliptic. Tongue large, oval, entire and free behind, forming a plicate pouch at the point of its poste-
new Genus of Frogs. 43
rior attachment. Palatine teeth forming a long transverse series narrowly interrupted in the middle. ‘T'wo denticulate transverse dermal folds in front of the pharynx. Tympanum hidden. Fingers free, toes webbed at the base, the tips not dilated; outer metatarsal bound together. Coracoids strong ; precoracoids very weak, ligamentous; no omosternum ; sternum a large cartilaginous plate. Diapophyses of sacral vertebra moderately dilated.
Colpoglossus Brooksti. (Pl. II.)
Habit very stout ; head strongly depressed, once and two thirds as broad as long; eye small, interorbital width three times the width of the upper eyelid. Fingers short, obtusely pointed, first shorter than second; subarticular tubercles indistinct ; a large, oval, inner metacarpal tubercle. ‘Toes short, blunt, with a very short basal web; subarticular tubercles feebly prominent; a rather large and very promi- nent inner metatarsal tubercle. “The tarso-metatarsal articu- lation reaches the eye. Skin of head and body granulate, of belly and limbs smooth. Yellowish above, elegantly marked with dark brown lines, which form a network on the sides and limbs; a )-( shaped dark brown, light-edged marking on the head and nape, each of the longitudinal branches bifureating in front and behind; two chains of small black spots, some with light centre, along the middle of the back ; lower parts white, throat with wrinkle-like transverse brown lines.
From snout to vent 50 mm.
A single specimen from Bidi, Sarawak, discovered by Mr. Cecil J. Brooks in a hole whilst prospecting, and pre- sented by him to the British Museum.
The discovery of a member of the family Dyscophide in Borneo is a very important addition to our knowledge, all the members of this natural group being inhabitants of Mada-~ gascar, with the exception of the Burmese Calluella guttulata. So many genera and species have been added to this family since the publication of the British Museum Catalogue (1882) that a complete list, such as is here appended, will be welcome to herpetologists and to students of geographical distribution.
I, Pupil vertical ; palatine teeth in long transverse series, A. Preecoracoids ossified ; tips of fingers and toes not dilated,
a, Sternum large. 1, Dyscophus, Grand. 1872.-~Madagascar, 1. zmsularzs, Grand. 1872, 2. Guineti, Grand, 1875, 3. Antongilt’, Grand, 1877,
44 On a new Genus of Frogs.
4, Grandidiert, Blgy. 1896. 5. Alluaudi, Mocq. 1901. 6. Sternum small. 2. Calluella, Stol. 1872.—Burma., 1. guttulata, Blyth, 1855, B. Precoracoids not ossified. a, Sternum large ; tongue forming a pocket behind; tips of fingers and toes not dilated. 3. Colpoglossus, Blgr. 1904.—Borneo. 1. Brooksti, Blgr. 1904. b. Sternum small; tips of fingers and toes dilated. 4, Piethodontohyla, Blgr. 1882.—Madagascar. 1. notosticta, Gthr. 1877. 2. inguinalis, Blgr. 1882. 3. brevipes, Blgr. 1882. II. Pupil horizontal. A. Palatine teeth in long transverse series. a. Preecoracoids ossitied ; tips of fingers and toes dilated. a. Fingers and toes free. 5. Mantipus, Peters, 1883.—Madagascar. 1. Hildebrandti, Peters, 1883. 8. Fingers and toes webbed at the base. 6. Platyhyla, Blgr. 1889.—Madagascar. 1. grandis, Blgr. 1889. 2. verrucosa, Mocq. 1901. b. Preecoracoids not ossified ; tips of fingers and toes not dilated. 7. Phrynocara, Peters, 1883,—Madagasear. 1. tuberatum, Peters, 1883. B. Palatine teeth in one or two small groups or absent ; preecoracoids ossified; tips of fingers and toes dilated. a. Two small groups of palatine teeth. 8. Platypelis, Blgr. 1882.—Madagascar. 1. Cowanii, Blgr. 1882. 2. pollicaris, Blgr. 1888. b. A single small group of teeth in the middle of the palate. 9. Cophyla, Bttgr. 1880.—Madagascar. 1. phyllodactyla, Bttgr. 1880. ec. No teeth on the palate.
10, Anodontohyla, F. Mull. 1892.—Madagascar. 1. Bowlengert, F. Mull. 1892.
EXPLANATION OF PLATE II.
Colpoglossus Brookst, upper view, natural size. a, open mouth (xX 2); 6, lower view of hand (X 2); ¢, sternal apparatus (x 13).
The Collections of William John Burchell. 45
IV.— The Collections of William John Burchell, D.C.L., in the Hope Department, Oxford University Museum.
I. Introduction. By Epwarp B. Poutton, D.Sc., M.A., Hon. LL.D. (Princeton), F.R.S., F.L.S., F.Z.8., F.G.S., President of the Entomological Society of London, Hope Professor of Zoology in the University of Oxford, Fellow of Jesus College, Oxford.
[Plate IIL]
WuEeEvn, in June 1893, I was first placed in charge of the Hope Collections of the University of Oxford my attention was at once arrested by specimens of insects and other arthro- pods collected in South Africa about ninety years ago, and much larger numbers from Brazil with dates going back about three-quarters of a century. I was struck by the precision and detail of the data and by the existence of numbers which evidently referred to a diary. Three manu- script note-books were eventually found in the Hope Library, and these showed that the material had been collected by the great naturalist William John Burchell, truly described by Swainson as “one of the most learned and accomplished travellers of any age or country—whether we regard the extent of his acquirements in every branch of physical science or the range of the countries he has explored’ (‘ Cabinet Cyclopedia’ of Dionysius Lardner, vol. Taxidermy &c., Appendix, p. 383: London, 1840).
The first necessity was to ascertain if the data were as accurate as they were full and elaborate. A single quotation from the Brazilian note-book throws much light upon this important question. From Oct. 6th to Nov. 16th, 1825, Burchell was upon an expedition into Minas Geraes from Rio de Janeiro. The following note refers to the beetles collected on four days towards the end of this journey :—
* All the Coleoptera of 3rd, 4th, 5th, and 6th have since been marked 4. 11. 25, as the different day’s collections being mixed in one paper could not be distinguished. They were, however, all caught in forests or on the edge of forests. Some other Coleoptera caught on these same days, but which were put up in separate papers and marked, are proper! distinguished by their labels, but those certainly of the 4th are marked 4. 11. 25, with the 4 underlined, and consist of only a few minute insects caught at night by the candle.”
It is obvious that the man who wrote that note was a man
46 Prof, E. B. Poulton— The Collections
to be trusted, and the immense numbers of his unpublished observations on natural history at once acquire the value of records by a trained naturalist with a fanatical love of truth for its own sake. Here, then, was the means of carrying back the detailed record of the occurrence of many thousands of species in two most interesting parts of the world, and to construct a trustworthy standard by which to measure the rate of future change; for one great justification of the immense funds which are expended on museums is that they will serve this very purpose for generations yet to come. The critical examination of the Burchell specimens proves that with ordinary care and the exclusion of light insects’ pigments will endure for probably an indefinite period. Many of these specimens have not had ordinary care during a part of their history, the African collection being especially attacked by Anthrent, probably between 1825 and 1830, when Burchell was travelling in Brazil. But even upon the most fragmentary of these the patterns are still quite distinct and have undergone hardly any change.
The collection, combined with the manuscript notes on labels and in the note-books, furthermore supplies a great body of observations on habits, instincts, &c. which are still im- perfectly known, and often altogether unknown, In many cases | find the records of interesting observations since made and published by others, such as the sound produced by the South-American butterfly, Ageronia feronia, described by Darwin in the “ Voyage of the ‘ Beagle’”’ (London, 1876, pp. 33, 34), or the habits of the driver-ant (/ctton) and leaf- cutting ant (Gcodoma), described by Darwin, Belt, Bates, &e.
When I first began to arrange for the publication of an account of the Burchell Collections at Oxford it was intended to prepare an introductory memoir upon the life of the great naturalist himself; but this proved to be too extensive an undertaking for these pages, and it is hoped that the “‘ Life” will appear as a separate work at no distant date. In the meantime a brief abstract of the chief facts which I have been able to bring together is set forth below as an introduction to the papers which will follow.
William John Burchell, the eldest son of a nurseryman at Fulham, was born about the year 1782. He received an excellent education, as is proved by the admirable style of his published works, the facility with which he wrote Latin, and the number of sciences with which he was intimately acquainted. His manuscript notes on South-African insects in the Hope Department are written on the blank sides of the pages of his French exercise-book—a history of Greece
of William John Burchell. 47
translated into French in 1794, when he was about twelve years old. Burchell was also an accomplished artist and musician. He must have had a remarkable constitution, for he enjoyed uninterrupted good health and vigour throughout his long and, with the exception of native attendants, solitary journeys. He laboured throughout the whole of the time with astonishing energy—collecting, observing, recording, sketching, and writing detailed journals. The details of his tragic end in his eightieth year also show that he possessed extraordinary resoluticn at that advanced age.
Burchell’s features at about thirty-four years of age are preserved in a drawing made by J. 8S. Cotman in 1816, the year after the South-African travels had come to an end. The drawing was etched by Mrs. Dawson Turner, the grand- mother of Sir Joseph Hooker. The portrait, of which there is a copy at Oxford, brings back to us Burchell in the full vigour of manhood. ‘The face is highly intellectual and indicative of strong purpose and resolution, yet singularly attractive, even winning. ‘The appreciation and description in his South-African travels of many a quaint incongruity shows that he possessed an ample fund of humour. His invariable breadth of view and justice are well seen in the calm discussion of the methods and results of missionary labours and his accounts of the shabby treatment he received from some of the Boers, in which he always warns the reader against coming to a too hasty conclusion as to the character of a whole people.
In 1805, when he was about twenty-three, Burchell was appointed ‘ Schoolmaster and acting Botanist’”’ at St. Helena by the East India Company, and he remained in the island for five years, until his departure for Cape Town in order to begin his South-African travels. He was elected a Fellow of the Linnean Society, Feb. 15, 1808. ‘The romance of his life happened in &t. Helena, and probably exerted a profound influence upon his character, explaining much that is difficult to understand, and especially the secretive barren period which followed his return from Brazil in 1830. His father had disapproved of Burchell’s engagement to a lady in Fulham, and had, perhaps, obtained the appointment in St. Helena, hoping that everything might be forgotten. But the two still corresponded, and Burchell persuaded the lady to come out and join him in the island. During the voyage someone on the ship—it is said, the captain—fell in love with her and married her. Burchell had always been a naturalist and collector, but it is probable that the terrible shock drove him into these pursuits and away from companionship with his
48 Prof. E. B. Poulton—The Collections
fellow-men, for consolation or, at any rate, oblivion. Natural history pursued in this spirit, especially when habits become fixed and deepened with advancing age, is only too likely to lead to the non-productive life of the recluse, poring for long years over his collections, jealously guarding them from the sight of others, and yet giving no account of them to the world.
We now enter on the next great period of his life, the five years (1810-1815) of splendid work in South Africa. The first part of his travels, discoveries, and observations are described in the classical ‘ Southern Africa’ (vol. i. London, 1822, vol. ii. 1824), covering the period between his landing at Cape Town on Nov. 26, 1810, and his departure from Litakun on Aug. 3, 1812. The work contains a large and excellent map, showing the whole of his route. He had intended to follow up these volumes by a complete account of the whole journey, but this was never accomplished, and the manuscript of his journal and other materials from which it might be written have not yet been found. The fine collection of insects which he made in St. Helena and South Africa was almost destroyed by neglect, probably during his absence in Brazil (1825-30), but hundreds of species can be named from the fragments preserved in the Oxford Museum. The botanical collections, now at Kew, did not suffer in the same way, and are in excellent condition.
Burchell remained in England during the ten years which intervened between his South-African and Brazilian journeys. He sowed in his garden at Fulham hundreds of South- African seeds and some from St. Helena, keeping a careful record, now preserved at Kew, of the dates at which they came up. On Sept. 30th, 1817, he presented forty-three skins of South-African quadrupeds to the British Museum, and the neglect of these specimens, many of them unique, was the cause of his quarrel with that Institution (‘ Southern Africa,’ vol. i. p. 883 &c., vol. ii. p. 386 &c.).
A letter to Sir William Hooker, dated March 31, 1819, shows the care he took to suggest appropriate names for the new species which he had discovered ;—‘‘ I should mention that it was my practice when on my travels to give such specific names to my plants as the view of them in their native place of growth naturally suggested, without attending to their being new or not, which | had not always on the spot time to ascertain; but my object in thus naming them was that on my return to England I should find all the new species with more appropriate names than an inspection of the dried specimens in the herbarium might probably suggest
of William John Burchell, 49
to me.” An examination of his Brazilian note-book proves that he adopted the same excellent method in his later travels.
In 1819 Burchell was called to give evidence before a Committee of the House of Commons on the question of emigration as a relief from pauperism. In his evidence, which occupied nearly three hours, he advocated the suit- ability of the Albany district in the easternmost part of Cape Colony. Ina few days the Committee reported, and a grant of £50,000 was voted for this purpose. Burchell then ampli- fied and published his evidence in a pamphlet, ‘ Hints on Emigration to the Cape of Good Hope’ (London, Aug. 1819). This was savagely attacked in the ‘ Quarterly Review ’ for the following November, and Burchell replied in a sheet of four pages bound into the first volume of his ‘ Southern Africa.’ Looking at the controversy from the standpoint of the present day, there can be no doubt that Burchell was entirely right and that the loyalty of the Grahamstown district, which has shone so conspicuously during recent years, is in large part the outcome of his wise advice.
More than all the work described above, the arrangement of his South-African collections and the preparation of the two volumes on South Africa occupied Burchell’s time until he began to get ready for his next great journey.
Of the five years in Brazil very little is known, mainly because Burchell published nothing after his return. Hooker’s ‘ Botanical Miscellany’ (vol. ii. 1831, pp. 128-133) contains some very interesting extracts from his letters to Sir William Hooker, and the life of Burchell in the ‘ Dictionary of National Biography’ (vol. vii. London, 1886, p. 290) also has an excellent short account of these travels.
Inasmuch as the Brazilian collections of insects &e. are far more extensive than the African, and are, considering their age and the vicissitudes through which they have passed, in excellent condition, the following papers will be chiefly con- cerned with them, and it becomes of the utmost importance to show the exact route traversed by Burchell. This is clearly shown by the map on the accompanying Plate III., prepared from the data obtained by Miss Cora B. Sanders, of Lady Margaret Hall, Oxford. The data were gained by a careful study of Burchell’s manuscript note-books at Oxford, and especially the Index to the Localities of the Plants and Insects. Miss Sanders was able to find many of the names which have disappeared from modern atlases by an exami- nation of the older maps of Brazil in the possession of the Royal Geographical Society. ‘he numerous smaller viliages, halting-places, streams, &c. mentioned in the manuscript
Ann. & Mag. N. Hist. Ser. 7. Vol. xiii. 4
50 Prof. E. B. Poulton—The Collections
note-books or upon the specimens themselves will always be described as between or near places which have been thus identified and are indicated upon the map. As regards Bur- chell’s two expeditions from Rio de Janeiro into Minas Geraés and the Organ Mountains, hardly any of the places mentioned could be found; but it is clear from the time occupied and the account of the work done that he did not travel far.
Following the exact data which Burchell always records, we find that he left Fulham at 9.30 A.M. on March 10, 1825, and sailed from Portsmouth at 9 A.M. on March 15th. The main outlines of the journey are set forth below in a tabular form copied from a paper gummed into one of his manuscript note-books in the Hope Department, viz. ‘‘ Index to the Lo- calities of the Plants in the Brazilian Herbarium &c., serving also for the Localities for the Collection of Insects &c.” The only modification of Burchell’s original table is the insertion of the two expeditions from Rio in their proper positions in time, instead of placing them at the end, and thus putting the three separated periods in Rio itself in juxtaposition.
PLACE. Dates, = ae 3 Monrus. Days.
Voyage... ce ee de eee eee 11 hopPontural: 56. -caariev: Ls e ae 2 0 Voyage... Pa a Reach Adu ay ph hale ih en a te Voyage... hectare Oy aseeiee aeeen ome ie. 4 Teneriffe . soe de see 6. 25 Vat Rie oe ra ae Voyage... earns ii iy Pune S35 Neniwalt yp: jie Rio de Janeiro........ Picuiee Minas Geraés ........ io a opt 1 4% Rio‘de Janeiror.... - 2) a ani a Organ Mountains...... . r ae 27 Rio de Janeiro........ | He a 364 Lo eceie my 16
* This number is given by Burchell. He probably deducted the days spent in travelling from and to Rio.
of William John Burchell. 51
TIME PLACE. DatEs. Monrus. Days.
Voyage from Rio...... Yee Mada sacs ir eis 3
ambgs 4 Moe sy cee Bae ae a 58 2 21 9) > WWiavha tinal ha
SCOPES fe cites s 3,588 4 8h9 2 ye a di 14
PAVOUIG ast h.eett fb wip ate omaAs ae | 3
PS Wed (rl OR Be mere 138 s ont 6 | 4 Waar. 95 7 oO” or © set a Ga
Travelling. ...:..-++-. 179, ee aoetilakis mi SERRE A
omen tt decay: ae ah 9 18 i ore ee EE ES PES
ABmapel LUGS ots. clase» ates cs i it 3 2 22
y 92 |
ONO GaL fo osc eco on i 20} 5 15
Travelling (Tucantins) . 110. a Sot 1 13 2 pe ee i) ae. 6. 29 Gg ee ee
Para ©) (ete lela: 'e ie n)0) 6) 86's 0) a) © 10. 2. 304 8 0
POW BEO e acteany dover evap) My) te asievarhe iu 15
Burchell landed at Dover on March 24th, 1830, and reached his home in Fulham on the following day.
The journey originally planned by Burchell was far more extensive. Thus he wrote to Sir William Hooker from Rio (July 8th, 1826) :—
“. . . It is at least my wish to visit the city of S. Paulo, and thence by land through the provinces of Goyaz, Cuyaba, and Matto Grosso into Peru, having the city of Luzco as my principal object; and after doing in Peru as much as my time (for my family prefer my being in England) and slender means will allow me to do, I should wish to proceed by land to Arequipa, Potosi, Salta, &c., &c., to Buenos Ayres, and thence to my home at Fulham. . .”
A letter nearly two years later to the same friend explains the change. It is dated from Goyaz, April 25, 1828 :—
Ak
52 Prof, E. B. Poulton—The Collections
“. . Thave kept my original plan always in view, and had advanced thus far on my way to Peru &c. when letters from Fulham overtook me, stating that my dear father’s health, from the infirmities natural to his age, was gradually declining, and that it was his wish and that of the rest of the family that I should return directly to England. Whatever regret I may feel at thus relinquishing my American travels, and whatever disappointment I may experience from a prema- ture return, I have no hesitation whatever in preferring filial duty to science and the gratification of my own inclinations. I have therefore greatly altered my plans, and instead of ending this journey at Beunos Ayres, shall, Deo volente, end it at Paré, where I shall embark for England.”
Burchell was not destined to see his father again, for Matthew Burchell died soon after this letter was written, on July 12, 1828.
An excellent brief account of the Brazilian journey is given in a letter to Sir William Hooker, written from Burchell’s home at Churchfield House, Fulham. Much of it is printed in ‘ Hooker’s Botanical Miscellany’ (vol. ii. 1831, pp. 128- 133). The original letter, together with the others which have been made use of on the present occasion, are preserved in the Herbarium of the Royal Gardens at Kew. The letter is dated Nov. 1, 1830 :—
“TY left England in March 1825, passed two months at Lisbon and in the vicinity : landed at Rio de Janeiroin July, where | continued making collections in botany, entomology, and geology, &c., till Sept. 1826, during which period I visited a part of Minas Geraés. While at Rio I made some drawings of landscape, among which was a panorama taken from a hill in the middle of the city ; many astronomical, philosophical, and geodetical observations. I finally quitted Rio in Sept. 1826, and proceeded by sea to Santos, where I remained three months, and then proceeded and took up my station in a solitary hut in the midst of forests at the foot of the great range of mountains, for the purpose of exploring them at leisure. My next station or headquarters was at the city of 8. Paulo, nearly under the tropic of Capricorn, where I remained about seven months, extending my excursions in various directions. Having there purchased a troop of mules and engaged the requisite muleteers, I travelled northward, and finally took up my station at the city of Goyaz, being the first and only Englishman who has entered that province. There I passed the rainy season of 1827 and made large collections, being detained there nine months, owing chiefly to the difficulty of finding the means of conveyance for my
of William John Burchell. 3
On
baggage. At length, resuming the road and still continuing Northward, I reached in November 1828 Porto-Real, on the great river Tucantins. Here I remained till the proper season for embarking, and, descending the stream, at all times rendered dangerous by numerous rocky falls, rapids, and whirlpools, I made considerable collections on ground over which no scientific traveller had ever passed. I completed a survey of the whole length of this voyage, fixed by numerous astronomical observations. Finally, I arrived at the city of Para in June 1829, and, while’ waiting till February for a convenient opportunity of embarking for England, added largely to my collections both in zoology and botany. Of this city I made a panorama, which, with that of Rio, L hope perhaps to succeed in getting engraved, together with my landscapes &c. Of insects I found from 16 to 20 thousand specimens (at a guess). Of birds I shot and preserved 362 species. In the other classes a proportionally smaller number. I am not aware of any part of my collections being lost, though I daily lament my inability to unpack them for want of room in the house. The space I require is large, and I have some hesitation in building on bishop’s land, unless it were possible to enfranchise it. I fear I shall lose much time before I am comfortably settled: nothing is more dis- tressing to me than thus to be forced to delay my labours in arranging my collections and rendering them useful to science. You, who are so great an example of industry, complain also of overwhelming collections, and feel the necessity of manual help. But I have nowhere beheld an herbarium so large as my own; and, added to this, I cannot bring my mind to abandon any branch of natural history for the sake of giving more time and attention to any one in particular ; although I know this is wrong and can never lead to perfection in any. Still the contemplation of the whole system of created objects is so fascinating that it is very difficfult to] turn away from all but a few.”
These latter sentences, together with the considerations mentioned on pages 47, 48, help us to understand Barchell’s unproductive later years. Living secluded in the midst of his vast collections, he wandered from one point to another without the stimulus to work out any one part thoroughly which contact with his brother naturalists would have supplied. Furthermore, he belonged to that class of men, much rarer now than formerly, who value and gloat over collections as collec- tions. His letters, even to his most intimate friends, such as Sir William Hooker, as well as many records preserved in his note-books, show that he jealously watched over the material
54 Prof. E. B. Poulton—The Collections
of his collections, and indicate that he suffered much anxiety on this account. His will, which was proved for probate at under £4000, also shows that he was right in the contention that he could not afford to employ assistance in the skilled mechanical work which was required, while his almost too- scrupulous care and attention to detail must have consumed an immense amount of his time. Sir William Hooker had evidently urged him to employ a curator or librarian, for Burchell’s letter of June 25th, 1835, contains the following passage :—“ After the consumption of so much of my property by my travels and the disinterested pursuit of science all the rest of my life, the obtaining of assistance by payment is quite out of the question.” Similar advice had been given and answered in the same sense five years before.
The degree of D.C.L. Honoris Causa was conferred upon Burchell by the University of Oxford on May 8th, 1834. Daubeny, the Professor of Botany, had given his inaugural address on May Ist, and the first lecture of his first course (on Vegetable Physiology) was delivered on May 8th. It seems probable that Burchell came to Oxford in order to be present, and that the occasion was selected for the conferment of the degree.
There is no doubt that Burchell expected a government pension and that he bitterly resented what he regarded as un- deserved neglect. Hence, to the other causes which operated to prevent productive work, we must add the brooding melancholy and the bitterness of a disappointed man, the * man with a grievance.
It is probable that he freely communicated his tdeas on this subject to his friend Swainson, and that the attack on the government for neglect of Burchell was a result of their intimacy. ‘These severe criticisms may be seen in Swainson’s article quoted on page 45. The same article is probably responsible for exaggerated statements, which have been con- stantly repeated, as to the condition of his collections and the assertion that they were never unpacked. It was probably an extreme way of indicating the injury which science was receiving because Burchell remained unassisted. But it was ceitainly exaggerated. In the note-books at Oxford there is the record of the different dates at which he accomplished the setting of the various groups of Brazilian insects. More- over, the beautifully written labels which nearly all specimens possess are very different from the hasty but distinctly legible notes made in the field. Many specimens still retain both labels, but generally the older ones have been discarded.
‘J’o this grievance was added the further sense of failure in
of William John Burchell, 55
that others were continually gaining credit for work which he had done but had not published. Thus he wrote to Sir William Hooker on Sept. 3, 1832 :—
“I am vexed almost to death at all my fine collections being thus shut up from me while [ am daily losing portions of the only reward a traveller has—that of his discoveries.
I trust that [in] future my work will make more show, at least to the world.”
A few years later the same kind friend seems to have made a great and probably a final effort to induce Burchell to publish his results. Burchell’s reply is dated June 25th, 1835 :—
“From the manner in which you express yourself with
regard to my botanical collections you appear to be under very erroneous impressions, for to say that I ‘ w7l/ not publish’ is quite the opposite to what has ever been my intention, and the almost only pleasure I had in my travels to alleviate the excessive toil of forming them was the anticipating of the gratification of publishing them at my return to Kurope, and of obtaining the satisfaction of being useful to science, and of securing the honor [spelt thus, according to his custom] due to my discoveries; and if I have been, and still am being, robbed of those honors by others, who, having less on their hands than I have, can run the publishing race with more expedition, I feel most sensibly the injury I sustain. Many circumstances have unfortunately concurred hitherto to tie up my hands, but I do and shall ever look to Natural History as a most delightful and congenial employment for my future rears.” : Probably owing to the combination of causes set forth above and their deepening effect as years went on, Burchell became more and more of a recluse, and kept his collections more and more from the sight of other naturalists. The climax was reached when he refused the request of his old friend to allow his son, Sir Joseph Hooker, to see the collec- tion of St. Helena plants, in order to help in the production of a work upon the flora of that island.
Towards the end of his life Burchell must have come to realize that his methods could lead to nothing. He committed suicide on March 23rd, 1863, in his eightieth year. It is stated by C. J. Feret, in ‘ Fulham Old and New’ (London, 1900), that he “ shot himself under the large cedar tree in front of Churchfield House. The wound not proving fatal, he terminated his existence by hanging himself in a small out-house at the back.”
Burchell’s collections were not specially mentioned in his
56 Mr. R. I. Pocock on a new will (dated March 2, 1841). Upon his death in 1863 they
came into the possession of his sister, Miss Anna Burchell, who offered the whole of them to the University of Oxford in the following year upon the condition “ that separate rooms shall be set apart for them, and that the whole be put ont, set up, and systematically arranged, and be called ‘The Burchell Collection’ or presented to the Museum by Wm. J. Burchell, Esq., D.C.L.” The Delegates of the Museum were unable to accept these conditions. A few months later Miss Burchell wrote (April 8, 1865) concerning “ the collec- tions in Zoclogy and Entomology,” “ I am still desirons, in accordance with what I believe to have been his [Dr. Bur- chell’s] wish, of presenting the same to the University of Oxford.” The only condition was “that the Collections should be distinguished as those of my late Brother.” This offer was gratefully accepted, and in a few weeks the collec- tions arrived. About the same time the immense Herbarium was offered by Miss Burchell to the Linnean Society, which was unable to accept it. A little later it was presented to the National Collection at Kew.
In drawing up this brief account of Burchell, as a preface to the description of his collections, I desire above all to acknowledge the kind help I have received from Miss Cora B. Sanders in the study of Burchell’s manuscript at Kew and Oxford, and of his collections in the Hope Department. It has been already mentioned that the map forming Plate IIT. is en- tirely due to Miss Sanders’sresearches. I have also received the kindest assistance and encouragement from Sir Joseph Hooker and also the authorities of the Royal Gardens at Kew. The Delegates of the Oxford University Museum have kindly given me access to their correspondence and minute-books.
II. Ona new Stridulating-organ in Scorpions discovered by W. J. Burchell in Brazil in 1828. By R. J. Pocock, i ZS, [Plate IV.]
Ur to the present time stridulating-organs are known with certainty to exist in three genera of scorpions, namely, the Oriental genus Palamneus, the tropical African and Arabian genus Pandinus, and the South-African Opisthophthalmus, all belonging to the family Pandinidz. ‘The certainty in these cases lies in the fact that in both Palamneus and Opisthophthalmus the hearing of the sound preceded the anatomical investigation which led to the discovery of the organ, and that in the species of Pandinus an organ exists
Stridulating-organ in Scorpions. 57
exactly similar in structure to that of Palamneus, although the rasp and the vibratile bristles occur upon different segments of the chele and legs of the first pair in the two genera. What is believed to be a stridulating-organ has also been found in certain South-African species of the genus Parabuthus, which belongs to a totally different family, namely, the Buthidz. Unfortunately in this instance there is no proof, based upon human perception of the sound emitted by the living animal, that the function of the organ described has been correctly interpreted. The tenability of the suppo- sition, however, is justified by the structure of the organ and by the distinctly audible stridulation it can be made to yield, when the appropriate movements, all capable of being per- formed by the animal itself, are induced by artificial means on a freshly killed or alcohol-preserved specimen.
In the Pandinide the stridulating-organs have been deve- loped in connexion with the anterior appendages. In Opisth- ophthalmus it consists of large foliaceous bristles on the inner (preaxial) surface of the basal segment of the cheli- cere, and the sound given out by the rubbing of these appendages together is in many cases supplemented by the sound produced by the catching of certain short, erect, stiff bristles on the dorsal side of this segment against the anterior edge of the carapace as the appendages are forcibly withdrawn beneath it.
In Pandinus and Palamneus it lies between the basal segments of the appendages of the third and fourth pairs, commonly called the chelz and first pair of legs, and consists of a finely papillate area and an area beset with short erect bristles exactly like those that are found upon the upperside of the basal segment of the cheliceree in Oposthophthalmus *.
In Parabuthus what is supposed to be a stridulating-organ is totally different both in structure and position. It is a finely granular or transversely ridged area upon the dorsal side of the first and second segments of the tail, possibly also upon that of the last tergal plate of the abdomen, and the stridulation above mentioned can be artificially produced by scraping the point of the sting over the roughened field in question T. A fairly similar but less differentiated system of granules, probably subserving the same end, is found upon the first segment of the tail in certain black North-African species of Buthus, namely, the Egyptian B. bicolor and the Algerian B. eneas f.
* See Pocock, Nat. Science, ix. pp. 17-25 (1896). + Pocock, Proc. Zool. Soc., March 1902, pp. 222-224. { Pocock, Ann. & Mag. Nat. Hist. (7) x. p. 374 (1902),
58 Mr. R. I. Pocock on a new
Apart from the legs, which are almost immovably welded by their basal segments to the sternal surface of the body, the chelicera, chele, and tail are, with one exception, the only organs in a scorpion susceptible of vigorous and rapid movement. ‘The one exception is the pectines. It is in connexion with these appendages that the stridulating-organ now to be described has been discovered *.
In the course of a recent study of Burchell’s manuscript ‘ Note-book of Brazilian Insects &c.,’ Professor Poulton found the following record under the date December 3rd, 1828 :—
“1274, Scorpio of a light redish [thus] brown. Legs and claws pale. Several of these were caught in my house. I found one feeding on a large blatta which it held close to its mouth with its claws. ‘ Lacrdia.’ Makes a noise between a hiss and a whistle, v. J. 81. 12. 28, with its pectiniform appendages.”
The word “ Lacrdia’’ evidently represents the native name of the species. Burchell always made a point of ob- taining such names whenever possible, and took the greatest pains in writing them clearly and inserting accents. The reference “ v. J. 31. 12. 28” apparently alludes to a Brazilian journal which has unfortunately not been found, It certainly did not reach either Oxford or Kew.
At once appreciating the interest and importance of the last sentence of the note, Prof. Poulton arranged for the collection to be searched for a scorpion bearing the number 1274. The specimen was soon found by Mr. W. Holland, and Prof. Poulton brought it to the Natural History Museum and asked me to determine it and to examine the pectines, to discover if possible the nature and situation of the stridulating- organ. ‘This I undertook with the greatest pleasure, and with the result that the accuracy of Burchell’s observation was substantiated to the full.
The specimen is a male and belongs to the Brazilian species that I described last year as Rhopalurus Borelli. Although dried, it is sufficiently well preserved to preclude all likelihood of error on this point ; but without the relaxation or removal of the pectines the structure of the stridulating-organ could not be investigated. ‘The examination necessary for this
* Reference may here be made to the suggestion of Landois (‘ Tierr- stimmen,’ pp. 22-23, 1874), that the pectines might be capable of emitting sounds by friction. ‘This idea, however, was not supported by facts, and, except that the guess has now been verified, it is on a par with Wood- Mason’s view that the prehensile teeth on the digits of the chele in Buthide might also be used for this purpose (Proc. Ent. Soc. London, 1877, p. Xix).
Stridulating-organ tn Scorpions. 59
purpose therefore was carried out upon the three spirit-
reserved examples the Museum possesses, namely, the type, an adult female, an immature specimen of the same sex, and an adult but badly preserved male.
Although only described seven months ago, this species has been known to me for ten years. Briefly told, its history and that of its allies is as follows :—In 1893 * I pointed out that two American species of Buthide identified with the Scorpio junceus of Herbst and Tityus agamemnon of C. Koch differ from their allies in the structure of the pectines and of the first sternal plate of the abdomen. The pectines are unusually broad in their proximal half, and the overlying area of the sternal plate is depressed, the grooves which ordinarily pass forwards and inwards from the inner extre- mity of the stigma being exceptionally deep and lying nearer to the middle line, so that they define a narrow, smooth, tri- angular area, standing at a higher level than the depressed lateral portion already mentioned. On the strength of these structural features the genus [Heteroctenus was established for these two species. It was also stated that Heteroctenus junceus differs from the form then referred to H. agamemnon in having the depressed area smooth instead of closely and finely but distinctly granular. Subsequently, as a result of the publication of Dr. Kraepelin’s monograph + on the scor- pions, it was found that these two species can scarcely be separated generically from the species described as Rhopalurus laticauda by Thorell and R. princeps by Karsch. Further- more, the description given by Kraepelin, presumably from an examination of the type of 7%tyuws agamemnon, proved my previous determination of agamemnon to be erroneous. I[ therefore redescribed the species so determined under the new name Borelliit, and at the same time attempted to show that the five species under discussion—namely, junceus, lati- cauda, princeps, agamemnon, and orellii—possess certain characters in common of sufficient systematic value to justify their separation from the series forming the genus Centru- roides, with which Kraepelin associated them, and to demand their recognition as a distinct genus for which the name Ehopalurus is available §.
The significance of the depressed sternal areas and of the expanded pectines in &, Borellit and L. junceus was always
* Journ. Linn. Soc., Zool. xxiv. p. 393.
+ Das ‘Nerr., Scorpiones et Pedipalpi, pp. 94-95 (1899). t Ann. & Mag. Nat. Hist. (7) x. p. 376 (1902).
§ Biol. Centr.-Amer., Arachn, Scorp. p. 37 (1902).
60 Mr. R. I. Pocock on a new
a puzzle, and would probably have remained unsolved, so far at least as I was concerned, had it not been for Burchell’s until now unpublished discovery of three-quarters of a century ago. Probably the absence of the granules on the depressed sternal area in F. junceus, suggesting as it did the secondary importance of their association with the sternal depressions and with the pectinal expansions in R. Boreddi’, coupled with the flexibility, comparative softness, and known sexual physiological significance of the pectines in these and all other scorpions, combined to conceal the true construction, which, thanks to Burchell’s observation, is now known to be assignable to the features in question.
How, then, is the sound described by Burchell as ‘‘ between a hiss and a whistle’’ produced? Without doubt by sweeping the pectines across the granular field on the overlying sternal plate (Pl. IV. fig. 2). When one of these organs is turned over it may be noticed that the teeth opposabie to the granular area are not parallel-sided, as is normally the case in scorpions; the distal edge is sinuous, presenting towards the apex of the tooth a very decided bulge, which shows up as a slightly thickened area as it catches and reflects the light. When examined under a half-inch objective, or even a lower power, practically the entire face of the tooth, and especially the bulging area, is seen to be covered with a multitude of fine striz lying parallel to the longitudinal axis of the tooth (PIV. fig. 3). That the structural modifications of the teeth above described are directly connected with the depression and granulation of the sternum is shown by the absence of such modifications in the teeth at the distal end of the series which lie beyond the granular area and sweep clear of it with the movement of the pecten. No doubt the expansion of the shaft of the pecten in its proximal half is correlated with an increase in the size of its muscles and of the surfaces to which they are attached to add force to the sweep of the organ.
Except for the apparent absence of the granules, the sternal depressions in L. junceus closely resemble those of A. Bo- rellit. I originally described these depressions as smooth ; this is only true relatively speaking. No granulation is visible under a lens of low power, and no roughness is per- ceptible with a pin-point ; but when scrutinized with a half- inch objective the entire surface of the depression is seen to be exceedingly minutely shagreened, so minutely as to suggest that the sound emitted must be much finer than that which the organ in &. Borellii gives out. Nor is this all the
Stridulating-organ in Scorpions. 61
difference between the two species. ‘The pectines in R. junceus are expanded exactly as in &. Borellii, and the distal edges of the teeth bulge in almost precisely the same way, but the differentiation of the striae is carried to a greater extreme. Along the edge of each tooth there is a distinct series of small tubercular elevations, which are largest where they cross the thickened bulging area, becoming smaller both above and below it. These elevations are very distinctly striated, and the strive appear to be practically restricted to them (PI. IV. fig. 4).
In R. laticauda, Thor., the granules on the sternite are relatively as coarse as in ft. Borellit, but the area is less depressed and less sharply differentiated both in front and towards the middle line than in that species. Also the poste- rior surfaces of the pectinal teeth are less visibly striated and the distal edges of those opposable to the granular area are straight and without the characteristic bulge so noticeable in R. Borellii and R. junceus. In all these features the organ in R. laticauda is less specialized than in the two species just mentioned.
The remaining species of Rhopalurus are unknown to me. Those who have had the opportunity of seeing and describing R. princeps have made no mention of any structural peculi- arities in the pectines or in the first abdominal sternum. According to Kraepelin, who has seen the typical examples, however, this species is nearly related to R&R. laticaudu. Hence it is permissible to suppose that it also possesses a stridulating-organ similar in its general features to the stridu- lator of that species. In the case of 2. agamemnon the last- mentioned author states that the pectines are expanded and the sternum grooved and depressed as in 2. junceus, but that the sternum differs from that of 2. gunceus in being distinctly granular on the median triangular area. This peculiarity, in which &. agamemnon holds a unique position in the genus, suggests that the median area in question constitutes an integral part of the stridulating-organ. Whether the de- pressed areas are granular or shagreened, or neither, is at present unknown.
Two other important facts connected with Burchell’s observation remain to be mentioned. ‘The first is the discovery of stridulating-scorpions in America: those in which sounding- organs are known or supposed to exist have hitherto been recorded from the Mediterranean, Oriental, and Ethiopian regions. The second is the announcement of the exact locality of R. Borellit. FR. princeps occurs in Hayti, R.junceus
62 - Mr. E.S. Russell on Depastrum cyathiforme.
in Cuba*, R. laticauda in Venezuela and Colombia, whereas the only examples of 2. agamemnon and R. Borellit hitherto known are labelled “ Brazil,’ without further particulars. Thanks, however, to Burchell, we are now aware that R. Borellit is found in the Province of Goyaz, in the upper valley of the Rio Tocantins or that of at least one of its tributaries. Burchell was at Porto Real (now Porto Nacional) when he made his note on specimen no. 1274. Burchell’s collection also contains another specimen of the same species (a female) bearing a label “ Body and legs redish. Between the boxes at our station at Sape. 15. 10. 28.” Referring to the Index we find that Burchell gives “S? Brigida” as his locality on Oct. 15, 1828. Sapé is mentioned on Oct. 14. The position is between Caval- canti, his resting-place on Sept. 30th, and Conceigio, which he reached on Oct. 18th, but apparently much nearer to the latter. A glance at Plate III. will show the positions of these two localities of &. Borellit.
So far as the function of the organ in these American Buthidee is concerned, it need only be said that since it is equally well developed in both sexes, and occurs also in immature forms, there is no reason to suppose that it has any sexual significance. Hence, like the stridulating-organs of other scorpions and of the spiders of the family Aviculariide, its significance must be regarded as purely aposematic.
EXPLANATION OF PLATE IV.
Fig. 1. Rhopalurus Borellit, Poc., 9, nat. size; drawn from typical example.
Fig. 2. Ditto. Ventral surface of anterior extremity of abdomen and of posterior extremity of cephalothorax, to show the granular areas on the first abdominal sternite, the pecten of the left side being removed.
Fig. 3. Ditto. Piece of the pecten seen from its dorsal side, to show the finely ridged stridulating area.
Fig. 4. Rhopalurus junceus (Herbst). Ditto.
V.—WNotes on Depastrum cyathiforme, Grosse. By E. 8. Russet.
[Plate V.] M. Sars, in 1846, was the first to describe and figure this
interesting little Lucernarian. He discovered it near Bergen and described it under the name of Lucernaria cyathiformis
* There are specimens in the British Museum labelled “ Mexico ” and “Brazil.” These localities, however, require confirmation,
Mr. E. 8. Russell on Depastrum cyathiforme. 63
as follows :— Semipollicaris, stipite disco circulari, repando sese affigente; corpore cyathiformi, margine dilatata, repanda circulari, integra (s. non in radios divisa) tentaculifera, ten- taculis sepissime in fasciculis 8 fere continuis, ad marginem corporis dispositis; organis generationis 8, binis approxi-. matis ’’ (Faun. lit. Norveg. no. 1, p. 26, tab. i. figs. 8-13).
Shortly afterwards it was found in great abundance by Mr. David Landsborough, Jun., at Southend, Arran, and also by Dr. Landsborough at Corriegils, Arran. The specimens were identified by Mr. Joshua Alder as Lucernaria cyathiformis, Sars, and he sent a drawing to Mr. George Johnston, who, on the strength of this drawing, incorporated the species in his ‘ Hist. of Brit. Zoophytes,’ vol. i. p. 475 (London, 1847).
Gosse (Synopsis Brit. Actiniz, 1858) then founded the genus Depastrum for specimens which he found at Weymouth, which he regarded as identical with the Lucernaria cyathi- formis of Sars. Next year some small specimens were found by Allman (Rep. Brit. Assoc. Aberdeen, 1859) in the Orkney Isles, which seem to have been immature specimens of Depastrum cyathiforme, Gosse. It does not appear to have been recorded at any other locality until found by Beaumont at Port Erin, Isle of Man (‘ Fauna of Liverpool Bay,’ iv. : Liverpool, 1895). He mentions also a specimen from Plymouth.
In the month of July 1903 I rediscovered Depastrum on the shore at West Bennan, Southend, Arran; and in August, while at the Millport Biological Station, near the Lion Rock, Millport, and also near the old castle on the east side of Little Cumbrae. The animal seems to have a wide distribution, and I have no doubt that a careful search would reveal its presence in many localities from which it is hitherto unrecorded.
I found Depastrum in large numbers under stones at about half-tide, and also farther out. It adheres very firmly to the underside or occasionally round the edges of fairly large stones, so firmly that it has to be scraped off with a knife. It is very local in its distribution, but generally abundant where it does occur, though at one locality in Little Cumbrae I found only a few scattered individuals, It is difficult to account for its local distribution, but in my experience it is never found in muddy localities nor in spots where there is much decaying seaweed. It occurs well up the beach, and appears to be quite a hardy form. In Arran my largest specimens were got near low-water mark, but at Cumbrae large specimens occurred more plentifully halfway
64 Mr. E. S. Russell on Depastrum cyathiforme.
up the beach. In its natural conditions it is almost always pendent, being incapable of supporting itself with stalk extended and erect, on the upperside of a stone. When watched carefully in confinement it is seen to turn the widely expanded bell-like umbrella in different directions, as if searching for food. It appears to be quite incapable of refixing itself after having been dislodged from its resting- lace.
The stalk is very contractile, as is also the rim of the umbrella. Four muscles, which extend up the teenioles (Pl. V., ¢m.), are the agents for contracting the stalk, while the margin is contracted by a circular muscle (cm.) which passes round outside the insertion of the tentacles, and in contracting pulls the margin well over the tentacles, leaving only a hole in the centre, through which the tips of some of the tentacles appear. I may here remark that it is only in partly contracted individuals that several rows of tentacles are seen; in fully expanded adult individuals there do not appear to be more than two rows. Haeckel, in his diagnosis of this species (‘System der Medusen’), describes it as having the tentacles in several rows. Furthermore, none of my specimens reach the dimensions noted by Haeckel (8-10 mm. for length of stalk, length of umbrella, and breadth of umbrella), the largest I have seen having a stalk only 7 mm. long, while the usual size of good-sized specimens is 4 mm. for length of stalk, 6 mm, for height of bell, and 5-6 mm, for breadth of same. ‘These specimens seemed mature, having well-developed gonads. .
There appear to be two forms of the species among my specimens—one as figured, the other with a much sharper distinction between stalk and umbrella, and with the breadth of the umbrella as great as, or even greater than, the height of the umbrella. ‘This latter seems to be the typical form, for Haeckel describes the umbrella as being almost as high as broad. ‘The measurements of a medium-sized individual of this latter form are:—Lenegth of stalk 3 mm.; height of uinbrella 4 mm; breadth of umbrella 4°8 mm. The smallest specimen I possess measures respectively 1 mm., L'l mm., and 1:4 mm.
The sexes are distinct, but, so far as I can make out, indis- tinguishable in external appearance. ‘lhe gonads are typically in four double rows, but I have a specimen with only three gonads and three lobes to the manubrium. Indeed, the animal is very variable, especially as regards the number of fascicles of secondary tentacles. The ova and spermatozoa are very minute and very numerous. I attempted five times
On a new Spider from Bounty Island. 65
in August to fertilize artificially, but failed each time, chiefly, I believe, on account of the immaturity of the spermatozoa.
In the stomach of Depastrum I have noted the remains of a small crustacean (probably a Copepod). When kept in confinement unattached to a stone they sometimes void a floccular mass, along with one or two phacelle, which looks like a portion of the stomach epithelium. ‘The tentacles also are apt to slough off. It is very difficult to kill them well expanded, but I have obtained good results by carefully narcotizing with 30 °/, alcohol.
VI.—On a new Genus of Spiders from Bounty Island, with Remarks on a Species from New Zealand. By H. R. Hoae, M.A., F.Z.S.
Proressor CHARLES CHILTON, of Canterbury College, Christchurch, New Zealand, kindly sent me recently some spiders obtained by Mr. L. Cockayne from the islands lying to the east and south-east of the New Zealand coast. Among these were some specimens found on the guano deposits of Bounty Island, situated about 9 degrees east of Dunedin (170° 30’ East longitude), between the better-known Anti- podes and Chatham Islands.
The spiders belong to the family Agalenide, and the well- developed colulus, front spinnerets close together, inner margin of the falx-sheath toothed and sloping, with fringe of incurved bristles on the outer, the upright maxille, and square lip show them to belong to M. Simon’s group Cybzeeze. Allied to the genus Lmmenomma, Sim.*, this species differs too materially to be included therein, so that 1 have formed a new genus to receive it.
PACIFICANA, gen. nov.
Differs from Hmmenomma in having the cephalic part of the cephalothorax convex and wide in front instead of not convex and slightly attenuate. The thoracic fovea quite short and shallow instead of long and deep. Rear row of eyes so recurved as to form an area as long as broad instead of about one half as long as broad. ‘Two teeth on inferior
* The single species for which M. Simon formed his genus Emme- nomma was found on the islands adjacent to Cape Horn (about 67° W. long). The two localities are therefore separated by over 120 degrees of longitude.
Ann. & Mag. N. Hist. Ser. 7. Vol. xiii. 5
66 Mr. H. R. Hoge on a
margin of falx-sheath instead of three; three on superior margin, About five pectinations on superior tarsal claws instead of about nine.
The trochanters of all four pairs of legs are slightly but clearly hollowed on the underside. This, with the mandibular fringe and shape of lip and maxille, breaks down the last quotable distinction between the Agalenidz, Pisauridx, and Lycoside.
Pacificana Cockaynt, sp. n. Yynt,
The colour of the cephalothorax is dark brown, the cephalic part being bounded by a pale yellow marginal stripe. A similar pale yellow area extends round the thoracic part almost to the margin, where there is again a narrow streak of brown. The mandibles are dark brown. Lip and maxille paler brown, yellow on the outer edges of the latter. Sternum pale brown on each side, with a longitudinal central yellow streak. ‘The legs and palpi are yellow, with brown rings, one near the anterior end of the femur, one on the patella, two on the tibia, two on the metatarsus, one at the anterior end of the tarsus. In the front pair the whole of the tarsus and metatarsus is brown. ‘The abdomen on the upperside has a series of transverse scolloped stripes yellow and black alter- nately. The underside is greyish yellow.
‘The shape of the cephalothoraz is a long oval, truncate at the slightly narrowed anterior end. The cephalic part is considerably raised above the thoracic; a short, shallow, longitudinai fovea extends from behind the cephalic part to, but not down, the rear slope.
The pattern of the eyes is quite unique. The front laterals are large, one and a half diameters apart, and one third of their diameter from the margin of the clypeus. Four small intermediate eyes one fourth of the diameter of the above are situated between them at the corners of a trapezium, the rear pair, their diameter apart, slightly above the line joining the upper edges of the laterals; the lower pair, rather farther apart, are below the line touching the lower part of the laterals. The lateral eyes of the rear row, rather more than their diameter apart, are about three fifths the diameter of the front laterals and the diameter of the latter away from their own median. 'The small front median eyes are their diameter from the margin of the clypeus.
The mandibles are nearly twice as long as the front patella, much kneed at the base, and taper to the anterior end, the fangs being rather long, slightly curved, smooth for the first half and striated longitudinally the second. ‘Tie
new Spider from Bounty Island. 67
falx-sheath is sloping and has two teeth on the inner margin near the upper end. There are three teeth below the fringe of bristles on the outer margin, the middle one being the largest.
The maxille are upright, convex, rounded in front, and broadest near the anterior end. The dp longer than broad, on a narrowed base, is rounded at the sides and broadly truncate in front.
Pacificana Cockaynt.
a, eyes, X 10; 4, spider, nat. size; c, epigyne, x 10.
The sternum is nearly twice as long as broad, truncate in front, and running to a point posteriorly.
The abdomen is oval, sparsely covered with short fine hairs,
The spinnerets two-jointed, tapering, the second joint quite short. ‘The inferior pair close together, the colulus broad and long.
The legs are moderately stout, the metatarsal and tarsal joints tapering to a rather fine point. The superior tarsal claws have about five pectinations at the basal end only, the
5%
68 Mr. H. R. Hogg on a
inferior claw being smooth. At the anterior end of the meta- tarsi is a ring of short incurved spines and four pairs of spines on the underside of the front two pairs. The tarsi are without spines.
There is a longitudinal seam along the front side of the cox, and the chitinous margin of the trochanters is slightly hollowed on the underside, the species in this respect, as in the mandibular fringe, approaching the Lycoside.
The measurements ( @) in millimetres are as follows :—
Long. Broad. Cephalothorax.... 95 34 in front. 5 Abdomen neeiicr: 12 63 Mandibles ...... 4 Tr. & Pat. & Metat. Coxe. fem. tib. & tars. MUS satone et reluiele winters ] 3 7 8 ig = 25 2 92 62 64 es = 2c 3 2 6. 52 62 = 193 4 22 ff 8 7 = 242 Palpi: co eho eee OF. Wea hb 2h Se ayes cle
There are one male (unfortunately wanting a moult) and four females.
From Wanganui, North Island of New Zealand, Mr. W. Gray was so good as to send me two small pieces of moss- covered bark, each a few inches square. On my first examina- tion I could see no reason of adequate interest to account for their having been sent so long a journey by post. It was only after careful search that I found the lids of no less than five nests of a little Migas spider, apparently that first described by L. Koch, M. paradoxus.
The doors of the nests fitted so closely, and, although com- posed of woven felt, so exactly resembled the adjoining bark and lichen as to be quite invisible on a casual inspection. The occupant of one nest had come out and was unhappily crushed, but the other four nests contained live females, one in each. The nests are little silken sacs wedged between interstices of the bark, about ? inch in depth and inch across the opening.
In the collection made by the ‘Challenger’ expedition, recently returned to the British Museum (Natural History) after a prolonged absence, is another specimen from Wellington, evidently the same.
Spider from New Zealand. 69
The legs in all the specimens are rather longer in propor- tion to the cephalothorax than the measurements given by L. Koch, but they agree closely otherwise with. his description of his type specimen from Auckland, and I have no reason to doubt their being the same, more especially as the legs are normally carried closely bent up and are not easy to measure.
As in all this group, the tarsi and metatarsi of the front two pairs of legs are flattened and abnormally short. The metatarsi are furnished with a double row of stout curved spines on the underside (in my paper, Proc. Zool. Soc. Lond. 1901, 11. p. 229, by a misprint this character is ascribed to metatarsus iv.).
The superior tarsal claws have one long pectination, with a few uneven rugations on either side.
The front row of eyes is straight, the rear row is slightly recurved,
Migas paradoxus, L. Koch. a, eyes, X 10; 4, profile, nat. size,
The cephalothorax and mandibles are yellow-brown; sternum, lip, and maxillee yellow ; abdomen black and rather - rugose above, dark yellowish grey below. The space in front of the genital aperture and spinnerets yellow.
The strongly recurved cephalic fovea and rather profuse bespining of lip and maxille (in female) are marked features.
I append measurements (in mm.) of one of Mr. Gray’s specimens, apparently adult, andof the still larger ‘Challenger’ specimen :—
Specimen from nest (W. Gray).
Long. Broad. Cephalothorax.... 33 25 in front. 91 Ly a PAU COMIGN cipicn ssf oi0 4 3 Mandibles ....+. 3 hor’, 2 vert’,
70 Mr. H. Schwann on new Forms of
Tr. & Pat. & Metat.
Coxe. fem. tib, & tars. MSGS. aii a as tale Hea 13 3 a 2 103 2 18 pa ee eh EST 3 1 1 92 —— gL 4, 13 1 2 pee a Palpion ss... Aenea 13 22 3 1 == 2 ‘ Challenger’ Expedition specimen. Long. Broad. Cephalothorax.... 45 33 in front. 42 Abdomen........ 6 4t Mandibles ...... hor’, 33 vert’. Tr. & Pat. & Metat. Coxe. fem. tib. & tars. LUIS SG Heart roo, tr is 2 5 4} 3 = 143 2, cae woe: or. Tr ne 3. 12 33 31 2 = ae 4. 2 et Ome aaa MPP ats hn ates lassievecass lz 3 23 1 83
Migas distinctus, Cambr., from the South Island, described as having a pattern of yellow spots on the back and having more widely separated eyes, will no doubt be distinct from the above; but Mr. Goyen’s Migas Sandageri, from Moko- hinou Islands, near Auckland, now that we know he means recurved by bent forward, would seem from his description to agree exactly with IM. paradowus of L. Koch. Mr. Goyen found the nests of M. distinctus in clay-banks; those of M. paradoxus and M. Sandageri are on the trunks of trees. It is interesting to note that M. Simon has found the nests of the allied South-African Moggridgea to be built both in the ground and on bark.
VII.—On new Forms of Anomalurus and Sciurus from Tropical Africa. By HAROLD SCHWANN.
AN examination of some of the more recent African accessions to the British Museum collection which I have been enabled to make with Mr. Thomas’s permission shows that the following forms require description.
Anomalurus Beecroftt argenteus, subsp. n.
General colour above silvery grey, more or less suffused with yellowish towards the middle line; basal portion of the
Anomalurus and Sciurus from Tropical Africa. (i:
hairs on the flanks darker than those on the body, producing an indistinct dark patch on the edge of the membrane; general colour of uader surface dirty grey; throat strongly suffused with “ orange-rufous,”’ passing into pinkish buff on the stomach and hind limbs; head silvery grey, cheeks and lower jaw silvery white, a white patch on the crown between the ears and a white band running along the shoulders; under- part of forearms and sides of stomach dirty white ; outer edge of membrane on upper surface behind forearms covered with stiff black hairs extending backwards for about 14 inches ; tail dirty grey.
Dimensions of the type (measured in skin) :—
Head and body about 385 mm.; tail 139; hind foot . (s. u.) 41,
Skull: greatest breadth 86; length of upper tooth- series 12°5. Hab, Abutschi, River Niger, about 150 miles from the coast. Diy. eM, no, 2. 12 10. 7. Collected: Web. 1902 by
A. J. Braham, Esq.
This subspecies differs very considerably from the type of Anomalurus Beecrofti from Fernando Po described by Fraser both in general colour and skull-measurements. ‘The latter, however, in this group are so variable, even among members of the same species, as to be of little value. As an example two adult specimens, both undoubtedly Anomalurus Beecroft?, differed by as much as 1°5 mm. in the length of the upper tooth-series. In colour A. B. argenteus differs from Anoma- lurus Beecrofti in being of a light silvery grey on its upper surface instead of ‘ yellowish grey.’ It is also much less suffused with rufous on its under surface.
Sciurus rufobrachiatus ruwenzori, subsp. n.
Allied to S. kenie, Neum.*, but with a certain amount of fulvous on its muzzle and feet and a pure white streak on the under surface.
General colour above ‘ olivaceous,” the hairs brown, speckled with “ochraceous,” without the marked rufous suffusion found in S. nyanse. Base of the hairs “ slate- grey.” Length of the underfur about 15 mm. and of the long hairs 25 mm. Under surface “creamy buff,” not sharply detined, gradually passing into the “ olivaceous” of the sides. Middle line of under surface with a sharply defined white streak about 4 inch broad extending from the inter- ramia to the inguinal region, its hairs white to their bases.
* SB. Ges. nat. Fr. Berlin, 1902, p. 176.
72 On new Forms of Anomalurus and Sciurus.
Top of muzzle dull fulvous, passing into “ olivaceous’’ on the crown. Cheeks and upper surfaces of the feet and fore- arms grizzled yellowish. Hairs of tail annulated with black and buffy yellow.
Dimensions of the type (measured in the skin) :—
Hind foot (s. u.) 51 mm.
Skull: greatest length 52; basilar length 40°5; greatest breadth 29°5 ; length of upper tooth-series 9.
Hab. Wimi Valley, Ruwenzori. Alt. 2400 m.
Type. Adult male. B.M. no. 95. 3. 5. 2. Collected 6th July, 1894, by G. F. Scott Hlliot, Esq.
In colour this subspecies is intermediate between S. r. ny- anse and 8. kenie, having less fulvous on the muzzle and limbs than the former and more than the latter.
Sciurus rufobrachiatus pasha, subsp. n.
Fur hardly so thick or so long as that of S. 7. nyanse ; length of long hairs on back about 21 mm. and of underfur 11. General colour above dark brownish, rather warmer than Ridgway’s “ bistre” ; base of the hairs slaty black. Flanks distinctly lighter than back; base of hairs “ slate-grey.” Under surface very thinly covered with creamy-white hairs, interspersed with a few black ones. Difference in colour between flanks and belly unusually conspicuous, with the line of demarcation well defined. An indistinct white patch on throat and chest, hardly constituting a streak. Top of muzzle and round orbits dull orange-buff. Fore and hind feet and outer side of forearms rich ‘ ochraceous rufous.” Underside of thighs and lower limbs sparsely covered with whitish-buffy hairs. ‘Tail like back for its basal two inches, the remainder annulated with black and dirty white.
Dimensions of the type (measured in the skin) :—
Head and body 249 mm.; tail 234; hind foot (s. u.) 49.
Skull: greatest length (c.) 50; basilar length 39; greatest breadth 31°5 ; length of upper tooth-series 10.
Hab. Bellima, Monbuttu.
Type. Adult male. B.M. no. 87. 12. 1. 31. Collected 13th July, 1883, and presented by Dr. Emin Pasha.
This subspecies, allied to 8. r. nyanse, is more strongly suffused with rufous on the back and base of tail, while it is of a much lighter colour on the feet and belly. 8. kaffensis, O. Neumann *, from the other side of the Nile, differs by “ die schéne rostfarbene”’ annulation of the caudal hairs. It may be mentioned that an allied form from Southern Nigeria
* Op. cit. p. 57.
On new Lycenide from Sierra Leone. 73
is also remarkable for the almost naked condition of its under surface, but is distinguishable by the absence of any rufous colour on the limbs.
The four members of the 8. rufobrachiatus group found in Central and Central East Africa may be distinguished as follows :—
A. Fulvous or reddish on muzzle and feet. a, A marked white streak along under surface .... 8. 7, rutvenzorit. 6. No white streak along under surface. a', Underside of forearms and thighs deep rufous colour ; belly well haired, dull buffy ...... Sir. nyanse. b'. Underside of thighs with no rufous suffusion ; belly thinly covered with whitish-grey hairs. S. 7. pasha. B. No fulvous colouring on feet or muzzle ........ .. SS. kenie.
VIII.— On new Species of Lycenide from Sierra Leone. By D. Cator.
I FEEL pretty sure that the Pseuderesie here described are not the only new ones that I have lately discovered, but I await further material, which I hope to find before very long.
They need a deal of hunting, as their haunts are in shady places and they are most difficult to capture on the winge— firstly, because of their sombre colouring on the underside and the small amount of colour above, so that they can be seen only at intervals whilst flying; and, secondly, because if not taken at the first attempt they will not probably give another opportunity, as they easily take fright. If, however, they can be seen at rest they can easily be caught if they are not too high up, but they need much looking for; they rest on twigs and creepers bare of leaves, but, excepting one or two species, seem to be distinctly uncommon.
Pseuderesia Bakeriana, sp. n.
3 .— Upperside. Fore wings black, outer margin faintly scalloped, inner margin up to beyond vein 1 orange from near the base to beyond the middle: hind wings orange, with very broad black posterior borders decreasing rapidly towards costa. Underside. Both wings greyish black, hind wings rather the paler of the two: fore wings with red irrorations on the costa, a squarish red patch on the costa beyond the cell, which is confluent with the red irroration up to the
74 Mr. D. Cator on new
posterior margin and which broadly occupies the whole of that margin; in this marginal red irroration is a short black macular stripe, lower part of cell and below vein 4 to inner margin spotless and paler: hind wings with three interrupted irregular transverse reddish stripes, bordering the third a broad blackish band, edged exteriorly by a band of fine reddish irrorations, beyond which is another dark band, followed again by fine reddish irrorations to the margin; fringes white, intersected with blackish.
?. Upperside. Both wings orange: fore wings distinctly more rounded than in male; costal edge, base, cell and rather beyond irregularly black, apex very broadly, outer margin broadly black: hind wings like the male. Underside. Fore wings (with outline of pattern as above) orange, fading into yellowish on the inner margin ; costa blackish, three dark spots in the cell and one larger beyond it ; apex very broadly finely irrorated with reddish on a blackish ground ; outer margin similarly irrorated, but less broadly: hind wings like the male, but paler and without reddish.
Exp. wings, ¢ 31-33, 9 30-32 mm.
I have much pleasure in naming this after my friend Mr. G. T. Bethune-Baker, who has assisted me so much in working out my captures. Found so far in March, April, May, and October, but probably flies from October to May, which covers the dry season in Sierra Leone.
Pseuderesia nigra, sp. n.
3 .—Upperside. Both wings entirely black, with white fringes tessellated with black. Underside. Fore wings black, shading into dark greyish on the inner margin; traces of three black spots in the cell, with a red one between the second and third; apex darkly spotted, in front of which is an oblique row of four red spots, two being below the apex and two on the posterior margin: hind wings grey, of a peculiar texture, the wing having the appearance of having been denuded of scales, with various black and red spots; the base of the wing is suffused with red, with a small black spot palely encircled below the costal vein near the base; in the cell are two black spots, palely edged, the small one at the base and the other large, directly below which is another large black one; on the costa is a large black patch reaching to the upper angle of the cell, beyond and touching which is a red irregular spot; transverse stripe from apex to inner margin very decided, composed of a black-spotted stripe edged externally by an equally decided red-spotted stripe, the two
Lycenidee from Sterra Leone. 75
central spots in each being confluent and very large, margin spotted with a black Junular stripe; the ground has a suffusion of red beyond and below the cell; all the red spots are very bright, approaching vermilion.
2? .—Upperside. Both wings bright ochreous: fore wings with costa broadly dark brown, very broadly dark brown from the end of the cell and tapering down the posterior margin to the anal angle; cell with three spots in it and several below; ground-colour suffused with brown below the cell almost to the inner margin; fringes brown, intersected with white: hind wings with costa broadly brown, posterior margin very broadly dark brown, increasing in width from the apex to the inner margin, base suffused with brown ; fringes intersected with white. Underside. Fore wings pale orange, base suffused with blackish ; costa blackish, cell with three spots, increasing in size, and one below the cell touching the second and third cell-spots ; apex as in the male, but the oblique orange spots are preceded by a very broad blackish band: hind wings ochreous grey, with pattern as in the male, only the red spots are replaced by orange ones.
Exp. wings, ¢ 34, 2 30 mm.
This species may prove to be a subspecies of P. vardegata, S. & K., but it is a beautiful and striking form. Besides the types described above, I have one male with a small orange patch on the upperside of the fore wing. Caught in February
and April. Pseuderesia fusca, sp. n.
3.—Upperside. Fore wings black, with white fringes intersected with black: hind wings black, with an orange- coloured costa increasing in width from the base to below the apex on the outer margin ; fringes whitish, intersected finely with black. Underside. Fore wings dark grey, with a small black dash closing the cell and asmall black spot at the origin of vein 2; beyond the cell a curved transverse row of small blackish spots, followed by a similar more obscure submarginal row: hind wings ochreous brown, witha small dark spot near the base below the costal vein, followed by three small oblique dark spots—one below the costa, one closing the cell, and one below the cell; beyond the cell is a transverse, fine, inter- rupted, blackish macular stripe from the costa to the internal vein, beyond which is the rather obscure posterior marginal row of blackish dots ; margin finely dark.
9 —Upperside. Both wings black: fore wings with a broad orange-yellow patch a third from the base on the inner margin to near the outer angle, extending obliquely across
76 Bibliographical Notices.
the wing to above vein 4, where it suddenly narrows and is
inversely oblique to the costa : hind wings like those of the
male, but not so dark. Underside. Ochreous grey, inner
marginal area of fore wings yellowish ; pattern as in the male,
but rather more distinct, owing to the lighter ground-colour. Exp. wings, g¢ 27-29, 2 26-28 mm.
Liptena albicans, sp. n.
Upperside. Both wings white: fore wings with the costal half slightly tinged with cream-colour ; costa finely blackish (rather wider near the base), apical area rather broadly dark grey to black at extreme apex: hind wings with fringe cream- coloured. Underside. Both wings whitish, slightly cream- coloured: fore wings have costa to costal vein pale orange- yellow, continued finely to the apex; on the costa close to the apex are three dark dots or lines, which, however, are not always present; outer margin orange-yellow, edged internally finely with black, intersected at the veins as far as vein 3, the fringe of this part also being black, inner marginal area pure white: hind wings with the posterior margin very finely cream-coloured, edged internally by a fine black line; fringes whitish.
Exp. wings 29-31 mm.
This species is near L. decipiens, Kirby, but the underside of the wings has no trace of any marginal band at all. It very often flies high among the trees, settling occasionally, and not, as arule, moving faraway. Found in March, April, and June.
BIBLIOGRAPHICAL NOTICES.
Catalogue of the Collection of Birds’ Eggs in the British Museum (Natural History). Vol. Lit. By Kueenr W. Oares and Capt. Savite G. Rem. London: Printed by Order of the Trustees of the British Museum. 1903.
TE present volume contains brief descriptions of the eggs of 907 species, ranging from the Parrots to the Bulbuls (Pycnonotide).
Though the greater part of the book had been written by Mr. Oates, he was, owing to protracted ill-health, obliged to relin- quish the work, a fact which we must all deplore. The Museum, however, is fortunate in having secured the services of Capt. Savile ‘eid for the completion of the remaining volumes.
Bibliographical Notices. 77
No change has been made in the method of treatment, which, as we have already remarked, seems to us wanting in fulness and to miss a great opportunity for suggestive generalizations. Perchance Capt. Reid may be induced to give us a general summary on the study of oology in the last volume. Nowhere is the need for such a summary so well exemplified as in the case of the treatment of the eggs of the Common Cuckoo.
This volume is illustrated by ten coloured plates, remarkable for their extreme beauty. The selection of the figures has obviously been most carefully made.
The Geological Structure of Monzoni and Fassa. By Marre M. Oertvie Gorpoy, D.Sc., Ph.D. 1902-03[1903]. 8yvo. 180 pages, with 14 photographs, 33 figures, 4 geological sections (black and white), 8 geological sections (coloured), 1 table of stratigraphical succession, 1 coloured geological map, and 1 reference contour and fault map. Edinburgh: Turnbull and Speers. London: Simpkin, Marshall, & Co.
Tis memoir is a ‘Special Part” of Vol. viii. of the ‘ Transactions of the Edinburgh Geological Society,’ published in 1903. The date of “* 1902” on the titlepage refers to the year when it was reud before the Royal Society, as stated in the Prefatory Note. According to the generally aecepted bibliographical and nomenclatorial rules only the date of publication can be taken for the chronological status of a book. An abstract having been printed elsewhere, the Royal Society, by its rules, could not itself print the paper.
The Alpine Range, as a whole, is well known as a region that has been subjected to repeated movements ; and, indeed, it cannot be positively said that the cracks in the rocks and their displacemeuts are even now in a state of absolute equilibrium. In the South Tyrol the elevated areas of Triassic strata, rugged and precipitous, are characterized by more or less isolated, rudely columnar or sharply peaked mountains, which have long been objects of wonder to the tourist and of study to the Geologist. To the former it has attractions in its picturesque aspects; but, if his reflections reach farther and deeper than the common notions of mystery and romance among the bizarre cliffs, peaks, and gorges, he may well desire to know the ‘‘ why and wherefore ” of their real history and outcome. This country has for a long time been carefully examined by many Continental Geologists, to whose published observations and de- scriptions Miss Ogilvie (afterwards Mrs. Ogilvie Gordon) has referred in several papers. Attention had, however, been especially drawn to the fossils of Saint Cassian &c. Difficulties, however, were found in determining the relationships of the strata and the fossils. Of late years the lady-student above mentioned directed her energies to the elucidation of the doubts and difficulties which seemed hitherto to be beyond solution. Aided and guided especially by the advice of Baron yon Richthofen among her Continental and of
a Bibliographical Notices.
Professor Lapworth among her British friends, Mrs. Ogilvie Gordon, D.Sc., Ph.D., entered more fully into her projected work in the Tyrol. After hard field-work, making important contributions to our know- ledge of Alpine Geology, both as to the arrangement of strata and the occurrence of fossils, she completed in 1901 the excellent geological map which accompanies the paper before us. This brilliant and solid geological work has been steadily continued and improved by the same lady, asshown by hercontributions to scientific periodicals *, with elaborate and trustworthy descriptions of the region in explana- tion of its complex structure.
In these researches Dr. Ogilvie Gordon has always kept in touch with the Continental Geologists working at the same problems.
The Triassic masses in this region consist largely of Dolomites ; and these are said by the Author to be isolated by faults. Folded by many successive creeping movements of the Earth’s crust, inter- sected by slip-faults and thrust-faults, they have also suffered much by local subsidences, and by repeated cross-faultings, with shear- planes and their crush-breccias.
The outlines of the mountains in some places have been likened to that of upraised coral-reefs; and, if really such, the dolomite condition would not be strange, for it is known that corals become dolomitized. Careful scrutiny, however, detects fossiliferous strati- fication in some of the dolomite masses, but whether due to shells or to beds (not reefs) of Coral on bases of calciferous Algals is not settled. :
Both volcanic and deep-seated igneous rock-matter play important parts in the make-up and physical character of the country. The igneous magma has come up to the fissures of weakness in the various rocks, either to spread out on the top or to lose itself in the cross- cracks or in the side-planes and cleavage-lines. They take