Section IV., 1802. [ 39 ] . Trans?, lioy. Soo. Canada.

111. — On iJic Corah nnd ConiUifor in Types of Pahtozoh Sti<da.

JJy E. J. CliAi'MAN, rh.l)., LL.J)., rroffssor ol' Miiieialof»'y aiuKicology iu the

University ol" Toronto.

(Kea.l June Iwt, Ivij.)

Among the Ibssil I'orms ol" post-pahi-ozoic diite refi-rred to the Anthuzoa or Corals
proper, very tew, it" any, occupy a debatable position. Many ol" the commonly admitted
paheozoic corals, on the other hand, are of more or less doubtlul character, and have thus
been placed by some authorities under other subdivisions. Whilst admitting their uncer-
tain atlinities, however, it seems impossible to separate these doubtful types on any really
satisfactory grounds from the more distinctly coralloidal forms, into which, indeed, they
appear to merge by almost insensible transitions. In this paper, therefore, the term
" coral " will include all the generally recognized coralloidal types of paheozoic
occurrence.

The great division of the CcKiiENTEKAi A, to which the corals belong, is usually sub-
divided into three leading classes : — Hydrozoa, Anthozoa, and Ctenophora. The latter
class, represented by a small number of pelagi'-, ciliated types, is an entirely aberrant
group, and is without any known fossil representatives. Living Hydrozoa are distin-
guished from existing Anthozoa by no very certain characters— many of the supposed
distinctions, at one time regarded as characteristic, having been shown of late years to
be more or less indefinite ; and it is extremely probable, as suggested by the late Professor
Agassiz, that the so-called " tabulated corals " should be referred, really, to this class —
the Hydrozoic Millepores forming the connecting link. These tabulated corals, again,
offer in many cases a complete transition into the tabulated Rugosa,' the assixmed tetra-
merous character of the latter beinii' iu many instances, as shown farther on, either with-
out foundation or entirely unrecognizable. It would seem advisable, therefore, in place
oi' three, to adopt //ce leading subdivisions in the classification of the Cadenterates gener-
ally, as in the annexed tabular synopsis :

A. — Without natatory cilia.

AJ — Stomach cavity completely identical with body cavity :

(i). — Without stony corallum :

Ct.ass I. Hydrozoa.
(«). — With calcareous (typically tabulated) corallum :

'As exemplified, for exainpli^, by the following scries: — FarnnieH—('i>lumnnna—Amplixuf — /uphr' ntis —
( 'yiithophiillum.

40 E. .1. CHAPMAN ON TIIH

CljAt-'S II. Hijdrucoralhi.
A.- — St'UiiacJi partially separated from body-cavity.
(t ) — Oro-aual orifice with eight fringed tentacles :

CiiASS III. Alcyimaria or Crossoro^ratla.

(ii) — Oro-iinal Drilice with luimerous simple tentacles. C'unillum e^ss^'lltially
iioii-tabulated, but with distinct septa ;

Cj>ASS IV. Zonntltaria or Anlhocordlht.
B. — AV4th natatory lilia :

CJiAS8 V. Ctenoplioni.

The present commuuicatiou refers essentially to the second of the above named
classes — that of the Hydrocorali-.\. This includes the Hydro-CoralJimc of Moseley,
together with the so-called TabuJata and Riigosa of other classifications. In the present
state of our knowledge it is necessarily to some extent a group of convenience, connecting
the Hi/dro:t>u with the typical corals. The Tubu/ata in many classifications are widely
separated from the Riigosa, and placed with the ordinary " Hexamerous Corals :" although,
from the absence or rudimentary character of septa in many of these forms it is not pos-
sible to tell whether the tentacles of the living animal were hexamerous or otherwise.
The Rugom are also for the greater part essentially tabulated forms ; and although,
commonly separated from the Tabulata under the name of Tetracoralla, the actual number,
of septa iu many cases is either indeterminable or exceedingly variabb'. As examples of
variation iu the number of septa in both the Tabulata and Rugosa, the genera Sli/lina,
Lamarck, Sti/loarnia, Edwards and Haime ; Helero/ihylfia, McCoy ; Diincanella. Nicholson,
and many others, may be cited. The descriptions of many Canadian species by the late
Mr. Billings, a most minute and trustworthy observer, also lubstautiate this point, and
prove that, although very convenient on paper, the distinction (except in certain well
marked cases) is virtually of impossible application. And again, we have no certainty
that the number of septa or septal markings really indicate the number of the tentacles
possessed by the living animal. In the living Mi/le/)ora, for example, the researches of
Moseley have shown the presence of eight tentacles, as in the Alcyonarians. If therefore,
as commonly assumed, the fossil HelioUtes be regarded as a closely related type, its twelve
septa or ' pseudo-septa '" are entirely misleading.

Many of the forms placed under this subdivision in the present synopsis — espei ially
those of the three first sections given below — may very probably belong to the Ahyonaria
or to the Bri/ozoa; but this view is entirely conjectural, and cannot at present be either
l>roved or disproved On the other hand, the strikingly tabulated structure, so charai-
teristic of the great majority of these forms, serves to unite them conveniently, and, in
the absence of negative evidence, naturally also, into a common oroup.

The class HviiROCOllAT.LA, as here adopted, may therefore be defined as follows : —
Hydrozoa or closely allied types with calcareous corallum. The cells of the latter eithor

CORALS AND CORALLIFOL'M TYPES. 41

(iutirely destitute of iuterual structures, or coutiiiiiiug in the great majority ol" oases : (1)
well developed tabula-, with or without radiating septa ; or (2), vesicular tissue with or
without tabuhe and septa ; or (3) a distinitly hexamerous or tetranierous system ol" septa ;
or (4) iudicatious of bilateral symmetry.

The class, as thus defined, may be subdivided provisionally, and especially for deter-
minative purposes, into eight leading sections, as in the following scheme :

Corallite cell entirely empty, i.e., witnout any internal structures :

v^ 1. Vacuata (Type form, Aiilopora.)
Septa absent or quite rudimentary ; tabuhe well developed :

§ 2. Tabtti:ata (Type form, Farosi/ts.)
Corallites united by a tubular or cellular c(i>nenchyme.

s^ 3. Cellulata (Type form, Fislulipora.)
Tabuhe and septa both present.

§ 4. Tabulo-Stellata (Type forms, Amplex/is, Zuphrenlis.)

TabuliB central only, surrounded by area of vesicular tissue. Septa well
developed :

§ 5. Vesicut.o-Stet>t,ata (Type forms, CyalhnphyUum, Lonsdalia.)

Tabula- entirely replaced by irregular vesicular tissue :

k 6. Vesiculosa (Type form, Cipiipfii/t/um.)

Cell provided with an operculum composed of a single valve or of sevt-ral valves :

§ 7. 0PERCUr..4.TA (Type form, Calceola.)
Cup or cell containing septa only ; tabula> absent :

§ 8. Inteqri-Stellata (Type form, Pelraia.)

§ 1. Vacuata : — The representatives of this section, distinguished essentially by their
tubular cells being entirely destitute of internal structures, an- of very doubtful position.
Very probably, they should be referred to the Alcyonaria or placed near the Serlulariaus
under the Hydro-medusa- : but nothing delinite, it is obvious, i-an be determined on this
point, and as the forms in question are of not uncommon occurrence in pakeozoic strata,
it has been thought advisable to retain them here, at least provisionally. They form but
one family, that of the Anlojwridir, (-ontainiug the following gimera :

Fam. Aiiloporida; : — Corallum composed of short, tubiiiar or funnel-shaped empty
cells, united in branching groups :

Sec. IV.. 1892. (I.

42 B. J. CHAPMAN ON THE

Aulopora, Goldf., Sil, Dev., C!aib.
Reptellu, Rolle, Sil., Dev.
Cladoconus, McCoy, Carb.

§ 2. Tabulata.

The corals of this section are distinguished by the presence of numerous transverse
tabular in their component cells, coupled with the absence of r idiating' septa, or the rudi-
mentary, imperfect character of these when indistinctly present. They have been
separated widely in many systems, and have been allotted partly to the Alci/onaria, partly
to the Bryozou, and in part also to the Riigoi^it generally. Whilst thu.s manifestly occupy-
iug an uncertain position, the two fundamental characters given above — namely, the
presenci' of tabuLe, with absence or merely indications of septa — are sufficiently definite
to warrant their retention in a common group, and to show at the same time their
relations to the coral forms of section -1. in which botli tabula' and well developed septa
are present. Th(! more typical i-epresentatives, including all the better known genera,
may be arranged under six families, as in the following synopsis:

Fam. 1. ClKclelidie : — No cceuenchyme Corallites, capilliform, or of small diameter,

C/uctetes, Fisch. (including MonlicuUpora and Slenopora), Lr.

Sil. to Trias.
Tetnidium, Satford. Lr. Sil.

Fam. '1. Bedianoidithc : — No cteneuohyme. C'orallites comparatively large, polygonal,
with imperforate walls.

Beaumontin, E. & H., Carb.

Fam. o. Favuaiddte : — No coenenchyme. Corallites, polygonal, round or crescented at
surface, with perforated walls.

G-roup A. Plant. — Tabulte horizontal or inclined :

Favosites, Lmk., Sil., Dev., Carb.
Columnipora, Nich., Lr., Sil.
Alveolites, Lmk., Sil., Dev.
Pachypora, Linds., Sil.
Nodulipma, Linds., Sil.

Group B. Timidi. — Tabula; convex or more or less vesicular :

Emmonsia, E. & H., Sil., Dev., Carb.
ML'helinia, de Kon., Dev., Carb.

Fam. 4. HalysUidie : — Corallites lubular, imperforate, connected only at opposite
sides, typically in chain-like aggregations. Tabuhe, horizontal : .:

Halysiles, Fisch., Sil,
Thecostegiles, E. & H., Dev.

CORALS AND CORALLIFOEM TYPES. 48

Fam. '). S//ringo/>orid(C : — Corallites tubular, imperforate, coniiocted by short trnns-
vprse processes or by lateral expansions. Tabuhe, funnel-shaped :

Sf/ri)/i:^oporn, Gold., ]")ev., Carb.
Hdinifo/iliijllum, Bill., Dev.

§ ^. CELTiULATA.

Tliis settion. like the last, comprises a series of coralloidal forms of very doubtful
l^osition, but conneeted with the Hydro-Coralla irenerally by the presence of numerous
tabuhe; and in one of the two families, into whi(^h they are subdivided, by distinct
septa, or " ps<'udo-septa." They are made up of capilliform or narrow tubular coralli-
ti's, traversed by septa, and connected by ccenenchyme, in itself composed of minute,
tabulated lubes, the whole .somewhat resembling the surface of a sponge in which the
( orallites represent the oscula. Some have been given to the Bryozon. Others, from their
supposed relations to the modern Heliopwa, have been referred to the Alcyonaria ; but
it seems better to leave them among the HyuRO-CoRAT,iiA until more certain evidence
is obtained of their true affinities. They may be subdivided into two families, with
gejiera as follows :

Fam. 1. Fistuliporidce : — Corallum compound, composed of minute corallites with
surrounding capilliform cconenchyme ; both tabulated, but without septa.

Fistulipora, McCoy, Sil., Dev.

Cullopora, Hall (scarcely differing from Fistulipnra), Sil., Dev.

Fam. 2. Heliolitidce : — Corallum compound, composed of small corallites separated by
a < ellular or finely tubular ca-nenchyme ; both tabulated ; the corallites showing twelve
short septa or pseudo-septa around their inner margin.

Heliolites, Dana, Sil., Dev.
Lijellia, E. & H., Sil.
Plasmopora, E. & H., Sil.
Thecia, E. & H , Sil.

§ 4. Tabulo-Stellata.

The corals of this section are characterized by the presence of both tabula3 and septa.
The tabulce extend in typical examples entirely across the corallite-cell, but indications
of an outer area of vesicular tissue are occasionally observable. The septa are marginal
or short in some cases, although always distinctly developed. In other cases they extend
into the centre of the cell, and form by their union a twisted pseudo-columella. The
typical representatives form three families, as in the annexed tabular distribution :

Fam. 1. FavistelHdte : — Corallum compound, with hexagonal or polygonal corallites
in close juxtaposition, much resembling Favosites ; but walls imperforate, and distinct
septa (short or long) always present.

Columnaria, Groldf., Lr. Sil.

Favistella, Dana, Sil. •

44 E. .1. CHAPMAN ON THE

Fam. 2. AmphrxkhB : — Corallitos tubular or reed-liko, typically with short or marginal
septa and extended tabulic.

Amplexus, Sowerby, Sil., Dev., Carb.

Calo/)Iu/l/um, Daua (int-ludiug Cynlho/ilii/l/oides), Sil.

ChonopJu/l/iim, E. & H., Sil., Dev.

Fam. 3. Zaphrentidic .— Corallum simple, turbinate or horn-like, with well developed
tabula? and septa.

G-roup A. — Connciilali (simple, horn-like forms) :

'/Aipln-entU. Rat', (including Anisophi/l/um and PenlaphyUnm), Up. Sil.,

Dev., Carb.
Ptj/co]ilii/llinn, E. & H., Sil., Dev.
Slreptelasma, Hall. Under this genus, the forms o{ Zaplirentix with marked

pseudo-columella may be appropriately placed. Sil.

G-roup B. — Columniferi (with distinct columella) :

Loplioph/jlliim, E. & 11., Dev., Carb.

Group ('. — Cniciferi (primary septa forming a distinct rcctangxilar cross) ;
Phri/ganophi/lfum, dc Kon., Carb.

Group D. — Bi-formes (septa showing a twofold mode of arrangement) :

MenophyUvm E. & H., Carb.

§ 5. Vesiculo-Stellata.

This section is more or less closely connected with section 4, but is distinguished
especially by the contracted dimensions of the tabula*, and the replacement of the outer
portion of these by an area of vesicular tissue. Hitherto, many genera of these corals
have been made to incli\de both simple and compound forms, as well as types of very
dissimilar couliguration ; but in the present distribution I have ventured upon an inno-
vation in this respect — believing, from the examination of numerous examples, that a
generic separation of the compound tesselated forms on the one hand, and the sub-
aggregated, reed-like and cylindrical types on the other, from the essentially simple forms,
is warranted on natural . .ounds, and is conducive to a more ready determination of
genera, as well as to clearer and sharper definitions.

In the present distribution these vesiculo-stellate corals are arranged under four
families, comprising : (1.) Cyat'ioplri/llida; distinguished by the absence of a columella
and by the absence of a distinctly cruciform arrangement of septa; (2.) Lonsd(iltd(B, in
which a distinct columella is present ; (3.) Staurida, characterized by the primary septa
forming a distinct rectangular cross ; and (4.) Halliadie, with distinctly twofold arran-
gement of septa.

a

■ CORALS AND (^OKALFilFORM TYPES. 48

Fam 1. Cyathnphi/lltdrr.

Group A.— Tesselalt : Compound vesioiilo-stellatc Ci/atlio/th!/lli(l(C composed of closely
uuitod corallites, forming mosaic-liko, tesselatod or asteroidal groups.

Ci/athogonium, E. J. C. ( = tesselati'd forms of Ci/ulliojilii/llitm ; also

Str- abodes and Acervularia), Sil., Dev., Carb.
HeHos;onium, E. J. C. ( = tesselated forms of Helv-pfiyl/um), Sil., Dev.
Phillipsaslrcsa, E. & H. (including Puchyphijlium), Dev., Carb.

Group B.—Aritndinacei : Vesiculo-stellato CyathophijUidfC, composed of roed-liki- or
tubular corallites, either .simple or in partially connected groups.

Cannophyllum, E. J. C ( = reed-like forms of Ci/alhophyUum), Dev., Carb.
Dipliy/iJiyllum, Lonsd. (including Slylaslrea), Sil., Carb.
Eridophyllum, E. k H., Sil., Dev:

Group C.—Corntculati: Simple vesiculo-stellate C//«Mf#/y//«V^'' of more or less conical
or horn-like form.

Cyaihophylhtm, Goldf. (including Dicophylhim and Compophylhtm), Sil.,

Dev., Carb.
Heliopliyllum (r .ig AntntliopltyUmii), Sil., Dev.

Blothrophyll .il.. Dev.

Chsiophyliwh, Dana, Sil, Dev., Carb.
{Pycnophylltm, Linds., Sil. 0
Strephodes, McCoy, Sil., Dev., Carb.

Fam. 2. Lonsdalidce.

Group A. — Aggregali: Compound forms of IfOrts</a/tcfe.

Lonsdalia, McCoy (including StyHdophylliim), Carb.

Lithostrolion (including Koninckophylhim in part, Petala.ris. Stylaxis, etc.),
Carb.

Group B. — Turbinati : Simple, more or less turbinated or horn-like forms of Lons-
dalid<v :

AxophyUum. E. & H., Carb.

Koninckophyllum (f) Nich. ( = simple forms oi LithoHrotion), Carb.

Fam. -3. Slaurida- : — Primary septa forming a rectangular cross :

Group A. — TesselaH: Compound tesselated forms of Staurida :

Stauria, E. & H., Sil.
Group B. — Turbinnli : Simple forms of Slauridcc :

Omphyma, Eaf., Sil.

46 E. J. CHAPMAN ON THE

Fam. 4. HalliHur. — Septa distinctly twofold in their arrangement :
Hallia, E. Ik H. (including AnlacophyUum) , Sil., Dev.

§ 6. Vesiculosa.

This section includes but one family, that of the Cifstiphijllidw, distinguished by the
corallite cell being entirely filled with vesicular tissui-. Radiating septa absent or quite
rudimentary :

Group A. — Ag'^reguli : Septa absent or quite rudimentary.

CystiiihyUoides (^aggregated forms (jf Cijalipliyllum, lypiHcd by C. iioiin:-
galHm, Billings), Dev.

Group B. — CornicuJiiti : Septa absent or rudimentary.
Cysft/ihylhim, Lons., Sil., Dev.

§ 7. Ol'EROTJTi.iTA.

The forms of this section comprise a small number of peculiar types in which the
corallite cell is furnished with a cover or operculum composed either of a single valve cr
of several valves. Radiating septa are mostly rudimentary, but are well developed in
one genus. Provisionally, the representatives of the section may be classed in a single
family, with subdivisions as shown below :

Fam CaheoHdd'.

Group A. — Arundimicei : Corallum made up of cylindrical or reed-like corallites.
Septa rudimentary.

Flelcheria, E. & H., Sil.
Rhizopora, de Kon., Garb.

Group B. — Univalvati: Operculum composed of a single valve. Septa rudimentary.

Calceola, Lam., Dev
Rhizophylhim, Linds., Sil.

Group C. — Tetravafvati : Operculum composed of four valves. Septa well developed.
Goniophyl/um, E. & H., Sil.

§ 8. Integri-Stell.4.ta.

In the representatives of this section, the internal tabulae, so characteristic of pahe-
ozoic corals generally, and the vesicular tissues which frequently accompany or replace
the tabuke, are practically unknown. Radiated septa, on the other hand, are well deve-
loped. The known genera present collectively, two distinct types of configuration— a
disciform, and a coniform type, respectively. A central columella is present in some
representatives of the coniform type, and is absent in others. Hence the Integri-SteUata
admit of a separation into three well characterized families, as in the following dis-
tribution : . ' . - -

CORALS AND CORALLlFOltM TYPES. 47

Fain. 1. Cyclinicld: : — Corallum simple, disciform.

Group A. — S/miferi: Septa represented by longitudiual rows ol' short spines.
Aainthocyclifs, Dyb., Sil.

Group B. — AUernali: Septa, alternating in length, regularly arranged.
Palmocyclus, E. & H., Sil.

Group C. — Sulcati: Cup with one or more septal furrows, towards which some of
the septa commonly incline.

Mitroci/cltis, Meek, Dev.
Campo/)hi/l/iim, E. & H., Dev.
Bari/iihyllam, M. Ik B.., iJev.
Hadrophi/llnm, E. & H., Dev.

Fam. 2. Pe/raiefe .--Corallum simple, conical, straight or curved, without columella.

Pelraia, Munst., Sil., Dev., Carb.
PolyccBHn, King., Pala'ozoic.
Kenophyllnm, Dyb., Sil.

Fam. 3. Cyatkaxonidic : — Corallum coniform, simple, a central columella present.

Cyitthdxoniit, Mich., Sil., Dev., Carb.
Duncanel/a, Nichol., Sil.
Duncania, de Kon., Carb.

48

K J. CHAPirjAN, ON THE CORALS, &c.

Ketrospect.

A. Subdivisions of tiie C'iklbxtbeata :

I. Hyilrozoa.
II. Hydrocoralla.

III. Crossocoralla.

IV. Anthocoralla.
V. Ctenophora.

B. Subdivisions of the Hydrocoralla :

? 1. Vacuata :
Fain. Aidopoiid:i (i . ij. Auloi)ora, Reptella, etc.)

i -'. Taeilaia :

Fam. ('hivtetidn: {c. <j. Cluetetes, Tetradiuai.

Fam. Beaumoiilidx (f. g. Beanmontia).

Fam. Fiiro!iilid:v :
Group I. I'lani (i: g. Favosites, -Alveolites, etc )
Group 2. 'J'uiiiidi (c. (j. Miclielinia, Emmonsia).

Fam. Ituhjsnlhhric. g. Halysite.s, etc.)

Fam. !<ijiirigiip(indx (<'. g. Syriiigopora, Haiuieo-
phyllum).
'i 3. Cellulata :

Fam. FiMvUpui-idx (t. g. Fistulipora, Callopora).

Fam. Hi iioli(id:r (e. g. Heliolites, Tiiecia, etc)
(! 4. Taiu i.o-Ste..lata :

Fam. Favisldlid ;i: («,. y. Columnaria, FavistoUa).

Fam. Ampltxidx{i.g. Amplexus.Calopiiyllum.elc.)

Fam. Zoplirerttid.i :

Group 1. ('"imcuhtli (e.g. Zai)lirenti8, Streptelas-
ma).

Group 2. Ciilvmnife ri (c. g. Lophopliyiluin).

Group 3. ('rucijcn (i: g. Phryganopliylluni)

Group 4. Biformcg {e. g. Menophyllum).

? r>. Vesiculo-Stbllata :
Fam. Cyathopliyllidif :

<iroup 1. TtfU'kiti (<■. y. C'yatliogonium, PliillipH-

astnea, Heliogonium).
Group".'. Annidifian! r. g. Cannopliylluni, l>i-

pliyphyllnm, Eridopliyllum).
Group)!, ('oniic'ilati (•■ g. C'yathopliyllum, Ile-

liophylluni, Clisiopliylhim, etc.).

Fam. Lonsdalid:i :

(jroup 1. Aggiegali (e. g. Lonsdalia, Litliostro-
tion).

Group 2. Turhitiali (f. ;/. Axopliyllum, etc.)
Fam. Staurida: :

Group 1. 'I\mluii (i . g. Stauria).

Group 2. Tuib'mnll {i.g. Onipliynia).
Fam. II'dHid.i ((>. g. Ilaliia).

^ (i. Vesiculosa :
Fam. ( gHiphyllkhe:

Group 1. Aggngati [e. g. ("ystipliylloidrs, etc.)
Group 2. O'liiicvkiti (f. ;/. Cystipliyllum).

? 7. Ol'EEClI.Al A :

Fam. CuJcc'jUdx :
Group 1. Amndiiiacei (c. g. Fletclieria, Rliizo-
pora).

Group 2. Unii-alntli {<-. g. Calceola, Rliizopliy'-

lum).
Group ". Tilraralvaii (c. g. Gouiopliylluni).

^ 8. Istegri-Stellata :
I'am. i'ljcliiiidii: :
Group 1. ^^piniffii (f. g. Acanthocydus).
Group 2. Alttmati {t\ g. Palicocyclus).
Groups. Siilcad (c ;/. Microcyclus, ("ami)0pliyl-
lum, etc.)
Fam. Fitraidx {,. g. Petraia, Polycsjelia, etc.)
Fam. Cyatlioxonidx {e. g. Cyatliaxonia, Duncan-
ella, Duncania.)