Catalogue of the madreporarian corals in the British Museum

:

377

CATALOGUE

Zool,

OF THE

MADREPORARIAN

CORALS

THE

IN

BRITISH MUSEUM
(NATURAL HISTORY).

VOLUME

The Genus

I.

MADREPORA.
BY

GEORGE BEOOK.

«•

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PREFACE.

The

first

attempt to get the Collection of Stony Corals in the British

was made

Museum

in the year 1876,

when the Trustees engaged

Dr. Briiggemann to prepare a complete Catalogue.

Unfortunately, the work of

into systematic order

this able

Zoologist was interrupted by his premature death in

various reasons could not be resumed for
additions were

of the

Naturalists

should be

made

specially

'

to

many

1878, and for

In the interval important

years.

the collection, of which the specimens collected by the

Transit of Venus,'

mentioned; then,

'

in

Alert,'

and

'

more recent

Challenger

years,

when

'

expeditions
the question

about the formation of Coral-reefs had been reopened, a considerable amount
of material was received from several Naturalists
this inquiry

:

especially

who took

special interest in

from Dr. Guppy, who collected in the Solomon Islands

from Mr. G. C. Bourne, who investigated the coral-reefs of Diego Garcia

Mr.

J.

who

J. Lister,

passed some years in Polynesia

;

;

from

from Mr. Bassett-Smith,

;

who, by the instructions of the Hydrographer of the Admiralty, thoroughly
searched the Tizard and
'

Kambler

'

and

'

Penguin

the Eamesvaram reefs.

Macclesfield
'

;

their

survey by H.M.SS.

and from Mr. Thurston, who paid several

The

Mr. Saville-Kent transferred

Banks during

present work was near
to the

Museum

its

visits

completion when,

to

finally,

the large collection which he had

formed on the Great-Barrier Reef and in Torres

Straits.


PEEPACE.

IV

From

these and other sources the collection in the

present extent, the

1104.

They

the total

The
to the

are

Museum

has grown to

its

number of specimens of the genus Madrejwra amountino-

to

described

number of

in

the

present volume

species of the genus

amounting

under 180

names

specific

to 221.

task of arranging these materials with the nomenclature most
conformable

preceding literature was surrounded by unusual

the Author was assisted by a series of

named specimens

difficulties

;

and although

of Klunzinger's

Red Sea

Madrepores purchased some years ago, by one species collected by
Haeckel in
Ceylon and described by Ortmann, and by the whole of
the
'

he would not have succeeded so well in his work

if

Challenger

'

types,

he had not, at considerable

personal sacrifice, visited the principal coUections on
the Continent with the object
of studying the types contained in
them.
The Museum, besides, is greatly

indebted to him for the

care

bestowed on the curatorial part

of the

work,

by which every specimen has been now rendered
avaUable for future study.

ALBERT GtJNTHER,
Brim

M^eur., ^.
June

H.,

IQth, 1893.

^''^'' '^ '^'' De^^aHmenf of Zoology.


INTRODUCTION.

The

number

total

of species referred to the genus

Madrepora

s.

up

to

1890,

present work was undertaken, appears to be 157, not including the

when the

nomina nuda of Valenciennes and other authors.

recorded by the

descriptions,

and I

am

In most cases the

various authors have been identified

not aware that any author has

type specimens contained in continental museums.

from published

compared together the

The American

not been redescribed or refigured since the original publication.
therefore, lapsed into a state of confusion,

and the

occurring in certain areas are quite unreliable.

make, as

far as possible, a

species already described.

It

be added

these must

To

twelve or fourteen others recently published by Eehberg,
species

s.

lists

types have

The synonymy,

of species recorded as

has thus been necessary to

renewed study of the type specimens of the numerous
It has not yet

been possible for

me

to study the type

specimens preserved at Washington and other places in the United States, and
thus I have only been able to judge of the characters and positions of the species
described

by Dana,

Horn, and Verrill from the more or

supplied by these authors.

Museums

of Paris,

The

Berlin, London, Strassburg, and Jena, whilst Klunzinger's

Museum

be found in the Berlin Collection.

visiting the Stuttgart

are based

complete data

types described by European authors are in the

types are, I believe, preserved in the
also to

less

Museum, and my

1

at Stuttgart,

but a complete

set is

have not yet had an opportunity of

notes on Klunzinger's

Eed Sea

Collection

on the study of the specimens identified by Klunzinger which are

contained in the Berlin and London Collections.


INTRODUCTION.

vi

am

I

who have given me

various gentlemen
its

progress.

may be permitted

1

M. Bernard

in Paris, Professors

and

Haeckel

Professors

To Dr. Weltner

am

I

facilities for

all

my

hearty thanks to the

study and assistance during

mention the names of Professor Perrier and

to

Mobius, von Martens, and Dr. Weltner in Berlin,

and

Kiikenthal in Jena,

Dr. Ortmann in Strassburg,

Berlin

my

glad to have this opportunity of expressing

of

whom

me

gave

further indebted for

every assistance in their power.

much

valuable information on the

which he has been kind enough to supply since

types

and

Doderlein

Professor

I

the

visited

collections.

It

was

suggestion

the

at

undertaken, and

constant

his

which he has given

me

As a

by

my

result of the

I

in critical points

present

work was

the readiness with

have been a source

have also pleasure in acknowledging the

comparison of the various European Collections already
it

necessary to reduce the

described by previous authors from 169 to 130.

new

the

friend Prof. Jeffrey Bell.

referred to, I have considered

the

that

progress and

its

and advice

me.

to

Giinther

Dr.

interest in

assistance

of great encouragement
assistance rendered

of

species described

by myself

— 91 in

To

number

this

of distinct species

number must be added

Short descriptions of 62 of these

all.

have already appeared in the 'Annals and Magazine of Natural History' for

December 1891 and 1892.

Some
others

The remainder

are

now

described for the

are based on specimens in the Collection of the British
are

founded on specimens referred

to

Museum,

previously described

time.

first

whilst

species by

various authors.

For some time the idea was entertained to include an account of the
species of

Madrepora

in

the present volume.

position and affinities of the fossil species,
existing forms

would be of

The

little

is

known

and a careful study of

special interest

the various species as reef-builders.

Very

fossil

as to the

their relations to

on account of the great importance of

subject

is,

however, a very

difficult

one

;

the type specimens are scattered, imperfectly described, and rarely figured, added
to

which

most

fragmentary.

of

the

Satisfactory

specimens which
material

is

have come

under

my

notice

not readily obtained, but even

if

are
that

INTEODUCTJON.
had been

available, the advantages to

vii

be derived from a study of

been doubtful, inasmuch as in the greater number of the
characters on which the classification of recent species

is

it

would have

fossil

forms the

based are not preserved

with sufficient clearness to admit of comparison.

The

Plates which illustrate the present

Morgan and Kidd from

Collotype process by Messrs.

For various reasons
species

it

in

the

As isochromatic

to represent closely allied

plates

were chiefly used, the

depth of colour of the various specimens accounts for the

variation in intensity of the figures on

the specimens

negatives taken by myself.

was often found impracticable

on the same Plate.

diff'erence

volume have been reproduced by the

are figured

is

also

some of the

Plates.

not constant, and

The

scale

on which

depends entirely on the

reduction necessary for each specimen or group of specimens in order to
10- by 8-inch plate.

fill

a

I trust that sufiicient detail will be found in the figures to

give a good idea of the habit of the specimens, and also, although necessarily to a
less extent, of the form, angle,

and variation of the

corallites.

GEORGE BROOK.



SYSTEMATIC INDEX.

GENEEAL REMARKS.
page

HisioKicAL

1

Morphology

7
15

Classification

DESCRIPTIVE PART.
Page

MADREPORA.
Page

EUMABEEPORA, Broolc
1.

L

murioata,

palmata, LamJc
prolifera,

Lamlc

Lamh

cervicomis,

Dana
M.-Edw.

2. secunda,
3. crassa,

4.

Sf

H

intermedia, Bronl-

5. gracilis,

Dana

6. lieteroclados,

Brook

7. attenuata, Brool:
8.

cyclopea,

9.

conigera,

Dana
Bana

10. smithi, BrooTc

11. efl3orcsceus,

Dana

12. vasiformis, BrooJc
13. orbicularis, Brook
14. acuminata, Verrill
15. pacifioa, Brook

Dana
Dana

16. arbuscula,
17. virgata,

Ehrh
Dana

45

23. nigra, Brook
24. valenciennesi,

M.-Edw.

Sf

H

Lamk

23

25. laxa,

23

26. multiformis,

25

27. multicaulis. Brook

26

28. ehrenbergi,

27
30

29. clathrata. Brook

30

31. decipiens.

31

32.

32

33. florida,

Ortmann
M.-Edw.

47
48

£^

H.

48
49
50

Brook

30. irregularis,

51

Brook

52
53
54

Brook

listeri,

Dana
florida, Dana

32
33

confluens,

33

34. tuberculosa,

34

35. austera,

34
35
37
37
38
39
40
40

36. abrotanoides,

37.

danffi,

Brook

M.-Edw.

54
Sf

H.

Dana

M.-Edw. Sf H.
M.-Edw. Sf E.

39. gravida,
affinis.

Dana

Brook

60

41. compressa, B.- Smith
42. pociUifera,

43. aspera,

55

56
56
57
58
59

Lamk

38. pharaonis,

40.

46
46

60

Lamk

61

Dana

62

18. tylostoma,

41

44. manni, Quelcli

63

19. robusta,

45. scabrosa, Quelcli

64

20. grandis. Brook

42
42

Dana

43

47. squarrosa, Ehrh

22. pulchra. Brook

44

47a, thurstoni,

21. formosa,

46. divaricata,

,

Dana

64
65
200

Brook
h


SYSTEMATIC INDEX.
Page
II.

Obontoctathus, Brook

Page

66

94. microclados,

66

95. aculeus,

H.
50. stigmataria, M.-Edw. 4- H.
51. subtilis, Klunz

67

96. glochiclados. Brook

68
68

97. Burculosa,

98. macrostoma, Brook

105

52. reticulata, Broolc

68

99. antbocercis. Brook

106

53. oligocyathus, Broolc

69

100. recumbons, Brook

106

54. ambigua, Broolc

70

101. hyacintbus,

Dana

70

102. conferta, Quelch

107
108
109
110

^

48. arabica, M.-Edw.
49. borealis, M.-Ecliv.

57. impUeata,

6f

Brook

55. complanata,

56. tortuosa,

H...

Dana
Bmia

58. pruinosa, Brook

^

Ehrh

Dana
Dana

71

103. delieatula, Brook

72
72

104. kenti, Brook

103
104

104
104, 200

105. bifaria. Brook

110

106. patula, Brook

Ill

73

107. latisteUa, Brook

59. nasuta,

73

108. polystoma, Broolc

112
112

60.

74

109. indica. Brook

113

75

110. sinensis. Brook

114

76

111. frondosa, Brook

114

77

112. elegantula, Ortmann

115

III. PoLTsTACHTS, Broolc

61.
62.

Dana
paxilligera, Dana

digitifera, Dana
efFusa, Dana

63. haimei,

M.-Edw.

64. retusa,

Dana

df

H.

65. decurrens, Elirh

Horn
capillaris, Klunz

77
78

IV. Lepidoctathits, Brook

115

66. tubigera,

79

113. imbricata, Ehrb

67.

80

114. millepora, Ehrb

116

68. diffusa, Verrill

80

115. convexa,

118

69. dilatata, Brook

81

116.

Dana
prostrata, Dana

119

81

117. squamosa, Broolc

120

71. nana, Studer

82

118. subulata, Daiui

120

Dana

83

119. spathulata, Brook

121

83
84
84

120. selago, Studer

122

70.

dendrum, B.-Sinith

72. tenuis,

73. africana, Brook

74. rosacea, Esper
75. disticha, Brook

121. cribripora,
122. cuspidata,

Dana
Dana

116

123
124

85
85
86
87

123. rubra, Studer

124

Dana

125

127. sarmentosa, Brook

81. tizardi, Brook

87
89

82. quelchi, Brook

76. appressa, Elirb
77. assinulis,

Brook

78. cymbicyathus, Brook
79. alliomorpha, Broolc
80. secale, Studer

124. exigua,

125. mirabUis, Quelch

125

126. studcri. Brook

126

128. hebes,

Dana

127
128

90

129. obscura, Brook

129

83. cerealis,

91

130. monticulosa, Brilgg

130

84.

92

Dana
spicifera, Dana

85. sj-mmetrica, Brook
86. pectinata.

Brook

94
95

V. IsoPOEA, Studer
131. palifera,

131

Lamk

131

87. tenuispicata, Studer

96

132. hispida, Brook

133

88. candelabrum, Studer

96

133. securis,

133

89. patella, Studer

97

1.34.

Dana
cuneata, Dana

97
99

135. pHcata, Brook

134
134

90. corymbosa,
91. cytherea,

Lamk

Dana

92. annata. Brook

100

93. arcuata. Brook

102

VI. TnopoBA, Brook
136. nobilis,

Dana

135
135


SYSTEMATIC INDEX.

XI

Page
137. canalis, Qttelch
138. dactylophora, Brook

139. eurystoma, Klunz
140. fruticosa, Brook
141. tubicinaria,

Dana

142. botta;, Brool-

136
137
137
138
139
139

^

184. rousseaui, M.-Edw.

171

H.

172

Brook
186. elseyi, Brook

185.

exilis,

172
173
173

VIII. Rhabdoctathus, Brook

187. hemprichi, Ehrh

143. amblyclados, Brook

140

188. tubulosa,

144. diversa, Brook

141

189. hystrix,

Brook

141

190. sjTingodes, Brook

146. gemmifera, Brook

142

191. carduus, Darui

178

147. samoensis, Brook

143
144
145

192. striata, VerriU

178

193. rosaria, Daiui

179

14.5.

spectabilis,

148. scherzeriana, Briigg
149. humilis,

Dana

150. brueggemanni, Brook

14-5

151. ortmanni, Brook

147

Dana

Ehrh

175

DarM

176
177

Dana

pygmsea, Brook

179
180

damosa, Brook

180

rosaria,

181

147

194. hydra, Brook

148

195. orientalis, Brook

182

196. confraga, Quelch

182

197. clavigera, Brook

183

156. seriata, Ehrh

148
148
149

157. pyramidalis, Klunz

150

158. canaliculata, Klunz

151

152. acervata,
1-53.

klunzingeri, Quelch

154. ocellata, Klunz

155. cophodactyla, Brook

1.59. bullata,

Brook

160. globiceps,

Dana

161. platycyathus, Brook

184

IX. Tbachtlopoea, Brook
198. echidnaea,

Lamk

184

Dana

151

199. echinata,

152
153

200. subglabra. Brook

185
186

201. longicj-athus, M.-Edw.

Sf

H.

187

153
154
155

202. proeumbens, Brook

188

Dana
204. granulosa, M.-Edw.

188

156
157

205. rambleri, B.-Smith

189

206. rayneri, Brook

191

207. speciosa, Quelch

191

168. bseodactyla, Brook

158
158

169. leptocyathus, Brook

159

X. DisncHocTATHTJs, Brook

170. brevicoUis, Brook

159

162. botryodes. Brook
16.3.

calamaria, Brook

164. australis, Brook
165. plantaginea,

Lamk

166. erythrsea, Klunz
167. gnppji, Brook

Vn.

203. horrida,

210. inermis. Brook

161

211.

172. ce3-lonica, Ortm

162

212. angulata, Quelch

173. tumida, VerriU

163
164

176. loripes, Brook

177. cancellata, Brook
178. pumila, VerriU

179. violacea, Brook
180. valida,

Dana

181. microphthalma, VerriU
182. polymorpha, Brook

183. forskali, Ehrl

165
165
166

92

194

160

174. glauca, Brook

1

209. tenella, Brook

Klunz

175. concinna, Brook

189

H.

192
193

208. elegans, M.-Edw.

CoirocTAiHUs, Brook

171. variabilis,

4-

6f

H.

194

parilis, Quelch

195

Species incertce sedis.

213. coronata, R<-hh

196

214. dichotoma, Rehh

196

166
167
168

215. gonagra, M.-Edw.

168
169
170

Sf

H.

.'

196

216. papillosa, Rehh

197

217. parvistella, VerriU

197

218. philippinensis, Rehh

197

219. teres, VerriU

198

220. turgida,

Va^l

198



CATALOGUE
OF

MADREPORARIA.

GENERAL REMARKS.
HISTORICAL.

The name Madrepora

appears to have been

dictionaries) is madre, Ital.,

madreperla, mother-of-pearl,

The

derivation usually given

and the Greek
is

'

precise

The Century ' and other
The

Trwpo?, or possibly 7r6/3o?.

Zoophytes, which he classes together as Fori.
pora, Retepora, Frondipora, Tubulara, &c.

{cf.

Imperato* describes and

evident.

its

term was originally employed, does not appear

significance, or rather the sense in wliich the

to be generally understood.

used by Imperato in 1599, but

first

This group

is

affinity to Ital.

figures

a

number

divided into Millepora,

His work was published at a time

when

of

Madre-

naturalists

were anxious to show that the animal and vegetable kingdoms meet in a common area, the
Imperato is therefore especially
sea, and there produce an intermediate type of structure.
concerned to prove the " animal " nature of his Madrepora, and in doing so makes use of

which are quoted by Donati and appear to indicate the association of ideas
which led to the use of the word. The following quotations from Donati's work f (French
"Madrepora" " les
edition, 1758) are a free translation of the original of Imperato :— In
several expressions

nouvelles additions se forment en consistance de Pore, et de substance charnue

on

Madrepora, amas epais de Pores

animaux marins comme

calcareous

cups] n'est

les

abeilles

croit

dans les gateaux de

t

(Napoli, 1599)

'

Dell' Historia naturale

'

Essai sur I'histoire naturelle de la

'

Mer

oii

La

cire

se forment des

tubulaire [the

"Madrepora."

in

Dendrophyllia, probably D. ramea.^ln

the

qu'un reservoir d'animaux."

Imperato's work appear to represent a species of
*

que c'est une m^re

;

The

figures

of

Latin transl., Colonise, 1695.

Adriatique

'

(La Hayo, 175S).


GENEEAL EEMAEKS.

I
word

test the

replaced by

is

Madripora

spelled

717)^ but in the explanation of the figures the

(p.

i

is

In the Latia translation of Imperato's work, published in 1695, the word
" Madrepora " is always translated " Porus matronalis." It is clear from the above quotations
that Imperato regarded what we now speak of as the " corallum" as a stony "nurse" in the
e.

porous cups of which animal polyps undergo their development, and " stony mother " appears
to

indicate

borrowed

the

among

this,

When

meaning intended.

other words, from Imperato and gave

There

a proper Latin construction.

and Linnaeus applied

Linnaeus established his binomial system he
it

a generic value without adopting

no doubt that the word

same group of Zoophytes

to the

it

is

as

is,

in the

instance, Italian

first

Imperato had done.

As, however,

the term was originally used to indicate the " maternal " character of the " stone " rather than
its

porosity,

appears that the root should be referred to the Greek Trwpo?,

it

sideration of the

Linnean genus Madrepora, further

disposed

of.

Linnaeus established three genera

Corals.

The

first,

now

Tubipora

(= Tubulara,

difficulties arise

among what

which are not so

are popularly

known

as

in

its

remains the third genus, Madrepora, which, for practical purposes,
the whole of the Madreporaria or Zoantharia Sclerodermata.

from time to time subdivided the Linnean genus Madrepora,

The question then

genus has the name Madrepora been retained

must be

that, strictly speaking,

in the original

species of

Linnean genus.

species which

So

?

until,

may be

taken to include

what part of the original

can ascertain, the only just reply

is

M.

Systema Naturae

'

muricata, and this, so far as I can ascertain,
as

now

understood.

The

considered species 39,
ever, quotes Pallas's

Lamarck, and possibly

of

viz.

M.

M.

xii.

Ed.

xii.

the

name Madrepora

is

'

Systema Nature,' we

here referred

now

the only

M.

cervicornis,

M.

muricata.

Linnaeus, how-

synonym, and was probably correct in doing so j in that

appears as Tubipora in Ed. x.
is

is

the 33rd

It should be stated that Esper

Turbinaria and not to Madrepora

attention to the 10th edition of the

muricata in Ed.

by others.

infundibuliformis, as a variety of

crater as a

case the species belongs to

also

'

name has disappeared

specific

from our nomenclature and has been replaced by M. palmata, M. flabellum,

all

There

up to the present time, over

arises, for

far as I

is

none of the species now referred to Madrepora were included

comes within the genus

and M. prolifera,

Stony

Subsequent investigators have

In the twelfth edition of the

Madrepora described

easily

present restricted sense,

although by no means entirely in the original one, belongs to the Hydrocorallinae.

400 new genera have been proposed.

and

to the con-

Imper.) includes the "organ-pipe" coral and

The second, Millepora,

referred to the Alcyonaria.

stone,

i. e.

When we come

the English pronunciation of the word altered accordingly.

if

now, we turn our

If,

find that the species

to Millepora, whilst

Thus,

s. s.

named Madr.

Madrepora infundibuliformis

we take the 10th

edition as our guide,

applied to a genus of corals which were referred

by Linnaeus

to Millepora, and only on the publication of the 12th edition were the limits of the genus

How

extended so as to receive them.
Pallas

is

not certain.

In the

'

far Linnaeus

may have been

influenced by the views of

Elenchus Zoophytorum ' Pallas quotes " Systema Natm-ae,

Ed. X.," in the synonymy, and divides the genus Madrepora into 7 sections,

viz. simplices,

catinatae, conglomeratic, aggregatae, dichotomae, vegetantes,

In the

anomalae, he arranges three species,

M.

muricata,

and anomalee.

M. porites, and M. foliosa.

con-

last section,

It is interesting


GENEEAL EEMARKS.

S

to note tbat, according to the accepted classification, these three species belong to three allied
genera,

Madrepora

viz.

Linnseus's

Lamarck

s. s.,

work the three

The

for the

The genus,

limits than those at present recognized

known

Ehrenberg, in 1834, was the

Lamarck

not characteristic and

is

to him, but

first

in nomenclature,

equally

is

by him, had much narrower

as restricted

and did not include the

Trachylopora, &c., species of which were

instead of following

When

group of species represented by M. muri-

genus given by Lamarck

definition of the

the publication of the 12th Ed. of

were placed in the genus Madrepora.

Madrepora and instituted a number of new ones, he

name Madrepora

applicable to the genus Anacropora.

Porites, &c.

On

and Montipora.

in 1808 subdivided the genus

unfortunately retained the

cata.

Porites,

species in question

divisions, or subgenera, Isopora,

were referred to Astrma, Oculina,
genus

to give the

its

present limits, but

he proposed a new name Heteropora for the

genus, on account of the distinction between axial and radial corallites which leads to the
characteristic

mode

of colony-formation.

All the species which came under his notice, and

which really come within the genus, were, with one exception, correctly referred to

by

it

It appears, however, that Blainville * had in 1830 applied the name Heterojiora

Ehi'enberg.

to a genus of fossil Polyzoa

rank of a family, there

is

and

;

as the

name

is still

and has even been raised to the

in use,

no doubt that Heteropora, Ehrenberg, cannot stand.

The paper

in

which Ehrenberg proposed the name, although not published until 1834, was communicated
to the Berlin
'

Academy on March

3rd, 1831, and probably at that time the last volume of the

Dictionnaire des Sciences Naturelles

Dana and

all

'

had not come under

subsequent investigators have in

effect

his notice.

adopted the

name Madrepora

as

synonymous with Heteropora, Ehrb., and have extended the limits of Madrepora, Lamk.,
accordingly.

The question

as to the justification of the use of the generic

name Madrepora

in

its

present sense rests, then, with Lamarck, and the conclusion arrived at will depend on whether

the 10th or the 12th edition of the

'

Systema Naturte

'

is

taken as the starting-point.

I

am

aware that in the rules for Zoological Nomenclature adopted by the British Association, the
12th edition

taken as the starting-point

is

;

concerned, I see no sufficient reason for doing

but, at any rate, so far as the Zoophytes are
so.

any very accurate knowledge of Zoophytes, and
attitude towards his

name was then

retained the

name

original genus.

however, he has been followed by

institution of a
s. s.

it

is

clear that

new name would not
as a

generic

custom rather than on priority

a change

t

Of. F. J.

A

it

to

all

some

to

have

typical section of the

subsequent authors, with the

made now which would

like Holothuria,

We

involve the

must therefore regard

depends for

its

justification

on

f.

* 'Dictionnaire des Sciences Naturelles,' vol. Ix. 1830,
Bell, "

The genus

Lamarck ought not

serve any good purpose.

name which,

case.

and the type of the genus which now bears

in its present sense, but to have applied

As,

unjustifiable to adopt an

any other

in

referred to Millepora; evidently, then,

exception of Ehrenberg,

Madrepora

in the 10th edition,

me

appears to

work which would not be tolerated

Madrepora was established
that

it

Linnseus could not be expected to have

Test Case for the

Law

of

p.

381.

Priority," Ann. Mag. N. H. 1891,

vol. viii. p. 108.

B 2


;

GENERAL EEMAKKS.

4

In an enumeration of the works in -which species of the genus Madrepora
recorded

M.
'

Pallas did not describe

muricata.

varieties,

viz.

:

—

a.

ramosa,

corymbosa, and

/S.

M.

Pflanzenthiere/ figures several varieties of

M.

Lamarck, in 1816

name.
the

which appears to be

rosacea,

name M.

distinct,

7.

an incrusting

muricata and also another which he

named

9 new species of Madrepora and dropped altogether

*, described

No new

muricata, Linn., to which, indeed, he does not refer in the synonymy.

appended of 7

list is

Esper, in his

variety.

but had been overlooked until Studer revived the

recent species are added in the second edition of Lamarck's

but a

are

s. s.

who recognized only one species^ viz.
any new species, but divided M. muricata into three

necessary to begin with Linnseus,

is

it

fossil species

of

work edited by Milne-Edwards,

doubtful affinity described by Goldfuss and

Defrance.

Ehrenberg, in his

new

;

'

Red-Sea

Corals,' recognized 21 species, 13 of

which are described as

the remainder are referred, though sometimes erroneously, to species already described

He

by Lamarck.

regarded Astrcea puhinaria and A. microphthalma, Lamk., as synonyms of

Madr. palmata, Lamk., and referred Oculina
{Heteropora, Ehrb.).

It should

be understood that, although Ehrenberg's work

on the Red-Sea Corals, several species of Madrepora are included in
collected in the

the 'Astrolabe,'
Fiji, all

Madrepora

echidruea, Lamk., to the genus

it

is

nominally

which were not

Red Sea. In the account of the Zoophytes collected during the voyage of
Quoy and Gaimard enumei-ate 4 species of Madrepora from Tongatabu and

of which are referred to species already described by Lamarck.

Unfortunately the

descriptions and figures given are so imperfect that subsequent authors have foimd

impossible to identify the species

up the task

Next

;

almost

it

even Milne-Edwards, who had access to the collection, gave

as hopeless.

in order follows Dana's

work on the Zoophytes

of the

North-American Exploring

Expedition, probably the most important work on recent Corals which has yet appeared.

work contains

descriptions of

of which are described as

new

;

M.

new

the majority of the

Duchassaing, in 1850, published a

Madrepora are enumerated.

The

64 species of Madrepora which were observed by the author, 53

list

species are figured in the Atlas.

of Radiata from the Antilles in which three species of

plantaginea, Lamk., was probably included in error, and was

omitted from the author's later works.

In 1860 the 3rd volume of Milne-Edwards and Haime's

'

Coralliaires

'

appeared, in

which the species of Madrepora described by previous authors are ai-ranged and
eighteen

new

been published, and the
usefulness

is

No

species are also described.
'

Coralliaires

'

is

work of

classified

so comprehensive a character has since

therefore

stiU

a necessary handbook

;

but

its

considerably interfered with by the shortness and insufiSciency of the diagnoses

and the almost complete absence of

illustrations.

It is

probably owing to these conditions

that subsequent investigators have so frequently failed to recognize the species described

by

Lamarck and Milne-Edwards.
In 1860

also Valenciennes contributed a short

paper to the

'

Comptes Rendus' on the

* References to the systematic works are given under each species in the synonymy.

GENERAL EEMARKS.
corymbose varieties of Mff<^r-e/?om, in whicli he introduced several new names ; but as no proper
descriptions accompany them, they have been regarded as nomina nuda by subsequent authors.

Horn,

in 1861, described briefly three

new

species

been recoo-nized as distinct by his successors.

;

but only one of them,

published by Duchassaing and Michelotti in 1861, three
described, all of

which are very imperfectly characterized.

species in question are simply form-variations of already

new

So

species of

This

list

Museum

M.

tubigera,'h&s

Madrepora are

far as can be ascertained, the

known West-Indian

Verrill's contributions to the subject consist chiefly of three papers.

of species sent from the

viz.

In the account of the Zoophytes of the Antilles

species.

The

first is

a

list

of Comparative Zoology to other institutions in exchange.

includes 36 species of Madrepora, three of which are new.

In

his later

work on the

of
Polyps and Corals of the North-Pacific Exploring Expedition, 12 species are recorded, 7

new and two or three others are referred doubtfnlly to species previously
None of the new species of Madrepora described in either this or the
described by Dana.
In
still uncertain.
preceding paper are figured, and the exact position of some of them is
which are described

as

the Catalogue of Deep-Sea

Corals

contained in the

Pourtalt!s records only three species, all

Korallen,'

figures

Museum

from the West Indies.

of

Comparative Zoology,

Haeckel, in his

'

Arabische

three, possibly four, Red-Sea species of Madrepora under the name

Heteropora, Ehrb.

Briiggemann, in 1877, recorded a new species of Madrepora collected by Haeckel in the
Red Sea and also gave Mauritius as a habitat for M. gonagra, Ed. & H. but the identification
In the following year he also included M. laxa,
in the latter case is probably not correct.
;

Lamk., in a

list

name does not

of corals

from Singapore.

The

species described

by Briiggemann Tinder

this

agree with Lamarck's types, but, probably owing to the fuller description

which he supplied, subsequent determinations nearly always refer to Briiggemann's species
the
and not to the true M. laxa, Lamk. In 1879 Briiggemann published a report on
described

which is
Corals of Rodriguez, which includes a list of 11 species oi Madrepora, one of
as new.

the voyage of
Studer's account of the Stony Corals collected in the Pacific Ocean duiing
the

'

Gazelle

'

The name M. secale is
includes 33 species of Madrepora, of which 5 are new.
M. plantaginea, Dana, which has generally been held to be distinct from

also proposed for

the species described under that

name by Lamarck.

Singapore, the same author gives a

list

In a later paper (1880) on the Corals of

of 23 species, none of which are new, but figures are

given of two or three of the more obscure species.

The subgenus Isopora was proposed by

Studer in this paper.

An
feature

an excellent
important work by Klunzinger on the Red-Sea Corals appeared in 1879,
The work has a further value from the
of which consists of the photographic plates.

fact that the

author

made

and gives more
a careful study of Ehrenberg's type specimens
work includes descriptions and figures of 24 species

accurate descriptions of his species.

The

of Madrepora, of which 17 are recorded

as

new.

This work

is

excellent, but sufi-ers in

some

M.-Edwards, which, it must be admitted,
respect from a lack of acquaintance with the types of
purposes o£ identification. In 1880 the
are, in many cases, not sufficiently diagnosed for


6

GENEEAL EEMAEKS.

•

Report on the Florida Reefs by Loiiis Agassiz was published by

his son, the special interest

of which, for our present purpose, centres in the three excellent plates of

M.

cervicornis,

In the

'

M. palmata,

and M. prolifera.

Marine Fauna of Mauritius and

by Mobius, Richters, and

Seychelles,'

v.

Martens,

seven species of Madrepora are recorded by Mobius from Mauritius, which were identified by

Haacke

Fauna
'

six of the species also occiu- iu the

;

of Ceylon

Sea.

S. O. Ridley, in a paper

on the Coral

H. O. Forbes

gives, in his

two species of Madrepora.

(1883), records

Wanderings

Naturalist's

Red

in the Eastern Archipelago

(1885), a

'

of Corals collected in

list

the Keeling Islands, which were determined by Ridley and Quelch, in which two species of

Madrepora are recorded, one

of which

was described by Dana under that name

;

is

named M.

the specific

orhipora,

Dana,

name intended

var.

;

but no species

probably cribripora.

is

The 'Challenger' Report on Coral Reefs by Quelch appeared in 1886, and in it fifty
species of Madrepora are recorded, eleven of which are new.
In the same year Duncan
described a collection of Corals from the Mergui Archipelago, which included eight species
of Madrepora,

all

of which are referred to

known

species.

In 1888 Rathbun published a catalogue of the species of Madrepora in the United-States
National Museum. The-list includes 59 species in all; the type specimens of 48 of the new
species described by Dana and also 6 of Verrill's are in the Collection.
The types of the
remaining species of Madrepora described by Dana appear to have been
rare instances

(e. g.

M.

digitifera)

This collection also contains the type of

Madrepora are recorded.
the Strassburg
is

The

Museum.

first,

descriptive papers in

published in 1888,

is

a

list

secale,

which numerous species of

of the collection of Corals in

In this paper 44 species of Madrepora are enumerated, only one of

described as new, but in the case of 8 others the reference to

with considerable doubt.
Corals

M.

(= M. plantaginea, Dana, non Lamk.).

Ortmann has published two important

which

excepting in the

where the species were described from specimens already

in the collection of other institutions.

Studer

lost,

made by Haeckel

The second paper

is

new

species

is

made

devoted to a description of a collection of

in Ceylon, the types of

recorded, 5 of which are described as

knowu

which are in Jena

to science.

In Faurot's

27 species are here

;

'

Report on the Red

Sea Mission,' published in 1888, 4 species of Madrepora are recorded, 3 from the Gulf of

Aden and

1

from Kamaram Island.

In 1890 Bassett-Smith described a collection of Corals from the China Sea (Macclesfield

and Tizard Banks).

In

described as new, but a

this
fifth,

paper 30 species of Madrepora are enumerated, 4 of which are

which

is

considered new,

in this collection is the fact that all the

30 fathoms.

Fauna

of the

new

is

not named.

An

interesting feature

species were obtained at depths between 23

and
In the same year Thurston published a paper on the Fisheries and Marine

GuK

of Manaar, in which the occurrence of 3 species of Madrepora

is

recorded.

Since the systematic portion of this work was completed, a paper by Dr. Rehberg, " Ueber
neue und wenig bekannte Korallen," has come into my hands ; it was published in November


GENERAL EEMARKS.
appearance of

last shortly before the

Museum^ but

my

I did not see a copy of

it

7

second contribution on the new species in the British

until the

end of December.

on a study of the German Collections, and especially of that

in

Rehberg's work

Hamburg

is

based

but he has also

;

studied the Berlin Collection, and specimens from the collections in Kiel, Lubeck, and

A long

Bremen.

of species

list

already pointed out, such

and

I

given according to their geographical distribution, but, as

unless accompanied by a complete revision of the

lists,

new

This paper also includes descriptions of 10

are not reliable.

number

is

of other species.

I

species,

synonymy,

and notes on a

have not seen the specimens described and recorded by Rehberg,

therefore do not feel justified in fully criticizing his results, nor in including the species

my

recorded by him in

To some

synonymy, excepting in a provisional way.

names

rectification of specific

extent a

Rehberg describes a new

necessary between us.

is

species,

M. incrustans, one specimen of which is in the Berlin Museum this specimen is referable to
M. plicata, a species which I described in 1891, and I therefore presume that M. incrustans
The name M. edwardsii is proposed for M. echinata, M.-Ed. & H. (non
is a synonym.
Dana), from Luzon. I do not know if Rehberg has studied the Paris tyj)e, but I did not
;

myself note any difference between the specimens referred to Dana's species by Milne-

Edwards and the

species

which

I regard as

M.

echinata,

Dana, but there

the figure given by Milne-Edwards does not represent Dana's species.
list

(p.

33) a species

this species

referred to as "

is

from Mauritius, which T described
Again, plate

name

M.

symmetrica, n. sp.

given in the text, and I therefore cannot say

is

iv. fig.

10

is

M.

if not, it

;

name

name has

for a different species

priority,

—Palau."

if it is

No

description of

M. symmetrica

no species

new

to a

doubt that

the same as

must be regarded

spinosa, n. sp., but

Rehberg gives the name M. coronata

in the text.

I have also used the same

paper ; Rehberg's

1891

in

referred to

is little

In the geographical

is

as

a

nomen nudum.

described under that

species

from Madagascar.

from the Great-Barrier Reef, 1892

and I have therefore changed the name of M. coronata,

mihi, in the present work.

The views
no means

as to

synonymy which we derived from

identical.

I understand that Dr.

which I saw later, and I

may

a study of the Berlin Collection are

Rehberg was unable

also state that as in

to find

many of

no case did I find the original

by

the specimens
labels

on the

specimens, I had to spend considerable time in referring to the original labels and comparing

the specimens with the descriptions so as to

make

sure that the specimens which I studied

were really the types.

MOEPHOLOGY.
Skeleton.

—Duncan * has given an account of the structure

of Madrepora.

The

first

species described

with long slender proliferous branches, and
species, in

* "

which the corallum

On

pp. 181-191.

the

Hard

is

of the corallum in three species

was not determined, but

may be taken

as

is

an arborescent form

an example of quick-growiug

not thickened by a secondary deposition of carbonate of

Structures of some Species of Madrepora," Ann.

Mag. N. H. 1884,

vol.

xiv.


;

GENEEAL EEMAEKS.

8
lime, excepting in

The

older parts.

tlie

of mural tissue, and

and costate externally;

corallite-wall is very perforate

The

the costse often project as trabeculae.

wall of

young

corallites consists of only

In older corallites spinules are always present on the free edges of the

down.

one layer

provided externally with plain or finely serrate costse for some distance

is

costse,

and

these are largest round the bases of the corallites, where they form a basis of lax tissue which
is

the

stage in exogenous growth, and occasionally assists in the production of buds.

first

The formation

new

props on which a porous layer of

costse the supports of the

new

in axial corallites the costse assist in the formation of a
lateral spinules fi'om the free edges.
is

In transverse sections of

new

;

The

septal cavity, are dense.

old the

is

first,

and the

In old and

by sending out

layer of tissue

corallites of

the circles having been, in turn, outside walls

If the corallite

costse.^'

spinules act as

roof.

moderate thickness

seen to consist of " concentric circles of thin calcareous structure

radiating pillars

or

—The

:

formed in such a manner that the old wall

tissue is

fonns the floor of a chamber, and the spinose

the wall

manner

of mural tissue takes place in the following

.

.

separated

.

second, or third circles of tissue, next to the

costse are imperforate,

and as a rule the septa

Buds from

also.

a radial corallite are formed from trabeculse which arch over and form a low hood.
costse

next appear on the outer

these

arise first

cavities of the

medium

the

margin, and

bud and

may

No

" except in a very iadirect manner and through

axial corallite exists,

Budding takes place remote from the

M.

In the second species described, probably a variety of
gascar, there are

communication between the

from sclerenchyma remote from the wall of the

arise

numerous immersed

The

hood, and after elongation the septa appear

siu-face of the

as linear series of spines directed inwards.

of the dermal structures.

by

radii either spinules

corallites.

calicular

corallite."

cytherea, Dana,

from Mada-

These are closely arranged, and their walls

are well defined in longitudinal section, and well-developed trabeculse pass completely across

Between neighbouring

the polyp-cavities.

corallites

successive layers or storeys of laminte separated

On

the

connecting structures are

by rows of small,

irregular,

and short

in

pillars.

the upper surface between the immersed corallites numerous minute, broad-based, sharp-

pointed spinules occiu", arising from a perforated calcareous lamina

;

this is precisely the

condition of the successive layers below the surface, and new layers are evidently here added
in the

same manner

The
from the

on the

as

corallite-wall.

third species described
description,

material investigated, I

am

is

referred by

inclined to think

any case the species serves to
corallum

is

Duncan

to

M.

granulosa, Ed.

and from small carmine-stained fragments which

compact.

The surface

very few perforations, and

is

it

more nearly

illustrate the structure of
is

related to

and

are closer together.

but their apertures are

M.

&

H., but judging

believe to be the

secale, Studer.

here veiy dense, and consists of a stout lamina with

clothed with knobbed spinules in place of costse.

In

In

slow-growing varieties in which the

section concentric laminae, as in other cases, are separated

represent former spinules.

I

this case,

In transverse

by radially disposed bars which

however, the laminse are

much

thicker and denser

Exceedingly narrow tubes lead from the polyp-cavities to the surface,
difficult

to detect, as they are

Little need at present be added to the results of

surrounded by the bases of spines.

Duncan.

Evidently the condition of the


GENEEAL EEMARKS.
may be

surface of the corallum

9

young

corallites is fenestrated

—that

is

to say,

longitudinal thicker bands connected together by a delicate and perforate lamina

bands may be considered

costse,

as they project

beyond the intervening

development the wall increases in thickness, and the
costag

become

consists of

it

the thicker

;

In later

parts.

become spinose

costae

altogether, the surface-layer gradually

lost

In some

generally taken as a guide to the density within.

species, however, the thin wall of

still later,

;

becomes denser and

is

the

simply

lu such species the costulate and simply echinulate conditions coexist

clothed with spinules.
in different parts of the

same colony, and the denser surface of the older

corallites does

not

As growth

correspond with the primitive density of the earher-formed layers of the wall.

proceeds, however, a further deposition of carbonate of lime takes place beneath the surface of

the corallum, so that a branch which

is

quite porous near

apex

its

may become

nearly solid

towards the base.

Before leaving this subject,

term

costse in the case of

synapticulse, and

it

appears desirable to discuss shortly the application of the

The porous

Madrepora.

therefore not a theca, as

is

corallite-wall is essentially

differs

it

both in structure and origin.

thecate Madreporaria, and, indeed, in non-thecate types also,

it

is

In the

wall.

and position with the septa, and are usually

regarded as the distal extremities of septa which pass beyond the thecate wall.
theory of the origin of a theca

In

usual to apply the term

is

"costse" to the longitudinal ridges which mark the outer surface of the
majority of cases the costse correspond in number

composed of

If G. v.

KocVs

correct, and considerable evidence has already been collected

In

in its favour, the costse should morphologically be considered as the distal parts of septa.

Madrepora

the genus

the so-called costse

They bear no regular

undoubtedly do not come under

number

relation to the septa either in

occur.

Under

these circumstances

it

category.

or position, and in point of

time appear before the septa and are also present on the walls of

more than two septa

this

corallites in

which never

does not appear desirable to

continue the use of one term for two perfectly distinct structures, and in the systematic
descriptions I have endeavoured to indicate the condition of the wall

by the use of such terms

as fenestrate, striate, striato-echinulate, rugose, &c.
It is often stated as characteristic of

more prominent than the other primary
is

that condition confined to the genus.

corallites the outer directive

In other

species the

which case the

septum

columella

;

may

septa

;

but this

all

is

directive septa

(c/. p.

by no means always the

In Madrepora one frequently

broad and the other

primary septa of both axial and

bilateral

five

11) are

case,

nor

finds that in radial

primaries narrow and equal.

radial corallites are of equal breadth, in

arrangement of parts in the polyp

importance of the directive septa.
axial corallites

is

Madrepora that the

is

not indicated by the relative

Again, in some specimens the six primary septa of the

meet together

and fuse together so as to form a

in the axial fossa

in other specimens only the directive septa

become confluent.

subject to similar variations to

Montipora the relative importance of the primary septa

is

those which occur in Madrepora.

corallites are

In Anacropora the

false

In Anacropora and

prominent, and the

it is

not until the absence of an axial

corallite is observed that the generic distinction is realized.

In this genus the directive septa

branches resemble those of Madrepora so closely that

c


GENERAL REMARKS.

10
are, as in
all

more

In Montipora the septa are sometimes

Madrepora, usually broader thau the others.

some

or less rudimentary and trabeculate, but in

lamellar, and the directive septa may then be very broad or

A

in the middle line.

bilateral

arrangement of parts

species they are undoubtedly

may even become

fused together

thus as well-marked by the directive

is

septa of Anacropora and some species of Montipora as in Madrepora.
Ridley in 1884* discussed the mode of budding in Marfrepora and
Colony -formation.

—

Montipora, and considered that there

a fundamental diSerence between the two types,

is

dependent on the termiuality or non-termiuality of the
Isopora, Studer, a subgenus of Madrepora,

but that
is

it

more or

a

centrifugal,

i.

e.

new buds

arise

below the axial

undifferentiated ccenenchyma and

He compared
buddiug

in

pointed out that

In Madrepora the budding

abundant trabeculate ccenenchyma.

less

He

In both Madrepora and Montipora there

provided with several instead of one.

is

distal corallite.

not without axial corallites as bad been supposed,

is

new buds

corallite.

In Montipora the apex consists of

are added above those existing,

i.

e.

is

the

same

as in the

genus Montipora.

centripetally.

The mode

the condition to determinate or indeterminate inflorescence.

Anacropora

is

of

Ridley therefore suggests

the foundation of two subfamilies, Madreporinse and Montiporinse, with characters based on

this distinction.

centrifugal and centripetal do not appear to express accurately

The terms

the precise modes of budding to which they are applied, and

it

would probably have been

had Ridley employed the botanical terms determinate and indeterminate to express the
In foliate species of Montipora the budding is
distinction in the case of branching species.

better

centrifugal,

not centripetal, seeing that new corallites are added at the periphery.

branched specimens of Madrepora the buds arise around, and

elongate corallite forming the axis of each branch, and extending from

the apex, where

it

always projects more or

This

less.

In

are indirectly connected with an

corallite,

its

point of origin to

often spoken of as the parent

diameter than the others, and often exhibits a better-developed
usually termed the apical coraUite (" Endkelche " by the Germans), but

corallite, is usually of larger

series of septa.

It is

axial corallite seems
in surface- view,

is

much more appropriate

;

the part of

only an insignificant part of

its

it

which

is

" apical," and recognizable

whole length.

Although the types of budding indicated by Ridley form an essential distinction between

Madrepora and Montipora, the type
by

the living colony during its

manner

to those of Montipora.

iu the first instance),
tissue

new

characteristic of

growth

is

confined to branches formed

in other situations the buds are formed iu a similar

In specimens which form incrustations (and

all

are incrusting

corallites are added peripherally from an undifferentiated mass of

which projects beyond existing

in length

;

Madrepora

corallites.

and thickness, and indicate the

first

It

is

only when certain of the corallites increase

formation of branches by the development of

buds around them, that the form of budding characteristic of Madrepora comes into operation.
Frequently both types of budding take place at the same time in one colony
branch-formation, the other to marginal or basal extension.

One not

;

the one leads to

infrequently meets

* " The Classificatory Value of Growth and Budding of the Madreporidse," Ann. Mag. N. H. 1884,
vol. siii. pp.

284-21)1,

pi. xi.


GENEEAL EEMAEKS.

11

with specimens in which a colony of a younger generation forms an incrustation over the
In such cases new corallites are added from

branches of a dead colony of the same species.

a marginal mass of undifferentiated tissue until the apex of the dead branch

only

when independent growth

later,

probable that the immersed

begins,

is

the

mode

of budding changed.

which frequently occupy the

corallites

A

further point remains

mode

peculiar

which appears to

Madrepora

of budding in the genus

— there

colony termed " patrio-ramose " by Dana

word, excepting at points where the colony

on

ccenenchyma

It also

mode

As

important.

seems

between

of budding.

a result of the

—which leads to the formation of a type of
is

no ccenenchyma in the true sense of the

incrusting.

The

radial corallites are arranged

and the space between them

the branches at variable intervals,

consist of

is

me

reached, and

lines of fusion

adjoining branches are formed by the primitive and not by the specialized
to be noticed,

is

usually considered to

is

but these intervals really form part of the thickened wall of the axial

;

around which the radial corallites are developed, and the trabecular network of which

corallite

they are composed

is

not precisely comparable with the interzooidal ccenenchyma of Turbi-

naria, for example, which

is

a true secretion of interzooidal tissue,

and not of the walls of the

zooids themselves.

The Septa and

their Relation to Tentacles.

of the various species

is,

number may be regarded
less developed.

as 12, viz., 6 primaiy ones

Quelch recorded that

making 24

a third cycle of septa,

—The number of septa present in the

within certain limits, subject to considerable variation.

in

in

M.

all.

and 6 of a second

cycle,

list,

and

it

which

is

typical

usually

mirabilis, Quelch, certain of the corallites possess

This has hitherto been the only instance on record

of the occurrence of a third cycle of septa in the genus Madrepora.

add 2 or 3 other species to the

corallites

The

is

I

am now

enabled to

of interest to note that the third cycle

is

present

usually, but not invariably, only in certain of the corallites which are nearly or completely
It appears reasonable to expect the polyps in

immersed.

instead of 12 mesenteries,

and

it

such cases to be provided with 24

would be interesting to ascertain the precise manner

in

which the additional mesenteries are formed.

The

number

septa are generally most fully developed in the axial corallites, but even there the
in

many

In other cases 2 or 4 of the second cycle

species never exceeds 6.

may

be present, but by far the greater number of species have the second as well as the primary

The members of

cycle complete in the axial corallites.

or subequal, or
others.

more

rarely the directive septa

In the radial

corallites there

is

may

the primary series

may be

equal

be more prominent than any of the

almost every variation from the apparent absence of

septa to a development as complete as that of the axial corallites or in certain cases even

more complete.

The usual order of development

the remaining members of the primary

Of

the second cycle one

the outer directive

is

is

that the directive septa appear

reached.

A

;

first,

then

but not always simultaneously.

frequently note the presence of two septa

—before the others appear

before the final stage
occurs,

may

is

series are added, usually

— one on

each side of

may be

three or four

and in other cases there

third cycle of septa, in the rare instances in which

it

usually confined to radial corallites, but in one species occurs in the axial corallites.

Whilst in axial

corallites the

most usual arrangement

is

for the

primary septa to be subequal,

c2


GENERAL REMARKS.

12

in the radial corallites the directive septa are most frequently better developed, either stouter

In certain groups of species, however, the outer

or broader, than the other primaries.
directive
it

is

septum

is

more important than the inner

form characters of value

now proposed

several of the subdivisions

and in case only one septum

the species were

present

In

for the classification of species.

associated together on general

first

grounds, and without any regard to the condition of the septa

and

;

yet, in

the relative importance of the septa will be found nearly constant— e.

g.,

In certain species there appear to be two types of radial

Trachylopora.

is

Apparently the number, but more particularly the

invariably the outer directive.

relative importance, of the septa

;

almost every case,

Lepidocyatlms and

In the

corallites.

usual more or less prominent one the septa are naturally least developed in the young buds

near the apex, and there

normal condition

is

a gradual increase in importance in older corallites until the

attained.

is

It

may

be that the condition of the septa

subject to

is

little

variation whether the corallite be prominent or immersed, situated near the apex or near the

lu certain species, however, some of the immersed corallites of the

base of the corallum.

upper surface of the corallum have quite a different development of the septa to the

Sometimes the septa in these immersed

prominent ones.

or better developed

always the case

is

—

clear

axial corallites, or,

from the fact that such

corallites

more septa than the

are

M.

In Dana's figures of the axial polyps of

In Agassiz's

shown of equal length.
prolifera are figured,

and

M.

importance of the directive septa

cribripora and

Florida Reefs

'

is

'

M.

drawn

me

M.

and

far as I

am

aware,

state the usual condition of the tentacles of the radial polyps in

specimens which came under his notice.

There seems reason

generally of equal size, from the fact that he figures
case of

M.

With regard

aspera.

one of the tentacles
directive septum,

is

much

which

embryo of Astroides.
and Barrier-Reef

:

M.

is itself

Madrepora

as such without

comment

is

long and

having thin labellate

flexible.

calicles

;

in the

prostrata he calls special attention to the fact that
is

situated over the outer

That

this

experience will be rendered clear from the following

— " Among the species of Madrepora there are some in which one of

of the polyps

in the living

broader and stouter than the remaining septa.
his

which

Dana does not anywhere

to suppose that they were

longer than the others, and that this

arrangement was not usual in
quotation

to

them

This

septa.

species the tentacles of the axial polyps are always of equal length, even in species in

So

not

and

that in his experience of the Torres- Straits

this does not hold good for the radial polyps.

is

cervicornis

and narrower secondary

in accordance with Lacaze-Duthiers's figures of the

Mr. Saville-Kent informs

them

aspera the tentacles

alternately large

the polyps of

in both cases the tentacles are

small, corresponding evidently to the broader primary

arrangement

may

from that characteristic of the upper surface.

distinct

in the case of the axial corallites the relative

usually associated with a variation in the size of the tentacles situated over

known.

not

is

on the under surface of prostrate or vasiform specimens

show a condition of the septa quite

is

corallites often contain

this

on the other hand, may show an almost complete absence of septa.

Similarly, prominent

Whether

more numerous

corallites are

a condition which might be attributed to age; but that

the tentacles

This was observed in a cespitose species {M. prostrata)

but whether

it

belongs or not to

all

the horizontally-growing


GENEEAL EEMAEKS.
with similar ealicles remains

species

be determined.

13

to

This character

may

hereafter

lead to a subdivision of the genus and a separation of the species having labellate ealicles

{M. conigera,

cytherea, spicifera,

efflorescens,

subulata, twrbinata,

and convexa)

possessing one tentacle which

is

specimens that the outer directive septum
inner

;

M.

cervicornis are represented as

louger and stouter than the others

;

the arrangement of the

from an examination of a number of

I find

only represented in the axial corallites.

is

surculosa, millepora, prostrata,

group."

as a distinct

Agassiz's 'Florida Reefs' the radial polyps of

Ill

septa

Injacinthus,

sometimes, but not always, broader than the

is

both directives are broader than any of the other septa.

In the case of M. prolifera

Agassiz's figure shows the radial as well as the axial polyps to have the tentacles arranged in

correct

minute

any case

in

:

which characterizes nearly

detail

lithographic

is

not a good one, and apparently not drawn with the same view to

is

it

questionable whether this figure

It appears

two cycles, alternately large and small.

representations of

the other figures

all

—which

are, indeed, the best

Madreporaria which have come under

my

My

notice.

doubts as to the accuracy of the figure in question are further based on the following
considerations
(2)

:

—

(1) I

do not consider

M.

prolifera specifically distinct

from M.

cervicornis

;

the septa of the radial corallites have the same relative importance and are subject to the

same variations

as in

M.

cervicornis

(3)

;

there

apparently no other recorded instance of

is

the alternation of long and small tentacles in the radial polyps of any species belonging to

the genus.
I conclude,

from the general remarks of Klunzinger on the structure of Madrepora, that

he regards the presence of an elongate tentacle in the radial polyps as a usual condition

usually better developed than the inner ("

is

he refers to the association of this condition

He

bilaterally.

and

After calling attention to the fact that the outer

he does not mention any other arrangement.
part of the wall (" Riickenwand ")

;

Avith

two broad

septa,

Bauchwand "),

which divide the

corallite

then goes on to state that, associated with this condition, we find one of the

tentacles corresponding to one of the principal septa broader than the others, but that this

In the descriptions of the Red-Sea species he only

only occurs in such bilateral corallites.
refers twice to the occurrence of

an elongate tentacle,

viz. in

M. corymbosa and M.

cytherea.

In the former instance he refers to the colour of the long tentacle in a manner which appears
to indicate that in his opinion such a long tentacle
species of

Mr. Saville-Kent has kindly shown me proof copies of the

Madrepora.

illustrating his forthcoming

work on

figures of the radial polyps of

tentacle

is

shown.

M.

'

am

that he paid
its

more

of

M.

hebes

;

M.

from Dana's

and in each case an elongate
list

of species for which the

All the species which are

known

to have one

cervicornis, belong to the group indicated by
list is

probably to be accounted for by the fact

attention to habit than to the form of the corallites in assigning the species

position in his classification.

labellate

liebes,

aware, completes the

tentacle larger than the others, excepting

The absence

M.

prostrata and

This, so far as I

plates

The Great Barrier Reef of Australia,' which contain

condition of the tentacles has been recorded.

Dana.

of general occurrence amongst the

is

but the term

is

The

used in a

radial corallites of

much wider

M.

hebes are not, strictly speaking,

sense in Dana's work than in the present