Acta Geologica Polonica, Vol. 48 (1998), No.4, pp. 377-386
General trends in predation and parasitism upon inoceramids
PETER 1. HARRIES & COLIN R. OZANNE
Department of Geology, University of South Florida, 4202 E. Fowler Ave., SeA 528, Tampa, FL 33620-5201 USA. E-mail: [email protected]
ABSTRACT:
HARRIES, P.I. & OZANNE, C.R. 1998. General trends in predation and parasitism upon inoceramids. Acta Geologica Polonica, 48 (4), 377-386. Warszawa.
Inoceramid bivalves have a prolific evolutionary history spanning much of the Mesozoic, but they dra matically declined 1.5 Myr prior to the Cretaceous-Tertiary boundary. Only the enigmatic genus Tenu ipteria survived until the terminal Cretaceous event. A variety of hypotheses have attempted to expla in this disappearance. This study investigates the role that predation and parasitism may have played in the inoceramids' demise. Inoceramids show a range of predatory and parasitic features recorded in their shells ranging from extremely rare bore holes to the parasite-induced Hohlkehle. The stratigraphic record of these features suggests that they were virtually absent in inoceramids prior to the Late Turonian and became increasingly abundant through the remainder of the Cretaceous. These results suggest that predation and parasitism may have played a role in the inoceramid extinction, but more rigorous, quantitative data are required to test this hypothesis further.
Probably the C011lmonest death for many animals is to be ous-Tertiary boundary (KAUFFMAN 1988). The mar eaten by something else. ked demise of such an important group has led to the C. S. Elton (1927) formulation of a wide range of hypotheses to explain this phenomenon. To date, most of these explana INTRODUCTION tions have hinged on global change brought on by cooling, alteration of oceanic circulation, or general The inoceramid bivalves arose in the Permian environmental degradation (e.g. KAUFFMAN 1984; and by the Cretaceous were dominant constituents of 1988; KAUFFMAN & al. 1992; MACLEOD 1994; FI level-bottom communities in a wide variety of shel SCHER & BOTTJER 1995; MACLEOD & HUBER 1996). fal settings. Although they are well-known inhabi This overview aims to investigate initial data in sup tants of oxygen-depleted, marine facies, they also port of a fUlther hypothesis in this debate: the role of thrived in a wide range of well-oxygenated environ predation and parasitism. ments, including shallow-water settings (e.g. KAUFF MAN & HARRIES 1996). Hence, the group and even some species appear to have had very broad ecologi EVIDENCE OF PREDATION AND cal tolerances that are also minored in their broad PARASITISM IN INOCERAMIDS geographic distribution. Despite their abundance and diversity, all inoceramids, except for species of Te Evidence for parasitism on bivalve shells ge nuipteria, disappeared 1.5 Myr prior to the Cretace- nerally comprises a variety of pits, scars and addi- 378 PETER J. HARRIES & COLIN R. OZANNE tional calcitic secretions which modify the shell burial compaction, or simply the differential morphology. Predation signatures on molluscs are degradation of the organic matrix that held the typically confined to those predators that either prismatic layer together. Although consistent bore through or bite and crush the shell. Because breakage patterns have been documented in inoceramids are very rarely bored by predators, some predator-prey relationships (e.g. CADEE most evidence for predation intensity derives 1968), this has not been seen from most available from sublethal (rehealed) injuries. The study of inoceramid data. HATTIN (1975) has suggested these features as a proxy for predation intensity is that Ptychodus was an extremely efficient less than ideal (VERMEIl 1982); it is difficult to di processor of inoceramids. He hypothesized that rectly relate predatory failure to predatory suc their predation and excretion of bivalve material cess. Despite the limitations, it is the best indica was responsible for the common inoceramid tor available. Common predatory and parasitic fe prism-rich calcarenites in mid-Cretaceous atures found on inoceramid valves are described Western Interior sequences. However from the briefly below. numerous specimens investigated from this inter val (strata spanning the Cenomanian-Turonian boundary), only a single occurrence of sublethal Boreholes injuries has been documented (see PI. 1, Fig. 1; W ALASZCZYK 1992). Unless these predators were The line of evidence most commonly used to almost 100% efficient, which seems extremely evaluate the role of predation in the fossil record unlikely (see discussion in VERMEIJ 1982), there of invertebrates is the presence of boreholes. In should be evidence of failed prey SUbjugation. most cases they are produced by gastropod pre Additionally, SAGEMAN (1996) in his study of dators, especially naticids and muricids (SOHL Western Interior calcarenites, suggested that they 1969). Unlike predatory processes that result in formed "through winnowing by storm events fragmentation, where unequivocal causes are dif during relative sea-level fall and condensation ficult to determine, boreholes can be readily in due to starvation during subseqent rise" (p. 891). terpreted. Because the borer leaves its signature Hence, the primary control appears to be relative in the morphology of the borehole, this predator sea-level position and not a function of predatory prey system has the advantage of identifying not efficiency. only the obvious prey, but the predator as well. Inasmuch as predators are rarely completely Typically, completed boreholes are interpreted to efficient and effective attackers of their prey, one represent a successful predatory attack, in which of the ways in which shell attacks can be estima the prey was successfully subjugated, killed, and ted is through the relative abundance of repaired consumed. Bored shells also have the advantage shells. Because bivalves precipitate calcium car of generally being preserved as reliably as pris bonate throughout their lives, earlier ontogenetic tine ones (but see Roy & at. 1994); therefore, events are preserved in their shells. Therefore, population dynamics and estimates of such previous attacks on the shell that produced suble aspects as predator efficiency can be reconstruc thal injuries have the potential to be recorded, ted. The geologic record is replete with evidence especially those events that resulted in injuries to of boring in a wide variety of groups and by the mantle. Numerous inoceramid specimens a spectrum of predators. To date, only a single show evidence of injury to the mantle and shell horizon in the Lower Maastrichtian is known to (PI. 1, Fig. 4; PI. 2, Fig. 2b). These features have contain inoceramids bored by gastropod preda a variety of morphologies, but they generally tors (SCHOPF & HARRIES, in prep.) consist of mantle damage which then is reflected by aberrant growth following the attack. Some of these attacks can be quite localized (PI. 1, Figs Shell breakage and repair 1-2), while in other cases the attack may be more severe and substantially affect the shell morpho Unfortunately it is difficult, if not impossible, logy (PI. 1, Figs 3-4 and PI. 2, Fig. 2b). In cer to determine whether most fragmentation in ino tain instances, the predator may leave a fairly ceramids was due to the action of predators or distinctive bite (e.g. KAUFFMAN 1972), but in the whether it simply reflects shell breakage due to majority of cases it is difficult to precisely deter wave and current activity prior to burial, post- mine the predator. GENERAL TRENDS IN PREDATION AND PARASITISM UPON INOCERAMIDS 379
Fecal pellets and regurgitated masses times referred to as "mud blisters" (HOFSTETTER 1965), form when the mantle is irritated and ad Another common mode of predation on mari ditional nacre is precipitated over the irritant. In ne invertebrates, especially among fish, is to its most extreme form, this results in pearl forma swallow bivalves and other benthic organisms. In tion, but in most cases it simply results in the se a study of stomach volumes of modern hogfish, cretion of additional nacreous layers. RANDALL & W ARMKE (1967) determined that 42.6% of the stomach contents were bivalves. Clearly, certain fish can playa significant preda Parasitic tracks tory role on bivalves, although extrapolating the results of these modern studies directly to the One of the most evident types of paraSItIc fossil record is lenuous. Rays also have the abili scars left on modern bivalves is the presence of ty to ingest large quantities of bivalves (GRAY & polydorid worm borings. Polydora produces al. 1997). Unfortunately, stomach contents of a distinctive U-shaped boring, but because the predators are rarely preserved in the fossil boring occurs following shell precipitation, it do record. Instead, paleontologists must rely on pre es not disrupt the mantle and later shell forma served fecal pellets (coprolites) or regurgitation of tion. However, where the parasite resides betwe shell masses to reconstruct predation patterns of en the shell and the mantle, dramatic changes in the past. One of the main stumbling blocks to this shell precipitation can occur. Some inoceramid approach is the difficulty in determining which parasites clearly disrupted the mantle's ability to predator produced the various defecated or dis precipitate calcium carbonate, while in other ca gorged masses present in the fossil record. Shell ses, such as the formation of a Hohlkehle (di masses composed of inoceramid remains are rela scussed below), the inferred parasite was covered tively common, but their origin is questionable. by nacre. In the former case, the shell contains SPEDEN (1971) has documented several types pronounced, but often very localized, deformities of shell masses from the Albian-Cenomanian? (see I v ANNIKOV 1979). Clarence Series in New Zealand very similar to Pennsylvanian shell masses attributed to fish pre dation (ZANGERL & RICHARDSON 1963). The most "Hohlkehle" likely candidates to have produced these fecal or regurgitated masses are fish, sharks, rays, or ma One of the common features found in Santo rine reptiles, such as mosasaurs. Given the com nian through Maastrichtian inoceramids, and mon co-occurrences of inoceramids and Ptycho which may be present in Coniacian and potential dus teeth (KAUFFMAN 1972, COLLOM 1991), these ly even in Jurassic inoceramids (MORRIS 1995), durophagous sharks would seem to be the leading is a pronounced internal rib or H ohlkehle, that candidate for many Cretaceous localities. How initiates behind the beak and is oriented towards ever, the absence of Ptychodus teeth from New the posterior margin (PI. 1, Fig. 5 and PI. 2, Fig. Zealand sequences (CRAMPTON, pers. comm., 2a). The Hohlkehle has been the source of con 1998) suggests that fish may be more important siderable debate. WHITFIELD (1880) used this in that region. feature as the critical character in identifying the subgenus (or genus of some authors) Endoco stea. Some recent workers have followed "Bubbly" nacre WHITFIELD'S convention and used the internal rib as the diagnostic character (MACLEOD 1994, Features which may be indicative of the man MORRIS 1995). TOOTS (1964), following tle responding to parasites potentially living be ROEMER'S (1852) and HEINZ'S (1928) positions, tween the shell interior and the mantle are the suggested that the Hohlkehle was the result of presence of "bubbles" on the internal shell surfa a specialized parasite feeding on material expel ce. They can differ substantially in shape and si led by the exhalent current. He based his conclu ze ranging from very small features confined to sion on five lines of evidence summarized here: the nacreous layer (PI. 2, Fig. 1) to larger featu 1) the ridge commences at variable distances res which probably influenced the external, pri from the beak; 2) the ridge abruptly terminates in smatic layer as well. These shell features, some- some specimens; 3) the nacre in specimens with 380 PETER 1. HARRIES & COLIN R. OZANNE a Hohlkehle is thicker - a common response to paper follows SEITZ'S (1967, p. 37) conclusion the presence of parasites; 4) the ridge is a hollow that the Hohlkehle has no taxonomic value. cavity; and 5) the surface of the hollow is orna A modern analog to this type of parasitism oc mented in a manner suggesting the encasement of curs in modern freshwater mussels. COKER & al. an organism or at least living tissue. SEITZ (1966; (1921) have documented a feature that is similar 1967) has a thorough discussion of the problem in terms of the consistency in position and the as well as a plethora of morphologic, stratigra well-developed scar produced on the affected phic, and facies distribution data of this feature. shells (see their plate XII). Because of its regular SEITZ (1967) also noted that including all the ino ity and its spatial repetition regardless of substra ceramid morphotypes with a Hohlkehle within te, they attribute the feature to an unknown a single genus was problematic. The Hohlkehle is parasite rather than due to mechanical injury. present in a very broad spectrum of inoceramids Although the shell scar produced is not identical (e.g. including Platyceramus, Cladoceramus, to the Hohlkehle, it nevertheless suggests that pa Cordiceramus, Selenoceramus, and Endocostea rasites can preferentially inhabit a given location species; SEITZ 1967) that, in the absence of the on a shell. feature, would be grouped as separate genera. A final line of observation that suggests the feature is a response to parasites is that rare spe STRATIGRAPHIC TRENDS cimens have the Hohlkehle only on one valve, whereas the other shows no evidence of it. To determine the efficacy, efficiency, intensi Hence, if the inoceramid were disarticulated, ty, and the temporal distribution of predation and some workers would place the two valves from parasitic types on the inoceramids, it is critical to a single specimen into separate genera. This investigate the stratigraphic as well as geogra phic patterns. The data summarized below are (see Text-figJ), for the most part, solely qualita tive and represent culling material from the lite Mya rature as well as from unpublished data from the .,j ... :--Abundant evidence for predation here may be substantially altered. and parasitism on inoceramids The earliest incidence of predation on inocer en ~ 75 ::J amids was documented by SPEDEN (1971) from 0 ~ t predator-bored inoceramids LU (S chopf and Harries, in prep.) the Albian (?Cenomanian) of New Zealand. He () CAMPAN. they thrived in black-shale environments (KAUFF mechanisms are potentially interrelated and MAN & SAGEMAN 1990) where potential predators clearly could have produced major environmen appear to have been relatively rare. Inhabiting re tal and ecological disruptions. The question gions where predators are excluded or rare can be remains, however, whether these would have a decisive factor in determining the habitats ava been sufficient to cause extinction of the inocera ilable to a species. Experiments on extant Mytilus mids. Given the length of their evolutionary edulis show that if they were protected from pre history (from the Permian to the Late Cretace dators, such as starfish, they could thrive in envi ous), the inoceramids experienced many of the ronments where under normal conditions they same environmental changes invoked by the would be excluded by predation pressure (W AR extinction mechanisms outlined above. Certainly, REN 1936). If the inoceramid predators moved the changing oceanic and atmospheric conditions into environments that they previously were in the Late Cretaceous may have influenced the unable to inhabit (or if they developed durapha inoceramids, especially if they were forced to gous ability within those environments), inocera live in environments favored by predators as mids would have been fairly defenseless prey. black-shale habitats became restricted. These The strength of their prismato-nacreous shell events compounded by the radiation of duropha may have afforded them some protection, but the gous crustaceans may have played an important evidence of increased predation suggests that its role in the inoceramids' demise. effectiveness in warding off predators may have Even if the predation itself were not responsi been waning. ble for the ultimate extinction of the various ino Another factor is that predation and parasitism ceramid clades which disappeared in the Late on prey or host organisms play important roles in Maastrichtian, it may have reduced population regulating the abundance of taxa and community sizes, changed community structure, and altered structure (e.g. DODD & STANTON 1981). From the geographic distribution to the point where smal study of modern ecosystems it is known that a ler-scale perturbations, that under normal cir distinctive relationship exists between predators cumstances would have simply led to reduced and their prey. The cropping pressures exerted by population sizes, could become the proximate predators regulates the abundance of prey which causes of extinction. SIMBERLOFF (1986) has in turn controls the predator population. In the outlined a range of different small-scale pertur vast majority of cases, the balance between preda bations which could lead to extinction of species tors and their prey ensures the prey's, and hence limited to small populations and geographic ran the predator's, survival. However, if a predator is ges. If the inoceramids were being cropped by in not dependent on a primary prey source, there is creased predation, weakened by increased parasi the possibility that overcropping and potentially tism, affected by changing oceanographic para extinction could occur. Parasites can also make meters including the loss of their favored habitat, their hosts more vulnerable, hence the inferred in the cumulative effects may have been more than crease in parasitism, which occurred simultane the group, with the exception of Tenuipteria, ously with increased predation, potentially had could withstand. profound effects upon inoceramid populations. Further potential evidence in support of pre Despite the dominance of the inoceramids du dation as a factor in Late Cretaceous biotic dyna ring the Cretaceous, they (except for taxa within mics is seen in ammonite data. In their study of the enigmatic genus Tenuipteria) disappeared abnormalities in Late Cretaceous scaphitid am approximately 1.5 Myr prior to the Cretaceous monites from the Western Interior Basin, LAND Tertiary (K-T) boundary (KAUFFMAN 1988). Se MAN & WAAGE (1986) investigated a range of in veral hypotheses have been presented to explain juries found in this group. One of their goals was the Late Maastrichtian disappearance of the ino to evaluate the level of predation intensity in sca ceramids: 1) the first step in the K-T mass extinc phitid ammonites approaching the Cretaceous tion (KAUFFMAN 1988); 2) the disappearance of Tertiary boundary. Their analysis covered a var abundant low-oxygen habitats during the latest iety of ammonite biozones from the Turonian Cretaceous (FISCHER & BOTTJER 1995, KAUFF through the Maastrichtian, although it was focus MAN & al. 1992); and 3) a change in Late Creta sed on evidence from the Fox Hills Formation. ceous oceanic circulation (BARRERA 1994; 1997; They showed that throughout in strata below the MACLEOD & HUBER 1996). These hypothesized Fox Hills Formation, sublethal injuries occurred 384 PETER J. HARRIES & COLIN R. OZANNE in approximately 10% of the individuals found Acknowledgments (values varied from a minimum of 4.0% in Sca phites leei III macroconchs to a maximum of This overview would not have been possible without 11.1 % in Scaphites hippocrepis I microconchs). the assistance of numerous colleagues. We are very In the lower Trail City Member of the Fox Hills grateful to Bill COBBAN who not only suggested that Formation, the incidence of injury is 10.4%; a va this topic warrented further investigation, but also for lue virtually identical to the earlier values. In the the specimens he loaned to us from the USGS. We are overlying Timber Lake Member, the incidence extremely appreciative of the insights and unpublished rises to 30%. LANDMAN & WAAGE (1986) inter data supplied by Christopher COLLOM, James CRAMP preted these data as a function of several poten TON, Will ELDER, Cris LITTLE, and Ken SCHOPF. Further tial factors, including: 1) a facies change; 2) an thanks are due Christopher COLLOM, James CRAMPTON, overall decrease in prey abundance resulting in E. G. KAUFFMAN for their thorough and insightful increased predation pressure on those present; reviews of an earlier draft of this manuscript. and 3) the ability of these scaphitids to survive injury. 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The Mesozoic marine revolution: paleoecological history of two Pennsylvanian Evidence from snails, predators and grazers. black shales. Fieldiana, Geology Memoir, 4, Paleobiology, 3, 245-258. 1-352. PLATES 1 - 2 ACTA GEOLOGICA POLONICA, VOL. 48 P.l. HARRIES & C.R. OZANNE, PL. I PLATE 1 1 - Mytiloides labiatus (SCHLOTHEIM), Lower Turonian, left valve; arrow shows localized, healed attack (from W ALASZCZYK 1992); x 1 2 - Cremnoceramus crassus (PETRASCHECK), Middle Coniacian, left valve; arrow shows healed attack (University of Colorado Museum [UCM] 1722); x 0.5 3 - Inoceramus n. sp.?, left valve; arrow shows substantail shell deformation from a sublethal attack (United States Geological Survey [USGS] D1983); x 1 4 - Inoceramus pacficus WOODS, Upper Coniacian, left valve; note the dramatic change in direction of ornamentation following the severe attack (see arrows) (from CRAMPTON 1996); x 0.5 5 - "Endocostea" typica WHITFIELD, Upper Campanian, articulated specimen, with pronounced Hohlkehle (see arrows); there is also an area of shell deformation near the anterioventral margin (additional arrows); x 1 All specimens whitened with ammonium chloride ACTA GEOLOGICA POLONICA, VOL. 48 P,J. HARRIES & C.R. OZANNE, PL. 1 ACTA GEOLOGICA POLONICA, VOL. 48 P.J. HARRIES & C.R. OZANNE, PL. 2 PLATE 2 1 - Fragment of Inoceramus proximus TUOMEY, Upper Campanian; arrows denote "bubbles" formed on the nacreous layer; x 1 2a-2b - Cataceramus balticus?(BoEHM), left valve; specimen has a Hohlkehle as well as significant shell deformation following geniculation (see arrows)(USGS D3137); x 1 All specimens whitened with ammonium chloride ACTA GEOLOGICA POLONICA, VOL. 48 P.J. HARRIES & C.R. OZANNE, PL. 2