Geological Society, London, Special Publications Parasites in Emsian-Eifelian Favosites (Anthozoa, Tabulata) from the Holy Cross Mountains (Poland): changes of distribution within colony M. K. Zapalski Geological Society, London, Special Publications 2009; v. 314; p. 125-129 doi:10.1144/SP314.6
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Parasites in Emsian –Eifelian Favosites (Anthozoa, Tabulata) from the Holy Cross Mountains (Poland): changes of distribution within colony M. K. ZAPALSKI Faculty of Geology, Warsaw University Z˙wirki i Wigury 93, 02– 089 Warszawa, Poland and Laboratoire de Pale´ontologie stratigraphique FLST and ISA, UMR 8157 ‘Ge´osyste`mes’ du CNRS. 41, rue du Port, 59046 Lille cedex, France Present address: Institute of Paleobiology, PAS, Twarda 51/55, 00-818 Warszawa, Poland (e-mail: [email protected]
) Abstract: Organisms of unknown biological affinities, assigned to the genus Chaetosalpinx, are known to infest Palaeozoic tabulate corals and stromatoporoids. Analysis of distribution of these parasites, performed on Emsian– Eifelian material of Favosites goldfussi (Anthozoa, Tabulata) from the Northern Region of the Holy Cross Mountains (Poland), shows that parasites were absent in the early astogenetical stages, and that during astogeny both the absolute number of parasites per colony and the number of parasites per polyp were increasing. The latter can reach 2.7 parasites per polyp. Preferred settling places are in corallite corners (junction of three individuals), but dense infestation also produced settlement in the corallite walls (between two individuals). Probable causes of the increase are insufficient protection by host’s cnidae, insufficient immune system response, and parasite ability to adapt to the host’s defences.
Endobionts belonging to the genus Chaetosalpinx (and other closely related genera such as Helicosalpinx) inhabited coralla of various tabulate corals (Tapanila 2005). They were originally considered as their commensals (e.g. Oekentorp 1969), but Stel (1976) suggested that they may be parasites. A recent analysis (Zapalski 2004, 2007) has shown that the relation between them and their hosts was parasitic rather than commensal. Howell (1962) proposed placing them among serpulids, but the biological affinity of these endobionts remains unknown. Although these organisms were often described and illustrated (e.g. Oekentorp 1969; Plusquellec 1968a, b; Tapanila 2002), very little attention was paid to their distribution within the host’s corallum. Tapanila (2004) analysed Ordovician Helicosalpinx infesting Calapoecia and Columnopora tabulates, and stated that there is no particular pattern in the distribution of the endobionts within the host’s corallum. This paper attempts to answer the question of whether the number of parasites increased in relation to the number of individuals during astogeny, or remained constant. Such a study has never been undertaken before. The increase of the number of parasites in relation to the number of polyps could be an effect of insufficient protection by host nematocysts, and/or insufficient immune system response. This analysis uses Emsian–Eifelian material of Favosites goldfussi d’Orbigny infested by Cheatosalpinx ferganensis Sokolov from the northern part of the Holy Cross Mountains, Poland.
Material and methods The analysed material consists of three coralla of Favosites goldfussi infested by Chaetosalpinx parasites (infested coralla are rare and therefore the material is limited). They come from the vicinity of Grzegorzowice in the Łysogo´ry Region (Northern Region) of the Holy Cross Mountains (for a map see Zapalski 2005; for description of this material of Chaetosalpinx see Zapalski 2007). The Emsian –Eifelian mudstones of Grzegorzowice Beds (serotinus–patulus conodont zones according to Malec & Turnau 1997; see also Halamski & Racki 2005) contain numerous rugose and tabulate corals (Stasin´ska 1954, 1958; Zapalski 2005) and other macrofossils (Pajchlowa 1957). Three specimens are analysed here. Two were collected in the late 1940s to early 1950s and used by Stasin´ska (1958) for taxonomic work on favositids; the third was collected by the present author in summer 2006. The collection of A. Stasin´ska is dispersed, and therefore it is not possible to conclude how many coralla are infested and what is the ratio between infested/uninfested coralla. The material is partly housed at the Faculte´ libre des Sciences et Technologies, Lille (GFCL), and partly at the Institute of Palaeobiology, Polish Academy of Sciences, Warsaw (ZPAL). Coralla were sectioned perpendicularly to the axis of corallum growth. Each corallum provided several thin sections, spaced 5–12 mm (altogether
From: KO¨ NIGSHOF , P. (ed.) Devonian Change: Case Studies in Palaeogeography and Palaeoecology. The Geological Society, London, Special Publications, 314, 125–129. DOI: 10.1144/SP314.6 0305-8719/09/$15.00 # The Geological Society of London 2009.
M. K. ZAPALSKI
19 thin sections); additionally acetate peels were made in order to record changes in morphology (altogether seven peels). On each thin section the number of corallites and parasites (Chaetosalpinx) was counted; parasite prevalence (P) was calculated as follows (calculation method introduced here): P ¼ Tp =C where C stands for number of complete crosssections of corallites in the thin section and Tp for the total number of parasites in the thin section. The ‘prevalence’ in epidemiology is a term indicating ratio of infested individuals to total number of individuals (e.g. Palm & Klimpel 2008; Vathsala et al. 2008). The formula presented here is adapted to colonial animals, where several individuals are parasitized at once and parasites are shared between individuals. The analysed material is rather poorly preserved: central parts of coralla are calcified, and precise observations cannot always be performed. Counted numbers of parasites should be treated as approximate, and probably the true P values are slightly higher (in recrystallized parts of coralla the corallites can be counted, but the Chaetosalpinx tubes are invisible).
Table 2. Parasite infestation in specimen ZPAL T.II/17* Distance from bottom of corallum Thin section no. (mm) ZPAL T.II/17a ZPAL T.II/17b ZPAL T.II/17c ZPAL T.II/17d ZPAL T.II/17e ZPAL T.II/17f
13 19 25 29 32 37
36 161 317 248 312 83
7 21 192 345 332 227
0.194 0.130 0.606 1.391 1.064 2.735
*Morphology: discoidal; size: 41 (height) 56 82 mm.
Patterns of parasite distribution with in the host’s corallum In these cross-sections the parasites do not display any general preference for location in either the centre or peripheries of the corallum, but there is variation of the location of parasites relative to individuals of the host. In the early astogenetic stages parasites prefer settlement in the corners of corallites (at which three individuals join). Conversely, in the late astogenetic stages, they seem to occupy both corners and walls between adjacent individuals.
Serial thin sections
Any comparison of the parasite infestation in fossil and modern cnidarians will remain more or less speculative, as the interactions between colonial hosts and their parasites are poorly understood (Hill & Okamura 2007). Nonetheless it is worth trying to understand what might be the factors controlling the infestation. The distribution of the Chaetosalpinx parasites within both of the analysed coralla shows three important features: (1) absence or scarcity of parasites during the early stages of astogeny; (2) increase in the total number of parasites during astogeny; (3) increase in number of parasites per corallite (Tp) during astogeny. The increase in the number of parasites does not seem to be unusual, as long as the number of individuals in the colony increases. Newly growing coral individuals created further space for the parasites to infest the colony. The decreased number of parasites in certain thin sections is caused by sectioning a smaller area of corallum (as is the case in the sample ZPAL T.II/17 in sections close to the corallum surface, see Fig. 2) or recrystallization, excluding some areas of the thin section from counting (where it was possible to count the corallites, but not the parasites, as in the case of sample GFCL 2174, see Fig. 2). Therefore, as stated above, the
The Tp values calculated for each thin section for two coralla are given in Tables 1 and 2. The third corallum (specimen ZPAL T.II/54) is hemispherical, measuring 25 (height) 40 46 mm, and displayed only two parasites (Chaetosalpinx) in the late stages of growth. The gradual increase in the number of parasites during astogeny in the two specimens is illustrated in Figures 1 and 2. Table 1. Parasite infestation in specimen GFCL 2174* Distance from bottom of corallum Thin section no. (mm) GFCL 2174a GFCL 2174b GFCL 2174c GFCL 2174d GFCL 2174e GFCL 2174f GFCL 2174g GFCL 2174h
10 23 31 40 47 54 58 65
– 107 171 255 250 321 414 369
0 11 52 188 160 192 242 419
0 0.103 0.304 0.737 0.640 0.598 0.585 1.136
*Morphology: bulbo-columnar; size: 71 (height) 52 66 mm.
PARASITES IN TABULATE CORALS
Fig. 1. Increase in number of parasites during astogeny, shown by transverse thin sections through a corallum. Pictures on the left show thin sections, and on the right the same thin sections with crosses marking the positions of Chaetosalpinx parasites. (a) Thin section GFCL 2174h, 65 mm from the bottom of the corallum (6 mm from the top); (b) thin section GFCL 2174d, 40 mm from the bottom of the corallum; (c) thin section GFCL 2174a, 10 mm from the bottom of the corallum. Note the strong increase in number of parasites. Grzegorzowice Beds, Emsian–Eifelian; Grzegorzowice, Holy Cross Mountains, Poland. Scale bar 1 mm.
true P values may be higher than the 2.7 calculated in the analysis. Research on modern bryozoans has shown that the high number of parasites in colonial animals is correlated with their low virulence (Hill & Okamura 2007); this may be similar to the discussed case. The increasing number of parasites per individual (parasite prevalence) during astogeny may be a sign of two different processes: (1) insufficient
protection by cnidae; (2a) insufficient response from the coral immune system or (2b) ability of the parasite to adapt to the immune response of the host, and therefore (2a). The insufficient protection by cnidae was probably the first factor allowing infestation. Cnidarians have no specialized immune cells (Kuznetsov & Bosch 2003), therefore the insufficient immune response seems to be highly possible; however, some modern corals do appear
M. K. ZAPALSKI
Fig. 2. Diagrams showing the increase of the total number of parasites (left) and the parasite prevalence (right) plotted over the corallum height (standardized). The decrease in the total number of parasites in specimen ZPAL T.II/17 is caused by the smaller area of thin sections, while the decrease in their number in specimen GFCL 2174 is caused by local recrystallization of specimen. Note the systematic increase of parasite prevalence during astogeny (right diagram).
to have a well developed self-defence system (Chadwick-Furman & Rinkevich 1994), at least in the terms of self/non-self recognition. The increase in the number of parasites per individual might indicate that in the investigated Favosites the immune system was undeveloped. Parasite adaptation to the general host defences is not uncommon among modern organisms (Schulenburg et al. 2007), but it is correlated with immune systems unable to create a new immune answer to the changing strategy of the parasite. Comparisons with parasites of modern corals are problematic. Digenean meacercariae, copepods and cirripedians are the most common parasites of recent scleractinians (e.g. Cheng & Wang 1974; Humes 1986; Ross & Newman 1995; SimonBlecher & Achituv 1997; Aeby 2007). These parasites are usually less numerous (several individuals within a colony) and much larger (Aeby 2003), and their distribution cannot be compared with that of Chaetosalpinx. The only modern analogue are parasites of Favia occurring between corallites; their dynamics of infestation remain, however, unknown (Rosen 1968). The preferred location of the parasite seems to be in the corners of corallites. On the other hand, when the infestation attains high Tp values, parasites seem to occupy more frequently the walls between adjacent individuals. This was probably an effect of prior occupation of the corners: the newcomers had to settle in the corallum parts that were not occupied yet. The absence of preferred zones of settlement within the corallum (e.g. in the centre or periphery) is similar to that observed by Tapanila (2004).
Conclusions The absolute number of Chaetosalpinx parasites within a single corallum increased during astogeny. The number of Chaetosalpinx parasites per individual in the host colony increased during astogeny and could reach 2.7 parasites per polyp; such a high infestation may indicate insufficient protection by cnidae and insufficient immune system response. Chaetosalpinx parasites did not prefer any particular location within the corallum (neither centre nor periphery). On the other hand, their placement in relation to individuals is usually in corallite corners (where three individuals join), but they also occur in the walls between individuals (where two individuals join); the latter situation seems to be more common in late astogeny stages. I wish to express gratitude to Bruno Mistiaen (Lille) and Stefan Schro¨der (Ko¨ln) for valuable comments on the manuscript. Adam T. Halamski (Warszawa) and Benoıˆt Hubert (Lille) discussed an early draft of the text. I am very grateful to John Brenner (Wokingham) who improved my English. The Foundation for Polish Science is thanked for the scholarship funding.
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