morphometrics and palaeoecology of the coral …...the strata containing agetolites were reported to...

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Alcheringa: An Australasian Journal of Palaeontology

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Morphometrics and palaeoecology of the coralAgetolites from the Xiazhen Formation (UpperOrdovician), Zhuzhai, South China

Ning Sun, Robert J. Elias, Suk-Joo Choh, Dong-Chan Lee, Xun-Lian Wang &Dong-Jin Lee

To cite this article: Ning Sun, Robert J. Elias, Suk-Joo Choh, Dong-Chan Lee, Xun-Lian Wang &Dong-Jin Lee (2016) Morphometrics and palaeoecology of the coral Agetolites from the XiazhenFormation (Upper Ordovician), Zhuzhai, South China, Alcheringa: An Australasian Journal ofPalaeontology, 40:2, 251-274, DOI: 10.1080/03115518.2016.1111071

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Morphometrics and palaeoecology of the coral Agetolitesfrom the Xiazhen Formation (Upper Ordovician), Zhuzhai,South China

NING SUN, ROBERT J. ELIAS, SUK-JOO CHOH, DONG-CHAN LEE, XUN-LIAN WANG and DONG-JIN LEE

SUN, N., ELIAS, R.J., CHOH, S.-J., LEE, D.-C., WANG, X.-L. & LEE, D.-J., February 2016. Morphometrics and palaeoecology of the coral Agetolitesfrom the Xiazhen Formation (Upper Ordovician), Zhuzhai, South China. Alcheringa 40, 251–274. ISSN 0311-5518.

Agetolites is a peculiar cerioid coral possessing traits of both tabulates and rugosans. The presence of numerous mural pores has been consideredby some workers to indicate a relation to tabulates, although an affinity to rugosans has also been proposed based mainly on well-developed septathat alternate in length. Agetolites is by far the most common colonial coral in the Upper Ordovician Xiazhen Formation at Zhuzhai, in the JCY(Jiangshan–Changshan–Yushan) triangle region of South China. It occurs in various lithofacies representing a wide range of depositional environ-ments. Five species are recognized and verified by cluster analysis, discriminant analysis, descriptive statistics and bivariate plots: A. yushanensisLin, 1960, A. raritabulatus Lin, 1960, A. waicunensis (Lin & Chow, 1977), A. oculiporoides Lin, 1960 (including A. huangi Lin, 1960 as a juniorsynonym) and A. maxima (Lin & Chow, 1977). Agetolitids from the JCY triangle described previously under the generic name Agetolitella areherein transferred to Agetolites. In the Xiazhen Formation at Zhuzhai, A. waicunensis and A. maxima are restricted to the lower part of the forma-tion, whereas A. yushanensis occurs in the upper part. Agetolites raritabulatus and A. oculiporoides have greater stratigraphic ranges in the lowerand upper parts of the formation. The size and shape of Agetolites coralla at Zhuzhai are considered primarily to be intrinsic characteristics of thespecies but in some cases also seem to be related to lithofacies; large and domical coralla are dominant in calcareous mudstone facies, whereassmall coralla with various forms are commonly found in reef or reef-related facies. In the latter facies, spatial competition is indicated by commonoccurrences of both Agetolites encrusting stromatoporoids and stromatoporoids encrusting Agetolites.

Ning Sun [[email protected]], School of Earth Sciences and Resources, China University of Geosciences, Beijing 100083, PR China; Robert J.Elias [[email protected]], Department of Geological Sciences, The University of Manitoba, Winnipeg, Manitoba R3T 2N2, Canada; Suk-JooChoh [[email protected]], Department of Earth and Environmental Sciences, Korea University, Seoul 136-701, Korea; Dong-Chan Lee[[email protected]], Department of Earth and Environmental Sciences, Chungbuk National University, Cheongju, 361-763, Korea; Xun-LianWang [[email protected]], School of Earth Sciences and Resources, China University of Geosciences, Beijing 100083, PR China; Dong-Jin Lee*[[email protected]], Department of Earth and Environmental Sciences, Andong National University, Andong 760-747, Korea. *Also affiliated withCollege of Earth Science, Jilin University, Changchun 130061, PR China. Received 13.6.2015; revised 10.10.2015; accepted 19.10.2015.

Key words: Agetolites, corals, morphometric analysis, palaeoecology, systematics, Late Ordovician, South China.

AGETOLITIDS are a group of problematic LateOrdovician cerioid corals having features of both tabu-late and rugose corals. Several workers have consideredthem to be allied with tabulates, owing to the presenceof numerous and large mural pores (Sokolov 1955, Lin1960, Yu & Zhang 1963, Kim 1966, Lin & Chow1977, Hill 1981, Lin et al. 1984, Lin & Huang 1986,Oliver 1996, Wang 1997). An affinity with rugosanshas also been proposed, based mainly on septal patternsand other morphological characteristics, such as an axialstructure of septal lobes or a columella (Kim 1974, Xuet al. 1999). However, a recent study revealed that theseptal pattern in Agetolites Sokolov, 1955 from Zhuzhaiin South China is variable and can not be considered anindication of rugosan affinities (Sun et al. 2014).

In spite of their systematic and phylogenetic signifi-cance, agetolitids have been the subject of very few

detailed studies since Sokolov (1955) first documentedAgetolites. Three genera of agetolitids were recognizedby Hill (1981, pp. F559–F561): Agetolites, AgetolitellaKim, 1962 and Somphopora Lindström, 1883. Theidentification of Agetolitella in certain regions, such asSouth China, is doubtful (Lin & Chow 1977, p. 144).The Silurian Somphopora is considered to be a theciidtabulate (Sokolov 1962, p. 329). Hemiagetolites Lele-shus, 1963 was originally interpreted to be phylogeneti-cally intermediate between the tabulate PaleofavositesTwenhofel, 1914 and Agetolites (Leleshus 1963) but isnow considered to be a junior synonym of Agetolites(Hill 1981, p. F559).

Eight more genera of agetolitids have been erectedbased on Chinese material, raising the potential forconsiderable taxonomic confusion: Agetolitinus Deng &Li, 1979; Paragetolites Lin in Li & Lin, 1982;Subagetolites Li in Li & Lin, 1982; HemiagetolitinaLin, 1986; Agetolitoides Lin, 1986; Paragetolitella Lin,1986; Subagetolitoides Lin, 1986; and Hemiagetolitella© 2016 Australasian Palaeontologists

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Lin & Huang, 1986. Morphological parameters used todefine these genera are vague and inadequate, makingtheir taxonomic validity questionable. There is a clearand present need for further research to clarify the con-fusion in agetolitid classification.

The type species of Agetolites, A. mirabilis Sokolov,1955, was originally described as a tabulate coral fromthe ‘lower Llandoverian’ of the Chingiz Range inKazakhstan. Kim (1966), however, speculated that thedeposit from which the genus was reported by Sokolovis Upper Ordovician. Agetolites has been reportedfrom localities in numerous areas: Kazakhstan(Sokolov 1955, Popov et al. 1999); Tajikistan andUzbekistan (Hill 1981); northeastern Russia (Koren &Sobolevskaya 2008); Jiangxi (Lin & Chow 1977), InnerMongolia (Lin et al. 1984), Ningxia and Xinjiang (Lin& Huang 1986, Wang 1997) in China; North Korea(Pak 1983a, b, Kim 1990, An & Ma 1993, Om et al.1993); central New South Wales (Semeniuk 1970) andnorthern Queensland (Hill 1981, Webby et al. 2004) inAustralia; and Alaska (Oliver et al. 1975) and doubt-fully western Utah (Pandolfi 1985) in North America.The strata containing Agetolites were reported to beUpper Ordovician except for a location in northernQueensland, which Hill (1981, pp. F560, F561) identi-fied as Upper Ordovician or questionably lowerSilurian. The latter occurrence is probably UpperOrdovician (lower Bolindian; Webby et al. 2004).

The JCY (Jiangshan–Changshan–Yushan) triangle,located on the border of Jiangxi and Zhejiang provincesin southeastern China, has long been known as the classicregion for study of the Ordovician System in South China(Zhang et al. 2007; Fig. 1A). Despite the abundance ofcorals within the Upper Ordovician in this region, theyhave received surprisingly little attention compared with

other invertebrates, such as brachiopods (Zhan et al.2002). Only a few preliminary investigations of coralshave been carried out in the JCY region (Lin 1960, Yu &Zhang 1963, Lin & Chow 1977) and a comprehensiveand modern systematic treatment is overdue. Fifteen spe-cies of Agetolites and ten species of Agetolitella havebeen reported from Upper Ordovician strata including theSanqushan Formation (Lin 1960, Lin & Chow 1977),which is considered to be the seaward equivalent of theXiazhen Formation (Zhang et al. 2007, fig. 2.2).

Agetolites is by far the most common genus of colo-nial corals within the Xiazhen Formation in the presentstudy area at Zhuzhai, at the southern end of the JCYtriangle (Fig. 1A, B). This genus occurs in diverselithofacies deposited in a wide range of depositionalenvironments. The principal aims of this study are: (1)to re-evaluate morpho-taxonomic characteristics previ-ously used in studies of Agetolites, and apply multivari-ate analytical methods for distinguishing andcharacterizing species; and (2) to document and assessthe palaeoecological distribution of Agetolites in theXiazhen Formation.

Geology and stratigraphyOrdovician carbonate successions are well exposed nearthe border between Jiangxi and Zhejiang provinces, inthe JCY triangle of southeastern China (Zhang et al.2007; Fig. 1A). Numerous Upper Ordovician reef com-plexes have been described from the shallow lagoon tomarginal reef of the Sanqushan Formation and itslandward equivalent, the Xiazhen Formation (Bian &Zhou 1990, Yu et al. 1992, Webby 2002, Li et al.2004, Zhang et al. 2007). The shallow platform carbon-ates of these formations grade into contemporaneous

Fig. 1. A, Map of China showing location of the South China Plate and enlargement of the JCY (Jiangshan–Changshan–Yushan) triangle regionshowing location of the study area at Zhuzhai (grey square) and locations of previously reported type specimens (small circles). B, Geological mapof the study area and sub-sections ZU 1, ZU 2 and ZU 3 (modified after Lee et al. 2012).

252 NING SUN et al. ALCHERINGA

fine-grained clastics of the Changwu Formation (Chenet al. 1987, Li et al. 2004). The Xiazhen Formationwas first defined by Lu et al. (1976) and mostly studiedaround Tashan, where the type section of the formationis located (Chen et al. 1987, Zhan & Fu 1994, Zhanet al. 2002; Fig. 1A).

The Xiazhen Formation at Zhuzhai in Jiangxi Pro-vince, from which specimens of Agetolites forming thebasis of this study were collected, has recently been re-measured and described in detail by integration of litho-logical and palaeontological data (Lee et al. 2012;Fig. 1A, B). This study follows Lee et al. (2012), whopresented a revised stratigraphy of the formationexposed at Zhuzhai. Comparison of lithological andpalaeontological data of the exposures (designatedtherein as sub-sections ZU 1, ZU 2 and ZU 3, separatedby Quaternary deposits) demonstrates that stratigraphicintervals of the sub-sections overlap (Figs 1B, 2). Theexposure of the formation at Zhuzhai is about 190 mthick (Fig. 2). Four informal members are recognizedwithin the formation: the lower limestone member,lower shale member, middle mixed-lithology memberand upper shale member (Lee et al. 2012; Figs 1B, 2).Corals were not found in the lower shale member. Allof sub-section ZU 3 correlates with the upper part ofZU 1, and identical species of Agetolites occur in thecorrelative interval (Lee et al. 2012, figs 8, 9).

The depositional environments of the coral beds inthe Xiazhen Formation are interpreted as back reef tomarginal platform settings (Lee et al. 2012). No fossilsthat could indicate a precise age (e.g., graptolites orconodonts) have been found in the formation, but trilo-bites and brachiopods suggest that it is mid-Ashgill(Zhan et al. 2002, Zhang et al. 2007). Based on coralsand a proposed correlation with the ChangwuFormation, Zhang et al. (2007) estimated the XiazhenFormation to be of early to late Katian age (middleAshgill).

Material and methodsCoralla from the Xiazhen Formation at Zhuzhai aresuperficially well preserved, but recrystallization andsilicification commonly obscure their internal structures.Over 500 coralla of Agetolites were collected, including108 relatively well-preserved specimens used for multi-variate analysis. Of these specimens, 107 are from sub-sections ZU 1 and ZU 2 (Table 1), and one is fromtalus at sub-section ZU 3. Also used in the analysis aresix type specimens of Agetolites species designated byLin (1960) from the Upper Ordovician at Shiyanshan,which is located 5 km northeast of Zhuzhai (Fig. 1A).These include the holotypes of A. huangi Lin, 1960 andA. raritabulatus Lin, 1960, and paratypes of A.oculiporoides Lin, 1960 and A. yushanensis Lin, 1960deposited in the Geological Museum of China in

Fig. 2. Lithostratigraphic columns showing relevant portions of theXiazhen Formation (Upper Ordovician) at sub-sections ZU 1, ZU 2and ZU 3, Zhuzhai, South China (modified after Lee et al. 2012; forcomplete columns and correlation, see Lee et al. 2012, figs 4, 8).Specimens examined in this study were collected from intervals A3–A12 at ZU 1, A1–A3 at ZU 2, and the upper part of ZU 3-I. LLM,lower limestone member; LSM, lower shale member; MMM, middlemixed-lithology member; USM, upper shale member. S, shale; M,mudstone; W, wackestone; P, packstone; G, grainstone; F, floatstone.

ALCHERINGA ORDOVICIAN CORALS FROM CHINA 253

Mem

ber

Sub-section

/stratigrap

hic

interval

Lithofacies

Relativeenergy

level

Orientation

,grow

thform

,an

dsize

ofcoralla

Agetolites

species/number

ofspecim

ens

ZU

1ZU

2A

BC

DE

Upper

shalemem

ber

A12

Shale,lim

estone–shale

alternation

includingpartially

developedsm

all

reefs,no

dularcalcareous

mud

ston

e

Low

tomod

erate

Transpo

rted

andin

grow

thpositio

n;mostly

hemispherical

with

few

branching;

largeand

small

112

1

Middlemixed

litho

logy

mem

ber

A11

Peloidalwackestoneto

packstone

Moderate

Transported;hemispherical;sm

all

1A10

Peloidalwackeston

eto

packstone,

coralfloatstone

with

peloidal

packston

eto

grainstone

matrix

Moderateto

periodically

high

Mostly

transported;

hemispherical;

mostly

small

61

A9

Peloidalwackeston

eto

packstone,

limestone–shale

alternation

Low

tomod

erate

Transpo

rted

andin

grow

thpositio

n;hemispherical;sm

all

21

A8

Peloidalpackston

eto

grainstone,

coralfloatstone

with

peloidal

packston

eto

grainstone

matrix

Mod

erateto

high

Transpo

rted

andin

grow

thpositio

n;hemispherical

andsome

branching;

small

71

2

A7

Peloidalwackeston

e(topackstone)

Mod

erateto

period

ically

high

Transpo

rted

andin

grow

thpositio

n;hemispherical

andsome

branching;

small

2

A6

Calcareousmudstone

Low

tooccasionally

moderate

Overturnedandin

grow

thpo

sitio

n;largehemisph

erical

tosm

alllenticular

19

A5

Nod

ular

calcareous

mud

ston

eLow

Not

ingrow

thpo

sitio

n;hemispherical;largeandsm

all

55

1

A4

Bioturbated

wackestone

Low

(tomoderate)

Not

ingrow

thpositio

n;hemispherical;large

1

A3

A3

Coral

floatstone,peloidal

&bioclastic

grainstone,shale

Mod

erateto

high

Transpo

rted

andin

grow

thpositio

n;hemispherical;largeand

small

0+2

2+11

2+5

A2

Floatstoneto

fram

estone,peloidal

packston

eto

grainstone

Mod

erateto

high

Mostly

ingrow

thpo

sitio

n;hemispherical,lenticular,andfew

branching;

commonly

containrod-

shaped

stromatoporoids;sm

all

65

41

Low

erlim

estone

mem

ber

A1

Calcareou

smud

ston

eLow

Not

ingrow

thpo

sitio

n;hemispherical

andirregular;large

andsm

all

19

Table1.

Distribution,

orientation,

grow

thform

andsize

ofAgetolites

coralla

intheXiazhen

Formation(U

pper

Ordov

ician)

atZhu

zhai,Sou

thChina.A,A.yushan

ensis;B,A.raritabu

latus;

C,A.waicunensis;D,A.oculiporoides;E,A.maxima.


Beijing (Table 2). The holotypes of A. oculiporoidesand A. yushanensis are, however, unavailable and oneparatype (IV-3460) of A. yushanensis was not usedowing to inadequate preservation for precise morpho-logical comparison with our specimens. The typespecimens of A. multitabulatus Lin, 1960 and A.breviseptatus Lin, 1960 are inadequately preserved forinclusion in our study or could not be located and areprobably missing. All type specimens of species of Age-tolites and Agetolitella erected by Lin & Chow (1977)from the JCY region were supposed to be deposited inthe Institute of Geology, Chinese Academy of Geologi-cal Sciences, and later moved to the GeologicalMuseum of China, but we were unable to locate themat either institution, and they are probably missing.

The coralla of Agetolites are predominantly massive,domical or bulbous in growth form (Fig. 3A–I), butpyriform, tabular and even branching forms are alsopresent. They are composed of prismatic corallites withwell-developed septa that may alternate in length(Fig. 4A). Tabulae are mostly complete (Fig. 4B, arrows1–3) and flat (Fig. 4B, arrow 1), with a few concave orslightly convex cases (Fig. 4B, arrows 2, 3), but someare incomplete (Fig. 4B, arrow 4) or irregular (Fig. 4B,arrow 5). Mural pores are common (Fig. 4B, arrow P),usually connecting two or three corallites (Fig. 4A,arrows 2, 3) and rarely four to even five corallites(Fig. 4A, arrow 4). Common walls are slightly crenu-lated, with a median suture (Fig. 4A, B, arrow M).

Multivariate morphometricsA preliminary visual examination of the coralla led usto suspect that there is a great range of morphologicalvariation among the Agetolites specimens (Figs 4, 5). Inorder to incorporate these variations into systematics ofthe species, five corallites were selected for analysisamong the largest 10 percent of corallites (based ontheir tabularium area) in a transverse thin-section orpeel from the mature part of each corallum (Lee &Noble 1988). Five morphological characters were mea-sured for each corallite using an image-analysis system(image-Pro plus 5.0; Table 3; Fig. 6): tabularium area(V1) is defined as the area occupied by the polyp,excluding the space filled by the septa; wall thickness

(V2) is defined as the minimum thickness of the com-mon double wall between corallites; lengths of longsepta (V3) and short septa (V4) are averages for the dis-tance from the distal end to proximal end in the com-mon wall, based on measurements of the relatively longand short septa, respectively; and corallite perimeter(V7) is defined as the length of the median suture inthe common wall bounding the corallite. V6 is a countof the total number of septa. Three characters are deter-mined by calculations. V5 is a ratio of the length oflong septa (V3) to length of short septa (V4). V8 andV9 are ratios of corallite perimeter (V7) to the length oflong septa (V3) and short septa (V4), respectively. Araw-data matrix of 108 coralla plus six type specimensby nine morphological characters (V1–V9) was con-structed (see Appendix 1). Each coordinate is the aver-age value of the measurements of five corallitesselected from each corallum; the exceptions are fourcorallites from holotype IV-3454 and three corallitesfrom paratype IV-3465, owing to inadequate preserva-tion of other corallites. A multivariate analysis to dis-criminate closely related species and examine theirintra- and interspecific variations was performed basedon the principal-component score matrix obtained fromthe raw data set for the nine morphological characters.

Frequency histograms and correlation analysis of thenine characters for 114 coralla (including type speci-mens) were performed to examine the completeness ofthe raw data and relationships among the characters.Some of the characters are not independent, so aprincipal-component analysis was performed, and a clus-ter analysis was conducted using the first three principal-components score matrix. The unweighted pair-groupmethod using arithmetic average was employed to gen-erate a dendrogram of clusters of coralla, and each clus-ter was considered to represent a morphospecies. Theidentification of morphospecies was then determined byquantitative and qualitative comparison with the typespecimens that were available and suitable for study, andwith published descriptions and illustrations of otherdescribed species. Discriminant analysis was performedto test the statistical validity of the morphospecies andtype specimens recognized by cluster analysis. Finally,descriptive statistics and bivariate plots were performedto examine intra- and interspecific variation.

In order to trace morphological variations duringastogeny, serial transverse peels oriented perpendicularto the central growth axis were prepared from 12 cor-alla, with up to 403 sections per corallum. In addition,115 transverse and six longitudinal thin-sections of 115coralla were prepared to compare skeletal developmentand general morphology within and among the coralla.All transverse sections illustrated herein are oriented asthey appear viewed from above the corallum. Describedand figured specimens collected for this study aredeposited in the Palaeontological Collections, NanjingInstitute of Geology and Palaeontology, ChineseAcademy of Sciences in Nanjing (NIGP). Type

Specimen Identification Reference Designation

IV-3452 A. oculiporoides Lin (1960) ParatypeIV-3454 A. huangi Lin (1960) HolotypeIV-3463 A. yushanensis Lin (1960) ParatypeIV-3464 A. yushanensis Lin (1960) ParatypeIV-3465 A. yushanensis Lin (1960) ParatypeIV-3466 A. raritabulatus Lin (1960) Holotype

Table 2. Six type specimens of Agetolites species from theUpper Ordovician at Shiyanshan in the JCY triangle region ofSouth China, used for multivariate analysis. All specimens arein the collection of the Geological Museum of China in Beijing.


specimens with catalogue numbers having the prefix IVare in the collection of the Geological Museum ofChina in Beijing.

Frequency histograms

Frequency histograms of the nine morphological charac-ters for the studied coralla and type specimens confirmthat the overall range of morphological variation is

considerable (Fig. 7). The variation of four out of thenine characters is apparently continuous (V2, commonwall thickness; V3, length of long septa; V5, ratio ofthe length of long septa to length of short septa; V7,corallite perimeter); the Shapiro–Wilk test indicates thatthese characters are normally distributed at the signifi-cance level of 0.01. The variation of the other five char-acters does not deviate much from the normaldistribution, as suggested by their skewness. A further

Fig. 3. Field photographs of Agetolites in different stratigraphic intervals of the Xiazhen Formation at Zhuzhai, South China. A, Arrows point tosmall coralla in growth position on a grainstone surface in interval A2; B, C, Overturned large coralla on a calcareous mudstone surface in A1; D,E, Overturned coralla on a calcareous mudstone surface in A6; F, Small coralla on a grainstone surface in A8 (reefal facies); G, Arrows point totransported coralla of Agetolites with other colonial corals in A8; H, Arrow points to a corallum of Agetolites encrusted by a stromatoporoid inA8; I, Arrows point to small rod-shaped stromatoporoids encrusted by Agetolites in A2. Diameter of coin = 20 mm (A–C, I), 25 mm (F, H); scalebar on pencil = 40 mm (D); length of hammer = 26.5 cm (E); length of pencil = 13.5 cm (G).


analysis is, however, required to determine whetheronly one species contributes to the overall distribution.

Correlation and principal-component analyses

To examine the linear relationship between each pair ofmorphological characters, Pearson correlation coeffi-cients were calculated for the raw-data matrix (Table 4).The coefficient matrix shows that six out of the ninecharacters are strongly related to each other at the sig-nificance level of 0.01 using the t-test: tabularium area(V1), length of short septa (V4), number of septa (V6),corallite perimeter (V7), ratio of perimeter to the lengthof long septa (V8) and ratio of perimeter to the lengthof short septa (V9). These characters, except for thelength of short septa, are positively related to eachother. Among them, tabularium area and coralliteperimeter display the strongest positive relationship.

A principal-component analysis was conducted toconvert the original morphological characters into majorcomponents. Eigenvalues and their associated eigenvec-tors of the nine morphological characters were calcu-lated from the correlation matrix of the raw-data matrix,and then three principal components accounting for87.2% of total variance were extracted (Table 5). Thefirst principal component (Prin 1), which accounts for68.8% of the variance, is weighted heavily on six mor-phological characters: tabularium area (V1), length ofshort septa (V4), number of septa (V6), coralliteperimeter (V7), ratio of perimeter to the length of longsepta (V8) and ratio of perimeter to the length of shortsepta (V9; Table 5). Among these characters, the ratioof perimeter to the length of short septa possesses thehighest eigenvector of Prin 1. The second principalcomponent (Prin 2), which is weighted heavily on theratio of the length of long septa to the length of shortsepta (V5), accounts for 10.1% of the variance (Table 5).The third principal component (Prin 3), which isweighted heavily on the length of long septa (V3),accounts for 8.3% of the variance (Table 5).

Cluster analysis and identification of species

In order to distinguish species represented by the col-lected coralla, a cluster analysis was performed for thefirst three principal components (Fig. 8). The resultingdendrogram yields five major clusters separated at a rel-ative average distance of 15. Three of the five clustersare considered to correspond to Agetolites yushanensis,A. raritabulatus and A. oculiporoides, respectively,because they include the type specimens of those

Fig. 4. Transverse and longitudinal thin-sections illustrating morphological characteristics of Agetolites from the Xiazhen Formation at Zhuzhai,South China. A, A. raritabulatus, NIGP PZ162434; arrow M points to common walls with median suture; arrows 2–4 point to corner pores con-necting two, three and four corallites, respectively. B, A. raritabulatus, NIGP PZ162437; arrows M and P point to common walls with mediansuture and corner pores, respectively; arrows 1–5 point to complete, concave, convex, incomplete and irregular tabulae, respectively. Scale bar = 5mm.

Abbreviation Description Unit

V1 Tabularium area mm2

V2 Common wall thickness mmV3 Length of long septa mmV4 Length of short septa mmV5 Ratio of V3 to V4V6 Number of septa no.V7 Corallite perimeter mmV8 Ratio of V7 to V3V9 Ratio of V7 to V4

Table 3. Morphological characters used in this study ofAgetolites Sokolov, 1955.


species. The three paratypes of A. yushanensis are allwithin the same cluster. The holotype of A. raritabula-tus and the paratype of A. oculiporoides occur in differ-ent clusters, but the cluster containing A. oculiporoidesalso includes the holotype of A. huangi. Further com-parison of the latter two species supports the interpreta-tion that A. huangi is a junior synonym of A.oculiporoides (see Systematic palaeontology). Theremaining two clusters do not incorporate type speci-mens. Based on morphological similarity, coralla inthose two clusters are identified as A. waicunensis (Lin& Chow, 1977) and A. maxima (Lin & Chow, 1977),which were originally described from the middleSanqushan Formation (Upper Ordovician), 3.5 km westof Tashan, near Zhuzhai (Fig. 1A; see Systematicpalaeontology).

Discriminant analysis and descriptive statistics

In order to verify each of the five morphospecies, acanonical discriminant analysis using the original ninemorphological characters was performed. Two canonicaldiscriminant functions were obtained, which account for70.3% and 19.8% of the variance. The plot of the twofunctions (Fig. 9) demonstrates that three of the fivespecies are distinctly separated from each other: Ageto-lites yushanensis, A. waicunensis and A. maxima.Although partial overlapping is evident for the mor-phospace occupied by A. raritabulatus and A. oculi-poroides, they are regarded as separate species based ontheir different centroids.

Descriptive statistics of the nine morphologicalcharacters were calculated. The average value for most

V1 V2 V3 V4 V5 V6 V7 V8 V9

V1 r 1.000 –0.446 –0.539 –0.681* 0.494 0.662* 0.976* 0.912* 0.866*p <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001

V2 r 1.000 0.406 0.499 –0.409 –0.433 –0.407 –0.461 –0.524p <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001

V3 r 1.000 0.717* –0.273 –0.404 –0.516 –0.809* –0.721*p <0.001 0.003 <0.001 <0.001 <0.001 <0.001

V4 r 1.000 –0.840* –0.581 –0.676* –0.765* –0.914*p <0.001 <0.001 <0.001 <0.001 <0.001

V5 r 1.000 0.465 0.508 0.440 0.751*p <0.001 <0.001 <0.001 <0.001

V6 r 1.000 0.683* 0.638* 0.647*p <0.001 <0.001 <0.001

V7 r 1.000 0.912* 0.869*p <0.001 <0.001

V8 r 1.000 0.917*p <0.001

V9 r 1.000p

Table 4. Simple correlation matrix of nine morphological characters (V1–V9; see Table 3 for abbreviations) selected from 108 cor-alla of Agetolites from the Xiazhen Formation (Upper Ordovician) at Zhuzhai and six type specimens from the Upper Ordovicianat Shiyanshan, South China (see Table 2) [r, Pearson correlation coefficient; p, probability by t-test (H0: Rho = 0, Prob. > |r|)].*Values showing strong correlation between morphological characters at significance level 0.01 using the t-test.

Prin 1 Prin 2 Prin 3

V1 0.903* 0.252 0.242V2 – 0.591* 0.294 0.328V3 – 0.731* – 0.254 0.581*V4 – 0.900* 0.299 0.144V5 0.693* – 0.662* 0.169V6 0.741* 0.028 0.319V7 0.901* 0.250 0.292V8 0.939* 0.320 – 0.076V9 0.980* – 0.059 – 0.015Eigenvalue 6.189 0.912 0.747Proportion of variance 0.688 0.101 0.083Cumulative proportion of variance 0.688 0.789 0.872

Table 5. First three principal components (Prin 1–Prin 3) in principal-component analysis of 114 coralla and nine morphologicalcharacters of Agetolites (V1–V9; see Table 3 for abbreviations). *Values showing that V1–V9 are heavily weighted on Prin 1, V5on Prin 2 and V3 on Prin 3.


characters appears to be distinctly different from onespecies to another (Table 6). However, the ranges ofvalues for most characters partially overlap among thespecies (Fig. 10). Bivariate plots with r2 (r, Pearsoncorrelation coefficient) were prepared to estimate vari-ability of the characters among species and to aid indistinguishing the species (Fig. 11). For example, on aplot of tabularium area (V1) and corallite perimeter(V7; Fig. 11A), Agetolites yushanensis is distinctly sep-arated from A. maxima; the morphospaces occupied bythe other three species strongly overlap each other. Aplot of the ratio of the length of long septa to length ofshort septa (V5) and the ratio of perimeter to the lengthof short septa (V9) shows that A. yushanensis, A.raritabulatus and A. maxima are distinguishable fromone another (Fig. 11B). Based on the relationshipbetween corallite perimeter (V7) and the ratio ofperimeter to the length of long septa (V8), the five spe-cies are clearly distinguishable (Fig. 11C). However,based on the relationship between corallite perimeter(V7) and the ratio of perimeter to the length of shortsepta (V9), only A. yushanensis is distinctive(Fig. 11D). The descriptive statistical results demon-strate that A. yushanensis and A. maxima are morereadily distinguishable than the other three species.

Palaeoecological distribution ofAgetolitesIn order to examine the facies distribution of the spe-cies of Agetolites, each corallum was judged to beautochthonous or allochthonous according to the orien-tation and position in which it was found, its degree ofabrasion, and its relation to the enclosing sedimentarydeposit. The size and shape of coralla, the associatedfauna and growth relationships of the corals, especiallywith stromatoporoids, were also considered.

In the Xiazhen Formation, species of Agetolitesoccur in various facies represented by a wide range ofdepositional environments. The general association ofAgetolites, tabulate corals including halysitids and heli-olitids, solitary and colonial rugose corals, and stro-matoporoids in many beds probably indicates broadlysimilar ecological tolerances. However, rugose coralsare rather uncommon in the formation and stromato-poroids are more dominant in pure limestone reeffacies, where Agetolites and other corals are less com-mon to almost entirely absent.

Stratigraphic intervals in which the coralla of identi-fied Agetolites species were collected are summarized(Table 1). All coralla used in this study were collectedfrom 12 Agetolites-bearing intervals in sub-sections ZU1 and ZU 2 (Fig. 2). Sub-section ZU 3 is laterallyequivalent to the upper part of ZU 1, and the samespecies of Agetolites occur in the correlative strata (seeLee et al. 2012, figs 8, 9). Agetolites is not found inshale facies of the formation.

Agetolites

yushan

ensis

Agetolites

raritabu

latus

Agetolites

waicunensis

Agetolites

oculip

oroides

Agetolites

max

ima

Max

.Min.

Avg

.SD

Max

.Min.

Avg

.SD

Max

.Min.

Avg

.SD

Max

.Min.

Avg

.SD

Max

.Min.

Avg

.SD

V1

8.57

3.08

5.78

1.27

15.36

7.83

11.29

2.20

19.55

10.24

14.44

2.60

20.97

6.68

13.41

4.67

27.71

17.57

21.52

3.66

V2

0.32

0.22

0.26

0.02

0.29

0.20

0.24

0.02

0.29

0.16

0.23

0.04

0.31

0.23

0.27

0.03

0.28

0.18

0.23

0.02

V3

1.10

0.89

0.95

0.05

1.00

0.77

0.91

0.07

0.86

0.62

0.74

0.05

1.18

0.89

1.03

0.08

0.86

0.67

0.76

0.06

V4

0.69

0.46

0.55

0.06

0.44

0.24

0.36

0.06

0.43

0.22

0.32

0.06

0.54

0.30

0.43

0.08

0.43

0.29

0.33

0.04

V5

2.07

1.42

1.73

0.15

3.21

1.95

2.60

0.38

3.09

1.86

2.35

0.34

3.22

1.96

2.46

0.42

2.59

2.01

2.30

0.18

V6

18.75

15.40

16.41

0.83

20.00

16.00

18.04

0.91

20.00

17.00

18.22

0.83

20.00

17.20

18.37

0.82

21.00

18.00

19.17

1.05

V7

13.86

8.37

11.56

1.47

17.41

13.37

15.11

1.11

19.06

14.42

16.63

1.28

20.70

12.33

16.21

2.45

22.71

18.57

20.35

1.53

V8

14.21

9.09

12.19

1.31

19.93

14.23

16.78

1.83

28.03

18.86

22.76

2.69

23.28

12.30

15.83

3.12

33.90

23.09

27.08

3.52

V9

27.53

15.57

21.06

2.41

60.00

30.19

43.95

9.44

83.55

39.20

53.68

11.33

69.00

24.18

39.97

14.21

79.29

49.09

62.16

7.75

D4.41

2.66

3.68

–5.54

4.26

4.81

–6.07

4.59

5.29

–6.59

3.92

5.16

–7.23

5.91

6.48

–

Table6.

Descriptiv

estatisticsof

nine

morph

olog

ical

characters

(V1–V9;

seeTable

3forabbreviatio

nsandun

its)basedon

thecoralla

andtype

specim

ensof

five

speciesof

Agetolites

(max.,maxim

um;min.,minim

um;avg.,average;

SD,standard

deviation).For

comparisonwith

measurements

ofcorallite

diam

eter

repo

rted

inprevious

stud

ies,

Dis

avalueof

corallite

diam

eter

(mm)calculated

forconv

enienceas

V7/π(i.e.,assumingthecorallite

iscircular,which

isacloseapprox

imationforhexago

nalto

octago

nalcorallitesin

thesize

rang

eof

this

stud

y).


Three of the five species of Agetolites in theXiazhen Formation have restricted stratigraphicdistributions (Table 1). Agetolites waicunensis and A.maxima occur in the lower part of the formation(intervals A1–A6), whereas A. yushanensis is restrictedto the upper part (intervals A6–A12), with overlap of A.waicunensis and A. yushanensis only in interval A6. Onthe other hand, A. raritabulatus and A. oculiporoidesshow relatively longer stratigraphic ranges in the forma-tion.

On bedding surfaces within Agetolites-bearing inter-vals, the coralla can be grouped into three types ofoccurrences, listed in order of decreasing abundance:(1) coralla in growth position on calcareous mudstonelayers (interval A2 in sub-section ZU 2; Fig. 3A); (2)coralla mostly toppled or overturned on calcareousmudstone or marl layers (notably interval A1 in sub-section ZU 2 and A6 in ZU 1; Fig. 3B–E); and (3)transported coralla (interval A3 in sub-sections ZU 1and ZU 2, and A7–A12 in ZU 1; Fig. 3F, G) andabraded coralla (interval A8 in sub-section ZU 1 and itsequivalent in ZU 3).

Although the size and shape of Agetolites coralla areprimarily considered to be species-characteristic fea-tures, they also appear to be related to lithofacies. Largecoralla of A. waicunensis and A. maxima, usually domi-cal in form and up to 50 cm in diameter and 40 cmhigh, commonly occur in calcareous mudstone facies ofinterval A6 (Fig. 3E) in sub-section ZU 1 and intervalA1 (Fig. 3B) in sub-section ZU 2 (Table 1). Smallercoralla of A. raritabulatus and A. yushanensis, usuallyless than 15 cm in diameter and 10 cm high, with dom-ical, pyriform or, in some cases, branching forms, arecommon in reef or reef-related facies of interval A2(Fig. 3A) in sub-section ZU 2, and intervals A8(Fig. 3F) and A12 in sub-section ZU 1 (Table 1).Within these reefal intervals, selection of suitable sub-strates commonly resulted in coralla of Agetolitesencrusting stromatoporoids (Fig. 3I), together with stro-matoporoids encrusting Agetolites (Fig. 3H).

Two species, Agetolites waicunensis and A. maxima,commonly occur in fine-grained calcareous mudstonesof interval A6 in sub-section ZU 1, and interval A1 insub-section ZU 2, respectively, which are interpreted asrelatively low-energy deposits. Agetolites yushanensis iscommon in coarse-grained grainstone facies of intervalsA7 to A10 in sub-section ZU 1, representing moderate-energy deposits. Agetolites raritabulatus and A. oculi-poroides occur in strata that were deposited in a widerrange of conditions. These include lower-energy depos-its (A. raritabulatus in intervals A4, A5, A9 and A12of sub-section ZU 1; A. oculiporoides in interval A1 ofsub-section ZU 2, and A12 of ZU 1) and moderate-energy deposits (A. raritabulatus in intervals A2 andA3 of sub-section ZU 2, and A8 and A11 of ZU 1; A.oculiporoides in interval A2 of sub-section ZU 2, andA8 and A10 of ZU 1). The depositional energy leveland interactions with other sessile biota apparently were

important controlling factors in the distribution of thespecies of Agetolites.

Systematic palaeontologyPhylum CNIDARIA Hatschek, 1888Class ANTHOZOA Ehrenberg, 1834Subclass TABULATA Milne-Edwards & Haime, 1850Order FAVOSITIDA Wedekind, 1937Suborder FAVOSITINA Wedekind, 1937Family AGETOLITIDAE Kim, 1962

Agetolites Sokolov, 1955

Type species. Agetolites mirabilis Sokolov, 1955 fromthe ‘lower Llandoverian’ in the Chingiz Range of Kaza-khstan. Kim (1966), however, speculated that thedeposit from which the genus was reported by Sokolovis Upper Ordovician.

Emended diagnosis. Corallum hemispherical, nodular,pyriform or cylindrical, cerioid; corallites prismatic;common walls with sinuous median suture; septa alter-nately long and short thin plates with long axial partsspinose; tabulae horizontal, convex or concave, com-monly complete, in some cases incomplete or irregular;pores at or near angles between corallites, rounded andnumerous, commonly opening simultaneously into threeadjacent corallites, but may open into two, four or evenfive adjacent corallites, pores may occur rarely on facesof corallite walls.

Discussion. The above diagnosis largely follows Hill(1981, pp. F559, F560), but with additions to the state-ments regarding tabulae and pores based on the presentstudy.

According to Hill (1981, p. F561), Agetolitella isdistinguished from Agetolites by having septa that donot alternate in length and by having pores on thefaces of corallite walls and at or near the anglesbetween faces. Species from South China that wereassigned to Agetolitella by Lin & Chow (1977) havealternating long and short septa, as is characteristic ofAgetolites. Lin & Chow (1977) claimed that wall poresare present in all their species of Agetolitella. Basedon their illustrations, however, the species range fromhaving no wall pores, to rare wall pores in some cor-alla, to rare wall pores in all coralla. This suggestsintergradation between Agetolites and Agetolitella withrespect to the development of wall pores. In two ofthe species described below from Zhuzhai (A. wai-cunensis, A. maxima), our relatively large collectionsshow that rare wall pores occur in some, but not all,of the coralla. We do not consider the sporadic devel-opment of rare wall pores to justify the assignment ofsuch species to Agetolitella rather than Agetolites. Spe-cies from South China that were previously assignedto Agetolitella are herein transferred to Agetolites(Table 7).


Our study of material from South China calls intoquestion the distinction of Agetolitella and Agetolitesbased on the position of wall pores, raising thepossibility that the former genus is a junior synonym ofthe latter. As currently understood, however, the natureof septa (whether equal in length or alternating inlength) remains a diagnostic character distinguishingthese genera. This should be tested by examining intra-and interspecific variability in material from otherregions, including the type species Agetolitella primaKim, 1962 from the Upper Ordovician of Uzbekistanand Agetolites mirabilis. Such an analysis is beyond thescope of the present study.

Agetolites yushanensis Lin, 1960 (Fig. 5A, B)

1960 Agetolites yushanensis Lin, p. 58, pl. VII, fig. 1a,b, pl. VIII, figs 1a, b, 2a, b, pl. IX, fig. 1a, b.

1963 Agetolites yushanensis Lin; Lin in Yu & Zhang,p. 217, pl. 68, fig. 3a, b.

1977 Agetolites yushanensis Lin; Lin in Lin & Chow,p. 140, pl. 25, fig. 2a, b.

Material. Twenty-nine coralla (NIGP PZ162402–162419, 162421–162431) from intervals A6–A10 andA12 at sub-section ZU 1 (Fig. 2; Table 1) and onecorallum from talus at sub-section ZU 3 (NIGPPZ162420), Xiazhen Formation (Katian) at Zhuzhai,South China; paratypes IV-3463, IV-3464, IV-3465 (Lin1960) from the Sanqushan Formation (Upper Ordovi-cian) at Shiyanshan, South China.

Emended diagnosis. Corallites polygonal in transversesection. Species average of corallite perimeter 11.56mm, tabularium area 5.78 mm2, length of short septa0.55 mm, length of long septa 0.95 mm, ratio of aver-age length of long septa to length of short septa 1.73,number of septa 16.41. Wall pores absent.

Description. Corallites polygonal in transverse section,adult corallites mostly hexagonal, shape of offsetsrounded or triangular (Fig. 5A; Sun et al. 2014, fig.3a–n). Corallum averages of corallite perimeter 8.37–13.86 mm, tabularium area 3.08–8.57 mm2, length ofsepta 0.46–1.10 mm, ratio of average length of longsepta to length of short septa 1.42–2.07, number ofsepta 15–19, wall thickness 0.22–0.32 mm (Table 6).Septa well developed, some long septa reach centre ofcorallite with curved distal ends (Fig. 5A). Corner porescommonly open to three corallites simultaneously, nowall pores evident (Fig. 5A). Tabulae mostly incom-plete, subhorizontal, slightly convex or concave, a fewirregular (Fig. 5B).

Discussion. The examined type specimens ofAgetolites yushanensis from the Sanqushan Formationat Shiyanshan (paratypes IV-3463, IV-3464, IV-3465;see Lin 1960) are quite different from one another incorallite size (tabularium area, V1; corallite perimeter,V7; corallite diameter, D; Table 8), suggesting sub-stantial intraspecific variation. All the type specimensdescribed by Lin (1960) are similar in corallite size to

Location Species References

Shiyanshan, Tashan Agetolites oculiporoides Lin (1960), Yu & Zhang (1963), Lin & Chow (1977)Shiyanshan, Tashan A. huangi Lin (1960), Yu & Zhang (1963), Lin & Chow (1977)Shiyanshan, Tashan A. multitabulatus Lin (1960), Yu & Zhang (1963), Lin & Chow (1977)Shiyanshan, Tashan A. yushanensis Lin (1960), Yu & Zhang (1963), Lin & Chow (1977)Shiyanshan A. raritabulatus Lin (1960), Yu & Zhang (1963), Lin & Chow (1977)Shiyanshan, Tashan A. breviseptatus Lin (1960), Yu & Zhang (1963), Lin & Chow (1977)Shiyang A. grandiformis Lin & Chow (1977)Tashan A. triangulatus Lin & Chow (1977)Tashan A. intermedius Lin & Chow (1977)Tashan A. crassus Lin & Chow (1977)Tashan A. aquabilis Lin & Chow (1977)Huibu A. hemiagetolitoides Lin & Chow (1977)Yanrui, Shiyanshan, Tashan A. rariseptatus Lin & Chow (1977)Tashan A. sinensis Lin & Chow (1977)Yanrui A. paleofavositoides Lin & Chow (1977)Tashan Agetolitella tenuis Lin & Chow (1977)Tashan A. xiazhenensis Lin & Chow (1977)Tashan A. crassiseptata Lin & Chow (1977)Tashan A. tashanensis Lin & Chow (1977)Tashan A. concava Lin & Chow (1977)Tashan A. jiangshanensis Lin & Chow (1977)Tashan A. maxima Lin & Chow (1977)Tashan A. waicunensis Lin & Chow (1977)Yanrui A. gracilis Lin & Chow (1977)Tashan A. micropora Lin & Chow (1977)

Table 7. Fifteen species of Agetolites and ten species of Agetolitella reported previously from the Upper Ordovician at Shiyanshanand from the Sanqushan Formation (Upper Ordovician) at other locations in the JCY triangle region, South China (see Fig. 1A).Species assigned to Agetolitella by Lin & Chow (1977) are herein transferred to Agetolites.


Fig. 5. Transverse and longitudinal thin-sections showing four species of Agetolites from the Xiazhen Formation at Zhuzhai, South China. A, B,A. yushanensis, NIGP PZ162423; C, D, A. waicunensis, NIGP PZ162455; E, A. oculiporoides, NIGP PZ162487; F, A. oculiporoides, NIGPPZ162490; G, H, A. maxima, NIGP PZ162501, arrow in G points to wall pore; scale bar = 5 mm.


our coralla from the Xiazhen Formation at Zhuzhai(Tables 6, 8, 9). Lin (1960), Yu & Zhang (1963) andLin & Chow (1977) mentioned that long septa inA. yushanensis extend nearly to the centre of thecorallite and have bent distal ends, as also observedin our coralla (Fig. 5A). Based on the results of mor-

phometric analysis and qualitative morphologicalcomparisons, we identify the coralla from Zhuzhai asA. yushanensis.

In our collection, corallite size is quite variable. Inintervals A9 and A10, average corallite sizes (V1, V7)are larger than in A6–A8 and A12 (Table 9). In addition,

Species Stratigraphic intervalicinterval V1 V2 V3 V4 V5 V6 V7 V8 V9 D

A. yushanensis A12 5.42 0.25 0.95 0.54 1.76 15.98 11.50 12.09 21.33 3.66T 7.40* 0.27 1.01 0.66 1.55 17.20 13.86 13.69 21.15 4.41A10 6.72 0.26 0.99 0.57 1.73 16.67 12.72 12.92 22.39 4.05A9 6.99 0.24 0.94 0.63 1.49 17.07 12.70 13.50 20.06 4.04A8 5.23 0.26 0.92 0.52 1.78 16.57 10.45 11.37 20.18 3.33A7 5.15 0.27 0.92 0.51 1.80 17.00 10.25 11.16 19.88 3.26A6 6.08 0.28 0.89 0.58 1.54 16.60 12.30 13.82 21.21 3.92

A. raritabulatus A12 9.27 0.26 0.94 0.41 2.30 17.00 14.51 15.40 35.44 4.62A11 9.16 0.26 0.97 0.40 2.43 16.00 13.98 14.39 34.95 4.45A9 9.33 0.23 1.00 0.42 2.38 17.80 14.91 14.95 35.50 4.75A8 9.92 0.28 0.98 0.40 2.48 18.00 14.01 14.23 35.33 4.46A5 12.47 0.23 0.86 0.29 3.02 18.28 15.89 18.49 55.73 5.06A4 12.77 0.23 0.89 0.30 2.94 17.60 16.09 18.09 53.18 5.12A3 of ZU 2 8.97 0.26 0.78 0.31 2.65 18.63 13.81 17.64 46.80 4.40A2 12.99* 0.23 0.94 0.38 2.48 18.30 15.64 16.64 41.13 4.98

A. waicunensis A6 14.14 0.25 0.74 0.32 2.35 18.09 16.59 22.59 53.12 5.28A5 13.61 0.23 0.76 0.30 2.56 18.00 15.93 20.97 53.74 5.07A3 of ZU 1 18.06* 0.17 0.75 0.28 2.77 18.00 18.48 24.98 69.91 5.88A3 of ZU 2 15.28 0.21 0.69 0.31 2.25 18.49 16.92 24.53 55.63 5.39A2 12.54 0.23 0.80 0.37 2.19 18.20 16.00 20.04 43.88 5.09

A. oculiporoides A12 9.37 0.27 1.11 0.52 2.13 17.40 14.48 13.08 27.85 4.61A10 6.68 0.30 1.00 0.51 1.96 19.00 12.33 12.30 24.18 3.92A8 10.33 0.29 1.04 0.48 2.19 17.90 14.39 13.86 30.40 4.58A2 17.06 0.24 1.06 0.38 2.81 18.65 17.84 16.89 47.69 5.68A1 20.97* 0.26 0.89 0.30 2.96 18.00 20.70 23.28 69.00 6.59

A. maxima A5 18.72 0.28 0.80 0.33 2.41 18.00 18.77 23.59 56.88 5.97A3 of ZU 1 24.12* 0.25 0.85 0.41 2.05 18.50 21.39 25.27 51.92 6.81A3 of ZU 2 18.46 0.21 0.75 0.31 2.44 20.28 19.02 25.22 61.48 6.05A2 17.79 0.24 0.75 0.32 2.37 19.33 18.85 25.21 59.63 6.00A1 23.36 0.24 0.74 0.33 2.26 18.81 21.20 29.12 65.67 6.75

Table 9. Average values of nine morphological characters (V1–V9; see Table 3 for abbreviations and units) of the five species ofAgetolites in stratigraphic intervals of the Xiazhen Formation (Upper Ordovician) at Zhuzhai, South China (see Table 1, Fig. 2; T,talus). For each species, an asterisk under V1 indicates the stratigraphic interval where tabularium area is largest. For comparisonwith measurements of corallite diameter reported in previous studies, D is a value of corallite diameter (mm) calculated for conve-nience as V7/π (i.e., assuming the corallite is circular, which is a close approximation for hexagonal to octagonal corallites in thesize range of this study).

A. yushanensis A. raritabulatus A. oculiporoides A. huangiP. IV-3463 P. IV-3464 P. IV-3465 H. IV-3466 P. IV-3452 H. IV-3454

V1 6.21 4.47 5.54 7.83 9.11 12.51V2 0.30 0.32 0.31 0.25 0.31 0.28V3 1.00 0.91 0.94 0.86 0.95 1.09V4 0.53 0.53 0.55 0.44 0.43 0.49V5 1.87 1.71 1.69 1.95 2.20 2.24V6 16.50 15.60 15.67 19.00 18.00 19.25V7 12.15 9.23 12.27 13.37 14.99 15.65V8 12.20 10.15 13.10 15.52 15.85 14.33V9 22.87 17.38 22.13 30.19 34.91 32.13D 3.87 2.94 3.91 4.26 4.77 4.98

Table 8. Average values of nine morphological characters (V1–V9; see Table 3 for abbreviations and units) for six type specimensof Agetolites species previously reported from the Upper Ordovician at Shiyanshan in the JCY triangle region, South China (H.,holotype; P., paratype; see Table 2). For comparison with measurements of corallite diameter reported in previous studies, D is avalue of corallite diameter (mm) calculated for convenience as V7/π (i.e., assuming the corallite is circular, which is a closeapproximation for hexagonal to octagonal corallites in the size range of this study).


the length of long septa (V3) is greater in intervals A9,A10 and A12 than in A6–A8 (Table 9).

In a comparison of the average values of morpho-logical characters, Agetolites yushanensis is distin-guished by its smaller corallite sizes (V1, V7), fewersepta (V6) and lower ratios of perimeter to septallengths (V8, V9) than the other four species of Ageto-lites recognized in the Xiazhen Formation at Zhuzhai:A. raritabulatus, A. waicunensis, A. oculiporoides andA. maxima (Table 6). Tabularium area (V1) and ratiosof corallite perimeter to septal lengths (V8, V9) are lessvariable in A. yushanensis than in those four species(Fig. 10A, H, I). Agetolites minor Lin, 1963, asreported from the Upper Ordovician at Shanyang inShaanxi Province, North China (Lin 1963, p. 120), hascorallite sizes (diameter commonly 2.5–3 mm) and sep-tal lengths (0.75–0.9 mm) that are within the range ofA. yushanensis (Table 6). However, A. minor is distinctin having more septa (18–20; Lin 1963, p. 120) than A.yushanensis (V6; Table 6).

Agetolites raritabulatus Lin, 1960 (Fig. 4A, B)

1960 Agetolites raritabulatus Lin, p. 58, pl. IX,fig. 2a, b.

1963 Agetolites raritabulatus Lin; Lin in Yu & Zhang,p. 218, pl. 68, fig. 5a, b.

1977 Agetolites raritabulatus Lin; Lin in Lin & Chow,pp. 139, 140, pl. 25, fig. 3a, b.

Material. Nineteen coralla (NIGP PZ162432–162450)from intervals A2 and A3 at sub-section ZU 2, and A4,A5, A8, A9, A11 and A12 at ZU 1 (Fig. 2; Table 1),Xiazhen Formation (Katian) at Zhuzhai, South China;holotype IV-3466 (Lin 1960) from the SanqushanFormation (Upper Ordovician) at Shiyanshan, SouthChina.

Emended diagnosis. Corallites polygonal in transversesection. Species average of corallite perimeter 15.11mm, tabularium area 11.29 mm2, length of short septa0.36 mm, length of long septa 0.91 mm, ratio of aver-age length of long septa to length of short septa 2.60,number of septa 18.04. Wall pores absent.

Description. Corallites polygonal in transverse section,adult corallites hexagonal to octagonal (Fig. 4A).Corallum averages of corallite perimeter 13.37–17.41mm, tabularium area 7.83–15.36 mm2, length of septa0.24–1.00 mm, ratio of average length of long septa tolength of short septa 1.95–3.21, number of septa 16–20,wall thickness 0.22–0.29 mm (Table 6). Septa welldeveloped, some long septa reaching centre of corallitewith curved distal ends (Fig. 4A). Corner porescommonly open to three corallites simultaneously, nowall pores evident (Fig. 4A). Tabulae mostly complete,subhorizontal or slightly convex or concave, fewincomplete or irregular (Fig. 4B). Frequency of tabulaein longitudinal section varies within and between cor-

alla, average 11 to 15 per 5 mm vertically in sparsebands, 15 to 21 in dense bands (Fig. 4B).

Discussion. Corallite sizes reported in all the typespecimens of Agetolites raritabulatus from the San-qushan Formation at Shiyanshan (diameters 3–4 mmin Lin 1960, p. 58; 3–4 mm in Yu & Zhang 1963,p. 218; 2.5–4 mm in Lin & Chow 1977, p. 139) areslightly smaller than in our coralla from the XiazhenFormation at Zhuzhai. However, the value of corallitediameter (D) calculated in the present study for theholotype of A. raritabulatus (4.26 mm; Table 8) isclose to the range of average values for coralla instratigraphic intervals at Zhuzhai (4.40–5.12 mm;Table 9). Lin (1960), Yu & Zhang (1963) and Lin& Chow (1977) mentioned that long septa in A.raritabulatus stretch nearly to the centre of thecorallite and in a few cases have bent distal ends.However, such long septa are not commonly evidentin the type specimens or in our coralla (Fig. 4A).Compared with our specimens, the holotype of A.raritabulatus described by Lin (1960, p. 58) has manyfewer tabulae (3–5 tabulae per 5 mm in sparse bands,7–9 in dense bands). However, considering the vari-ability seen in our collection (Fig. 4B), spacing oftabulae is not considered to be a reliable criterion todiscriminate species. Based on the results of a mor-phometric analysis (which included the holotype) andqualitative morphological comparisons, we identify thecoralla from Zhuzhai as A. raritabulatus.

Corallite size is quite variable in our collection. Theaverage tabularium area (V1) and corallite perimeter(V7) are larger in intervals A2, A4 and A5 than in A3,A8, A9, A11 and A12 (Table 9). In addition, a higherratio of the length of long septa to length of short septa

Fig. 6. Schematic transverse section of Agetolites, showingmeasurement of five morphological characters used in this study(V1–V4, V7; see Table 3 for abbreviations).


(V5) is apparent in intervals A4 and A5 than in A2,A3, A8, A9, A11 and A12 (Table 9).

Agetolites raritabulatus is distinguished from A.yushanensis and A. maxima in average corallite sizes(Fig. 10A, G), and from A. waicunensis and A. oculi-poroides in the average length of long septa (Fig. 10C).This species has the highest average ratio of the lengthof long septa to length of short septa (V5), comparedwith the other four species of Agetolites recognized inthe Xiazhen Formation at Zhuzhai: A. yushanensis, A.waicunensis, A. oculiporoides and A. maxima (Table 6).Agetolites raritabulatus has a higher average value for thenumber of septa (V6) than A. yushanensis (Table 6).

Agetolites waicunensis (Lin & Chow, 1977) (Fig. 5C, D)

1977 Agetolitella waicunensis Lin & Chow, pp. 148,149, pl. 31, figs 1a, b, 2a, b.

Material. Thirty-two coralla (NIGP PZ162451–162482)from intervals A2 and A3 at sub-section ZU 2, and A2,A5 and A6 at ZU 1 (Fig. 2; Table 1), Xiazhen Forma-tion (late Katian) at Zhuzhai, South China.

Emended diagnosis. Corallites sub-polygonal to polygo-nal in transverse section. Species average of coralliteperimeter 16.63 mm, tabularium area 14.44 mm2, lengthof short septa 0.32 mm, length of long septa 0.74 mm,ratio of average length of long septa to length of shortsepta 2.35, number of septa 18.22. Very rare wall poresin some coralla.

Description. Corallites polygonal in transverse section,adult corallites hexagonal to octagonal or rarely withnine to 11 sides, shape of offsets rounded or rectangu-lar (Fig. 5C; Sun et al. 2014, fig. 6a–l). Corallumaverages of corallite perimeter 14.42–19.06 mm,tabularium area 10.24–19.55 mm2, length of septa0.22–0.86 mm, ratio of average length of long septato length of short septa 1.86–3.09, number of septa17–20, wall thickness 0.16–0.29 mm (Table 6). Septawell developed, relatively short (Fig. 5C). Cornerpores numerous, commonly open to three corallitessimultaneously (Fig. 5C); wall pores very rare insome, but not all, coralla. Tabulae mostly complete,subhorizontal or slightly convex or concave, fewincomplete or irregular (Fig. 5D).

Fig. 7. A–I, Frequency histograms of nine morphological characters (V1–V9; see Table 3 for abbreviations; units shown in parentheses) for 108coralla and six type specimens of Agetolites (avg., average; s.d., standard deviation; skew., skewness; no., number of coralla).


Discussion. The type specimens of Agetolitella wai-cunensis were collected from the Sanqushan Formation(Upper Ordovician), 3.5 km west of Tashan (Lin &Chow 1977). The corallite diameter reported for thosespecimens (3.5–5.2 mm; Lin & Chow 1977, pp. 148,149) tends to be smaller than that for our coralla fromthe Xiazhen Formation at Zhuzhai but overlaps theirrange (D, 4.59–6.07 mm; Table 6). The corallite wallthickness and number of septa are similar between thetypes and our collection. Lin & Chow (1977) men-tioned that the length of long septa in A. waicunensis(<0.88 mm) is less than half the corallite radius, whichis also the case in our coralla (V3, Table 6; Fig. 5C).Septa in the types alternate in length, as in our speci-mens. Figures of A. waicunensis (Lin & Chow 1977)show that wall pores are very rare, as in some of ourcoralla. As explained previously (Discussion under Age-tolites), species with alternating long and short septaand with sporadic development of rare wall poresshould be included in Agetolites, rather than Ageto-litella. Based on comparisons with the original descrip-tion and illustrations, the coralla described here areidentified as Agetolites waicunensis.

In our collection, corallite size is variable. The aver-age tabularium area (V1) and corallite perimeter (V7)are larger in interval A3 of sub-section ZU 1 than inA2, A3 of ZU 2, A5 and A6 (Table 9). In addition, thelength of long septa (V3) is lower in interval A3 ofsub-section ZU 2 than in A2, A3 of ZU 1, A5 and A6(Table 9).

Agetolites waicunensis is characterized by the short-est average septal lengths (V3, V4), in comparison withthe other four species of Agetolites recognized in theXiazhen Formation at Zhuzhai: A. yushanensis, A. rari-tabulatus, A. oculiporoides and A. maxima (Table 6).

Fig. 8. Cluster analysis of 108 coralla (identified by NIGPAS num-bers) and six type specimens (prefix IV) of Agetolites, based on thefirst three principal-components score matrix. An arrow on left side ofspecimen number indicates a type specimen (see Table 2). Dashed lineindicates recognition of five species at relative average distance 15between clusters.

Fig. 9. Plot of canonical discriminant scores of 108 coralla and sixtype specimens of Agetolites, using the first two canonical discrimi-nant functions (percent of variance in parentheses). Centroid coordi-nates: A. yushanensis (4.493, –0.664), A. raritabulatus (–0.394,1.897), A. waicunensis (–1.809, –1.342), A. oculiporoides (–0.592,3.738) and A. maxima (–4.223, –0.790).


The following species described from the SanqushanFormation (Upper Ordovician) at Tashan in the JCY tri-angle (Fig. 1A) have corallite sizes that are similar toA. waicunensis (Table 6): A. triangulatus Lin & Chow,1977 (corallite diameter 3.5–5 mm; Lin & Chow 1977,p. 138) and A. intermedius Lin & Chow, 1977 (corallitediameter 3.7–5 mm; Lin & Chow 1977, pp. 138, 139).However, A. waicunensis is distinct in having thinnercommon walls and more septa (V2, V6; Table 6) thanthose species: A. triangulatus (wall thickness 0.3–0.5mm, number of septa 16; Lin & Chow 1977, p. 138)and A. intermedius (wall thickness 0.4–0.6 mm, numberof septa 16; Lin & Chow 1977, pp. 138, 139).

Agetolites oculiporoides Lin, 1960 (Fig. 5E, F)

1960 Agetolites oculiporoides Lin, p. 56, pl. I, figs 1a±c,2a, b, pl. II, fig. 1a, b.

1960 Agetolites huangi Lin, pp. 56, 57, pl. II, fig. 2a, b,pl. III, fig. 2a–c.

1963 Agetolites oculiporoides Lin; Lin in Yu & Zhang,p. 216, pl. 68, fig. 2a, b.

1963 Agetolites huangi Lin; Lin in Yu & Zhang,pp. 216, 217, pl. 68, fig. 6a, b.

1977 Agetolites oculiporoides Lin; Lin in Lin & Chow,p. 136, pl. 22, fig. 1a, b.

1977 Agetolites huangi Lin; Lin in Lin & Chow,p. 137, pl. 23, fig. 1a, b.

Material. Nine coralla (NIGP PZ162483–162491) fromintervals A1 and A2 at sub-section ZU 2, and A8, A10and A12 at ZU 1 (Fig. 2; Table 1), Xiazhen Formation(Katian) at Zhuzhai, South China; paratype IV-3452(Lin 1960), IV-3454 (holotype of A. huangi Lin, 1960),both from the Sanqushan Formation (Upper Ordovician)at Shiyanshan, South China.

Emended diagnosis. Corallites polygonal in transversesection. Species average of corallite perimeter 16.21mm, tabularium area 13.41 mm2, length of short septa

Fig. 10. A–I, Variation of nine morphological characters (V1–V9; see Table 3 for abbreviations) in five species of Agetolites (each corallum isrepresented by an empty circle; solid rhombuses and lines represent averages and ranges, respectively, for each species). 1, A. yushanensis; 2, A.raritabulatus; 3, A. waicunensis; 4, A. oculiporoides; 5, A. maxima.


0.43 mm, length of long septa 1.03 mm, ratio of aver-age length of long septa to length of short septa 2.46,number of septa 18.37. Wall pores absent.

Description. Corallites polygonal in transverse section,adult corallites hexagonal to octagonal (Fig. 5E).Corallum averages of corallite perimeter 12.33–20.70mm, tabularium area 6.68–20.97 mm2, length of septa0.30–1.18 mm, ratio of average length of long septa tolength of short septa 1.96–3.22, number of septa 17–20,wall thickness 0.23–0.31 mm (Table 6). Corner porescommonly open to three corallites simultaneously, nowall pores evident (Fig. 5E). Tabulae mostly complete,subhorizontal or slightly convex or concave, a fewincomplete (Fig. 5F).

Discussion. Except for more variable shapes of coral-lites in Agetolites oculiporoides (5–8 sides) than in A.huangi (5–7 sides) and corallites that tend to be smaller

in A. oculiporoides (diameter 4–5.1 mm) than in A.huangi (diameter 4.5–6 mm), these two species as origi-nally described from the Sanqushan Formation atShiyanshan and Tashan are remarkably similar in othergeneral morphological characteristics (Lin 1960). In ourmorphometric analysis, the type specimens of both spe-cies are grouped with our coralla from the Xiazhen For-mation at Zhuzhai (Figs 8, 9). Differences in theaverage values of quantitative morphological characters(V1–V9) for the type specimens (Table 8) are consistentwith variability in the range of values for coralla fromdifferent stratigraphic intervals in the Xiazhen Forma-tion (Table 9). Agetolites huangi is, therefore, consid-ered to be a junior synonym of A. oculiporoides, andour coralla are identified as the latter species.

In our collection, corallite size is quite variable. Theaverage tabularium area (V1) and corallite perimeter(V7) are much larger in intervals A1 and A2 than in

Fig. 11. A–D, Selected bivariate plots of morphological characters (V1, V5, V7–V9; see Table 3 for abbreviations) for five species of Agetolites(r, Pearson correlation coefficient). Each point on a plot represents a corallum.


A8, A10 and A12 (Table 9). In addition, a higher ratioof the length of long septa to short septa (V5) is evidentin intervals A1 and A2 than in A8, A10 and A12(Table 9). Nevertheless, plots based on the average val-ues of morphological characters for individual corallaassigned to this species show that there is continuitywithin the overall ranges of variability (Fig. 10).

Agetolites oculiporoides is characterized by higheraverage common wall thickness (V2) and averagelength of long septa (V3) than the other species of Age-tolites recognized in the Xiazhen Formation at Zhuzhai:A. yushanensis, A. raritabulatus, A. waicunensis and A.maxima (Table 6). Intraspecific variation of tabulariumarea is higher in A. oculiporoides than in those fourspecies (Fig. 10A).

Agetolites maxima (Lin & Chow, 1977) (Fig. 5G, H)

1977 Agetolitella maxima Lin & Chow, p. 148, fig. 6a,b, pl. 31, fig. 3a, b.

Material. Eighteen coralla (NIGP PZ162492–162509)from intervals A1–A3 at sub-section ZU 2, and A3 andA5 at ZU 1 (Fig. 2; Table 1), Xiazhen Formation(Katian) at Zhuzhai, South China.

Emended diagnosis. Corallites sub-polygonal to polygo-nal in transverse section. Species average of coralliteperimeter 20.35 mm, tabularium area 21.52 mm2, lengthof short septa 0.33 mm, length of long septa 0.76 mm,ratio of average length of long septa to length of shortsepta 2.30, number of septa 19.17. Rare wall pores insome coralla.

Description. Corallites sub-polygonal to polygonal intransverse section, adult corallites hexagonal to octago-nal or even with nine or ten sides, shape of offsetsrounded or rectangular (Fig. 5G; Sun et al. 2014, figs4a–o, 5a–l). Corallum averages of corallite perimeter18.57–22.71 mm, tabularium area 17.57–27.71 mm2,length of septa 0.29–0.86 mm, ratio of average lengthof long septa to length of short septa 2.01–2.59, numberof septa 18–21, wall thickness 0.18–0.28 mm (Table 6).Corner pores commonly open to three corallites simulta-neously, wall pores rare in some, but not all, coralla(Fig. 5G, arrow). Tabulae mostly complete, subhorizon-tal or slightly convex or concave, few incomplete orirregular (Fig. 5H).

Discussion. The type specimen of Agetolitella maximawas collected from the Sanqushan Formation (UpperOrdovician), 3.5 km west of Tashan (Lin & Chow1977). The corallite diameter reported for the species(4.5–6 mm; Lin & Chow 1977, p. 148) tends to besmaller than that for our coralla from the Xiazhen For-mation at Zhuzhai (D, 5.91–7.23 mm; Table 6). Thecorallite wall thickness and number of septa are similarin the type and in our collection. Lin & Chow (1977)mentioned that the length of long septa in A. maxima(<0.9 mm) is less than two-fifths of the corallite radius,

which is also evident in our coralla (V3, Table 6;Fig. 5G). Septa in the illustrated type specimen alternatein length, as in our material. Rare wall pores occur inthe type and in some of our specimens. As explainedpreviously (Discussion under Agetolites), species withalternating long and short septa and with sporadicdevelopment of rare wall pores should be included inAgetolites, rather than Agetolitella. Based on compar-isons with the original description and illustrations, thecoralla described here are identified as Agetolitesmaxima.

In our collection, corallite size is quite variable. Ininterval A1 at sub-section ZU 2 and A3 at ZU 1, coral-lites are larger than in A2, A3 at ZU 2, and A5(Table 9). In addition, a lower ratio of the length oflong septa to length of short septa (V5) is evident ininterval A3 at sub-section ZU 1 than in A1, A2, A3 atZU 2, and A5 (Table 9).

Agetolites maxima is characterized by higher averagevalues of corallite size (V1, V7), number of septa (V6)and ratios of corallite perimeter to septal lengths (V8,V9) than the other species of Agetolites recognized inthe Xiazhen Formation at Zhuzhai: A. yushanensis, A.raritabulatus, A. waicunensis and A. oculiporoides(Table 6). Agetolites sinensis Lin & Chow, 1977, fromthe Sanqushan Formation at Tashan, has some morpho-logical characters that are similar to A. maxima: corallitediameters of 4–6 mm, wall thickness of 0.25 mm andsepta that are 0.67–1 mm long (Lin & Chow 1977,p. 143, pl. 27, fig. 1a–c). However, A. sinensis is dis-tinct in having sporadic development of septa.

Discussion and conclusionsIn the JCY triangle region of South China, species fromthe Sanqushan Formation that were assigned toAgetolitella by Lin & Chow (1977; see Table 7) areherein transferred to Agetolites (see Discussion underAgetolites in Systematic palaeontology). These specieshave alternating long and short septa, as characteristicof Agetolites, rather than septa of equal length as inAgetolitella. Lin & Chow (1977) reported that wallpores are present in all of the species, as would beexpected in Agetolitella. Their illustrations, however,show that the species range from having no wall pores,as would be expected in Agetolites, to rare wall poresin some coralla, to rare wall pores in all coralla.This suggests that the development of wall pores isintergradational between these genera. In two of thespecies described herein from the Xiazhen Formation atZhuzhai (A. waicunensis, A. maxima), rare wall poresoccur in some, but not all, of the coralla. In our opin-ion, the sporadic development of rare wall pores doesnot justify the assignment of such species to Agetolitellarather than Agetolites. The possibility that Agetolitella isa junior synonym of Agetolites should be tested in afuture study by examining intra- and interspecific vari-ability of septa (whether equal or alternating in length)


in material from other regions, including the type spe-cies of both genera.

Multivariate analysis was found to be effective indiscriminating species of Agetolites in this study. Acluster analysis was conducted based on the first threeprincipal components for nine morphological characters,employing the unweighted pair-group method witharithmetic average. Five morphospecies were recognizedfrom the cluster analysis. Their taxonomic validity asfive species was verified by discriminant analysis,descriptive statistics and bivariate plots, and the natureof intra- and interspecific variation of the nine morpho-logical characters was determined. The five specieswere identified as A. yushanensis Lin, 1960, A. raritab-ulatus Lin, 1960, A. waicunensis (Lin & Chow, 1977),A. oculiporoides Lin, 1960 and A. maxima (Lin &Chow, 1977). Lin (1960) stated that A. oculiporoidesand A. huangi Lin, 1960 are remarkably similar in gen-eral morphological characteristics, except for more vari-able shapes of corallites in A. oculiporoides than in A.huangi and smaller corallites in A. oculiporoides than inA. huangi. However, based on both qualitative examina-tion of corallite shape (adult corallites hexagonal tooctagonal) and quantitative examination of corallite size(tabularium area 6.68–20.97 mm2, corallite perimeter12.33–20.70 mm; Fig. 10A, G; Tables 6, 8, 9), theranges of morphological characteristics of A. oculi-poroides encompass those for A. huangi and it isimpossible to separate these species. Thus, A. huangi isregarded as a junior synonym of A. oculiporoides.

Coralla of Agetolites in the Xiazhen Formation atZhuzhai were preserved in various conditions, whichwere determined mainly by lithology and depositionalenergy level. Coralla preserved in growth position wereusually contained in limestone deposited under low- tomoderate-energy conditions. High-energy conditionsresulted in fragmentation and transportation of coralla.The size of corallites and septal length are consideredto discriminate between Agetolites species.

The apparent high diversity of Agetolites species inSouth China is partly a consequence of taxonomic split-ting by previous workers (e.g., Lin & Chow 1977), andis also considered partly a result of high speciation ratesand endemism of the agetolitids. Ordovician shallow-water, marine invertebrate communities in China tend tobe geographically restricted and highly endemic, assuggested by Rong et al. (1999) and demonstrated forbrachiopods (Zhan & Cocks 1998) and graptolites(Chen et al. 2009). In each of the five Agetolites speciesat Zhuzhai, morphological characters related to corallitesize, septal length and number of septa show somedifferences among stratigraphic intervals in the XiazhenFormation (Table 9). In general, A. yushanensis ischaracterized by relatively small corallite size and well-developed septa, whereas A. maxima has relatively largecorallite size. Agetolites waicunensis has comparativelyshort septal length, whereas A. raritabulatus has the

highest average ratio of the length of long septa tolength of short septa. Agetolites oculiporoides differsfrom the other species in having the highest intraspeci-fic variation of tabularium area (Figs 4, 5, 10).

Agetolites yushanensis commonly occurs in stratarepresenting a wide spectrum of environments, includ-ing low- to moderate-energy (intervals A6, A9 and A12in sub-section ZU 1) and moderate- to high-energyconditions (intervals A7, A8 and A10 in sub-sectionZU 1; Table 1). With its relatively small coralla andlonger septa, A. yushanensis appears to have beenstructurally stronger than the other four species. Thissuggests that it was capable of remaining intact underhigher-energy conditions, perhaps contributing to itspredominance over other species in the intervals thatinclude high-energy deposits. Among the five species ofAgetolites, A. raritabulatus was widely distributed in awide spectrum of environments, including both low- tomoderate-energy (intervals A4, A5, A9 and A12 in sub-section ZU 1) and moderate- to high-energy conditions(intervals A8 and A11 in sub-section ZU 1, and A2 andA3 in ZU 2). Agetolites raritabulatus appears to haveadapted well to different environments. Agetoliteswaicunensis, the most common species, was distributedin environments including low- to moderate-energy(intervals A5 and A6 in sub-section ZU 1) andmoderate- to high-energy conditions (intervals A2 andA3 in sub-section ZU 2). It appears to have beencapable of living in low- to high-energy conditions, butwas especially dominant in moderate- to high-energyconditions. Agetolites oculiporoides inhabited both low-to moderate-energy (interval A12 in sub-section ZU 1,and A1 in ZU 2) and moderate- to high-energy condi-tions (intervals A8 and A10 in sub-section ZU 1, andA2 in ZU 2). This species also occurred subordinate toA. yushanensis in moderate-energy deposits. However,it appears that in such conditions, A. oculiporoides wasdisadvantaged compared with A. yushanensis (intervalsA8 and A10 in sub-section ZU 1). Agetolites maximalived in a wide spectrum of environments, includingboth low- to moderate-energy (interval A5 in sub-section ZU 1, and A1 in ZU 2) and moderate- to high-energy conditions (intervals A2 and A3 in sub-sectionZU 2). With its relatively large coralla and short septa,A. maxima appears to have favoured low-energyenvironments in which it could remain intact.

AcknowledgementsFunding from the National Research Foundation of Korea,through research grant no. NRF-2013R1A2A2A01067612and NRF-2014K2A2A2000787, is acknowledged. We areindebted to Liming Guan (Beijing Snow Birch PetroleumTechnology Company Limited) for assistance duringfieldwork. We thank two anonymous reviewers andthe editor, Stephen McLoughlin, for providing helpfulcomments.


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Specimens V1 V2 V3 V4 V5 V6 V7 V8 V9

162402 6.0800 0.2811 0.8903 0.5800 1.5350 16.6000 12.3000 13.8156 21.2069162403 4.0200 0.2500 0.9077 0.4716 1.9247 16.0000 8.8600 9.7609 18.7871162404 6.2800 0.2900 0.9257 0.5545 1.6694 18.0000 11.6300 12.5635 20.9739162405 3.0800 0.2442 0.9211 0.4962 1.8563 16.0000 8.3700 9.0870 16.8682162406 6.2700 0.2900 0.9065 0.5988 1.5139 18.3300 11.9100 13.1384 19.8898162407 5.7200 0.2400 0.9422 0.4562 2.0653 16.0000 10.7500 11.4095 23.5642162408 5.0300 0.2400 0.9216 0.4883 1.8874 17.6700 10.3700 11.2522 21.2369162409 6.1500 0.2666 0.9122 0.5099 1.7890 16.0000 11.2500 12.3328 22.0631162410 4.7700 0.2500 0.9120 0.5421 1.6823 16.0000 9.6200 10.5482 17.7458162411 5.6200 0.2824 0.9200 0.5459 1.6853 16.0000 10.8600 11.8043 19.8938162412 6.4000 0.2458 0.8999 0.5787 1.5550 16.3300 11.8300 13.1459 20.4424162413 7.5800 0.2400 0.9783 0.6892 1.4195 17.8000 13.5600 13.8608 19.6750162414 6.9900 0.2300 0.9628 0.5660 1.7011 16.5000 12.6300 13.1180 22.3145162415 7.4800 0.2625 0.9437 0.5200 1.8148 16.0000 12.8500 13.6166 24.7115162416 7.2100 0.2800 1.0973 0.6200 1.7698 16.0000 13.3000 12.1207 21.4516162417 5.8300 0.2697 0.9734 0.5655 1.7213 16.7500 12.0800 12.4101 21.3616162418 7.1100 0.2600 1.0403 0.6400 1.6255 18.7500 13.0100 12.5060 20.3281162419 5.7100 0.2676 0.9050 0.5143 1.7597 16.0000 12.4200 13.7238 24.1493162420 7.4000 0.2700 1.0127 0.6554 1.5452 17.2000 13.8600 13.6862 21.1474162421 4.1500 0.2500 0.9195 0.4700 1.9564 16.0000 10.4300 11.3431 22.1915162422 5.4700 0.2300 0.9777 0.5000 1.9554 16.0000 11.3800 11.6396 22.7600162423 4.3900 0.2200 0.8947 0.5353 1.6714 16.0000 9.9600 11.1322 18.6064162424 3.4700 0.2500 0.9300 0.5800 1.6034 15.4000 9.0300 9.7097 15.5690162425 6.6700 0.2600 1.0087 0.6200 1.6269 16.0000 12.2700 12.1642 19.7903162426 4.7400 0.2300 0.9112 0.4899 1.8600 16.0000 11.2200 12.3134 22.9026162427 6.4800 0.2800 0.9800 0.6100 1.6066 15.8000 12.2400 12.4898 20.0656162428 8.5700 0.2600 1.0529 0.5939 1.7729 16.7500 13.5900 12.9072 22.8826162429 4.2700 0.2712 0.9136 0.5400 1.6919 16.0000 10.5500 11.5477 19.5370162430 5.5800 0.2500 0.9215 0.4755 1.9380 16.0000 13.0900 14.2051 27.5289162431 5.8700 0.2723 0.9400 0.5600 1.6786 15.8000 12.7600 13.5745 22.7857IV-3463 6.2100 0.2958 0.9960 0.5313 1.8746 16.5000 12.1500 12.1988 22.8684IV-3464 4.4700 0.3243 0.9095 0.5308 1.7135 15.6000 9.2269 10.1450 17.3830IV-3465 5.5443 0.3100 0.9365 0.5544 1.6892 15.6700 12.2712 13.1033 22.1342162432 12.7700 0.2265 0.8894 0.3025 2.9402 17.6000 16.0882 18.0888 53.1841162433 11.7700 0.2396 0.9118 0.3000 3.0393 18.0000 15.5000 16.9993 51.6667162434 14.3700 0.2300 0.9538 0.3000 3.1793 18.0000 16.7000 17.5089 55.6667162435 13.4600 0.2186 0.8240 0.3000 2.7467 18.6000 16.4200 19.9272 54.7333162436 10.5600 0.2200 0.7992 0.2500 3.1968 18.0000 15.0000 18.7688 60.0000162437 12.1900 0.2193 0.8235 0.2800 2.9411 18.8000 15.8500 19.2471 56.6071162438 9.9200 0.2775 0.9848 0.3966 2.4831 18.0000 14.0100 14.2262 35.3253162439 9.3300 0.2300 0.9976 0.4200 2.3752 17.8000 14.9100 14.9459 35.5000162440 9.1600 0.2630 0.9716 0.4000 2.4290 16.0000 13.9800 14.3886 34.9500162441 8.6100 0.2900 0.9284 0.3900 2.3805 18.0000 14.3200 15.4244 36.7179162442 9.9300 0.2368 0.9557 0.4303 2.2210 16.0000 14.7000 15.3814 34.1622162443 11.4707 0.2711 0.8654 0.3800 2.2774 18.4000 16.4100 18.9623 43.1842162444 11.8800 0.2500 0.9879 0.3900 2.5331 17.4000 14.6300 14.8092 37.5128162445 15.3600 0.2200 0.9078 0.3800 2.3889 18.0000 17.4100 19.1782 45.8158162446 13.5100 0.1998 0.9655 0.3600 2.6819 20.0000 14.9200 15.4531 41.4444162447 11.2200 0.2200 0.9774 0.3600 2.7150 18.0000 14.8200 15.1627 41.1667162448 14.4700 0.2127 0.9605 0.4155 2.3117 18.0000 15.6500 16.2936 37.6655162449 9.3700 0.2485 0.7943 0.3795 2.0930 19.0000 14.0000 17.6256 36.8906162450 8.5600 0.2629 0.7712 0.2400 3.2133 18.2500 13.6100 17.6478 56.7083IV-3466 7.8300 0.2500 0.8614 0.4428 1.9453 19.0000 13.3700 15.5212 30.1942162451 18.3700 0.1784 0.8100 0.3300 2.4545 18.0000 18.5700 22.9259 56.2727162452 17.7499 0.1699 0.6800 0.2200 3.0909 18.0000 18.3800 27.0294 83.5455162453 16.0400 0.2398 0.7689 0.2800 2.7461 18.0000 17.0100 22.1225 60.7500162454 10.6500 0.2507 0.7600 0.3058 2.4853 18.0000 14.7800 19.4474 48.3322162455 14.7200 0.2364 0.7600 0.3000 2.5333 18.0000 15.8200 20.8158 52.7333162456 13.8100 0.2107 0.7562 0.3100 2.4394 18.0000 15.8400 20.9468 51.0968162457 12.8300 0.2200 0.7511 0.2900 2.5900 18.0000 16.1800 21.5417 55.7931162458 12.7900 0.2900 0.7189 0.3009 2.3892 18.0000 15.6500 21.7694 52.0106162459 11.8400 0.2157 0.7000 0.2500 2.8000 18.8000 15.3700 21.9571 61.4800162460 12.8900 0.2616 0.6700 0.2300 2.9130 18.0000 15.6900 23.4179 68.2174162461 12.7300 0.2424 0.7800 0.2909 2.6813 18.0000 15.8300 20.2949 54.4173

(Continued)


Appendix 1. (Continued).

Specimens V1 V2 V3 V4 V5 V6 V7 V8 V9

162462 16.3100 0.2460 0.7128 0.2875 2.4793 18.0000 17.4700 24.5090 60.7652162463 16.1600 0.2400 0.8025 0.4300 1.8663 18.0000 19.0500 23.7383 44.3023162464 13.8800 0.2600 0.7740 0.3873 1.9985 18.0000 16.5000 21.3178 42.6026162465 16.7600 0.2584 0.6858 0.3268 2.0985 19.0000 17.2900 25.2114 52.9070162466 13.9000 0.2691 0.7834 0.3992 1.9624 17.0000 16.5000 21.0620 41.3327162467 13.1800 0.1918 0.8563 0.3900 2.1956 20.0000 16.5400 19.3157 42.4103162468 13.1000 0.1813 0.8108 0.3651 2.2208 18.0000 15.2900 18.8579 41.8789162469 14.5000 0.2372 0.7828 0.3150 2.4851 18.6000 16.7600 21.4103 53.2063162470 10.2400 0.2900 0.7600 0.3700 2.0541 17.2000 15.8000 20.7895 42.7027162471 11.6800 0.2695 0.7867 0.3982 1.9756 17.2000 15.6100 19.8424 39.2014162472 16.6400 0.1884 0.6500 0.2700 2.4074 20.0000 17.3700 26.7231 64.3333162473 19.3500 0.1635 0.6800 0.2800 2.4286 20.0000 19.0600 28.0294 68.0714162474 17.5519 0.2187 0.7014 0.3015 2.3264 19.3300 18.6700 26.6182 61.9237162475 11.6302 0.2829 0.7400 0.3500 2.1143 17.2000 14.8700 20.0946 42.4857162476 19.5500 0.1716 0.7500 0.3700 2.0270 20.0000 18.3100 24.4133 49.4865162477 17.6201 0.1800 0.6516 0.2312 2.8183 18.0000 17.9300 27.5169 77.5519162478 11.4200 0.2717 0.7426 0.3700 2.0070 17.7500 15.8000 21.2766 42.7027162479 15.5270 0.2117 0.6225 0.3002 2.0736 18.0000 17.1162 27.4959 57.0160162480 14.8171 0.2000 0.6931 0.2529 2.7406 17.5000 16.7041 24.1006 66.0502162481 12.9331 0.1993 0.6965 0.3751 1.8568 18.0000 15.8500 22.7566 42.2554162482 11.0267 0.2701 0.6915 0.3598 1.9219 17.6000 14.4200 20.8532 40.0778162483 10.6600 0.2900 1.0852 0.5400 2.0096 17.2000 14.4000 13.2694 26.6667162484 9.9900 0.2875 0.9948 0.4214 2.3607 18.6000 14.3849 14.4601 34.1360162485 6.6800 0.3000 1.0021 0.5100 1.9649 19.0000 12.3300 12.3042 24.1765162486 9.3700 0.2700 1.1073 0.5200 2.1294 17.4000 14.4800 13.0769 27.8462162487 20.9700 0.2555 0.8891 0.3000 2.9637 18.0000 20.7000 23.2820 69.0000162488 16.7800 0.2300 1.1762 0.4500 2.6138 18.6000 18.4500 15.6861 41.0000162489 18.0200 0.2600 0.9871 0.3400 2.9032 18.0000 18.3300 18.5695 53.9118162490 14.9100 0.2497 1.0474 0.4200 2.4938 20.0000 16.3900 15.6483 39.0238162491 18.5100 0.2312 1.0303 0.3200 3.2197 18.0000 18.1900 17.6551 56.8438IV-3452 9.1094 0.3068 0.9461 0.4295 2.2028 18.0000 14.9935 15.8477 34.9092IV-3454 12.5105 0.2800 1.0919 0.4870 2.2421 19.2500 15.6487 14.3316 32.1329162492 25.8300 0.2500 0.8311 0.3958 2.0998 19.0000 21.6700 26.0739 54.7499162493 22.4100 0.2400 0.8631 0.4300 2.0072 18.0000 21.1100 24.4583 49.0930162494 18.7200 0.2811 0.7956 0.3300 2.4109 18.0000 18.7700 23.5923 56.8788162495 20.5500 0.2402 0.6900 0.3144 2.1947 20.0000 20.0384 29.0412 63.7354162496 22.8800 0.2319 0.7347 0.2950 2.4905 19.2500 21.9200 29.8353 74.3051162497 21.4400 0.2400 0.7076 0.3491 2.0269 20.0000 20.7900 29.3810 59.5531162498 19.7900 0.2100 0.8371 0.3814 2.1948 20.0000 19.3300 23.0916 50.6817162499 27.7100 0.2329 0.6900 0.3233 2.1342 18.0000 22.6400 32.8116 70.0278162500 25.5800 0.2447 0.8201 0.3612 2.2705 18.0000 21.6200 26.3626 59.8560162501 27.3223 0.2529 0.6707 0.3152 2.1279 18.0000 22.7000 33.8452 72.0178162502 17.9900 0.2625 0.7996 0.3092 2.5860 18.0000 19.0400 23.8119 61.5783162503 26.9800 0.2280 0.6700 0.2864 2.3394 18.0000 22.7100 33.8955 79.2947162504 17.7866 0.2400 0.7478 0.3161 2.3657 19.3300 18.8500 25.2073 59.6330162505 19.0700 0.2100 0.7600 0.3136 2.4235 20.0000 19.0900 25.1184 60.8737162506 17.6870 0.2273 0.7296 0.3137 2.3258 20.4000 18.7000 25.6305 59.6111162507 19.5900 0.1814 0.7700 0.3200 2.4063 20.0000 19.5100 25.3377 60.9688162508 18.3700 0.2112 0.7700 0.3000 2.5667 20.0000 19.2200 24.9610 64.0667162509 17.5702 0.2400 0.7419 0.3000 2.4730 21.0000 18.5700 25.0303 61.9000

Appendix 1. Raw data matrix for Agetolites from South China: 108 coralla from the Xiazhen Formation (Upper Ordovician) at Zhuzhai (identifiedby NIGP PZ numbers) and six type specimens from the Upper Ordovician at Shiyanshan (prefix IV; Geological Museum of China) by ninemorphological characters (V1–V9; see Table 3 for abbreviations and units).


morphometrics and palaeoecology of the coral …...the strata containing agetolites were reported to...

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