全 文 ：　　1997-6-2收到。
论四射珊瑚征珊瑚属 Charactophyllum
Simpson ,1900
王训练 　陈建强 　王鸿祯
(中国地质大学　北京　100083)
　　提 要　对 Charactophyllum 作了全面的回顾整理 , 记述了本属的骨骼构造 , 在此基础上讨
论了其定义范围和时空分布及有关的分类问题。 Charactophyllum 的隔壁为板状隔壁而非包珊
瑚型隔壁 ,由波状弯曲的单列单晶榍组成 , 单晶榍整体呈半扇形排列而非扇形排列 ,隔壁光滑或
发育微弱的榍凸(carinae)而无典型的横耙状榍凸(yardarm carinae),鳞板半扇形排列而不是扇形
排列 。本属分布仅限于北美和华北的晚泥盆世早期。 Charactophy llum 隔壁中单晶榍由放射状
排列的不规则短轴晶柱和晶针组成 ,横列构造鳞板和横板均由晶板型层状骨骼组成。本属骨骼
微细构造的重要特征具有奥陶-志留纪与石炭-二叠纪四射珊瑚之间的过渡性质。表明四射珊瑚
骨骼微细构造演化趋向稳定 ,是追索演化的重要线索 ,在较高级别的分类中具有重要意义。
　　关键词　Charactophy llum 　四射珊瑚　骨骼构造　分类　泥盆纪
ON Charactophyllum Simpson ,1900
(DEVONIAN RUGOSE CORAL)
Wang Xun-lian ,Chen Jian-qiang and Wang Hong-zhen
(China U niversity of Geosciences , Beij ing 100083 , China)
Abstract　This paper gives a comprehensive review o f Charactophy llum , with description of
its skeletal macrostructures and microstructures and discussion on its diagnosis , scope , geological and
geog raphical distribution as w ell as its tax onomic position.The septa of this genus are laminar instead
of amplexoid , composed of sinuous uniseriate monacanthine trabeculae arranged in inwards and up-
w ards directed half-fans , and are smooth to faintly carinate but without typical yardarm carinae.The
dissepiments are globose and arranged in many row s to form half-fans.This genus is confined to early
Late Devonian (Frasnian) in age and to Nor th America and North China in distribution.The
monacanthine trabeculae in the septa consist of radially arranged irregular brachycolumns and needles ,
w hile the lateral skeletons , including tabulae and dissepiments , are composed of slabs instead of typical
flakes.The skeletal microstructures of Charactophyllum are of a transitional nature between Early
Palaeozoic and Carboniferous-Permian rugose co rals , indicating stable evolutionary trend of skele tal
第 37卷第 1期
1998 年 1 月
古　生　物　学　报
ACTA PALAEON TOLOGICA SINICA
Vol.37 , No.1
Jan., 1998
microstructure of rugose corals , w hich is a guiding clue of impo rtant taxonomic significance to tracing
the evolution of higher groups.
Key words:　 Charactophy llum , rugose co ral , skeletal structures , taxonomy , Devonian
Charactophy llum has been w idely used for Devonian rugose co rals f rom North America ,
Europe and Asia , but different kinds of rugose co rals have been referred to this genus due to the
inadequate know ledge of the type species and divergent view s on definition of the genus.Pedder
(1982)and M cLean(1993)made a detailed review of Charactophyllum based mainly on illus-
trations f rom Smith (1945),Wang (1950)and Watkins (1959)of the type species Cam-
pophyl lum nanum .Wang Xunlian(1987)described abundant Charactophyllum f rom the up-
per part of the Frasnian Low er Xipingshan Fo rmation in Zhusilenghaierhan area , Nei Mongol
(Tex t-fig.1).This paper attempts to give a comprehensive review of Charactophyl lum , with
descript ion of its skeletal macrost ructures and microst ructures and discussion on i ts diagnosis ,
scope , geolog ical and geog raphical dist ribution as w ell as it s taxonomic position.
Text-fig.1　Map show ing collection localit y of Charactophyl lum
1　Systematic Palaeontology
All specimens described here were collected from the upper part of the Frasnian Low er
Xipingshan Formation in Zhusilenghaierhan area ,Nei Mongol.
The following abbreviations are used in the present paper for measurements of rugose
23第 1 期 王训练等:论四射珊瑚征珊瑚属 Charactophy llum Simpson , 1900
corals:
No ,number of specimens;N , number of septa;D ,diameter of corallum(mm);Dt , diameter of tabularium(mm);
Lf , length of major septa(mm);Ls ,leng th of minor septa(mm);Wd , width of dissepimentarium(mm);Nd , number
of dissepiments(row);Nt , number of axial tabulae within a limited ver tical distance(mm).
Class Anthozoa Ehrenberg ,1834
　Subclass Rugosa Milne-Edwards et Haime ,1850
　　Order Columnariida Soshkina ,1941
　　　Suborder Cyathophyllina Nicholson , 1889
　　　　Family Disphyllidae Hill ,1939
Subfamily Charactophyllinae Wang ,1995
1972　Family C haractophyllidae Pedder , p.698.
Type genus　 Charactophyllum Simpson ,1900 ,pp.209 ,210
Diagnosis　Disphy llidae w ith septa composed of charactophylloid-type trabeculae.
Discussion　Three types of septal trabeculae may be distinguished in the mo re than 40
genera included wi thin Family Disphy llidae by Wang and Chen(1989).The first type is repre-
sented by Gurievskiella w ith t rabeculae arranged in regularly vertical fans , and may be called
phacellophy lloids.The second type is represented by Disphyllum with trabeculae g rowing in-
wards and upw ards at a nearly constant low ang le to horizontal , and arranged in regular half-
fans;this type has been termed the disphy lloid trabeculae or disphylloid type(Wang Xunlian ,
1995 ,p.30).The third type is represented by the charactophylloid t rabeculae.Since the septal
trabeculae in different shapes and arrangements reflect changes in configuration of the top of
polypal invagination , and are of important taxonomic signif icance , it is reasonable that Pedder
(1972)regarded those genera w ith septa composed of charactophy lloid t rabeculae as an inde-
pendent family.However , the charactophylloid t rabeculae on the w hole arranged in half-fans are
similar to disphylloid t rabeculae in arrangement;in addition , t ransit ional t rabeculae seem to ex-
ist between typical charactophylloid and disphylloid types in some genera.For this reason , the
authors consider that , except the genera wi th septa composed of phacellophlloid trabeculae ,
which should belong to the Phacellophyllidae Wedekind , 1922 , the genera put in Family Dis-
phyllidae by Wang and Chen(1989)may be subdivided into two subfamilies:Disphyllinae and
Charactophyllinae , the former including genera with septa composed of disphylloid trabeculae ,
and the latter w ith septa of charactophylloid t rabeculae.
Geological and geographical distribution Middle and Late Devonian;North America ,Eu-
rope and Asia.
Genus Charactophyl lum Simpson ,1900
1900　 Charactophyl lum Simpson , pp.209 , 210.
non 1924　 Charactophyl lum Simpson , Fenton and Fenton , p.25.
1940　Ptenophyllum Wedekind , Stumm , p.16.
24 古　生　物　学　报 第 37 卷
non 1940　 Charactophyl lum Simpson ,S tumm , p.59.
1945　 Charactophyl lum Simpson , Smi th , p.17.
1946　 Charactophyl lum Simpson , S tainb rook , p.415.
non 1949　 Charactophyl lum Simpson ,S oshkina , pp.90 , 91.
1950　 Charactophyl lum Simpson ,Wang , p.219.
non 1951　 Charactophyl lum Simpson ,S oshkina , p.68.
non 1952　 Charactophyl lum Simpson ,S oshkina , pp.90 , 91.
1956　 Charactophyl lum Simpson , Hill , p.280.
non 1957　 Charactophyl lum Simpson , Ivaniya , p.63.
1959　 Charactophyl lum Simpson ,Watkins , p.80.
non 1960　 Charactophyl lum Simpson ,Spasskiy , p.52.
non 1960　 Charactophyl lum Simpson ,Zheltonogova and Ivaniya , p.402.
non 1962　 Charactophyl lum Simpson ,S oshkina et al., p.11.
non 1963　 Charactophyl lum Simpson ,Altevogt , p.15.
non 1966　 Charactophyl lum Simpson ,Gunia , p.303.
non 1975　 Charactophyl lum Simpson ,S ytova and Kaplan , p.70.
non 1977　Heliophyl lum (Charactophyl lum)S impson , Spasskiy , p.108.
non 1978　 Charactophyl lum Simpson , Birenheide , p.84.
non 1979　 Charactophyl lum Simpson , Rozkow ska , p.29.
1981　 Charactophyl lum Simpson , Hill , pp.267 , 268.
1982　 Charactophyl lum Simpson , Pedder , p.562.
non 1983　 Charactophyl lum Simpson ,Cao et al., p.78.
non 1983　Charactophyllum Simpson , Zeng and Cai , p.141.
1987　 Charactophyl lum Simpson ,Wang X.L., p.100.
1993　 Charactophyl lum Simpson , M cLean , p.109.
Type species　 Charactophy llum nanum Hall et Whitf ield ,1873
Diagnosis　Solitary , cy lindrical and ceratoid coral.In early stages , major septa w ithdraw n
from the axis , becoming thinner in dissepimentarium and st rongly dilated in tabularium;both
cardinal and counter septa indist inct f rom other major septa , while minor septa shorter than
major ones.Dissepimentarium narrow , composed of concentric dissepiments.In subsequent on-
togenetic stages , septal dilation g radually reduced ,minor septa become long and dissepimentari-
um w idened.In late stages major septa long , reaching to or nearly to the axis and dilated in tab-
ularium in various degrees;minor ones rest ricted to dissepimentarium .Septa consisting of sinu-
ous uniseriate monacanthine trabeculae (charactophy lloid type) arranged in half-fans as a
w hole.In t ransverse section monacanths usually deviat ing f rom septal axis in g row th direction ,
and monacanthine trabeculae prot ruding f rom the sides of septa to fo rm indistinct carinae.
Monacanthine t rabeculae composed of irregular brachycolumns and a few of needles , with their
sides covered by slabs.Isolated axial ends of monacanthine trabeculae projecting outw ards to
form denticulate inner edges of septa.Sides of septa smoo th , or only faintly carinate.Dissepi-
mentarium mainly comprising concentric dissepiments , with a few herringhone st ructures.Dis-
sepiments composed of slabs parallel to the surfaces of dissepiments.Wide tabularium occupied
by variable tabulae w ith similar skeletal microst ructures as dissepiments.
25第 1 期 王训练等:论四射珊瑚征珊瑚属 Charactophy llum Simpson , 1900
Discussion　While erecting Campophy llum nanum , Hall and Whitfield (1873 , pp.232 ,
233)gave only a very brief description without any illust ration of the holotype.Simpson(1900 ,
pp.209 ,210)founded Charactophyl lum on the basis of Campophyl lum nanum in considera-
tion that the genus has essentially the same st ructure as Campophy llum except for the carinae
on the sides of septa , and pointed out that the difference between Charactophyl lum and Cam-
pophyl lum is the same as that betw een Heliophyl lum and Cyathophyl lum .Evident ly , Simp-
son believed that Charactophyl lum has typical yardarm carinae as Heliophy llum , but there is
not any sign of carinae in the line drawing of Ch .nanum(Simpson ,1900 ,p.210 , f ig .28).An-
other species , Ch .radiculum , illust rated by Simpson(p.210 , fig.29)bears features resembling
those of carinae.This Middle Silurian species w as later put under Tryplasma by Stumm(1952 ,
p.482 ,p1.125 , figs.1-9)based on its septal features.Neither Hall and Whit field(1873)no r
Simpson(1900)had clarified the nature of Ch.nanum .
Fenton and Fenton (1924), Smith (1945), Stainbrook(1946)and Watkins (1959)de-
scribed Ch.nanum from its typical locality and horizon respectively.This species illustrated by
Smith(1945 , p.17 , pl.1 , f ig s.7 ,8;pl.31 , figs.1a-i)resembles those by Watkins (1959 , p.
82.pl.16 , figs.13-20)in interio r features and ontogenet ic development;the form shown by
S tainbrook(1946 ,p.415 ,pl , 60 , fig.4)has only one t ransverse section , which is likely a junio r
member of the species represented by the specimen of Watkins(1959).The forms ment ioned
above are all dif ferent f rom that illust rated by Fenton and Fenton (1924 ,p.26.p1.1 , figs.1-
3).
Wang (1950)studied the septal microst ructures of the specimen in comparison w ith the
specimen shown by Smith (1945)in Pl.1 , f ig s.7 ,8 , holding that the septa of this species con-
sist of sinuous trabeculae , which were termed the charactophylloid type of t rabeculae by Pedder
(1972 ,p.698).
According to law of prio rity , the specimen of Fenton and Fenton(1924)should be used to
improve the definition of Charactophyl lum nanum ,but it is in distinct in characteristics , on-
togeny , and skeletal microst ructures.The specimen of Stainbrook (1946)w ith only one t rans-
verse sect ion is in suf ficient to give the definition.However , the specimens of bo th Smith
(1945)and Watkins (1959)bear dist inct character , well known ontogeny and skeletal mi-
crost ructures , and thus have been accepted by most authors (Hill , 1956 , 1981;Pedder , 1982;
McLean , 1993).For this reason in the present paper the defini tio n of this species is based on the
specimens of Watkins(1959)and Smith(1945), together w ith the material f rom Upper Devo-
nian of Zhusilenghaierhan area , Nei Mongol.
Smith(1945)stated that Ch .nanum has amplexoid septa ,which how ever have not been
accepted in general by later autho rs.These are laminar instead of amplexoid septa in the genus
as suggested by the excellent serial sections of Watkins(1959)and Smith(1945)as well as our
abundant specimens.The denticulation on the inner edges of the septa results f rom the axial
ends of the t rabecuale projecting out of the inner edges.
26 古　生　物　学　报 第 37 卷
Based on coral specimens f rom the former Soviet Union ,Soshkina(1949 , pp.90 ,91)added
to the defini tion of Charactophyl lum as follow s:
“Major septa not reaching to ax is , yardarm carinae as well-developed as those in Helio-
phyl lum , and trabeculae forming septa arranged in fans.”
This supplement is different from the o riginal defini tion.The species Ch .antiquum and
Ch.elongatum described by Soshkina , as no ted by Hill (1969 , p.11), should belong to
Gurievskiel la owing to thei r skeletal macrostructures and microst ructures.
Simpson (1900)and Soshkina (1949 , 1951 , 1952)considered Charactophyllum to have
yardarm carinae as in Heliophyllum .In fact the typical y ardarm carinae in the latter are formed
by the t rabeculae themselves(Wang , 1950 , pl.6 , fig.40;Sorauf and Oliver , 1976), while the
“carinae” of the former are caused by the dilation of trabeculae and their alteration wi th at tenu-
ated parts betw een adjacent trabeculae , or in some cases by the axial ends of the trabeculae pro-
jecting out of the septal edges.Therefore , the carinae in Charactophyllum are usually much
w eaker than those in Heliophy llum , and sometimes may be indistinct.
Pedder(1972 ,p.698)established Family Charactophy llidae based on charactophylloid t ra-
beculae in the septa.Charactophyllum is similar to Pseudozaphrent is , Temnophy llum ,
Temnophyl lum(Truncicarinulum), Alaiophy llum and Chostophyl lum included in Characto-
phyllidae in some aspects.In Pseudozaphrentis the septa are bilateral in arrangement , and the
major ones are regularly wedge-shaped and dilated in the median portion;these dif ferent iates it
from Charactophy llum .Temnophyl lum differs from Charactophyl lum in the presence of w ell-
developed stereozone and the absence of septal dilation in tabularium.Temnophyl lum (Trunci-
carinulum)may be dist inguished from Charactophy llum by the common absence of septal di-
lation in tabularium and by the presence of alternatively dilated setpa in transverse section
w hich may form numerous rows of inner w alls.Alaiophyllum may be distinguished from
Charactophyllum by its short major septa ,well-developed stereozone and at tenuate axial ends of
major septa , while Chostophyl lum is dif ferent from Charactophyl lum in having a pronounced
cardinal fossula in early stages , and sho rter major septa and absence of dilation at septal inner
ends in late stages.
Other genera close to Charactophyl lum in skeletal macrost ructures include Thoulelasma ,
Glossophy llum , Ceratophyllum , Houershanophyl lum , and Qiannanophy llum .
Thoulelasma ,erected by Pedder(1983 , p.226)for Early Devonian species f rom the Blue
Fio rd Formation of Sourthwestern Ellesmere Island ,Northwest Territo ries , is similar to Char-
actophyllum ,but dif fers in having a moderately long cardinal fossula and pinnately arranged
septa in cardinal quadrants as well as fine and apparent ly monacanthine t rabeculae arranged in
half-fans in septa.
Houershanophy llum Yu et Liao in Kong and Huang (1978 , p.54), based on H .involu-
tum Yu et Liao in Kong and Huang (1978 ,p.54 ,p1.17 , figs.1a-f)f rom the Eifelian Long-
dongshui Format ion of Guizhou Province ,China , resembles Charactophyllum in septal dilation
27第 1 期 王训练等:论四射珊瑚征珊瑚属 Charactophy llum Simpson , 1900
in tabularium ,but it may be dif ferentiated f rom Charactophy llum by the cry stal fascicles in the
septa and the pronounced cardinal fossula.
Qiannanophy llum Kong in Kong and Huang (1978 ,p.56)with Q .dushanense Kong in
Kong and Huang (1978 , p.56 , pl.17 , figs.3 , 4)as its type species is similar to Characto-
phyl lum in septal dilation , but differs from the lat ter in having a pronounced cardinal fossula
and pinnately arranged septa in late stages.
Glossophy llum was proposed by Wedekind (1924 , pp.77-78 , figs ,109-112)on the ba-
sis of G.dohmi Wedekind(1924 ,pp.77-78 , figs.109-112).Birenheide (1969 ,p.40)chose
the specimen illustrated by Wedekind in fig .112 as the lectotype;in 1978 , he put the other
three sections in Ceratophyl lum salmense (Wedekind).Since no figures or descriptions of a
longi tudinal section are available , the septal t rabecular structures of the type species still remain
unknown.As noted by Pedder (1982 , p.563;1983 ,p.227), the genus cannot be used with cer-
tainty at the present time.
Ceratophyl lum was founded by Gurich (1896 ,p.163)for Givetian species from the Holy
Cross M ountains , Poland , but the material w as destroyed during the Second World War and has
never been figured.Fedorow ski(1967)chose f rom Gurich s collection a neotype , in which the
trabeculae in the septa are arranged in asymmetrical fans wi th very narrow outer half-fans and
relatively much wider inner half-fans(Fedorow ski , 1967 ,pl.2 , f ig s.3 , 7).Hill and Jell(1970 ,
p.55)believed that the t rabeculae of C.typus are possibly rhipidacanthine.In addition , this
genus has a narrow dissepimentarium , bilaterally arranged septa and well-developed cardinal fos-
sula.All these may serve to dist inguish the genus f rom Charactophy llum .
Forms assigned to the genus:
Charactophyllum nanum (Hall et Whitfield), 1873 , pp.232 , 233.Frasnian , Iow a ,
U.S.A.
Charactophyllum praecursor Stainbrook ,1946 ,pp.415 ,416.Frasnian , Iow a ,U.S.A.
Ptenophyl lum sp.undet.Stumm , 1940 ,p.59.Frasnian ,Nevada , U.S.A.
Charactophyllum regulare Wang X.L.,1987 , p.101.Frasnian ,Nei Mongol ,China.
Charactophyllum ningxiaense Wang X.L., 1987 , pp.101 , 102.Frasnian , Nei Mongol ,
China.
Rejected forms:
Charactophyllum radiculum Simpson ,1900 , pp.209 ,210.
Charactophyllum sp.undet.Stumm ,1940 , p.59.
Charactophyllum sp.Smith ,1945 ,p.18.
Charactophyllum ant iquum Soshkina ,1949 ,pp.91-93.
Charactophyllum elongatum Soshkina ,1949 ,pp.93-95.
Charactophyllum lateseptatum Soshkina , 1951 ,p.69.
Charactophyllum spongiosum Soshkina ,1951 ,p.71.
Charactophyllum equchii Ma ,1956 ,p.43.
28 古　生　物　学　报 第 37 卷
Charactophyllum neospongiosum Ivaniy a ,1957 ,p.63.
Charactophyllum simplex Zheltonogova et Ivaniya ,1960 ,p.402.
Charactophyllum armenicum Soshkina et al., 1962 ,pp.313 , 314.
Charactophyllum lotzei Altevog t ,1963 ,pp.17 ,18.
Charactophyllum heterophyl loides Gunia ,1966 ,pp.303 ,304.
Charactophyllum proli ferum Gunia ,1966 ,p.304.
Charactophyllum balchaschicum Kaplan in Sytova and Kaplan ,1975 ,p.70.
Charactophyllum spongiosum Birenheide ,1978 ,p.85.
Charactophyllum vesiculosum Cao in Cao et al.,1983 ,p.78.
Charactophyllum qingheense Zeng et Cai ,1983 ,p.141.
Geological and geographical distribution 　Frasnian;North America (Stumm , 1940;
Smith ,1945;S tainbrook ,1946;Watkins ,1959),North China(Wang Xunlian , 1987).
Charactophyl lum nanum (Hall et Whitfield),1873
(Pls.Ⅰ -Ⅳ;Pl.Ⅴ , f igs.1-4;Text-fig.2)
T ext-fig.2　Scat ter diagram show ing number of major septa versus diameter in Charactophyllum nanum
1873　Campophyllum nanum Hall et Whi tfield , p.14.
non 1900　Charactophyllum nanum (Hall et Whitf ield), Sim pson , pp.209 , 210 , fig.28.
non 1990　Charactophyllum nanum (Hall et Whitf ield), Fenton and Fenton , p.26, pl.1 , figs.1-3.
1940　Ptenophyllum sp.undet , Stumm , p.16 , pl.8 , fig.6.
1945　Charactophyllum nanum (Hall and Whitf ield), Smi th , pp.17 , 18 , pl.1 , fi gs.7-8 , pl.31 , figs.la-i.
1946　Charactophyllum nanum (Hall et Whi tfield), S tainbrook , pp.415 , 416 , pl.60, fig.4.
1950　Charactophyllum nanum (Hall et Whi tfield), Wang , p.242 , pl.7 , fig.44.
1959　Charactophyllum nanum (Hall et Whi tfield), Watkins, p.82 , p.16 , f igs.13-20.
non 1962　Charactophyllum cf.nanum (Hall et Whit field), McLaren , Norri s and McGregor , p.11 , pl.4 , f ig.4.
non 1963　Charactophyllum nanum (Hall et Whitf ield), Altevogt , pp.15-17 , pl.1 , figs.1a-c.
1987　Charactophyllum nanum (Hall et Whi tfield), Wang , X.L., p.101 , pl.2 , f igs.4-7;pl.3 , fig.1.
29第 1 期 王训练等:论四射珊瑚征珊瑚属 Charactophy llum Simpson , 1900
1987　Charactophyllum ningxiaense Wang X.L., pp.101 , 102 , pl.3 , figs.3 , 4.
Table Ⅰ 　Measurements o f specimen
No N D Dt Dt/D Lf Ls Wd Nd Nt
84066 38×2 25.8 12.9 0.50 12.1 6.2 6.5 12
84190 33×2 24.2 13.5 0.56 10.1 6.1 5.2 8
84191 38×2 24.1 11.2 0.47 10.8 6.2 6.2 12 10/ 12
84196 37×2 23.4 13.2 0.56 10.1 5.2 5.1 10
84218 36×2 23.8 12.8 0.54 11.9 6.0 6.1 8 8/ 7
84219 34×2 25.8 12.3 0.48 13.0 7.8 5.5 12 11/ 10
84223 42×2 33.2 18.6 0.56 16.0 9.3 8.5 10 10/ 18
84226 37×2 26.3 14.2 0.54 11.2 7.4 7.6 11 7/ 8
9617 42×2 28.7 14.9 0.52 12.6 7.3 7.4 13 16/ 14
9614c 36×2 27.8 16.9 0.61 9.7 6.6 6.4 11 7/ 16
96107a 38×2 31.4 15.7 0.50 13.4 7.3 8.9 15
96107b 38×2 35.4 13.1 0.37 17.2 11.7 11.9 16
96107c 40×2 38.6 17.2 0.45 17.4 12.0 12.1 16 3/ 6
96109a 35×2 25.2 15.1 0.60 10.2 7.4 6.2 15
96109b 35×2 24.8 14.7 0.59 9.8 6.3 5.6 12
96109c 37×2 29.6 13.8 0.47 12.3 9.3 7.6 15
96109d 37×2 28.8 14.6 0.51 12.2 9.4 9.2 17 12/ 19
96112a 32×2 12.8 11.2 0.88 4.7 1.4 1.7 4
96112b 15.3 13.3 0.87 9.1 2.6 1.7 5
96112c 39×2 29.4 18.4 0.63 11.3 8.4 8.2 12
96112d 39×2 25.0 12.9 0.52 10.9 7.2 7.9 16 19/ 18
84244 39×2 35.3 18.2 0.52 13.4 9.0 7.6 12 16/ 11
84208 35×2 32.5 16.9 0.52 12.6 8.0 8.3 16 18/ 16
84232 36×2 27.9 13.4 0.48 12.3 8.3 8.5 17 3/ 9
84065 38×2 26.2 13.9 0.53 12.2 8.3 8.1 17 11/ 12
84199 33×2 22.0 14.8 0.67 7.2 4.1 4.1 9 7/ 9
84202b 36×2 19.2 11.1 0.58 9.1 3.2 4.1 7 12/ 12
84202a 30×2 14.9 12.0 0.81 3.7 2.0 1.5 3
84217 36×2 22.6 15.0 0.66 10.9 6.6 6.7 8
　　Diagnosis　Charactophy llum element with bilaterally arranged septa , shorter cardinal
septum and longer counter septum as well as irregular septal dilation which are considerably
v ariable in shape and thickness in tabularium.
Description　Corallum cylindrical and ceratoid (Pl.Ⅰ , fig.1a;Pl.Ⅱ , f ig s.1a , 1b),
ranging f rom 12.8 to 38.6mm in diameter (Tex t-fig.2).Largest specimen over seen more
30 古　生　物　学　报 第 37 卷
than 10.28cm in length.Irregular g row th w rinkles and grow th lines w ell-developed.Corallum
ceratoid in early stages , and cylindrical in late stages , becoming g radually smaller upward in di-
ameter in latest stage.Caly x steep-sided , deep , cup-shaped , with base usually somewhat flat o r
slightly concave.Narrow calicular platform occasionally occurring in some specimens.
Ex ternal w all thin , usually damaged.Septa in tw o orders , ranging f rom 30×2 to 42×2
in number.In early stages , major septa commonly ret reated f rom the axis , leaving an axial re-
gion(Pl.Ⅰ , f ig s.1b ,1c ,2a;Pl.Ⅲ , fig.7), relatively thin in dissepimentarium , and st rongly
dilated in i rregular shape in tabularium where they may connect laterally w ith each other.In
some specimens major septa weakly dilated in tabularium (Pl.Ⅰ , f ig .2a).Both cardinal and
counter septa not differentiated f rom other major septa.Minor septa short , less than half as
long as major ones.Tabularium broad , more than 4/5 of individual s diameter.Narrow dis-
sepimentarium composed of 3-5 row s of concentric dissepiments;herringbone dissepiments oc-
casionally present.
In late stages , majo r septa long , reaching to or nearly to axis , bilateral in arrangement.
Cardinal septum sho rter in poorly developed cardinal fossula;counter septum usually longer
than adjacent major ones.Major septa relatively thin in dissepimentarium , and irregularly dilat-
ed in tabularium , with their inner ends possibly ro tated about axis or joining each other (Pl.
Ⅳ, fig.1a)o r coated by lamellar skeletal thickening.Septal dilation variable in thickness and
shape , even w ithin dif ferent levels of a single individual(Pl.Ⅰ , figs.1d ,1e).In some speci-
mens only part of major septa dilated in tabularium.(Pl.Ⅰ , fig s.1d ,2b;Pl.Ⅱ , fig.2a;pl.Ⅴ,
fig.1a).Minor septa w ell developed , longer than half of majo r ones , confined to or slightly
penet rating dissepimentarium , and usually no t dilated at their axial ends.Sides of septa usually
smooth or wi th w eakly developed carinae.Zigzag carinae possibly developed in outer parts of
dissepimentarium in some individuals (Pl.Ⅱ , fig.2a).Septa composed of sinuous uniseriate
monacanthine trabeculae (charactophy lloid type)consisting of irregular brachycolumns(Pl.Ⅳ,
figs.2-4)and a few needles(Pl.Ⅲ , figs.2-5 , 8 , 9).In t ransverse and tangential sections
boundaries betw een trabeculae not easily recognizable , and the brachycolumns and needles in
pinnate arrangement f rom the axis to the sides and upper inner edges of septa , parallel to each
o ther or grouped into tuf ts(Pl.Ⅲ , fig.9).In longi tudinal section boundaries between trabecu-
lae clear (Pl.Ⅴ, fig s.1b , 2-4);trabeculae 300-500μm in diameter , comprising irregular
brachycolumns.Brachycolumns 5-12.5μm long and 2-3μm in diameter , with length/diame-
ter ratio from 2.5∶1 to 4.2∶1;needles 15-25μm in leng th , ranging f rom 2.5-3μm in diame-
ters , with leng th/diameter ratio f rom 5∶1 to 8∶1 , distinct in outline.Axial ends of t rabeculae
projecting over inner edges of septa , making them appear denticulate.In a few specimens , in-
ner edges of septa possibly appearing as spines on the upper surface of tabulae(Pl.Ⅳ, figs.1b ,
2-4).Septal spines consisting of nearly vertically growing trabeculae.Three vertically over-
lapping t rabeculae with unclear boundaries(Pl.Ⅳ, fig.3).Sides of septa covered by slabs(Pl.
Ⅲ , figs.4 ,6).Dissepimentarium w ide , about 1/4 of individual s diameter , consisting mainly
31第 1 期 王训练等:论四射珊瑚征珊瑚属 Charactophy llum Simpson , 1900
of concentric dissepiments.Herringbone dissepiments present where minor septa retreated.
Lateral dissepiments occurring occasionally .Dissepiments composed of slabs , which are paralel
to the surfaces of dissepiments or slightly curved , w ith concave side facing the upper skeleton
surface.
In longi tudinal section the dissepimentarium consisting of 7 -17 row s of dissepiments ,
which are inclining w ith the inclination angles increasing from outer to inner part;outer part
g lobose , while the inner part relat ively elongated and nearly vertical.Tabularium about half of
the corallum in diameter , occupied by complete or incomplete tabulae.Tabulae much more
variable , even within a single specimen.Axial tabulae flat , concave or slightly convex , with
3-11 countable wi thin a vertical distance of 10mm.Periaxial tabulae inclining inw ard at vari-
able angles to horizontal.Lamellar skeletal thickening of tabular surface common.Tabulae
composed of slabs (Pl.Ⅳ, figs.1b , 5-7)which are irregular in shape and parallel to tabular
surfaces , 10-15μm in diameter and 2-4μm in thickness , with diameter to thickness ratio of
about 4.2∶1.
Charactophyllum regulare Wang X.L., 1987
(Pl.Ⅴ , f igs.5-7;Pl.Ⅵ)
1987　Charactophyllum regulare Wang X.L., p.101 , pl.2 , figs.1-3;pl.3 , fig.2.
Table Ⅱ　Measurements o f specimen
No. N D Dt Dt/D Lf Ls Wd Nd Nt
84198 37×2 26.4 13.2 0.50 10.1 6.9 6.4 12 12/ 13
84209 38×2 28.2 18.2 0.65 11.3 6.1 5.2 7
84225 44×2 36.6 17.6 0.48 18.2 11.1 10.2 15
84247 36×2 22.3 14.2 0.64 9.1 6.0 6.0 8
　　Diagnosis　Charactophy llum element w ith nearly radially arranged septa and regular
fusiform septal dilation in inner part of dissepimentarium and outer part of tabularium.
Description　Solitary , cylindrical o r trochoid co ral.In early stages size and features re-
maining unknown because of poor preservation.Grow th w rinkles and g row th lines well-devel-
oped.Specimens observed w ith diameter ranging f rom 22.3 to 36.6mm .External w all thin ,
less than 0.5mm in thickness.Septa in tw o orders , nearly radially arranged , ranging f rom 36
×2 to 42×2 in number.Majo r septa long , nearly ex tending to axis , relatively thin in dissepi-
mentarium , regularly dilated in fusiform in inner part of dissepimentarium and outer part of
tabularium , w ith axial ends usually coated by lamellar skeletal thickening(Pl.Ⅵ , fig.1).Car-
dinal septum shorter than other major septa , while counter septum not easily distinguishable
from other major ones.M inor septa w ell-developed , longer than half of the major one , confined
to or slightly penetrating dissepimentarium , thin or only slight ly dilated at thei r axial ends.
Sides of septa smooth or w ith weakly developed carinae , which may make the septa appear
32 古　生　物　学　报 第 37 卷
zig zag in dissepimentarium (Pl.Ⅵ , fig.1).Septa composed of sinuous uniseriate monacanthine
trabeculae.In transverse section t rabeculae pinnately arranged from septal axis to sides(Pl.Ⅴ,
figs.6 ,7), composed of irregular brachycolumns in irregular arrangement(Pl.Ⅵ , f ig s.2-7).
Brachycolumns 5-10μm long , and 2-3μm in diameter , w ith a length/diameter ratio of about
3∶1.Dissepimentarium consisting mainly of concentric dissepiments.Herringbone structures
observable w here mino r septa ret reated.In outer part of dissepimentarium lateral and herring-
bone dissepiments usually developed betw een adjacent major septa and minor septa.Dissepi-
ments consist ing of slabs(Pl.Ⅵ , f ig s.2 ,8 ,9)parallel to the surfaces of dissepiments , flat o r
slightly curved , with concave side facing upper surface of dissepiments , 13-15μm in diame-
ter , and 2-3μm in thickness , w ith a diameter/ thickness ratio of about 5.6∶1.
In longitudinal section dissepimentarium composed of 8-15 rows of dissepiments ,which
are globose in outer part of dissepimentarium , steeply inclining , w ith inclination ang le increas-
ing f rom outer to inner part.Dissepiments in inner part of dissepimentarium elongated and
nearly vertical.Tabularium about half of corallum diameter , occupied by complete or incom-
plete tabulae.Axial tabulae variable , convex or concave;periaxial tabulae inclining in varying
deg rees.Upper surface of tabulae commonly covered wi th lamellar skeletal thickening.Tabulae
composed of slabs w hich are f lat and parallel to tabular surfaces.
Discussion　This species is similar to Charactophyl lum nanum in some aspects , but dif-
fers in having nearly radially arranged septa and regular fusiform septal dilation in inner dissepi-
mentarium and tabularium .This species resembles Pseudozaphrent is di f f icile in the form of
dilation of major septa in tabularium , but it may be distinguished from the latter w hich has bi-
laterally arranged septa and inner w all formed by septal dilation at the boundary betw een dis-
sepimentarium and tabularium.
2　Skeletal microstructures of Charactophyllum and its significance in
evolution
As mentioned above , the septa of Charactophyllum are composed of charactophy lloid type
trabeculae.The septal t rabeculae consist of radially arranged irregular brachycolumns and nee-
dles , while the lateral skeleton , including tabulae and dissepiments is composed of slabs.
The skeletal microst ructure of Charactophyllum is of a t ransit ional nature f rom Early
Palaeozoic to Carboniferous-Permian rugose co rals.
1.Lamellar skeletons are dominant in Ordovician and Silurian rugose corals , but reduced
in Charactophyl lum , and completely disappeared in Carboniferous and Permian Caniniida , in
w hich the skeletons consist entirely of fibrous st ructure , including tabulae and dissepiments.
2.The evolutionary t rend of lamellar skeleton is mainly recorded by variations in diameter
and thickness of fundamental elements as well as the diameter/ thickness ratio (Tab.Ⅲ).The
33第 1 期 王训练等:论四射珊瑚征珊瑚属 Charactophy llum Simpson , 1900
Ordovician and Silurian fundamental elements of lamellar skeleton are typical flakes , which are
large and thin in shape , w ith a small longer/ shorter diameter ratio , and a large diameter/ thick-
ness radio(>5∶1).In Charactophy llum the fundamental elements of lamellar skeletons are
small and thick in shape , w ith a longer/ shorter diameter ratio of more than 2∶1 and a diame-
ter/ thickness ratio of about 4.2∶1.Surfaces of fundamental elements of lamellar skeleton usual-
ly are not everyw here easily observable.Fo r this reason , the measured longer and shorter diam-
eters can hardly reflect any true si tuation.However , the thickness of lamellar fundamental ele-
ments can be easily observed , and these diameter/ thickness ratio may be used as a cri terion to
distinguish f lakes from slabs.Evident ly , the lamellar skeletons in Charactophyl lum are mainly
composed of slabs w hich may be represented by the lamellar fundamental elements with a diam-
eter/ thickness ratio of less than 5∶1.
Table Ⅲ　Measurements of fundamental elements of skeletal microstructures in some genera(μm)
Order
Genus
Cystiphyllida
Maikot tia
(C hen , 1997)
Gyalophylloides
(Chen and Wang , 1986)
Columnariida Caniniida
Charactophyllum
(This paper)
Aulophyllum
(C hen and Wang , 1986)
flakes and slabs
diameter 32.4 10.1 12.5
thickness 3.8 1.5 3.0
ratio 8.5∶1 6.7∶1 4.2∶1
brachycolumns
length 6.7 5.7 8.8
diameter 4.0 2.3 2.5
ratio 1.7∶1 2.5∶1 3.5∶1
needles
length 20.0 32.3
diameter 2.8 4.0
ratio 7.1∶1 8.1∶1
Age S1
1 S 1
2 D3 C 1
　　3.The evolut ionary t rend of fibrous skeletons show s an increase in the length/diameter ra-
tio of fundamental elements.From Ordovician to Permian , the fibrous skeleton developed from
g ranules , brachycolumns to typical needles.The Ordovician and Silurian fibrous skeleton of ru-
gose corals is dominantly composed of granules and brachycolumns;the fibrous skeleton of
Charactophyllum is dominant ly composed of irregular brachycolumns , with a few sho rt and
thick needles , while the Carboniferous and Permian fibrous skeletons are dominantly composed
of thin and long needles.
All these mentioned above show that the skeletal microstructure of rugose corals bears a
stable , prog ressive development tendency , and serves as a guiding clue to t racing the evolution
of higher groups.
34 古　生　物　学　报 第 37 卷
3　Conclusions
1.In Charactophy llum the septa are laminar instead of amplexoid , composed of sinuous
uniseriate monacanthine trabeculae arranged in asymmetric half-fans instead of erect fans.Sep-
tal sides are smooth , or only faintly carinate w ithout typical y ardarm carinae.The dissepiments
are arranged in half-fans.
2.The septal trabeculae consist of radially arranged irregular brachycolumns and needles
and the lateral skeletons are composed of slabs , including tabulae and dissepiments.
3.The skeletal microstructure is of a transi tional nature f rom Early Palaeozoic to Carbonif-
erous-Permian rugose corals , indicating that the skeletal microst ructure of rugose corals bears a
stable and progressive development tendency of important taxonomic significance providing a
guiding clue to t racing the evolution of higher groups.
4.This genus is confined to early Late Devonian(Frasnian)in age and to North America
and North China in dist ribution.
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Explanation of plates
　　All corals illustrated here w ere collected f rom the upper part of the Frasnian Low er Xipingshan Formation in Zhusileng-
haierhan area , Nei Mongol, China.All specimens are stored in China Universi ty of Geosciences , Beijing , China.Magnificat ion
of SEM figures is indicated by bar scale , U=μm.
Plate Ⅰ
1.　Charactophyllum nanum (Hall et Whi tfi eld), 1873.Cat.no.96112.
1a.exterior view s(before sect ioning), ×1;1b-1e.transverse sections, ×3;1f.longitudinal sect ion , ×3.
2.　Charactophyllum nanum (Hall et Whi tfi eld), 1873.Cat.no.84202.
2a , 2b.t ransverse sections , ×3;2c.longitudinal section , ×3.
3.　Charactophyllum nanum (Hall et Whi tfi eld), 1873.Cat.no.84066.
3.t ransverse sections , ×3.
Plate Ⅱ
1.　Charactophyllum nanum (Hall et Whi tfi eld), 1873.Cat.no.96107.
1a.exterior view s(f ront), ×1;1b.exterior views(lateral), ×1;1c.transverse sect ion , ×2;1d.longitudinal sect ion ,
×2.
37第 1 期 王训练等:论四射珊瑚征珊瑚属 Charactophy llum Simpson , 1900
2.　Charactophyllum nanum (Hall et Whi tfi eld), 1873.Cat.no.84119.
2a.t ransverse sect ion , ×2;2b.longitudinal section , ×2.
3 , 4.　Charactophyllum nanum (Hall et Whi tfield), 1873.Cat.no.9616.
3.micrograph of septa t ransverse section , ×30.
4.micrograph of septa t ransverse section , ×90 , enlarged area A in fig.3.
Plate Ⅲ
1-6.　Charactophyllum nanum (Hall et Whi tfield), 1873.Cat.no.84216.
1a.t ransverse sect ion , ×3;1b.longitudinal section , ×3.
2.inner ends of septa in t ransverse sect ion.
3.enlarged square in fig.2 , show ing septal t rabeculae composed of needles parallel to each other and grow ing f rom septal
axi s to sides and upper inner edge of septum.
4-6.septal microst ructures in longitudinal sect ion.
4.upper lef t part of the f igu re facing upper inner edge;square show ing nearly vertical fibrous skeletons in axial zone(upper
left), and lamellar skeletons with slabs in lateral zones of septum (low er right).
5.enlarged area A in fig.4 , show ing needles and i rregular brachycolumns in axial zone.
6.enlarged area B in f ig.4 , showing nearly horizontal or inclining lamellar skeletons w ith slabs in lateral zones.
7-9.　Charactophyllum nanum (Hall et Whi tfield), 1873.Cat.no.9614a.
7.t ransverse section of early stage , ×3.
8.t ransverse section of septa , upper part of the figure facing inner edge.
9.enlarged square in fig.8 , show ing fibrous skeletons w ith needles parallel to each other.
Plate Ⅳ
1-7.　Charactophyllum nanum (Hall et Whi tfield), 1873.Cat.no.84219.
1a.t ransverse sect ion , ×3;1b.longi tudinal section , ×3.
2-4.septal microst ructures in longitudinal sect ion.
2.enlarged area A in fig.1b
3.enlarged square in fig.2 , showing 3 nearly vertical overlapping monacanthine t rabeculae in axial zone and lamellar skele-
t on w ith slabs in lateral zones.
4.an enlarged single trabecula in fig.3 , show ing brachycolumns and their arrangem ent in monacanths and dist inct tuf ts
composed of brachycolumns.
5-7.microstructures of tabulae in longitudinal sect ion.
5.enlarged area B in f ig.1b , show ing lamellar slabs parallel to each other and to su rfaces of tabulae.
6 , 7.enlarged square in fig.5 , showing shape and arrangement of slabs.
Plate Ⅴ
1-4.　Charactophyllum nanum (Hall et Whi tfield), 1873.Cat.no.84226.
1a.t ransverse sect ion , ×3;1b.longi tudinal section , ×3.
2-4.enlarged area A in fig.1b , w ith upper lef t part of the f igu re facing upper inner edge , showing boundaries betw een
trabeculae and i rregular brachycolumns and needles in t rabeculae.
5.　Charactophyllum regulare Wang , 1987.Cat.no.84198.
5a.t ransverse sect ion , ×3;5b.longitudinal section , ×3.
6 , 7.　Charactophyllum regulare Wang , 1987.Cat.no.84247.
6.t ransverse section of septa , show ing uniseriate t rabeculae.
7.enlarged lef t septum in fig.6.
38 古　生　物　学　报 第 37 卷
Plate Ⅵ
1.　Charactophyllum regulare Wang , 1987.Cat.no.84225.
1.t ransverse section , ×3.
2-9.　Charactophyllum regulare Wang , 1987.Cat.no.84198.
2.t ransverse section of septum;low er part of figure show ing septal outer end and upper part showing di lated inner end.
3.enlarged area A in fig.2 , show ing fib rous skeletons with brachycolumns in variable di rections;brachycolumns nearly
vert ical in upper and low er part s and parallel to each other in cen tral part of the f igure.
4.enlarged area B in f ig.2(dilated inner end of septum).
5 , 6.enlarged square in fig.4 , show ing b rachycolumns parallel to each other.
7.enlarged longitudinal section of septa , show ing fibrous skeletons w ith brachycolumns in variable direct ions and grouped
into tuft s.
8.enlarged t ransverse section of dissepiments in area C in fig.2 , show ing lamellar skeletons w ith slabs parallel to each other
and to the su rfaces of dissepiments.
9.enlarged square in fig.8 , show ing small slabs wi th irregular edges.
39第 1 期 王训练等:论四射珊瑚征珊瑚属 Charactophy llum Simpson , 1900