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Remarks on the pattern of septal insertion in rugosecoralsGuangxu Wang, Ian G. Percival & Rongyu LiPublished online: 25 Feb 2015.

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Remarks on the pattern of septal insertion in rugose corals

GUANGXU WANG, IAN G. PERCIVAL and RONGYU LI

WANG, G., PERCIVAL, I.G. & LI, R., XX.XXXX.2015. Remarks on the pattern of septal insertion in rugose corals. Alcheringa 39, xxx–xxx. ISSN0311-5518

Well-preserved specimens of the Hirnantian (latest Ordovician) rugose coral Lambeophyllum? corniculum He from the Yangtze Platform of SouthChina, clearly show how catasepta (minor septa) are inserted, confirming the model proposed by Weyer in the 1970s. Our observations indicate theinsertion of counter lateral septa and their neighbouring catasepta on the counter side takes place in exactly the same manner as that of the subse-quent metasepta and catasepta. We propose abandoning the use of the term counter lateral septa. Therefore, exclusion of the pair of counter lateralsepta reduces the number of protosepta from six to four.

Guangxu Wang [gxwang@nigpas.ac.cn], State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology,Chinese Academy of Sciences (CAS), 39 East Beijing Road, Nanjing 210008, PR China; Ian G. Percival [ian.percival@trade.nsw.gov.au],Geological Survey of New South Wales, 947–953 Londonderry Road, Londonderry, NSW 2753, Australia; Rongyu Li [lir@brandonu.ca],Department of Geology, Brandon University, Manitoba R7A 6A9, Canada. Received 7.11.2014; revised 13.1.2015; accepted 14.1.2015.

Key words: rugose corals, septal insertion, major septa, minor septa, catasepta, protosepta, metasepta.

RUGOSE CORALS are characterized by their serialseptal insertion in four quadrants (Hill 1981, Scrutton1997). However, uncertainty remains regarding severalspecific aspects of septal insertion, particularly how theminor septa (now known as catasepta, sensu Ezaki1989) are inserted. Resolution of this issue has signifi-cant implications for the ontogeny of rugosans (e.g., thenumber of protosepta), and their phylogenic relation-ships with other coral groups (Oliver 1980, 1981).

Oliver (1980, 1981), Hill (1981) and Scrutton(1997) reviewed the various models proposed for cata-septa insertion. The arguments distinguishing thesemodels stem from the fact that evidence for differingpatterns of septal insertion has been obtained indirectly,from serial sections, calices and septal grooves on theepitheca. Some authors (e.g., Weyer 1972b, 1974, 1997,2007, Oliver 1980, 1981, Scrutton 1997) have sug-gested that septal grooves may play a significant role inunderstanding septal insertion, because septal expressionmay be unclear or misleading in thin-sections or calices.Fedorowski (1991), however, argued that septal groovesdo not provide useful evidence for septal insertion,because they are normally missing from the tips ofrugosans, and because mismatches in the appearance ofgrooves and septa are present. Therefore, one solutionto this debate lies in the study of well-preservedspecimens of early rugose corals that lack horizontalelements, allowing the pattern of catasepta insertion tobe examined directly.

Silicified specimens of the Late Ordovician rugosanLambeophyllum? from the Yangtze Platform of SouthChina satisfy these criteria. They clearly show howcatasepta are inserted and provide the basis for discus-sion concerning the concept of metasepta and thenumber of protosepta.

Historical reviewKunth (1869) first established the general pattern ofseptal insertion in rugose corals, arguing that metaseptaand catasepta appeared alternately in each of four quad-rants in a cardinal direction (Fig. 1A). Weyer (1974),however, postulated a different model, referred to asretro-alternate insertion, in which each metaseptum firstproduced a new cataseptum on its counter side, andthen a new metaseptum on its cardinal side (Fig. 1B). Itis worth noting that long before Weyer’s proposal,Faurot (1909) proposed a similar pattern, which unfor-tunately has been largely overlooked by most subse-quent workers. Vollbrecht (1928) interpreted a newmetaseptum to arise on the cardinal side of an existingmetaseptum and then a cataseptum to develop on thesame side in the loci between these two metasepta(Fig. 1C). However, subsequent observations have notsupported this idea (Weyer 1972b, 1974, 1997, 2007,Hill 1981, Fedorowski 1991, Scrutton, 1997). In afourth model (Fig. 1D) designated the Zaphrentis-typeof Hill (1935), the appearance of catasepta in rapid suc-cession has been explained by septal stunting and isnow considered to be merely a modification or variationof the normal plan (Hill 1981, Scrutton 1997).© 2015 NSW Government

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Current uncertainty, therefore, concerns only the firsttwo of these models. Some authors regard the latter(proposed by Weyer 1974) as the only type (Oliver1980, 1981), whereas Fedorowski (1991) consideredKunth’s (1869) pattern as the most generalized andWeyer’s scheme as one variant of that.

Materials and methodsSilicified specimens identified as Lambeophyllum? cor-niculum He, 1978 were extracted using dilute hydro-chloric acid (ca 10%) from limestone of the uppermostKuanyinchiao Formation at Hetaowan and Shichang,Renhuai, northwestern Guizhou province in southwestChina (Fig. 2A). This coral is associated with anotherrugosan species Paramplexoides breviseptatum Wang &Zhan, 2014, brachiopods of the Dalmanella testudina-ria-Dorytreta longicrura Community, gastropods andcrinoid debris, with an age constrained to the lateHirnantian (Rong & Li 1999). According to Rong & Li(1999), the lower–middle Kuanyinchiao Formation con-tains the Hirnantia brachiopod fauna, whereas the over-lying basal Lungmachi Formation yields graptolitescharacteristic of the Normalograptus persculptus orParakidograptus acuminatus biozones of latest Ordovi-cian age (Fig. 2B).

Although development of catasepta is variable,among the dozen specimens of Lambeophyllum? cornic-ulum examined, most clearly show the pattern of septalinsertion in the uppermost part of their calices wherenew septa appear. Two specimens, in which the patternof catasepta insertion is best shown in all four insertionpoints, were chosen for this study (Fig. 3A, J).

In the subsequent discussion regarding the terminol-ogy of rugosan septa, we adopt the view of Weyer(2007) who differentiated between morphological terms(i.e., major and minor septa) and morphogenetic terms(i.e., protosepta, metasepta and catasepta). Note alsothat we regard the thicker of the two new septa formedby splitting as the ‘parent’. Specimens are deposited inthe Nanjing Institute of Geology and Palaeontology(NIGP).

Insertion of catasepta inLambeophyllum? corniculum He, 1978In each of the two studied specimens, there are fourinsertion points where new septa are serially inserted ina cardinal direction, i.e., one on each side of the cardi-nal septum (Fig. 3D, K), and one on the counter side ofeach alar septum (Fig. 3E, G, L, M).

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Fig. 1. Schematic diagrams showing possible patterns of insertion of catasepta (minor septa) on the inner wall of solitary rugose corals. A, Alter-nate insertion of metasepta and catasepta, according to Kunth’s rule; B, Retro-alternate insertion of catasepta, deduced from Weyer (1974); C,Retro-alternate insertion of catasepta, modified from Vollbrecht (1928); D, Zaphrentis-type insertion of catasepta, according to Hill (1935). Notethat only one cardinal quadrant is shown. C, cardinal septum; A, alar septum; ml–3, metasepta in order of insertion; cl–3, catasepta in order ofinsertion. Arrows indicate the direction in which new septa arise.

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At the four insertion points of each specimen, vari-able development of the youngest septum can beobserved. In general, two types of distinctive insertionstages are recognized. The first type is a new metasep-tum that arises from a pre-existing metaseptum on itscardinal side (Fig. 3C, H). The other type is a catasep-tum that initially projects from the counter side of apre-existing metaseptum (Fig. 3F).

Both types jointly reflect the general pattern of sep-tal insertion of catasepta, which can be readily recog-nized based on our specimens. Each metaseptum firstdeveloped a new cataseptum on its counter side, andthen a new metaseptum appears on its cardinal side.

DiscussionOur observations verify the pattern of catasepta inser-tion postulated by Weyer (1974), which is also referredto as retro-alternate insertion (Weyer 1972b, Oliver1980, 1981) or Weyer’s model (Hill 1981). This pro-posal has considerable merit as a logical explanation forthe distinction between metasepta and catasepta, inwhich a new cataseptum appears on the counter side ofa preceding metaseptum, whereas a new metaseptumdevelops on the cardinal side. Apart from Lambeophyl-lum? examined in this study, another convincing exam-ple of this pattern is the late Silurian–Early DevonianSutherlandinia Weyer, 1972a (Weyer 1974). For thisreason, we regard Weyer’s (1974) model as the mostgeneral one, or probably the only type, rather thanmerely a variant as suggested by Fedorowski (1991).

This pattern of catasepta insertion as verified fromthe specimens of Lambeophyllum? also sheds light onthe concept of metasepta and the number of protosepta,

which will be discussed below. Consideration of thesepoints necessitates reinterpretation of the generalsequence of septal insertion in rugosan corals (Fig. 4).

Definition of metasepta

The term metasepta was introduced by Duerden (1902)only for major septa following the insertion of protosep-ta, based on the assumption that rugosan major septa(i.e., protosepta and metasepta) correspond to entosepta,and the minor septa (i.e., catasepta) correspond to exo-septa in comparison with living corals. Although thisview has been widely accepted (e.g., Hill 1981,Scrutton 1997), other authors have argued that metasep-ta developed from catasepta (according to Kunth’s law),thus implying that all septa that developed followingthe insertion of protosepta should be included in themetasepta (e.g., Flower 1961, Wright 1969).

By applying Weyer’s model of catasepta insertion(based on evidence from Lambeophyllum? from SouthChina), it is easy to make a distinction between meta-septa and catasepta not only theoretically but practically,as noted above. Thus, we agree with Weyer (2007) thatthe term metasepta should refer only to the major septafollowing the insertion of protosepta, and that the mor-phogenetic term catasepta, introduced by Ezaki (1989),should refer to minor septa (a morphological term).

Number of protosepta

Although almost all coral workers accept that the cardi-nal and counter septa are protosepta, the total numberof protosepta recognized by various authors varies fromtwo (i.e., cardinal and counter septa; Hudson 1935,

(A) (B)Fig. 2. Locality (A), shown by black triangles and stratigraphic occurrence (B), indicated by arrows, of Lambeophyllum? corniculum He, 1978studied in this paper.

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Wright 1969, Fedorowski 1997, 2009a, b), to four (withan additional two alar septa; Gordon 1906, Brown1909, Jull 1965, Ezaki 1989, Ezaki & Yasuhara 2004,2005) or six (including also a pair of counter lateralsepta; Duerden 1902, Carruthers 1906, Hill 1956, 1981,Scrutton 1997). Thus, there are actually two basic ques-tions involved, both of which are reviewed below.

Should the two alar septa be considered protosepta?Hudson (1935) first suggested that only the cardinal andcounter septa be considered protosepta, probably in

view of the close relationship of the alar septa to subse-quently developed metasepta as discussed by Wright(1969). This concept was further supported byFedorowski (1997, 2009a, b), who assumed that theentocoelic arrangement of mesenteries probably corre-sponded to the alar sclerosepta of living Scleractinia.However, as demonstrated herein, there is a clear dis-tinction between the alar septa (together with the cardi-nal and counter septa) and subsequent metasepta interms of the manner of septal insertion. In contrast tothe alar septa, insertion of metasepta commonly takes

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(J)Fig. 3. Pattern of insertion of catasepta in Lambeophyllum? corniculum He, 1978. A–H, NIGP 161475; A–B, Oblique lateral and calical views ofthe corallum; C, H, Diagrams showing the first type of the youngest septum, a new metaseptum that arises from a pre-existing one on its cardinalside (redrawn from Fig. 3D, G respectively); D, View of portion of calyx showing septal insertion on both sides of cardinal septum; E, G, View ofportion of calyx showing insertion of new metasepta and catasepta on the counter side of each alar septum in the counter quadrant; F, Diagramshowing the other type of the youngest septum, a cataseptum that first projects from a pre-existing metaseptum on its counter side (redrawn fromFig. 3E). I–M, NIGP 161476; I–J, Calical and oblique lateral views of the corallum; K, View of a portion of calyx showing septal insertion onboth sides of cardinal septum; L–M, View of a portion of calyx showing insertion of new metasepta and catasepta on the counter side of each alarseptum in the counter quadrant. Notation as in Fig. 1. Black triangles indicate septal insertion points. Numbers before m/c represent the insertionorder of metasepta and catasepta in insertion points. Note that only the first four septa (cardinal, counter and a pair of alar septa) are consideredhere as protosepta, and we advocate that the term ‘counter lateral septa’ (commonly symbolized as KL) be abandoned, as these represent the firstpair of metasepta (m1).

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place with retro-alternate insertion of catasepta. More-over, the alar septa played an important role in typicalseptal arrangement in rugose corals, which also warrantstheir recognition as protosepta.

Should the pair of counter lateral septa be recognizedas protosepta? The principal argument for inclusion ofa pair of counter lateral septa as protosepta is the pres-ence of a clear six-septal stage in the early growth ofcorallites (Duerden 1902, Carruthers 1906, Scrutton1983). This is not convincing because the appearanceof counter lateral septa following the insertion of thefirst four septa can be regarded as a result of accelerateddevelopment (e.g., Gordon 1906, Brown 1909).Scrutton (1997) argued that the absence of cataseptabetween the counter septum and a pair of counter lateralsepta in many cystiphyllids may serve as evidence sup-porting inclusion of the latter as protosepta. In fact, thestunting phenomenon of catasepta seems to provide asufficient explanation for this (Hill 1981). On the otherhand, the absence of catasepta in these loci is probablynot common in other rugosans. In Lambeophyllum?, forexample, catasepta are clearly present in the locibetween counter and counter lateral septa (Fig. 3B, I).Other well-illustrated examples from the literatureinclude Lambeophyllum profundum (Conrad, 1843)from the Blackriveran (upper Sandbian) of North Amer-ica (Brown 1909, p. 55, figs 1–5), Muenstraia franco-nica Weyer, 2001 from the late Silurian of Germany(Weyer 2001, p. 78, fig. 1), the Early Devonian M.thuringica Weyer, 2001 from Germany and Morocco(Weyer 2001, p. 78, figs 2, 3), and several species oflate Palaeozoic zaphrentoid genera including CaniniaMichelin, 1840 (in Gervais 1840), Ufimia Stuckenberg,1895, and Zaphrentites Hudson, 1941 (Fedorowski2010, p. 376, fig. 1).

On the contrary, according to Flügel (1975) andFedorowski (1991), counter lateral septa resemble thesubsequent metasepta in all aspects, and should beconsidered the first pair of metasepta in the counterquadrant. In our interpretation (and in accord withWeyer’s model), the first catasepta in the counter quad-rant appear to ‘split’ from the counter lateral septa,hence there is no significant difference between thecounter lateral septa and subsequent metasepta in termsof the insertion of their closely related neighbouringcatasepta. Thus, we fully support the exclusion of thepair of counter lateral septa from the four protoseptaand further suggest that the traditional concept of coun-ter lateral septa be abandoned (Fig. 4).

Concluding remarksThe pattern of insertion of catasepta is clearly expressedon the inner wall of the calyx of the well-preservedHirnantian rugosan Lambeophyllum? corniculum He,1978 from the Renhuai area of South China. Twoprincipal points may be summarized as follows:

(1) Retro-alternate insertion of catasepta (=minor septa)postulated by Weyer (1974) is supported by ourobservations. This allows a clear distinction bothpractically and theoretically between metasepta andcatasepta, indicating that Weyer’s model is probablywidely applicable to all rugosans (and may be theonly type). Such a distinction also supports the con-cept of metasepta proposed by Duerden (1902),who restricted the term to the major septa devel-oped after insertion of the protosepta, and furtherunderpins the introduction by Ezaki (1989) of themorphogenetic term catasepta as a substitute forminor septa.

(2) Confirmation of Weyer’s model also allows theseparation of protosepta and metasepta, which canbe distinguished by whether the latter are accompa-nied by catasepta. It seems clear that only the firstfour protosepta (cardinal, counter and the pair ofalar septa) are valid, and that the counterlateral septa should be regarded more properly asmetasepta.

AcknowledgementsRong Jiayu of the Nanjing Institute of Geology andPalaeontology (NIGP) kindly provided the studiedspecimens. Zhan Renbin (NIGP) reviewed an earlierversion of the manuscript. Thanks to Anthony Wright,Gregory Webb, Dieter Weyer and Stephen McLoughlinfor helpful comments, which have greatly improved thispaper. Financial support for this study is from theNational Natural Science Foundation of China[41221001, 41290260 and J1210006] and the State Key

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Fig. 4. Idealized sequence of septal insertion in rugose corals. Nota-tion as in Fig. 1. Note that only the first four septa (cardinal, counterand a pair of alar septa) are considered here as protosepta and that theterm ‘counter lateral septa’ is abandoned (see text for discussion).

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Laboratory of Palaeobiology and Stratigraphy. IanPercival publishes with permission of the Director, Geo-logical Survey of New South Wales. This paper is acontribution to IGCP Project 591—The Early to MiddlePaleozoic Revolution.

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