POST-PALEOZOIC PATTERNS IN MARINE PREDATION: WAS T~REA ~SQZOIC AND CENOZOIC MARINE PREDATORY REVOLUTION? SALLY E. WALKER" CCARTON E. BRETTZZ 'Department of Geology, University of Georgia, Athens, Georga 30602 USA Qepartmex1.t of Geology, University of Cincinnati, Cincinnati, Ohio 45221-0013 USA

AB~~CT-Mesozoicand Cenozoic evoltttion ofpredators involved u series of episodes. Predaaors rebounded rather rapidly csfier the Permo-Tsa'ussic extiacsion and by the Middle Triassic a variety of new predator guilds had appeared, including decapod crustaceans with crushing claws, shell-crushing sharks and boay>fish as well as marine reptiles adapted for crushing, smashing, and piercing shells. While several groups (e.g., placodoats, nothosaurs) becam extinct in the Late Triassic crises, others (e.g., ichthyosaurs) survived; and the Jurassic to Early Cretaceous saw the rise ofmalacostmcaa cmtaceam with cmhing chelae and prerkadoiy vertebrates-in particuhl; the marine cr-ocodiliuns, ichthyosaurs, dpless'osau~s.fie iate Crercrceor~~suw unprecedented levels of diversiiy of marimp redaceom vertebrates including pliusautids, plesiosaurs, admosasauw. The great Cretaceous- Tertiaiy extinction decimted marine ~ptiles.Howeves most invertebrate and$shpredaroly groups survived; and during the Paleoge~le,predatoly benthic invertebrates showed a spurt of evol~stionwith neogastropods and new groups of decapods, while the teleosts and neoseluchian shark7 both underweadparallel rapid evolutioaa~yradiations; these were joined by newpredcltoy guilds of sea birds and marine rnamls. Thus, although escalution ix so??wtimes casr as un ongoing "rrms race,'' in achtalih. the predatoly record shows long interludes of relative stability puncturuted by episodes of abrupt biotic reorganization during and afer mass extinctions. This patbem suggests episodic, but generally increasing, predation pressure on marine organisms through the Mesozoic-Cenozoic interval. Howevel; review of the Cenozoic record ofpredation suggests that there are not uarambiguow escaletuly treads in regard to antipredalo~shell architecmre, such as conchiolin and spines; nor do shell drilling and sheU repair data show a major increase fi-ont the Lute Meso~oicthrough rhe Cenozoic. ,Mosl' durophrrgozis groups are generalists, and thus it may be thaf they had a diffuse ejffect on their invertebrate prey.

"As evolutionists, we are charged, almost by INTRODUCTION definition, $0 regard historical path ways as the essence of our subject. We cannot be indiflerent to THE CONCEPT OF predator-prey escalation the fact that similar resulbs can arise by different is, j, large measure, an outpowth of the extensive historical routes." -4ouid and Vrba, 1982 studies of Venneij (1977, 1987) on the so-called "Mesozoic Marine Revolution." This term might "This is not to say that selection is not seem to imply that a dramatic development of important, but that its invocation is not justifled marine predators was initiated at the Triassic; a until the rule of C~UPZCEin the opem~iolzof a continuous intensification of predator-prey basically srochastic universe is ruled out." relationships has been envisaged. In actuality, the --Schram, 1986 Mesozoic and Cenozoic evolution of predators involved a series of episodes. In this paper we "A science grows only as it is willing to document the diverse predatory guilds of the question its assumptions and expand its Mesozoic and Cenozoic, especially vertebrates that approaches. " -Hickman, 1980 putatively devoured invertebrate prey, with comments on their modes of feeding and possible PALEONTOLOGICAL SO CIETY PAPERS, V 8,2002 impact on potential prey, focusing on benthic as Gould and Vrba (1982) have recognized, there invertebrates. Predator guilds (e.g., marine reptiles) are a number of historical and non-adaptive routes during the Mesozoic are surprisingly similar to by which specific aptations may ultimately arise those in the Cenozoic (e.g., marine mammals), in organisms, and such may also be the case with except that the players have changed. Despite the certain antipredatory strategies. setbacks of mass extinctions, the diversity of Despite its length, this paper is not an exhaustive predators remained at a nearly constant proportion review. However, we did strive both to provide a from the Late Triassic to mid-Cretaceous broad overview of Mesozoic and Cenozoic predators (Bambach and Kowalewski, 1999). However, and their potential prey, and, perhaps more during the jast 110 million years (Late Cretaceous- importantly, to demonsbate that there are alternative Neogene), predators diversified faster than the rest ways to think about these predatory patterns. of the fauna (Bambach and Kowalewski, 1999). We also examine the patterns of predation in TRIASSIC PREDATORS post-Paleozoic shell structure (e.g., conchiolin and AND PREDATION spines), shell repair, and shell drilling. However, data from drilling and shell repair thus far do not show All marine. benthic ecosystems were profoundly unambiguous escalatory trends. For the sake of altered by the Pem-Triassic extinction (Fig. 1). argument, escalation is not a continual trend horn Many Paleozoic predators were eliminated, including the Mesozoic to the Cenozoic; rather, within each most phyllocarids, platyceratid gasbopsds, gmiatite era it is dependent on the suite of predators and prey. arnrnonoids, and myprimitive lineages of sharks. In the final section of the paper we reconsider Other active predatory groups preferentially made the Mesozoic Marine Revolution hypothesis it though this bottleneck, including the hybodontid proposed by Vemeij (1 977,1978, 1987), and ask sharks and the root-stocks of Mesozoic mstaceans questions that may guide future research: Is the and ammonoids (Jholl et al., 1996). pattern of putative morin invertebrates strictly Gmtmpods and Bivalves.-Varied archaeo- and related to predation, or might there be other mesogastropod taxa rediversified in the Triassic. hypotheses that could explain armor in organisms? Nonetheless, records of gastropod drilling predation Is there evidence that most predators are specialists are surprisingly rare in this period (Kowdewski et on particular prey and thus migk cause extreme al., 1998).However, in a few instances, drillers seem selection in invertebrates who then respond with to have had a significant impact (Fursich and various escalating strategies (c.g., spines, Jablonsk, 1984; Kowalewski et al., 1998). The first chonchialin) to mitigate the increased predation naticid-like mesogastropods (Ampullifla) are pressure? Does diffuse selection from generalist known from this time (see Fig.'7) (Fiirsich and predators cause antipredatory armor to arise in a Jablonski, 1984; Newtofl1983; Kowalewski ec al., number of groups of invertebrates? If most 1998). Given the rarity of naticid-like boreholes predators are generalists, as appears to be the case from the Late Triassic to the mid-Cretaceous based on the evidence amass4herein, then perhaps (Albian), it has been suggested that predatory Lhere was not a sufficiently intense selective force drilling was relatively ineffective and largely Iost to produce a major "sea change" in antipredatov during the Triassic, only to be evolved again, armor in any one group of marine invertebrates, successfully, during the Cretaceous (Kow alewski et especially in past-Paleozoic organisms. Thus, for al., 1998). Predatory septibranch bivalves also examplc, durophagy may not necesszrily mean that originated at this time (Skelton et aI., 1990). a predator ate molluscs; durophagous dentition Ammonoids.-Ammonoids were nearly could also indicate the eating of crustaceans, other extinguished by the Perrno-Triassic crises. hard-shelled prey, or even soft prey (e.g., Plotkin However, the ceratitic ammonoids staged a rapid eta!., 1993; Wilga and Motta, 2000). Perhaps also, rediversificiation in the Triassic. Like other WALKERAVD BR.77'-POST-PALEOZOIC PATTERhrS IN MARTNE PREDATION

FIGURE I-Ranges of various taxa of Mesozoic and Cenozoic durophagous (hard-shell crushing) predators. Thin lines: present, but of limited abundance; thick lines: abundant; broken lines: possibly present but rare as fossils. ammonoids, the ceratites are assumed to have been isopod, and amphipod crustaceans appeared in the predaceous, although data are very sparse. Stomach Late Paleozoic, but they did not diversify or crop contents of ammonoids are very rare, but siegifIcantly until the Jurassic (Briggs and Clxkson, when they are found provide important evidence 1990). Four out of 27 PaIeozoic families survived for trophic relationships. One specimen of an Fmly into the Mesozoic, and only a few groups are known Triassic ammonoid (Svalbardiceras) had from the Triassic (J3riggs and Clarkson, 1990). ostracodes and foraminiferans among its gasbic Various lobster groups evolved in the Triassic contents and may have been a predatory nektonic (Table 1). Their appendages indicate that they were carnivore (Westemam, 1996, p. 675). durophagous, but modern lobsters feed on a wide Crustacera and 0stracodes.-The most variety of prey and are not specialists on molluscan important post-Paleozoic groups of decapod, prey. Ostracodes are known to be predators on TABLE 1-Geographic and temporal ranges and feeding preferences of crustacean predators.

Taxon Geographic range Specialist feeders on specific Geologic range of durophagous and depth range invertebrate prey? anatomical features (molariform Geologic Range (shallow, < 70 rn; claws, mouth parts, etc.) deep, > 70 m)

lnfraorder Brachyura Boreal to temperate Generalists Unknown Family Portunoidea (bivalves, like oysters and mussels; Carcininae scavengers; fish, plants, benthic Portuninae invertebtrates, jellyfish, epibionts on Zostera, tunicates) Cretaceous {Maastrichtian)-Recent . ..------. . - - .. . . .- .-- Family Xanthidae Temperate to tropical Generalists Menippe mercenaria is the most (mangrove detritus, other plants and famous shell-crusher of this group; the Cretaceous-Recent detritus, small crustaceans, oysters, genus Menippe originated in the Middle (SChram, 1986) barnacles; M. mereenaria eats molluscs, to Upper Eocene (Glaessner, 1969), and like oysters, polychaetes, barnacles and M. mercenaria is known only from the other crustaceans) Pleistocene (Rathbun, 1935) ------. - - - .--- - -.

Farnilv Calamiidea, , South Pacific, Caribbean, Generalists The shell-peeling Calappa flammea is North America, Europe known from the Oligocene (Rathbun, Eocene-Oligocene to Miocene; 1930; Ross et al., 1964) Recent (Glaessner, 1969); Cretaceous-Recent (Schram, 1986) -. .------. -. Decapod crustaceans Generalists Unknown (Order Dscapoda) (earliest decapod is Palaeopalaemon, Schram, 1986)

Late Devonian to Recent (Schram, 1986); Perrno-Triassic to Recent (Williams, 4 996) - -- . -- Stomatopods Chiefly tropical shallow Specialists Rapto rial mouthparts, Devonian- (Order Stomatopoda) to deep water; rare in Recent; folding raptorial thoracopods, in Extant: 350 spp, 66 genera, T 2 temperate zones, none pa[eostornatopods, Carboniferous; families in polar regions gonodactyloid shell smashers, ?Cretaceous, Upper Miocene to Recent; Devonian to Recent Squilllds, ?Cretaceous to Recent -. - - , - - - -- (Table 1, cont.)

Family Cancridae Chiefly boreal and Generalists (sea urchins, polychaefes, Molariform teeth and shell-crushing temperate; shallow and scavengers, molluscs) claws (Miocene to Recent) Eocene to Recent deep water (Schram. 19861 ... .. - ...- - . -. . . ? ------, --- Family Parthenopidae Temperate to tropical Generalists, omnivores, detritivores; one Gore and Soto (1979, p. 67) state that species known as a shell-crusher parthenopids are omnivores and Eocene to Recent (Parthenope horrida ) detritivores; Debelius (1999, p. 259) (Schram, 1986) figures Parthenope horrida eating a pufferfish; Yermeij 1978, p. 40, states that Parthenope (Daldodia ) horrida is a shell-crusher in the lab Family Majidae Generalists Eocene to Recent (Schram, 1986) . .-- .- . .- Family Grapsidae Boreal, temperate to Generalists tropical (sea urchins) Eocene to Recent {Schram, 1986) -. . . - - .- lnf raorder Palinuroidea Temperate to tropical; Generalists Family Pal~nuridae deep and shallow water {rnollwsks, sea cucumbers; Williams, 1984) (Lobsters) ?Lower Triassic, Mid-Triassic to

Recent (Williams, 1996) - - -. ------...- . --- A- -- Superfamily Nephropidae Temperate zones, North Generalists Molariform teeth on major crusher claw as in Homarus (Cretaceous to Recent; Family. Nephropidae. Atlant~c,Europe; shallow (molluscs, hydro~ds,crustaceans, (Clawed marine lobsters) and deep water (many lobsters, polychaetes, brittle stars, Glaessner, 1969) migrate) mussels, limpets, Lobster molts, chitons, Permo-Triassic to Recent byozoans, scallops, oysters, sea urchins, (Williams, 1996) seaweeds; Williams, 1984; Lawton and Kavalli, 1995) - -. - - .-- -. .- .. - Family Scyllarjdae Temperate to tropical Generalists? (Spanish Lobsters) (?scyphozoan medusae; Williams, 1984) ?Lower Cretaceous; ?lower Eocene to Recent (Williams, 1996) PALEONTOLOGICAL SOCIETY PRPERS, 1! 8,2002 polychaete annelids, and scavengers on dead placodonts (Figs. 1, 2.1-2.3) evolved from polychaetes, fish, and squid (Vmier et al., 1998). unknown diapsid reptilian ancestors (Benton, 1993, They have serrated appendages that act like knives 1997). The Triassic placodonts, sister gxoup to d (or sandpaper) to abrade their food. Based on their other Sauropterygia, have. members that are feeding appendages,the Early Triassic cypridinids interpreted to have been benthk predators on hard- (mdpossibly the Late Ordovician myodocopids) shelled invertebrate prey. Placodus, for example, may have been predators or scavengers on had pachyostosis (complete covering of the cheek cephalopod carcasses (Vannier et al., 1998). by dermal bone), which added weight to the jaw Chonda'cthyes.-The long-lived hybodontids and thus may have functioned as an adaptation for flourished in the Tiassic and became the dominant durophagy (Rieppel, 2002). Additionally, the Jurassic sharks (Maisey, 1982). Hybodonts procumbent and chisel-shaped premaxillary and possessed varied dentition, ranging from high- dentary teeth may have functioned to pick off cusped impaling teeth to low-crowned crushers, invertebrates from their substrate, which were then indicating rathcr generalized predatory diets crushed with the postefior tooth plates before they (Majsey, 1982); they gave rise to swimming, were swallowed (Westphal, 1988). Biomechanically, piscivorous sharks, as well as pavement-toothed the tooth plates of Phcodm were positioned in such forms. The Rybodont sharks may have &sen in a way as to enhance crushing, but not increase load the Devonian, but they underwent skong adaptive to the jaw (Rieppel, 2002). The basal stock of radiation during the Triassic (Maisey, 1982). Placodm already had large crushing tooth plates Suuropterygian clade.-The sauropterygian md procumbent premaxillary teeth (Rieppel, 20021, clade (Figs. 2,6; Table 2) contains the Triassic stem indicating that durophagy was an ancestral condition groups such as placodonts, pachypleurosaurs, in this group. The few durophagous taxa of nothosaurs, and pistosaurs, and the Jurassic- placodonts may have had an impact on the Cretaceous crown groups hown as plesiosaurs, rediversifying molluscan communities of the pliosaurs, and elasmosaurs (Reppel, 1999). Very Triassic, but they became extinct in the major crises little is known about the feeding mechanics of toward the end of that period. Triassic stern-group sauropterygians, which Not dl placodonts had dentition indicating that secondarily became aquatic from thelr terrestrial they ate benthic hard-shelled prey. More derived ancestors (see Rieppel, 2002). Feeding underwater, cyamodontids (Pk~coc~lysand Psepbdem)lack as the stem-group sauropterygians did, required a premaxillaq and anterior dentary teeth, and may suite of anatomical and behavioral adjustments that have picked up benthic soft-substrate invertebrates had to allow for their adaptive radiation into early (like crustaceans) through suction action (Rieppel, Mesozoic seas (Fkppel, 2002). Suction feeding 2002). Another basal cyamodontoid, Henodus, has appears to be the most efficient hydrodyamic way much reduced crushing dentition, and may have had to solve the underwater feeding dilemma (Lauder, baleen that was used in sieving benthic invertebrates. 1985); however, "quick snapping bites" at the air- Hertodm is thus interpreted to have been a bottom- water interface (or underwater) are also used, feeder-perhaps an herbivore or omnivore-but it especially by crocodilians (Reppel, 2002). Triassic was not durophagous (Reppel, 2002). sauropterygians covered all styles of feeding, and Pachyp1eurosauria.-PachypZeu~osaurs thus have littIe overlap in hypothesized feeding @g. 2.7) were swimming reptiles with long heads skategies. The varied nearshore habitats in the and interlockjag lower and upper sharp teeth Middle Triassic, with Iagoond basins interspersed presumably for the capture of fish (Benton, 1997). among reef habitats, may have accounted for the Pachypleurosauria are considered to be the sister trophic-functionaI diversity of stem-group group of the Nothosauroidea, or the sister taxon to sauropterygians (Xieppel, 2002). all other Eusauropterygia (composed of nothosaurs Placodonts.-During the Middle Triassic, the and plesiosaurs; Rieppel, 20023. Pachypleurosaurja WALKER AhD BRETT-POST-PALEOZOIC PATTERMS IN MNEPREDATION

FIGURE 2-Triassic predatory reptiles. 1-3, Placodont reptile Placodus. 2,3,Lateral and palatal views of skull; note spatulate incisors and broad "pavement" teeth in maxilla and palate. 4, Ichthyosaur Mixosaurus. 5,6,lch thyosaur Grippia; lateral and dorsal views of skull. 7, Nothosaur Pachypleurosaurus. Adapted from figures in Benton (I 997). are among he smallest of the sauropterygims: most they were not efficient in subduing vigorous prey attain a length of 50 cm; few attain lenghs of 120 (Rieppel, 2002). Pachypleurosaurs probably were crn (Carroll and Gaskill, 1985; Rieppel, 2002). pelagic predators that used suction and rapid closure Based on a relatively large tympanic membrane and or" the jaws to subduc soft-bodied cephalopods and limited bone ballast, these marine reptiles may have small fish (Sander, 1989; Rieppel, 2002). inhabited shallow, coastal, and estuarine waters Nothosaurs.-The Middle Triassic (Taylor, 2000). Pachypleurosauria (e-g., Nothosauroidea (up to 4 m in length) are a major Neusticosaum) had delicate jaws with homodont clade of the Eusauropterygia, members of which dentition; loading conditions of the jaw indicate that may have eaten fish, other sauropterygians, and TABLE 2-Cenozoic marine vertebrate predator guilds,

Marlne Reptile Geologlc Range; Feedlng Realm FedingType Putatlve Prey Gastrlc Evldence or Reference Biogeography other evidence - - .-- PlaccdcaUa Middle Trfassic Benthic Durophagous (basal and Hard-shelled sesslle to non- Riapple, 2002; (chiefly late Lower to derived cyamw'ontoids) to sessile invertebrates Massare, 7 997 Upper +rimsic of suction and sieving (cyamodonts); soft shelled Europe, North Africa, (Henodus ); crush gufld of invertebrates to plants Middis East; distribution Massare, 1997 (Henodus ) restncted to western periphery of Triassic Tefhys (Lucas, 1997) Pachypleurosaurs Middle Triassic Pelagic Suctjon feeding Soft-shelled invertebrates, Rleppls, 2002 cephalomis riddle Trlasslc Pelaglc Suctlon feeding to fish-trap Slrnosaums :hard-shelled Jwentile placodont In Rlepple, 2002 ch~efly);lata Early dentition invertebrates; all others fish, soft- Larimurus stomach Sanders, 1 989; riasslc to Uppar shell& invertebrates Tschanz, 1989 Triassic; endemk to Europeadhian continent (Lucas, 1997) Plstosaumldea MIddls Trlassic Pelagic Puncturing dantitlon Son-shelled Invertebrates; fish Rlepple, 2002 Plesiosaurs Late Triasslc (Rhaetlan) Pelaglc Needls-sha ed teeth; pierce Soft-shelled invertebmtes or Jurasslc forms with Riepple, 2002; to Late Cretaceous; I and II gull& of Maseere, fleshy prey: same may have cephalopod hoddets; Massare, 1997; lobal distrlbution in the 1997 strained small prey from water gastroliths; ?regurgitates Pollard, 1968; !wassic and (Cxyptoclidus ) thwght to be from Martill et al., Cretaceous, but had high lesiosaurs with ammonold 1994; Wetzel, endemism (Lucas,1997) Parvae and shells of 1m &TU\;~~S Cretaceous Pelagic PursuR prdators; Cut and Large, fleshy prey: other reptllas, Riepple, 2002; Pierce II guild of Massare, fish, and csphalopods Massare, 1997; 1997: shake fesdIng Martill et at., Is94 Efasrnosaurs Cretaceous Pelagic Robustteeth, pletce I and li Rlepple, 2002; guild of Massare, 1997 Massare, 1997 Ichthyosaurs Early Triassic to late Nearshore Triassic) Cut, pierce, smash, crunch, Large, fleshy pre ; soft prey; soft Gastrlc mass with Massare, 1987, Cenornanlan (Late to Pelagic (S urasolc and cwsh gullds of Massare prey wlth internaYhardpa&; prey caphalopod Iwoklets, most 1997; Pollard, Cretaceous); achieved to Cretaceous); some (I987,1997) with bony scales or hard, thin likely from belemnites or 1968; Kellsr, global distribution by the were deep divers to a exoskeleton; prey with a very othertypes, In many 1976; Motani st Middle Tnassic (Lucas, depth of 600 m to ths TrTasslc tcth osaurs had hard exterior ichthyosaur skeletons; a1.. 1999 1997) mesopelagic layer of hetemlont bien~tion coprolltes indicate fish heocearl suggesting ambush, Large, rear teeth of Middle remains such as the Liassic (Ophihalrnosaums ); generalist predators in Trlassic icthyosaurs nectonic or necto-benthic ' Shonisaums (Upper nearshore habitats; in (Phalardm, Omphalosaurvs) fish, Pholldophoys; Triasac, Nevada) Jurassic, mostly homdont are suggestive of mollusc- ?Jurassic cuttle-f~sh; was outer shalf or dentition suggesting pelaglc crushing (Massare and Callaway, ?rnar~nereptiles; wood; basinal ln d~stributionpursuit predators that 1990), however th~stype none found with ammonoid (Hogler, 1992) specialize in a certaln type disa peared by the Late Triassic: or belemnite shells In of prey (Massare and the /her Triassic Grippia gastric contents Callaway, 1990) may also have been durophagous (Lingham-Soliar, 1999) (Table 2, cont.)

Mosasaurs Late Cretaceous Pelagic, ?Benthic Ambush predators; most Large, fleshy prey, prey wkh hard Stomach oontents Indicate Kauffman and (Cenomanlan to end mosasaurs occupied the exterior; Globidens was arnrnono~ds,blrds, f~sh, Kesling, 1960; Cretaceous); most were Cut guild; Cwsh and cosmopolitan in distribution and is smaller mosasaurs; Martin and Bjork, endemic to one region Pierce I1 guilds also tllought to be durophagous Clidastes, had a rnadne 1987; Massare, (Lucas, 1997) occdrred (Massare,1997) (L~ngham-Soliar,7999) shark and a diving marine 1937 bird Hsspsrwmis In gastric Q Opportunlstic general- contents (Marttn and Bjork, ists (Massare, 1997) 1987) b Crocodiles Early Jurassic. Benthic Ambush predators; Teleosaurlds (teleosaurs):flsh, Crmdile 1~0thassociated Martilt, 1985, : to Recent gensrallsts, occup Ing the turtles, and ammonites; with turtle scutes in a 1986; Hua and Cut, Pierce I and llCrunch, Metriorjnchids (geosaurs): telaosaurid; Wetaut, 1997: b and Crush gullds of ammonites, belemnites, Metrioit~ynchuswith Martel[ et al., Massare, 1997; Globldens pterosaurs, and giant fish abundant csphalopd 19M only crush guild marine Leedsichihys hwklets; assoc~atedscales teptlle since Triasslc of Lepidotes assxlated (Massare, 1997) with skeletal elements of Stsnemurzls; Metriohynchus tooth embedded in giant fish Leads~chthys Sea Turtles Jurassic to Recent Benthic to pelagic Generalists to speclallsts One form tday feeds on No fossil svidence molluscs (Caretta ); one form Tn tfie Lais Cretaceous ma slso have fed on rnollusw (drayma, 1997); Others feed on sea grass, ]ellvfish, and crustaceans Hybodont sharks Benthic: scavengers to predators; Predators on surface-llving Belemnltes; one speclmen Poltard, 1968 .r3 nett*benth[c Asferncanthus kd arnrnonolds (H@dus ) wFh over 250 durophagous dentidon rostra in gastrlc contents {Jurassic) Chimaeras Jurassic Bsnthlc Martill et al., 1994 ~ekelachlan Jurasslc some with durophagous Martill st al., sharks dentition 1w PALEONTOLOGICM SOCIETY PAPERS, Y 8,2002 hard- and soft-shelled invertebrates in the pelagic Some Wddle Triassic ichthyosaurs with large redlm (Rieppel, 1998). Teeth of sorile nothosaus, rear teeth may have been rnolluscivorous &lassare such as Simosaurus, have a bulbous shape which and Callaway, 1990).ln the Jurassic and Cretaceous, may have had a somewhat durophagous function the dentition became homodont, indicating that they (Rieppel, 2002). Anatomical evidence indicates may have become specialized on pelagic prey that Simosaums had strong neck muscles and was (Massare and Callaway, 1990). Perhaps they were capable of rapid jaw opening; suction also may specialists on fish andot: soft-bodied cephalopads have been used to round up shelled ammonoids or (Sander, 1997; Massare and Callaway, 1990). fish (Rieppel, 2002). Nothosaum mirabilis had Gasfziccontents indicate that they myhave fed on specialized jaw adductor muscles, heterodont belemnite cephalopods, although no belemnite or dentition with procumbent fangs, and a very narrow amrnonoid shells have ever been found in and elongate skull (Rieppel, 20021. The heterodont ichthyosaurian stomach contents. Based on body dentition suggests most nothosaurs ate fish, dthough form, by the end of the Triassic, ichthyosaurs were the gastric contents of one nothosawid (jbiosaancs) hydrodynamically advanced and were very fast- contained placodonts and small pachypleurosaurs swimming animals angham-Soliar, 2001). (Sander, 1989; Tschaxz, 1989; Rieppei, 20023. They Pterosaurs .-The appear~mceof pterosaurs in may also have eaten soft-bodied invertebrates, such the middle of the Triassic Period (Benton, 1993) may as cephalopods. Simsaum may have eaten hard- have increased predation pressure on near-surface shelled prey Weppel, 2002). Some nothosaurs, nektonic organisms, including .Ersh and cephalopds. because of their large size, may have been at the top The long jaws and impaling spike-2ike teeth of of the food chain. 'I rhamphorhynchids and many pterosauroids suggsts Pistosauroidea.-The Triassic Pistosamoidea a piscivorous diet in these flying reptiles. gave rise to the plesiosaurs that were common in the Jurassic and Cretaceous seas. Some pistosaurians TRIASSIC BENTHllC had jaws simiTar to those of the putatively fish-eating ORGANISMS: nothosaurids (e.g., Nothosauna), and others, such ANTIPREDATORY RESPONSES? as Pistosaums, had narrow and elongated pincer- type jaws, that had less numerous and widely-spaced Varied morphological and behavioral feames heterodont dentition with maxwfangs (Rieppel, of benthic invertebrates have been interpreted as 2002). Punctlrring prey, rather than suction feeding, antipredatory adaptations pig. 3), although many may have been the modus operandi of these of these features may be merely exaptations creatures, and they may have fed on soft-shelled (seam Gould and Vrba, 1982). Triassic benthic pelagic invertebrates and fish (Rieppel, 2002). faunas are decidedly "no frills" relative to those Ichthyosauria.-Ichthyosaurs (Order Tchthyo- of the jate Paleozoic (Valentine, 1973). Shells are sauria) (Figs. 2.4-2.6; 4) are known from the Lower relatively thin and mainly lacking in spines. In Triassic to Cenomanian Pardet, 1994), but they addition, several groups of cemented bivalves- have only recently kenstudied in detail (Callaway, the ostreids, gryphaeids, plicatulids, and 1997a).Triassic ichthyosaurswere nearly as diverse terquernids-first became abundant on hard and widespread as Jurassic ichthyosaurs, but are substrates in the Triassic. Harper (1991) has notoriously affected by preservational bias demonstrated experimentally that predators avoid (Cdaway, 1997b; Sander, 1997). Because of this cemented bivalves when given a choice. preservational bias, little is known about the The evolutionary breakthrough of mantle evolution of dentition in Triassic ichthyosaurs. Most fusion in bivalves led to the rapid development of of them likely had heterodoot dentition, indicating infaunal clades in the earIy Mesozoic (Stanley, that they were generafist feeders in nearshore waters 1977). Mud- and rock-boring bivalves also first (Massare and Calloway, 1990). became common during this time (Seilacher, 1985; WMKERAND BREJT-POST-PALEOZOIC PmERNS IN AlXNNE PREDATION

FIGURE %Temporal ranges of various potential anti-predatory adaptive or exaptative strategies in various invertebrate groups; semi: semi-endobenthic (quasi-infaunal); burrow: burrowing endobenthic organisms. Thin lines: present, but of limited abundance; thick lines: abundant; broken lines: possibly present but rare as fossiIs. PALEOiVTOLOGICAL SOCIETY PAPERS, K 8,2002

Bottjer, 1985).Deep burrows and endolithic crypts disappeared, whereas smooth involute forms may have been particularly effective protection survived, and the first heteromorph appeared. from grazing predators, such as phcodonts. The Deep outer-shelf and upper-dope environments near-synchronous development of this strategy in from the Early Triassic of China contained both at Ieast three independent lineages of venerid smooth and costate ammonoids; deep basin (Skelton et d.,199Q) and myoid bivalves, as well ammomids were smooth-she fled, some with fine as the great increase in endobentbic and endolithic sculpture, many of which are interpreted to have anornaldesmatans-all during the klyTriassic- been pelagic (Westemam, 1996). Coarse scuIpture, suggests intensification of some selective pressure however, is thought to be commonly associated with (Fig. 3). Bottjer (1985) and Skelton et al. (1990) basin-slope habitaL drew attention to the coincidence ofthis downward Large pelagic predators, such as ichthyosaurs, push wid the Mesozoic Marine Revolution. plesiosaurs, placodonts, and turtles had evolved by However, MeRoberts (2001) has recently argued that the Late Triassic, and many are thought to have durophagous predators may not have been eaten ammonoids; ceratitic amrnonoids do not sufficiently abundant or widespread during the seem to show classic antipredatory defenses. Tflassic to account for the early radiation of However, the temporal trends, if any, of ceratite endobenthic strategies. It is possible that the shell injuries rem~nto be studied. antipredatory advantages of living cryptically were Echinodem,-Echino went through an merely a side-benefit of adaptation driven by other evolutionary bottleneck after the Permian pressures, such as competition (McRoberts, 200 1j . extinction, with at least five classes snrvivlxlg into The overall frequencyof shell repair, due either the EarIy Triassic (Sirnms, 1990). From Iow to predators or to physical factors in the diversity in the Triassic, echinoids and crinoids environment (see Cadke et al., 1997; Cadke, 1999; diversified in the Middle Triassic, but some clades Ramsay et zI., 2001) is also low during this be went extinct during the mid-Carnian. A second interval, with repair frequencies evidently even diversification occurred in the Norian and the Early lower than those of the late Paleozoic, as recorded Jurassic for both groups. Triassic crinoid forms re- by Vermeij et al. (1982) (Table 3). evolved "'passive" filtration systems like their Ammonoids.-Vemeij (1987) drew attention Paleozoic forebearers. Most post-Paleozoic crjnoids to the fact that surveys of ammonoid shell are thought to be anatomically similar to their architecture and traces of predation on cephalopods Paleozoic ancestors; however, Donovan (1993) and are critically needed for the whole Phanerozoic. A Oji (2001) provide evidence that the Mesozoic general view of amrnonoids suggests that their crinoids (especially the Jurassic fonns) were agiIe morphology is related to their pelagic, demersal, and could actively relocate-this hayhave provided or planktic lifestyle (Westermann, 1996, p. 689), a selective advantage as predation pressure increased rather than to antipredatory features. (Meyer, 1985). PseudopIaaktic pentacrinitids, Early Triassic ceratitic ammonoids from paracomatulids, and the true comatulids evolved in pIatfom environments are considered to have been the Late Triassic and occupied niches altogether chiefly nektonk in habit, although some planktonic different from their Paleozoic counterparts (Simm, and demersal forms occurred (Westemam, 1996). 1990). These new modes of life probably do not Offshore bituminous limestones of the Middle and reflect escalation; overall, predation on these Upper Triassic in Europe, North Arnerica, and China echinoderms is deemed to have been relatively low also contained coarsely costate to smooth ammonoid during the Triassic (Schneider, 1988). morphotypes, all of which were interpreted to be The two main ciades of echinoids, the pelagic (including some with planktonic lifestyles). Diadematacea and Echiaacea, diversrfied in the In the Late Triassic (Norim), however, most highly Late Triassic and Early Jurassic, but still retained sculpted evolute ammonoid morphotypes their mid-Paleozoic diversiv levels (Simms, 1990). WALKER AND BRETT-POST-PALEOZOIC PATTERNS IN MARliVE PREDATION

Nektonic Mobile Nektonic! Predators Nektobenthic Prey

+Seals, whales Marine birds

Mosasaurs + Hesperclnithines and + Teleosfs (abundant) lchthyornthine birds +Hesperonithines and lcthyomithine birds Pliosauroids + Neoselachian Sharks --+ Plesiosaurids + +Belemnites (common) +Teleosts (first) Ichthyosaurs + +"Holostean" fish "Holastean" fish

FIGURE 4--First appearances of major groups of Mesozoic and Cenozoic pelagic predators and potential prey. Arrows point to approximate time of first appearance of taxa.

The echinaceans developed carnivorous and of the evidence for vertebrate predation on Jurassic herbivorous habits, ancI others staxed to bore into prey is cjxcumstantid, based on overlapping faunal rock substrates. As noted, this latter could reflect compositions of predator and prey, interpretation of an antipredation strategy, but there is little tooth form, and attempts to match dental form with information available on predation-related injuries putative bite marks. Nevertheless, there is tantaking in these organisms degthe Triassic. evidence of predation. Fish (including sharks), JURASSIC PREDATORS ichthyosaurs and plesiosaurs are considered the dominant vertebrate predators (Figs. 1, 4; Much more is known about Jurassic predators Tables 1, 2). These organisms could function as chiefly because of the greater extent of both pelagic and benthic predators, so heir predatory epicontinend sea deposits compared to the Triassic. activities cannot be exclusively tied to either of these There are also many significant Lagerstatten from realms {see Martill, 1990). Alternatively, some the Jurassic. Still, information concerning predator- interprektions suggest that the dominant marine prey relationships for the Jurassic is limited. Much reptiles at this time were all pelagic predators TABLE 3--Shell repair in bivalves and gastropods from Mesozoic and Cenozoic Localties. Only papers that cite their raw data, or semblance of such, are included in this table. According to Vermeij (1 982, p. 233), the higher the frequency of scars, the greater is the role of the shell in protectingthe gastropods against lethal breakage; if selection for armor is weak, then the frequency of repair should be low. Thk exceptional preservation required to detect shell repair is rare, especially in Paleozoic and Mesozoic gastropods (Vermeij et al., 1981).

Taxonomic Group; Evidence of Shell CalcuIation of Shell RangelMeanlmedian frequency AgelLocality; Repair Repair Frequency Reference

Gastropods Broken edge is repaired; Number of scars ' 0.00-0.28/0.095*lno median tiny chips ignored; some per shell Upper Triassic; Northern Italy, St. had only body whorls n=558 indivs, I I spp. Cassian Group, dolomites; exposed, so not all shell species described areas were examined; "SEW calculated; differs from 0.08 reported in Vermeij et at., small shells examined 1981. If 2 spp. were excluded that had zero shell repair, then Vermeii et al.il9821 (dgrnrh) the mean frequency would be 0.12. Gastropods Jagged trace of the outer Number of scars no range/0.35"/no median* F lip where the latter has per shell w "Their Note 5 states that median frequencies are reported in N Cretaceous Ripley; species not been damaged and table I;text states mean frequencies are recorded. SEW is described subsequently repaired assuming these are mean frequencies here. Vermeij et al. (1981) 24 species - -- Gastropods As For Cietaceous Ripley Number of scars 0.1 9-0.33/0.40*/no median per shell -8 Miocene, all specles, (Gatun 1-3); *SEW calculation of 0.29; this is based on mean frequency for s species not described. all spp, from Gatun 1-3. 9

Verrneij et al. (1981) 19 species N A- . 8 Gastropods As for Cretaceous Ripley Number of scars 0.08-0.4410.28*/no median LU per she1 t Recent, all species (Panarnd, New *SEW caiculation: 0.28 is correct based on data presented. Guinea, Mindanao, Hamahera,

Guam) , Vermeij et al. (1981) (Table 3, cont.)

Bivlaves: Osteids, Exogyra, Repair scars that divert or Total number of repair No median recorded. =4 Pycnodon te cut across the normal scars on left valves Exogyra: 0.00-1 .I1 */0.4[4; Pycnodonte: 0.12-0.38/0.23 concentric growth larnellae divided by the total Combined (Exogyra and Pycnodonfe ) mean frequency: 0.34 $ Late Cretaceous (North Atlantic (including: scallops, ' sample size of repaired *Because more than one repair per shell was used, the B Coastal Plain, New Jersey} meandering clefts, divoted and uninjured valves frequency can exceed 1.0. repairs, and more x, Campanian: 0.I1 ; x, Maastrichtian: 0.44 b Dietl et al. (2000) extensive repairs including n =?528 or ?525 Exogrya spp., ?460 or ?450 Pycnodonte : irregular fractures) spp. for Cretaceous localities, total -7988; they state n = "> 8 1600 indivs," 7 spp. {Paleocene + Cretaceous) h Bivalves: Ostreids, Pycnodontes As for bivalves As for bivalves 0.07-0.09/0.0&; for all size classes, mean = 0.1 0 (from their 9 table 5) early Paleocene V Dietl et al. (2001) n = ?77, 1 sp. Pycnodonta dissmiEai-is % Gastropods Jagged repair scars on Number of individuals Excluding samples with 1 individual: 2 shells with one scarltotal 0.03-0.59/0.29/no median (their table 10, p. 349-3501 lower Pliocene; Albenga, Italy F individuals of that W Robba and Ostinelli (1975) species n = 3090 indlv.; 21 spp. W ------2 Bivalves: Anadara and Coxbula Jagged repair scars on As for gastropods 0.07-0.3510.21 only shells n=896, 2 spp. (corbulids had the highest shell repair lower Pliocene; Albenga, ltaly frequency at 0.35) Robba and Ostinelli (1975)

Gastropods: Terebrids Repairs were recorded Number of repaired Median frequency reported (their table 1y: Recent: n= 5735, 2 only if they extended 20% injuries divided by the 0.54; Pleistocene: n =I 10, 0.55; Pliocene, n = 314, 0.54; Eocene-Recent; troplcal to or more of the whole in a total number of shells In Miocene, n = 549, 0.57; Paleogene, n = 136, 0.47 subtropical spiral direction or if they the sample; frequsncy involved subjectively of repair is loosely * n must equal number of samples, but it: is not clear; stated 2 Vermeij et al. (1981) substantial breakage; correlated with the that samples with ten or more individuaEs were used; E excludes minor lip breaks number of species of frequencies of repair have remained unchanged from the kl she1[-peeling ca[appids Eocene to Present

Gastropods: Littorinids No information; appears to Frequency of snails with Many localities, here totaled together: 0.00-0.48/0.11 be large, jagged repair damaged shells Recent; cold temperate scars (his fig.-. 1) estimated for each n = 4593 Rafaelli (1978) population PALEONTOLOGICAL SOCIETY PAPERS, V 8,2002

(M~sare,1987). Importantly, the range in tooth form trophically complex functional groups similar to and function in Jurassic (and Cretaceous) marine those in the Middle Jmsic. Some ammonites may reptiles was at Ieast as great as that of modern marine have fed on both the plankton and the benthos, mammals (Massare, 1987). depending on food availability and benthic anoxia. Gast~-opo&.-Although naticid mes ogastropods Ammonoid forms at this time had costae or nodose (Fig. 6.4) existed in the Jurassic, boreholes are quite macroconch, and microconchs with horns on some rare. However, recent discovery of drjlled shelIs species; smooth shelled ammonoids were also proves that the capacity for drilling predation did common. Numerous records of ammonoid aptychi exist (Kowalewski et al., 1998). are reported frornthe body chambers of haploceratid Nauti1oids.-Nautiloids were very diverse in ammonites, bdicahng predation; and specimens of the PaIeozoic, but there were few nautiloids in the the Late Jurassic monoid, Neochetoceras, have ~esozoic(House and Senior, 1981). Nautiloids are aptychi of conspecific juveniles within their body thought to have continued with their Paleozoic chambers, indicating cannibalism (Westermann, predatory mode of life, perhaps scavenging or 1996, p. 676). A rare fi.d of a Saccocoma crinoid preying on crustaceans (Fig. 1).A nautiloid with a among the stomach contents of Physodocerm is complete jaw apparatus (rhynco'tites) is known known from the Solnhofen Limestone (Milson, from lithographic limestone, Upper Jurassic of 1994). Saccocoma is variously interpreted as either southwestern Germany (Diet1 and Schweigert, planktic or benthic in habit (Milson, 19941, and 1999). Modern nautiloids can repair their shells depending on the interpretation of the Life mode for (Meenahhi et al., 1974), although little is known Saccocoma, the ammonoid is interpreted as either a about shell repair in Mesozoic nautiloids. pl&c or a benthic feeder (the latter interprktation Ammonoids.--Shell shape in ammonoids is is favored by Westermm, 1996). sometimes used to infer directly whether or not Echinoderm Predators.-Living families of they were predatory. For example, large asteroids (e.g., Forcipulatida and Notomyotida) macroconchs of oxyconic forms are interpreted to have their roots in the Early Jurassic (Hettangian) be mobile predators (Westerma, 1996). Shell of Germany and Switzerland (Blake, 1993). shape, sculpture (Fig. 6. I), and size (especially for Complete asteroids are exquisitely preserved jn macro- and.microconchs) can also be explained pelletoidal calcarenite from this time period. by sexual dimorphism (Westemann, 1996). In Modern forcipulatids are known to prey on other terms of direct evidence, there is only one Early echinoderms, molluscs, barnacles, and many other Jurassic example of an ammonoid (Hildocerus) types of invertebrates. The presence of myarms with aptychi of juvende ammonids within its body in asteroids (e.g., solasteroids) susgests that they chambers (Westermann, 1996). were predators of active prep, such as other Middle Jurassic ammonoids appeared to asteroids. Predation on active prey by solasteraids occupy anumber of trophic functional groups, from most likely evolved in the Jurassic (Blake, 1993). planktonic to demersal forms that presumably fed Asteriids, in contrast continued to feed on on ostracodes and microgaswopods in algal mats molluscs and other benthic prey as their Paleozoic (Westemam, 1996), although there is no data on ancestors did. During the Jurassic, asteriids had gastric contents to co&m this. The lower Toarcim promitrent adambulacral spines that their rnodern Posidonia shale (northwestern Europe) is known descendants no longer have; it is thought that these to have clusters of fragmented harpoceratine spines functioned to trap prey (Blake, 1993). arnmonoids, presumably from cephalepod Decapods-Despite the common assumption predation (hhmann, 1975). In Urn, the stomach that shell-crushing crabs evolved during the Jwassic, contents from a harpoceratine indicate that it-preyed in reality, only one group of lobsters (the on small or juvenile ammonoids [hhmann, 1975). Nephropidae) is hornto have evolved during this Finally, Late Jurassic ammonoids had time. All other groups evolved during either the WUKERAND BRE7T-POST-PALEOZOIC PATTERNS LN MXRINE PREDATION

Paleozoic, Triassic, or Cenozoic (Table 1). Hermit containing partially ingested smaller fish. Jurassic crabs evolved in the Jurassic (Glaessner, 1969), and pycnodont reef fish developed deep-bodied while they may crush mollusc shells (Vermeij, morphologies. For example, Drsepedium (Fig. 5) 19871, it is diff~cultto assess their overall importance was a deep-bodied Jwassic marine fish with heavy aspredators on molluscan groups. Hermit crabs can ganoid scales, but with pebbIe-like teeth for be scavengers, carnivores, filter feeders, or crushing. Jurassic pycnodonts evolved batteries of debitivores (Sch,1986). rounded, shell-crushing teeth, plus specialized Chondricthyes.-The rapid radiation of sharks nipping teeth. A few py cnodontids even developed and marine reptiles (Figs. 1, 4) in the middle stout pavement teeth possibly for crunching corals; Mesozoic may have been triggered by the rise of rare specimens have been found with coral vast numbers of squids and actinopterygian fishes, fragments in the gut wohl, 1990). The general including semionitids and basal teleosts (Theis and morphology of these fishes overlaps with that of Reif, 1985). The advances of increased swimming deep-bodied platysomids of the Paleozoic and efficiency and maneuverability, and sensory ability many reef-dwelling Cenozoic teleosts. enabled the neoseIachians to pursue fast-swimming Fish with durophagous dentition, such as thin-scaled fishes and squids in nearshore Semionotidae (Lepidotes, Heterostrophus), environments (I'ackard, 1972). Pycnodontidae (Mesturus), as well as hybodont Hybodont sharks of the Triassic (Fig. 5.4) were sharks (Asteracanthusare) and chimaeroids largely supplanted by the expanding neoselachian (BrachymuIus, Pachymylus, Ischyodus), are sharks during the Late Jurassic. The evolution of thought to have been predators of ammonoids from highly flexible, hyostylic jaws clearly marked a the Middle Jurassic of the Lower Oxford Clay of new level of sophistication in shark predation England (Martill, I990). Many well-preserved (Maisey, 1996). In hyostylic suspension the upper ammonoid fragments are thought to be the result jaw is loosely articulated to the braincase and can of fish predation rather than physical factors by swung downward and forward on the (Martill, 1990). One ammonite specimen, a hyomandibdar bone. This enables sharks to thrust Kosmoceras, was found to have bite marks that the jaws forward and gouge out large chunks of were similar to the dental battery of the semionotid flesh from prey. This adaptive breakthrough fish, hpidotes macrocheirus (Mm,1990). fomented an adaptive radiation of sharks, which Sea ttkrtles.-Turtles are the only living continued through the present day. Modern sharks reptdes that are Mly adapted to a marine existence show varied feeding modes, incluhg grasping and (except for egg laying). Many fossil sea turtles swdIowin,o, suction feeding, cutting, goug,and are only known from their plastron and carapace crushing (Moss, 1977). One skongly modified (Nicholls, 1997). The earliest sea turtles are the clade from within the neoselachian shark Lineage Plesiochelyldae, possible predators that lived in is the highly successful Batomorpha: rays and shaltow, coastal waters. skates. These fnst appeared in the Late Jurassic Saumpterygim:Plefiosaurs mdplioscsu~s.- but diversified jn the Cretaceous. The dental plates The pIesiosaurs are thought to have diversified into of rays and chimaeroids of this type may be used two major grades during the Jurassic (Fig. 5; for digging up shelled invertebrate prey, and then Table 2): the short-necked forms as fast-swimming crushing them, leaving only fragments. pursuit predators (pliosaurs), and the long-necked 0steichthyes.-Among the Jurassic bony forms as lurking ambush predators (plesiosawoids fishes there is evidence for common piscivorous and elasmosam&). 0' Keefe (20021, however, has habits; for example, the famed Upper Jurassic called this an oversimplified view of their actuaI Sotnhofen Limestone provides many instances of morphological diversity. predator-prey interactions (Voihl, 1990). Most A cladistic analysis revealed that plesiosaurs "fossilized interactions" involve fish carcasses present a spectrum of body forms, and do not PALEONTOLOGICAL SOCIETY PAPERS, V 8,2002

FIGURE 5-Mesozoic predatory marine vertebrates. 1, Cretaceous ichthyodectid teleost Xi?hactinus. 2, Skeleton of the pycnodontid fish Proscinetes, Jurassic. 3, Skull of ichthyodectid teleost fish Cladocyclus, Cretaceous; Late Cretacous. 4, Hybodid shark, Hybodus. 5, Geologic distribution of marine reptiles, from [eft to right: ichthyosaurs, piesiosauroids, pliosauroids, teIeosaurs, metriorhynchids, and mosasaurs. 69,Skeletons of' Mesozoic marine reptiles: 6, Mosasaur Plofosaurus,Late Cretaceous. 7, Plesiosauroid Muraenosaums; Jurassic. 8, PIiosauroid Peloneslstus; Late Jurassic. 9, Ichthyosaur Ophfhalmosaunrs; Early Jurassic. 10, Cret8ceous foot-propelled diving bird Hesperornis. 11 , Skeleton of Creaceous marine bird Ichfhyomithyes.Figures 1-4, 10, 11 adapted from Benton (1 997); Figures 5-9 from Massare (1 987). WALKERAND BRETT-POST-PMEOZOIC PATTERNS IN MARINE PREDATION discretely fall into two basic shapes: from the long- carnivorous plesiosaurs (LEopleurodon,Pliosauaar) necked, small-headed elasmosaurs to the short- were considered to be at the top of the Middle necked, large-headed pliosaurornorphs [O'Keefe, Jurassic food chain, presumably feeding on fish and 2002). By the Late Cretaceous, these pelagic "naked" {without a shell} cephalopods (Md, masine reptiles were globally distributed (Rieppel, 1990). The ichthyosaur Ophbhamosaums was 1997). But the taxonomy of this group is still poorly thought to be a specialist on naked cephalopods, known because of the inadequacy of type material, while marine crocodilians (Metriorhynchus, and preservational problems such as skull-less Steneosauats) presumably fed on fish and naked skeletons (Carpenter, 19973. cephdopods (Martill, 1986%1986b, 1990). Massare The plesiosaurs (clade Plesiosauria) were (1987) examined the conical pointed teeth form of among the most diverse, geologically long-lived, some Jurassic ichthyosaurs and plesiosaurs, and and widespread of the Jurassic to Cretaceous concluded that the teeth functioned to pierce soft marhe reptiles, with a fossil record extending from prey. Fish from these deposits were either plankton the Triassic-Jurassic boundary to the Late feeders or fed on smaller fish, indcating that the Cretaceous (F'igs. 5.5, 5.7) (Carroll and Gaskill, Middle Jurassic had a highly complex marine food 1985; Rieppel, 1997). These large reptiles [up to web (Ma,.till, 1990). 15 m long) had long paddle-shaped limbs Ichthyosaurs.-Lower Jurassic localities from (considered hydrofoils), short tails, long necks, Europe (e.g., Lyme Regis, England; Holzmaden, needle-shaped conical teeth, and may have swam Germany) indicate that marine reptile guilds at this Iike modem sea lions (Godfrey, 1984; Carroll and time were dominated by a diverse array of Gaskill, 1985). Plesiosaurs are the only marine ichthyosaurs (Figs. 5.5,5.9) (Massare, 1987). Cmtric animals in which both forelimbs and hindlimbs contents from ichthyosaurs are hornfrom Lower performed as lift-based appendicular locomotion Jurassic localities in Europe (Pollard, 1968; Keller, (i.e., as hydrofoils; Stom, 1993). Pliosaslrs had large 1976; Massare, 1987). The majority of preserved skulls (up to 3 m) and jaws with large fang-like teeth food remains were cephalopod hooklets (Massare, (Taylor, 1992), and were capable of dismembering 1987, her table 1, p. 128). For example, preserved their prey (Taylor and Cruickshank, 1993). cephalopod hooklets (interpreted to be from The small-reIative skull size and neck length belemnites), fish remains, and wood were present of plesiosaurs, in addition to their dentition, in the gaslric contents from the small (< 3 m) Lower suggests that many ofthern may Rave fed on small Jurassic ichthyosaur Stenupte~gius(Keller, 1976). fish and soft-bodied cephalopods; some may have Putative phagmoteuthid cephalopods also were also strained the water for prey (Massare, 1987; preserved in the stomach contents of the small Rieppel, 1997). Their evolution may have been Lower Jurassic icthyosaur, Ichthyasaurm (Pollard, stimulated by the new abundance of larger 1968). No belemnite hardparts (besides hooMets) actinopterygian fishes and sharks in offshore have been reliably found in ichthyosaur gut marine environments. Plesiosaurs from the Middle contents (Massare, 1987; but see Pallard, 1968). Jurassic of the Oxford Clay also are thought to have In conbast to the Lower Jurassic, Middle to Late been specialists on soft-bodied cephalopods and Jurassic assemblages indicate a number of changes fish (Maaill, 1990). The gastric contents of one Late in the vertebrate predatory ensemble (Massare, Jurassic plesiosaur, Pliosaums brachyspondylus, 1987). Although the same functional feeding types included cephdopod hooklets parlo, 1959). Wetzel (based on tooth form and wear) were present, the (1960) has reported small ammonites in coprolites reptile groups were different, with pliosauroids and attributed to plesiosaurs. crocodiles dominating the assemblages, and with Case studies from the Middle Jurassic Oxford reduced ichthyosaur diversity (Massare, 1987). The Clay, Uaited Kingdom, provide a window into the Middle Jurassic cephalopod-eating ichthyosaur, marine trophic relationships of this time period. The 0~hthalamosaun6 (Fig. 5.9) is inferred to have PALEOiVTOLOGICAL SOCIETY PAPERS, V 8,2002 dived to depths of 600 m, based on an analysis of Pholidosauridae (Teleorhinus)are thought to have its eyes and bone condition (Motani et al., 1999). been piscivorous. Two groups of Wedryosaurids Ichthyosaurs are also thought to have regurgitated (Phosphatosawinae and Hyposaurinae) are hown: hardparts of indigestible food. In the Peterborough the phosphatosaurins had blunt teeth and robust quarry in England, Peter Doyle (unpublished, jaws, and are thought to have preyed upon Mes 2002) discovered "ichthyosaur regurgitates" of 160 and nautiloids; the hyposaurins, most common in million-year-old acid-etched juvenile befemnites. the Paleogene, had long slender jaws and pointed The acid-etched fossils indicate that they were once teeth and were probably piscivorous (Hua and sinthe ichthyosaur stomach. Buffetaut, 1997). Crocodilians are known to undergo Marine Crocodiles.-Little is known about the rapid changes in dental morphology in response to fossil history of marine crocodiles (Suborder environmental change related to dietary Mesosuchia) compared to other marine reptiles (Hua modification. It is thought that the piscivorous mode and Buffetaut, 1997). The earliest crocodiles of life became more common after the Cretaceous (Teleosadae) are known fiom the Early Jurassic. mass extinction, when ammonoids and hard- This group shows littIe adaptation for marine life, shelled marine reptiles were not as common and it is only because they are found in shallow (Denton et al., 1997). However, the extinction of marine deposits that they are infened to have been the dryosaurids in the Eocene is thought to have marine crocodilians Wua and Buffetaut, 1997). resulted from the expansion of whales, whch may Later in the Jmssic, these forms showed anatomical have competed with them for food {Hua and features that were more characteristic of life in Buffetaut, 1997). Crocodiles also regurgitate their marine conditions (e.g., stieamlined skull, reduction prey and such remains have been reported from in bony amor, and reduction of the forelimb). Some the Paleocene of Wyoming (Fisher, 1981 a, 1981 b) forms (Steneosaurus) had long, slender teeth and but not from the Cretaceous. may have been piscivorous. Other teleosaurids had CRETACEOUS PREDATORS blunt teeth, and more robust jaws, and are thought to have been durophagous predators on monoids The Early Cretaceous marked the beginnings or sea turtles (Hua and BuEetaut, 1997). of a major reorganization of marine predators, The Early Jurassic to Early Cretaceous including the rise of neogastropods, numerous Mehiorhpchidae include crocodilians with both cephalopod predators, and several new vertebrate long (Iongiroshe) and short fbreviroshine)snouts predatory gudds (Figs. 1,4-6).The Early Cretaceous that may reflect dietary differences (Hua and saw the radiation of large teleost fish (> 3 m in Buffetaut, 1997). Tkis group, because of its more length) md sharks, and the non-dominance of marine streamlined body form and a skull similar to reptiles (Massare, 1987). Massive shell-crushing mosasaurs, is thought to have been pelagic. The mosasaurs (e.g., Globidem) did not evolve until stomach contents of a brevirostrine form the Late Cretaceous. This major-specialized (Metriorhynchusj contained ammonites, belernnites, functional feeding type had been essentially absent pterosaurs (Rhamphorhy~zchw},and fie large fish throughout most of the Mesozoic, since the hedsichthys (Martill, 1986b). Metriorhynchm and extinction of Triassic placodonts (Massare, 1997). their ilk were probably lun_&g ambush predators Late Cretaceous marine reptiles were dominated that captured their prey by sudden bursts of by ambush predators, such as mosasaurs; marine swimming (Massare, 1987). fish (including sharks) were much more common Two other groups of marine crocodiles, the at this time and became more dominant Pholidosauridae (Lower-Upper Cretaceous components of the predator functional feeding boundary) and Dryosauridae (Upper Creatceous to guild than ever before in the Mesozoic. Marine late Eocene) had fiesh- and salt-water members (Hua reptiles such as plesiosaurs and ichthyosaurs were and BuEetaut, 1997). The marine species of minor components of the Cretaceous predatory WALXERAND BRETT-POST-PALEOZOIC PATTERNS IN MARLNE PREDATION

FIGURE 6-Representative Mesozoic marine invertebrate predators. 1, Ammonoid; note fluted ornamentation. 2, Hornarid lobster. 3, Belernnite; Belemnitella. 4, Naticid gastropod. 5, Brachyuran crab. 1, 3 from Tasch (1980); 2, 4, 5 from Robison and Kaesler (1987). food webs. In fact, ichthyosaurs and the among the neoselachian sharks (Benton, 1997). pholidosaw marine crocodiles became extinct in Gastropods.-The Cretaceous marks an the Early Cretaceous, and pliosaurids were rare important heof evolution in the predaceous shell- (Massare, 1997).Ichthyosaur extinction may have drilling gastropods. SeveraI groups appeared and been associated with the Cenomanian-Turonian or diversified during the Late Cretaceous and their boundary events, following a severe depletion in distinctive drilling traces (Oichnus) become their putative belemnite prey (Bardet, 1992). common at this time (Kowdewski et al., 1998). During the Cretaceous and Tertiary, the offshore Naticids (Figs. 6.4,7, 8.3) become abundant in the movement of fast-moving fishes and coleoids may Late Cretaceous as do their diagnostic boreholes have stimulated evolution of offshore hunting (Fig. 8.4) (see reviews by Kabat, 1990; Rowden~s~ PA EONTOLOGICAL SOCIETY PAPERS, V 8,2002

1993). The drilling frequencies in some Cretaceous Muricids (Neogastrapoda) also evolved in the samples equal or exceed those observed in early Late Cretaceous; predaceous muricids produce Cenozoic samples from the Gulf Coastal Plains characteristic cylindrical, non-chamfered bowholes (KelleyandHanson,1993,20Ol;KeUeyetal.,2001; (Fig. 8.3). Muricids form an ecletic gustatory see discussion below). These studies are possible group, ranging from herbivores to canion feeders; because naticids leave a unique type of counted however, most are shell drillers (Kabat, 1990). drillhole in scaphopad, bivalve, gastropod, and Shell drilling is most likely a pleisomorphic conspecific gastropod prey, as well as other behavioral wait within the Muricidae, although not organisms (Cder and Yochelson, 1968; SOH, allmuricid genera bore through hard exoskeletons 1969; reviewed by Kabat, 1990; Kowalewski, 1993). (Vermeij and Carison, 2000). In contrast to naticid

FIGURE 7-Diversification patterns of shell drilling through time: upper flgure shows diversity of drilling gastropod clades through the hlesozoic and Cenozoic eras: iower figure shows frequency of drilled prey per million years through the Phanerozoic.Adapted from Sohl(1969) and Kowalewski et al. (1 998). WALKER RiVD BRETT-POST-PALEOZOIC PATTERNS IN MARINE PREDATION boreholes, holes drilled by muricids are considerably most are scavengers (Schram, 1986). The majority less frequent in the Cretaceous than in most Eocene of the durophagous forms evolved in the Cenozoic, and younger samyles (Vermeij, 1987j. with just a few forms evolving in the Cretaceous Cepha1opods.-As in the Jurassic, there were (Table 1). The portunids and xanthids evolved in a host of belemnoids, ammonoids and nautiloids the Cretaceous, and today are generalist and present in the Cretaceous (Fig. 6). All of these were opportunistic feeders, occasionally eating hard- probably nektonic predators, though their food may shelled prey like molluscs. The slipper lobsters may have ranged from zooplankton and larvae to other have evolved in the Late Cretaceous, and they are cephalopods (Packad, 1972). Cretaceous octopi thought to feed on scyphozoans (Table 1). are also known. While modem nautiloids appear Chondricthy es.-The neoselachian sharks to be extensively drilled by octopods (Saunders et radiated during the Cretaceous. Cartilaginous shark al., 1991), no data exists for drilling predation on skeletons do not fossilize well, and consequently, Mesozoic nautiloids. As in the Jurassic, Cretaceous their teeth are used to infer their feeding behavior ammonoids are thought to have been carnivorous. {Shimada, 1997). Despite popular accounts that However, many species are thought to have eaten Cretaceous sharks were some of most voracious zooplankton (Ward, 1986). of all predators, it is still not clear whether their Stomatopods.--Stomatopods are known to attacks were on live or scavenged organisms. have extreme specialization in their limbs chat is Healed injuries are usually taken to be attacks on related to their predatory activities; no other major live prey, but these are rare in the fossil record. extant malacostracan group has such specialization. Necrosis around bite marks is also used to infer (Kunze, 1983). All stomatopods are obligate predatory shark attacks (Schwimcr et al., 1997). carnivores (Table I )-they eat only live prey-and Additionally, animals associated with shark use their large raptoria1 second maxillipeds for prey remains are usually interpreted as the shark's East capture (Kunze, 1983). These folding raptorial meal or as associative potential prey. For instance, thoracopods can be used in two ways: as either in the Late Cretaceous Niobrara Chalk, a lamiform smashing or spearing appendages. Folding raptorial shark {Cretoxyrh'hinarnmlelii) is accompanied by thoracopods are known from the Carboniferous well-preserved cartilagenous skeletal elements palaeostornatopods (Schram, 19691, and within the presumably from its last meal, the fish Xiphactinm Mesozoic forms. The extant superfamilies of audax (Shimada, 1997). stomatopods are thought to have originated in the Late Cretaceous lamniform sharks Cretaceous (approximately 100 Ma; Ahyorrg and (Cretoxyrhi~tu)up to 6 M in length attacked or Harling, 2000); however, the true fossil record of scavenged mosasaurs and perhaps plesiosaurs, ad, this group begins in the Cenozoic, and will be in turn, were themselves possibly attacked or discussed in that section. scavenged by other sharks (anacoracids; Shimada, Based on fossil mouthparts, specialization for 1997). Dental arcades of Cretoqrhina are similar the stomatopod's zealous carnivorous life style to those of modern predatory mako sharks, and, evolved very early, by the Late Devonian or Early not surprising!^, they belong to the Family Carboniferous, and the trend continued into the Lamnidae that includes the mako (Isurn), great Mesozoic (Sch,1979). Mouthpm shred the white (Carcharodon), and salmon shark (Lcunm) prey, and food is stuffed into the mouth, not unlike (Shhda, 1997). Although shark taxa are different the way an energetic, hungry teenager feeds. through geologic history, Late Cretaceous sharks' Undigestible shell and cuticular material is functional feeding capabilities in ecosystems show regurgitated. The regurgitated remains have not parallels to modern sharks (Shimada, 1997). been examined fkorn a taphonomic perspective. Direct evidence of shark predation on mosasaurs Decapods.--In contrast, to stomatopods, is very rare. Shimada (1997) discusses several decapods are not obligate carnivorous predators; reports of putative shark attacks on mosasam, either PMEONTOLOGICRL SOCIETY PAPERS, V: 8,2002

FIGURE 8-Traces of predation in fossil and Recent shells. 1, Shell of Cretaceous ammonite Placenticeras with rows of punctures, probably made by amosasaur. 2, ~ecent'gastropodshell (Fasciolaria) exhibiting peeling damage inflicted by a callapid crab, XI.3,4, Profile views of incomplete and complete gastropod drill holes in cross sections of bivalve shell: 3, cylindrical muricid drill holes; 4, ZypicaI naticid holes; note parabolic cross section and central boss in incomplete borehole. 5, Locations of most frequent drill holes in Recent bivalves from the Niger Delta. Redrawn from photographs in the following sources: 1, Kauffman and Kesling (1960); 2, Bishop (1975); 3, Reyment (1971); 4, Sliter (1971). figure modified from Brett (1990). with shark teeth embedded in bone, tooth marks mosasaws without evidence of healing are also slashed into the bone, or evidence of gastxic-acid reported (Hawkins, 1890). Shark bite marks are &o etchingonputatiYepreyitems.RothcbildandMartin known from elasmosaurid plesiosaur bones (1990) report on a shark tooth embedded in mosasaur mlliston and Moodie, 19 17; Welles, 1943). (Clihtes)bone, which subsquently was repaired Late Cretaceous galeornorph selachian shark and ultimately caused spondylitus. Bite marks on (Squalicorcx) are thought to have been scavengers WALKERAND BRm-POST-PALEOZOIC PATTERNS IN MARINE PREDATION par excellence in the eastern Gulf Coastal Plain Jurassic and Cretaceous (Benton 1997; see also and Western Interior of the United States Vermeij, 1987). These specialized elasmobranchs (Schwimer et al., 1997). All living ~eoselachian were adapted in large part for durophagous benthic sharks are carnivores, while galeomorph sharks predation. Rays evolved stout hypermineralized may prey on mollusks, crustaceans, and pavement plates for crushing hard-shelled prey, such vertebrates; Squalicorax is also thought to scavenge as crustaceans and molluscs (Fig. 9). Ray dentition many types of prey (see table I of Schwimmer, et is thus similar to the pavement teeth of Devonian d.,1997). Evidence of scavenging may include ptyctodonts and rhenanids, and late Paleozoic embedded teeth which do not show evidence of holocephalans, Triassic-Jurassic semionotid fish, wound healing or tissue necrosis, and shark teeth and Triassic placodonts. In all cases, crushing of associated with putatively scavenged remains hard-shelled prey is inferred, but of these groups (Schwimmer et al., 1997). Squalicorax, a certainly the batoid rays have been most successful. moderately sized shark at 3.5 m, had serrated Many rays, exemplified by the cow nose rays, are dentition of the "cutting type," whch may indicate capable of excavating shallow pits in sandy relatively diverse feeding strategies. Isolated substrates in pursuit of infaunal bivalve, gastropod, Sqnalicorax teeth, sometimes with associated bite polychaete, and other prey {see Fig, 12). Possible marks, were reported embedded in a decayed ancient ray pits have been reported from deposits mosasaur vertebra and a juvenile hadrosaur as old as Late Cretaceous (Howard et al., 1977). metatarsal (Schwimmer et al., 1997). Putative gut 0steichthyans.-The neoselachian radiation contents from Squalicorax include mosasaur limb saw its counterpart in the Cretaceous osteichthym bones (Dnrckenmiller et d.,1993). teleost fishes fig. 5.1-5.3). The achievement of Even fewer examples are known of shark improved buoyancy via swim bladders, attach on benthic invertebrates. Molluscan shells, development of deep bodies, and anterior placement such as those of inoceramid bivalves, are known to of pectoral and pelvic fins, represent coordinated have marginal edge fragmentation, and frequently adaptations for increased swimming efficiency and are preserved as fragmented remains sometimes maneuverability during the Jurassic Period. During associated with the putative shell-crushing shark, the Late Cretaceous, the development of hinged Ptychodm (Kauffman, 1972). One inoceramid maxillae-prernaxillae and highly protrusible mouths specimen, Inomram ternis, is described as having further gave rise to a new mode of suctorial predatory shell injuries on its left valve perhaps directly feeding. These adaptations in turn fostered a major stemming from Ptycbdm predation (Kauffman, adaptive radiation of neoteleost predators in the sea 1972). The right valve is uninjured, and Kauffman and in fresh water. (1972) explains that this lack of injury is compatible In the Cretaceous, Iarge basal teleosts clearly with the Life habits of the inoceramid, as the left dominated in the intermediate- to large-sized fish valve was more exposed. Speden (197 1) interprets eating predator guild. Many specimens of the large aggregations of fragmented inocemmid shells from Xiphactinus (Fig. 5.1) from the Cretaceous of North Late Cretaceous sites in New Zealand as evidence America have been found with ingested fish in the of regwgtated or fecal material from vertebrate body cavity. These include specimens hrnKansas predators. Inoceramids are usually found in quiet, with as may as ten fish in the stomach and a 4.5- deep-water settings, either in chalks or bhck shales meter specimen with a 1.6-meter related (Kauffman, 1972jthus, any information on their ichthyodectid fish inside (Benton, 1997)! S~pecimem potential predators would illuminate the little- of the pavement-toothed Tribodus from the known paleoecoIogy of deep-water fauna in the Cretaceous Santaea Formation of Brazil had Late Cretaceous. stomach contents that included shrimp and The batoid rays and skates f~stappeared in the fragmentary molluscan shells (Maisey, 1996). Early Jurassic and diversxed during the Late Advanced acanthornorph teleosts evolved PALEONTOLOGICAL SOC1ETY PAPERS, V 8,2002

erdontifomes 73CRUNCHc=' Orectolobifomes "CRUNCX"

Carcharhinifonnes "CRUNCH" Chlamydoselachiformes Hexancbrifome Echinorbinifomes

Cen~ophorifomes

FIGURE 9-Cladogram of chondrichthyes (sharks) showing the repeated evoiution of durophagy (indicated by "CRUNCH"; after Wilga and Motta, 2000).

Merdefense in lhe Late CretaceousrTertiary, extant, and the other omnivorous and herbivorous without substantial loss of mobility, in the form of groups (Cheloniidae) arose in the Aptian and are erectile fin spines. These adaptations may indeed still extant, having reached a diversity peak during have made the swallo~vingof whole prey sufficiently the Late Cretaceous (Hirayama, 1997). The difficult that individuals possessing longer, sharper Protostegidae were restricted to the Late fin spines were frequently spared andlor avoided Cretaceous. The Chelonidae and Dermochelyidae by experienced predators, thus driving adaptive survived the mass extinction at the end of the trends in neoteleosts (Patterson, 1944j. Cretaceous, while most other marine reptiles, with Sea tzkrt2es.-Although modern turtles are all the exception of the crocodiles, went extinct. The morphologicdy similar, Mesozoic sea turtles were skull of Late Cretaceous ~roto'ste~idaetdes is far more disparate (Hiray ma, 1997 j. There were similar to that of the modern freshwater up to three separate radiations of sea Mesin the rnolluscivorous turtle (Malayemys). Based on this Mesozoic WichoUs, 1997). The PIesiocheIyidae similarity, it may have fed on pelagic ammonoids evolved in the Jurassic. The second group (Hirayama, 1997). The Protostegidae were the (Pelomedusidae) is presently resbicted to fresh largest sea turtles hown, characterized by massive water, but in the Late Cretaceous and Early Tertiary, heads, like that of the late Campmian Archebn. This members of this group were present in shallow gigantism was shm-lived, as the Protostegidae went mm%e environments @Ticholls,1997). The third extinct before the end of the Cretaceous (EIjrayama, group (Chelonioidea) first appeared in the !ate Early 1997). The skulls of the Dermochelyidae are Cretaceous and includes the Demochelyidae, imperfectly known; however, it appears that the Cheloniidea, and the Pro tosteg idae (Hirayma, narrow lower jaw and other skeletal features suggest 1997). Of these, the jelIyfish-eating stock that the jellfl&-eating mode was developed during (Dermochelyidae) msein the Santonizn and is still the Cenozoic (Hmyama, 1997). WALKER AhD BREn-POST-PALEOZOIC PmERNS IN MARLVE PREDAIOAJ

Plesiosaurs.-Plesiosaurs are thought to have been top predators of Mesozoic seas, but there has been little evidmce to support this claim sat^ and Tanabe, 19983, The oldest firm evidence of predator-prey associations between ammonoids and plesiosaurs is from a Late Cretaceous (Upper Cenornanian) outcrop from Hokkaido, Japan. From this locality, Sato and Tanabe (1998) describe gastroliths, 30 isolated and disarticulated ammonoid jaws, a shark tooth, and molluscm shells from the putative gastric contents of a polycotylid plesiosaur. While the head of the plesiosaur was missing, comparable teeth in other polycotylids suggest that they were poorly adapted to crush ammonite shells, and may have swalIowed their prey whole. Plesiosaur gastric contents from the Early Cretaceous are known to include cephalopod jaws in association with gastroliths (Sato and Tanabe, 1998). Gastric residue fiom other Late Cretaceous plesiosaurs, however, had fish vertebrae, p terodacty 1 bones, and thin- shelled ammonites [Massare, 1987, her table 1, p. 128). Mosasaurs .-Mosasaurs originated and diversfied worldwide in less than 25 miIlion years during the Late Cretaceous (Fig. lo), but met thejx FIGURE 10-Temporal distribution of several untimely demise during the end-Cretaceous genera of mosasaurs (indicated by letters and their apparent ammonite prey, based on bitelcrush event the time extinction (r;l&amSoliar, 1999). By marks from Cretaceous deposits of the Western of their origin, the ichthyosaurs had gone extinct, Interior Seaway. From Kauffrnan (1 990). and only a few plesiosaur fadies were still extant (Lingham-Soliar, 1999). Not since the Triassic placodonts, had a reptile group so dominated the (Sheldon, 1997). Thus, even deep-water Late durophagous functional lifestyle (Massare, 1 9 87). Cretaceous ammonoids that were thought to use Mosasaurs, with elongated snouts and elongated, depth as a refuge against predation (Westemann, fusifom bodies, include the largest marine reptiles 1996) may not have been immune to their attacks, ever known (e.g., Msasaurus hoffmanni at over which may have fragmented the shells completely. 17 rn in length; Lingham-Soliar, 1998a). Because Evidence of deep diving in rnosasaurs comes fiom the Late Cretaceous sea levels were the highest avascular necrosis of their bones, indicating the recorded during the Mesozoic, these giant reptiles "bends"--decompression syndome (Martin and were more Likely te be preserved than were other Rothschild, 1989; Taylor, 1994). Some mosasaurs, Mesozoic marine reptiles, and so we have a better however, had pachy ostosis (bone thickening), which understanding of their habits. required that they increase lung volume to remain Bone microstructure and bone density are used neutrally bony ant; in turn, increased lung volume to infer the ecological distribution of rnosasaurs in means a larger rib cage, and thus more drag on the the water column (Sheldon, 1997). Reduced bone anidmaking it a slow swimmer (Sheldon, 1997). density of two c.omon mosasaurs (Clidastes and Mosasaws with pachyostosis (e.g., Platecappw) Tylosaum) inQcates that they lived at great depth usually lived in shallow waters, but even these forms PALEOIVTOLOGICAL SOCIETY PAPEm, I.! 8,2002

could dive to deeper depths as suggested by of crushing ammonite shells or other putative prey, avascular necrosis in their bones. Similarly, skeletal as well as how they crushed the shells. e.lernents of modern BelguIa whales have Such biomechanical studies would be pachyostosis, and consequently these whales spend beneficial at least in coming to some ~onclusion most of their time in shallow water. However, as to how some ammonites received large holes in Belgula whales also are susceptible to the bends if their shells; and indeed, Kase et al. (1998) they dive too deeply, and this is recorded in their performed such tests using modern Nautilus shells bones (Sheldon, 1997). Most deep-diving animals and a 'bosasaur robot" modeled after a putative usually do not suffer the bends, although some mosasaur predator of ammonites (Prognathodon turtles may show skeletal evidence of having had overtoni). S eilacher (1998) also performed decompression syndrome (Motani et d.,1999). biomechanical tests using steel pliers and nautdoid - Consequently, considering evidence from shells. While it can be argued that Nautilm shells dentition, body fom, thickness of skeletal are not malagous to ammonite shells with respect elements, and avascular necrosis of the bones, to biomechanical loading (Tsujita and Westemam, rnosasaurs are interpreted to have been top ambush 2001), it is still important to experimentally test predators that once foraged in lagoonal to open- the predatory hypothesis. ocean environments. Prey were swallowed whole, Numerous specimens of the ammonite, crushed, pierced, rammed, and shredded to name Placeaticeras, from he Late Cretaceous Pierre but just a few means of prey demise Ggharn- Shale and Bearpaw Formation of the western Soliar, 1998a, 1998b, 1999). The dentition of interior of North America, show putative mosasaur rnosasaurs was so divers'e that dentition patterns tooth marks (fig. 8.1) (Kauffman and &sling, fit all five of Massare's (1987) functional predatory 1960; Kaufhnan, 1990; Hexvitt and Westemam, groups for Mesozoic marine reptiles (i-e., cut, 1990) that have been reinterpreted to be limpet pierce, smash, crunch, and crush guilds), a homing scars that were enhanced by diagenesis functional feat accomplished in just a short (Kase et d.,1998). Kase et d.'s biomechanical tests evolutionary time period (Linghamsoliar, 1999). indicated that robot bite marks on live Nautih Because of these varied feeding modes, mosasaurs typically had jagged edges that did not show the most likely fed on an array of benthic and pelagic concentric cracks characteristic of putative bite organisms (Lingham-Soliar, 1999). marks in Placenticeras. The innermost nacreous The West African PLuridens walkeri, for layer was shattered in the experiment, whereas example, had broad-based and short tooth crowns internal shell layers under the putative mosasaur that are speculated to be powerful enough to have bite marks on Placenticeras were not (Kase et a]., crushed thin-shelled invertebrate exoskeletons 1998). They also found few examples of (Lingham-Soliar, 1998b). Globidens, from the Placendiceras with holes corresponding to Upper Cretaceous of Belg-rum, had rounded and mosasaur jaw shape. Consequently, they concluded deeply wrinkled mushroomshaped teeth that are that the holes in ammonites were limpet home thought to be specialized for crushing thick-shelled scars, and not mosasaur predatory bite marks. molluscs (LinghamSoZar, 1999). In fact, Globidem, Tsujita and Westermann (2001) rejected the along with another coeval mosasaur, Carirw&ns, findings of Kase et al. (1998) and provided further shows the most rernarkabIe durophagous crushing observations jn support of the mosasaurim origin dentition since the demise of the placodonts of the holes. In fact, they pointed out that some of (Lingham-Soliar, 1999). While the biomechanical the experimental robot-induced holes that Kase et importance of such dentition was discussed af. (1998) figured (e.g., their fig. 3b, p. 948) (Lingham-Soliar, 1999), it still. remains to be resembled those of putative mosasaurian bite marks independently verified using experiments whether on Placenticeras meek (Tsujita and Westermann, the many varieties of mosasaur teeth were capable 2001, fig. 3qb, p. 251). Further, they argued that WALKER AND BRETT-POST-PALEOZOIC PATTERNS IN MRINE PREDATION the biomechanical loading was oversimplified,and withrows of sharply pointed teeth, presumably for needs to be reanalyzed using various ammonite fish capture. Fish remains have been found in models in addition to testing various loading coprolites associated with Hesperonis (Benton, functions attributed to the jaws of rnosasaurs. The 1997, p. 273). Presumably, these ma filled the lack of exact matching between jaw shape and guild presently occupied by diving sea birds, putative bite marks is explained by the fact that although the Cretaceous orders were evolutionary like most marine reptiles, mosasaur jaws were not dead ends. One mosasaur specimen also contabs perfectly occluded; the lower jaw was loose enough ingested Hesperonis skeletal elements to pivot laterally. Further, while well-preserved Limpet fossils are found in the Pierre Shale, they ,JUFL4SSI~-CRETACEOUS ha& yet to be found in the Bear Paw Formation of BENTmC PREY AmTHEm Alberta. Thus, Tsujita and Westemam conclude POSSBLE ~T~~ATORY that putative predatory holes on ammonites may be only rarely associated with hpethome scars, RESPONSES and the vast majority are from mosasaur predation. Possible Behaviot-al Responses of No one has done a quantitative cornparision of the In,,efieb rates.- mg *e and the mk the the size of bite size of limpet home a host of from sponges to worms, scars, the diameter of the preserved limpets? and barnacles, bivalves independently an related it to the range of tooth sizes found in ability to bore ktostiff mud, rock, carbonate, contemporaneous mosasaurs. hardgrounds, shell substrates, and wood (Palmer, Gastric contents from mosasaurs include 1982; seilaCher, 1985; wdsonad palmer, 1990, cephdopodhooklets, fish, belehta, tdebones, 1992). Submarine crypts caverns and birds wassare, 1987, her table 1, p. 128). For pmeded a ~fugefor certain primitive groups, such example7 gastric a single a sclmsponges, many bryozoans, sedentary tube- of the mosasaur Qiohstes ~n*ded a marine d~~ dwellrng polychaetes, and pediculate brachiopods and a diving hpewmis martinand (Palmer, 1982; Wilson and Palmer, 1990). A BJQ& 1987). At least one squid gladius from the number of sedentary invertebrate groups persisted Pierre Shale ixhibits bite marks attributable to a ,, hard during heMesozoic. mosasaur (Stewart and C~enter,1990). Doll0 But these, ia particular, show allegedly strong (19 13) reported a broken test of the echinoid, ,tipredatory skeletal adaptations (Fig. 33: they are Hem@mmtes, between the teeth of the mQsasaur skongly cemented (oysters, corals, barnacles), have C@"i~~dens;andnumerous ammonites have tooth thick, heavy shells (e.g., rudists, oysters), marks, presumably from mosasaur predation camouflage, and spineslspicules or toxins. (Kauffman and Kesling 1960; Kauffman, 1990). A major decline in free-resting benthic Some of these tooth marks, however, may also be invertebrates occurred in the Mesozoic (relative to limpet homing scars on some specimens (aeet the Paleozoic). Quasi-infaunal forms, such as al., 3998). Ta date, no ammonites are known from grypheid and ex~gyridoysters, remained common mosasaur gashc contents (Mdand Bjork, 1987; on Mesozoic soft substrates, but these organisms Massare, 1.987),and this may be due to taphonomic were partially hidden and evolved thick, robust bias against the preservation of aragonititc ammonites shells (Fig. 3). Exposed epifaunal brachiopods, in gas~ccontents (Tsujita and Westemam, 2001). cords, and crinoids were greatly reduced or absent Sea and Shore Birds-Fbally, in the Late from shallow marine soft-substrate settings during Cretaceous, two orders of marine diving birds the Mesozoic (Thayer, 1983; Vermeij, 1987). evolved: the flightless, foot-propelled Thayer (1983) arguedthat this deche in epifaunal Hesperomithifomes and the swimming-winged suspension feeding may have been fostered by the Ichthyomithiformes. Both taxa had elongate beaks rise of deeply burrowing hfamal %ulldozers ," PALEONTOLOGICAL SOCIETY PAPERS, V 8,2002 especially siphonate bivalves, during the Mesozoic. heavier shells do not necessarily reduce predation Moreover, predator grazing ~&ybe a further cause mortality in queen conch, especially when predators for the decline of eprfaunal suspension feeders attack through the aperhrre, as do crustaceans and (Stanley, 1977). predato~yrnoVuscs (Ray and Stoner, 1995). Vermeij (1977) hypothesized that shell- Bivalves.-Much of the literature concerning breaking predation became a more important cause bivalve shell ornamentation in relation to predation of mortality and was a driving force in evolution has been based on largely cixcumstantial evidence from the Mesozoic to Cenozoic. He termed this (Fig. 3) (Harper and Skelton, 1993). Spondylid postulated major escalation of predator-prey bivalves provide an interesting example. These spiny interactions the "Mesozoic Marine Revolution" epifaunal bivalves appear in the Middle Jurassic and (MMR); in fact, most of the trends he described in show increasingly spinose shells up to the present adaptive morphology continued from the Mesozoic day (Harper and Skelton, 1993). However, these to the Recent, so the present discussion combines spines apparently do not increase shell strength evidence from both eras. (Stone, 1998; Carlson, pas. corn., 20001, but do Gmtropods.-Gaskopods show a number of increase effective size and make shells more =cult trends, probably in response to shell crushing/ to attack. Spines are also commonly worn off, and drilling predation pressure (Vermeij, 1977, 1983, it is not hown how this affects the survival of these 1987). These include a Merdecrease in umbilicate cemented groups (see Logan, 1974). and loosely coiled shells. The few remaining loosely Shell microstnrctures and the development of cdedgastropods lived within sponges (sil~@arids), sphes in some groups of bivalves may have or were cemented (vermetids), the latter sometimes originated in their Paleozoic ancestors (fable 5). forming largc aggregates, or "reefs" Wermeij, 1953). Additionally, changes in thichess and arrangement Other trends among Mesozoic gas~opodsinclude of shell microstructure may also be primarily increased proportions of thick-lipped shell apertures, controlled by water chemistry and temperature, slit-like apertures, varices, and spines or hobs. The rather than by predation. Some microstructures, simultaneous increase in these features suggests a however, may secondarily function to reduce crack common evolutionary pressure, presumably the propagation, such as cross-lamellar structures, and increase in. durophagous predation (Vermeij, 1983, increase abrasion resistance (Cnrrey and Kohn, 1987). However, as this review shows, most of these 1976). However, there are many ways to build cross- durophagous predators are generalist feeders, lamellar structures (Scheider and Carter, 2001). feeding on a variety of hard-shelled prey, and not Shell mic~ostructuresuch as spines, thickness of just molluscs. More work must be done to determine particular shell layers, and types of sheU layers may whether shell ornamentation really reduces reflect a phylogemtic constraint-Thickening of shell successful predation. Shell sculpture and varices margins through extensive inductural deposits may have been shown to decrease predation by sea stars be related to photosymbiosis, and not directly to and durophagous crustaceans in laboratory predation (Scheider and Carter, 2001). experiments (Donovan et al., 1999). Fish predation Cardiid (Jurassic to Recent) bivalve shell is also deterred by shell sculpture (e-g., stout spines) microstructure exhibits several evo1utjona1-ybends on gastropods (Palmer, 1979). However, field that may not be related to predation (Table 5). Some experiments demonskate that shell sculpture may Cretaceous cardiids evolved sbonger reflection of not always be a deterrant to predation for some the shell margins, and increased thickness or gaskopods my and Stoner, 1995). For queen conch secondary loss of the ancestml prismatic outer shell (Strumbus gigas), living in aggregations and layers. However, these changes appear to be related attaining large ovedsize was found to be more to water chemism and temperature. For example, important in deterring predation than was shell microsbxctllral convergences may be directly or scul.pture (Ray and Stoner, 1995). Longer spines and indirectly tied to ocean chemistry and temperature: WALKER AVD BRETT-POST-PALEOZOIC PATTERNS IN MMNE PREDATION cool climate may lead to thicker prismatic outer she71 shell-breaking predators commody occurred, for layers in some cardiid bivalves, similar to those of break mark5 are common in Jurassic and Cretaceous venerid bivalves (Scheider and Car%r, 200 1) . ammonites," but he does not provide data to support Cardiid bivalves are known for their spines. this statement. Data are needed on the number of Cardiid spines can be formed in different ways: 1) ammonoid shells with evidence of healed injuries, by a mantle that is strongly reflected over exterior and on whether this varies by environment of shell surfaces; 2) by extensions of the normal outer, deposition, and on shell ornamentation through the or outer and middle shell layers; or 3) by the Mesozoic. Equally important would be a periostracurn (formed on the undersurface of the comparative examination of healed scars on periostracum, and cemented to the shell exterior). microconchs versus macroconchs, and on dernersal Cemented periostracal granules or spines in versus more planktonic forms of ammonoids. Carboniferous astartids, and in three subfadies A few direct records of predation on monoids of cardiids (i.e., colpomyid and mytilid have beenreported. Several examples of ammonoids pteriomorhians and trigonioid paTaeoheterodonts), with smaller amrnonoid shells in their body suggest that periostracal mineralizing is chambers are cited above. Ammonite shell fragments plesiomorphic far the bivalvia, and is merely are known from fish feces from the Sahhafen retained by many anomalodesmatans (Scheider Limestone in Gemany (Schindewolf, 1958). An and Carter, 2001). Thus, some spine forms in these unknown marine reptile apparently left twenty Mesozoic and Cenozoic groups may be partly or possible bite marks on a specimen of the M~ddle Iargely the result of phylogenetic and physical Jurassic ammonoid Kosmceras gulielmi from the environmental contraints. Middle Oxford Clay, England (Ward and Amomids m Prey.-Ammonoids are known Hohgwoah, 1990). The bite marks are surrounded to have sublethal injuries from the Mesozoic that by an inclined ring of fractured shell, and because may not have affected their bouyancy as much as there was no sign of healing, the bites are considered sublethal injuries in nautiloids (Kroger, 2002). to have been fatal to the ammonoid (Ward and Unfortunately, little quantitative data exists for Hollingworth, 1990). It is also thought, because of shell repair in amrnonoids during this time. the diversity of predatory marine reptiles, fish, and Westemam (1996) suggests that ammonoids lived beIemnites, that amrnonoids may have lived in in deeper-water areas to avoid predators, esgecialIy deeper, slightly more oxygendeficient waters at hs in the Cretaceous. However, there is now extensive time (Western- 1996). Vermeij (1987, p. 283) evidence that marine reptiles were able to dive to reviewed the limited anecdotal information deep depths during this time. concerning shell repair on ammonoids and suggested Vemeij suggests that shell repair increases in that the incidence of shell repair was low in Early ammonoids during the Mesozoic, although he makes and Middle Jurassic ammonoids. a plea that more data be accumulated in. order to Tf benthic durophagous predators were preying really assess this claim (Vermeij, 1987, p. 283-284). on ammonoids, the ammonoid prey should show a To date, little if any data exist to analyze trend in antipredatory ornamentation and shell antipredatory features and predation on Mesozoic repair through the Mesozoic in accordance with monoids. Because shallow-water and deepwater the Mesozoic Marine Revolution theory of Vermeij forms were abundant, and because amrnonoids (1977, 1987). As is the case for the Triassic, little occupied many different habitats within those is hornabout antipredatory effects of ammonoid settings during the Jurassic and Cretaceous, they she11 shape and sculpture, although shell crushing would be ideal organisms by which examine masine reptiles, fish, and other cephalopods were environmental records of predation. quite diverse in the Jurassic and Cretaceous. Ward (1986, p. 818) states that there is Costae and spines in ammonoids have been "abundant evidence.. .suggeshg that predation by considered to be antipredatory (Logan, 1974; PALEONTOLOGICAL SOCIETY PAPERS, V 8,2002

Westermann, 1996). Costae presumably moderate ribbing, and very strong ribbing) strenthened the shell against predators (Checa and appeared to be similar for both time periods. Westermann, 1989; Westermann, 19901, yet there Essentially, there appears to be no difference in are numerous examples of smooth-shelled ornamentation between the Lower md Upper ammonoid groups Living contemporaneousIy in Cretaceous ammonoids, despite the origin and sdarhabitats. Spines and spine-like antipredatory evolution of durophagous mosasaurs in the Upper features of adult arnmonoids include spines on Cretaceous. Ward did not differentiate between ancyclocone monoids, and protuberances such benthic, planktonic, and pelagic ammonoids. as lappets, rostra, and horns on microconchs Ammonoid shell shape was also examined fiom (Westemam, 1996). the Berriasian to Maasbichtian, and little change was Ward (1981) argued, based on figures in the noted for coarsely ornamented ammonoids (Ward, Treatise of Invertebrate Paleontology, that many 1986, his fig. 3, p. 9). Non-streamlined (non- of the highly sculptured Cretaceous ammonoids ornamented) forms stay roughly the sarnc though evolved primarily as a defense against shell time, with slightly more in the Beniasian. Thus, it crushing predators. Ammonites showed Limited appears that shell ornamentation in ammonoids is trends in. sheU.,ornamentationduring the Mesozoic not a direct result of predation. relative to their Paleozoic counterparts (Ward, A great deal of work remains to be done on 1981). Ward documented trends toward increased testing various ammonoid shell forms in relation fluting and ribbing in ammonite shells during the to predation. For example, it would still be Jurassic and Cretaceous. beneficial to examine benthic versus pelagic There is only a slight increase in flne-to- ammonoids to determine if there is a dGference moderate shell ornament in Middle to Late Jurassic in she11 ornamentation between these two types. ammonoids compared to Early Jurassic ammonoids A shift to more offshore amrnonoid faunas in (Ward, 1981, his fig. 2, p. 98). Additionally, there the Late Cretaceous, and the extinction of nearshore is no difference in moderately coarse to very coarse North PacXc forms prior to the Maastrichtian shell ornamentation between-Early Jurassic and (Ward, 1986), may have resulted from increased Middle to Late Jurassic arnrnonoids ward, 1981). competition andlor predation. However, there-were The proportion of ammonoids with very coarse numerous offshore, deep-diving predators in the ornamentation stays the same through the Jwassic; Late Cretaceous (e-g., globally distributed moderate to strong ornamentationdoes increase by mosasaurs, sharks, and other fish} that may have about lo%, but then changes little throughout the preyed on pelagic ammonoids and other pelagic rest of the Meso7aic. Vemeij (1987) surmised fiom invertebrate fauna. this data that armor in ammonoids was no longer a The last ammonoids of the Late Cretaceous are successful anti-predation strategy after the Turonian. best known from continental slope deposits, and Some long-lived groups of arnrnonoids, such as the include nektonic and pIanktonic forms (Ward, Lytoceratidae and Phylloceratidae, remain 1987); curiously, it is the duroghagous nautilojds morphologically similar lhrough their geologic that survived the Cretaceous-Tertiary extinetion range, while other long-lived ammonoid families event, and not the pelagic, perhaps chiefly are morphologically diverse (Ward and Signor, planktivorous ammonoids of that time. This 1983). It is not known what causes morphological extinction may not be directly related to their stasis in some forms but not in others. planktic habit, but rather to the fact that that Ward (1981) found that 40% of Lower ammonoids had a planktic part of their early life Cretaceous, and approximately 42% of Upper cycle, whereas nautiloids had a benthic stage Cretaceous ammonoids had moderate to strong (Ward, 1986). However, nautiloids are dependent ribbing on their shells (his fig. 2, p. 98); all other on other invertebrates for food, including crustacea, shell surface types (i.e., no ornamentation, fine to which have a planktonic period in their life cycle WALKER AND BRETT-POST-PIPATTERNS IN MMNEPREDATION

Coleoids.-Other nektonic organisms gave up limited information concernkg predation on armor in exchange for more efficient swimming crinoids during the Jurassic (Schneider, 1988). The during the Mesozoic and Cenozoic (Ve-meij, 1983). offshore retreat of "'primitive" groups, such as Sofi-bodied coleoid cephalopods traded off external stallred crinoids, has been suggested to be a general armor for increased speed and mobility and evasive trend that might be related to increased predation defenses, such as sepia jnk for camouflage (Packard, pressure (Jablonski et a]., 1983; Vemeij, 1987; 1972; Lehmann, 1975). The majority of gastric Bottjer and Jablonski, 1988; Sablonslri and Bottjer, contents preserved in marine reptiles, however, 1990). For example, Meyer and Macurda (1 977) consist of hooklets from belcmnites, and few horn documented an offshore migration of stalked naked cephalopods. While the betemnites did not crinoids during the Jurassic. This onshore-offshore survive the Cretaceous mass extinction, the soft- pattern in crinoid distribution needs to be re- bodied coleoids did. They faced a renewed group examined in light of new data. of predators in the Cenozoic-the marine mammals. Most isocrinids (except for Pentacrinitidae) Despite two revolutions in their predators, their Iived in shaLIow waters until the Mid-Cretaceous, morphology has remained remarkabIy static. whereas in the Cenozoic these forms inhabited Decapods and 0stracodes.-Little is known deeper water Wottjer and Jablonski, 1988). about predation on decapods during the Cretaceous. Tn the Early Jurassic, the biggest evolutionary Evidence of drilLing predation jn Cretaceous to innovation in echinoderms appeared with the advent Recent ostracode assemblages from Texas includes of irregular echinoids (Simms, 1990). The flattened drillholes from juvenile naticid gastropods tests of these creatures are thought to Rave been an (Maddocks, 1988). It is not known if ostmodes aptation that provided greater stabihty within the developed antipredatory armor, although the subsbate. At the same time, the periproct moved incidence of drilling appears to increase from the away from the apex of the test, in accord with their ~retaceou; into the Tertiary, but then declines in sediment-eating habits. By the Middle Jurassic, the Holocene (Maddocks, 1988). It is thought that endobenthic irregular echinoids had evolved and smooth shells may be preferentially drilled, or at rapidly diversified. Today, their descendants least may make drillhales more discemable to the comprise nearly haK of all extant echinoids (Slmms, paleontologist Naddocks, 1988).'Because of their 3990). The abord spines and the anal sulcus of these abundance and worldwide distribution in a variety creatures (e-g.,Galexopygidae) were consistent with of environments, ostracodes would provide an their endobenthic lifestyle (Simms, 1990). The important database from which to test the various evolution of pencillate tube feet in these groups marine revolutions; but they remain little studied. allowed them to pick up finer sedimentary particles Echinodem.-Regional large deposits of via mucous adhesion (Simms, 1990). A peri-oral crinoid grainstones and packstones (encrinites) are tube foot also allowed them to expand into new not present after the Jurassic {Ausich, 1997). The tropbic realms. Was this endobenthic lifestyIe presence of regional en&tes since the Ordovician provoked by predation, or merely by the opportunity illustrated the domination of many shallow-shelf for better feeding conditions? It should be noted that environments by crinoids and other staked epibenthic echinoids were also diversifying at this echinoderms perhaps for millions of years up to time, with the Cassiduloids and their offshoots. the Jurassic (Ausich, 1997). Cornamlid crinoids Fish are the dominant predators of modern evolved rapidly from the stalked forms dwing the ophiuroids (konson, 1988). Little is hornabout Late Triassic-Early Jurassic (Meyer and Macurda, predation on Mesozoic ophiuroids, although the 1977),but their diversity remained fairly low (five rate of arm regeneration appears to be low for species) during the Jurassic. Modern comahllids Jutassic cornpared to Recent ophiuroids (Aronson, are hoynto be preyed upon by reef fishes (Meyer 1987, 199 3). However, there is no clear evidence and Ausich, 1983; Meyer, 1985). There is also that ophiuroids developed &predatory armor, as thek morphology has remained relatively the same CENOZOIC PREDATORS since their origin in the Ordovician (Aronson, 1991). It is possible that they developed better The Cretaceous-Tertiary extinction had a autotomization of their arrns, like some crinoids; devasting impact on pelagic ecosystems. Ammonites or maybe they developed into distasteful prey and belemoids, as well as Iarge vertebrate pdators, (Aronson, 199 1). The evolutionary radiation of the were decimated by this event. AIl of the marine spatangoids (and holasteroids) in the Cretaceous reptilian predator guilds, except sea snakes and sea may have been the result of the Jurassic innovation turtles, became exhnct during this crisis-including of pencillate tube feet, a feature shared with no mosasaurs, plesiosaurs, and ichthyosaurs, in addition other echinoid group (Simms, 1990). Early to the flying pterosaurs. This left only the highly Cretaceous spatangoids (Hemiaster elegans successful neoselachian sharks and teleost fishes in washitae) are reported to have drillholes from the vertebrate predator realm. Marine mammals parasitic gastropods (Kier, 198f ). Parasitic drillings emerged in the Eocene to essentially take over the are commonly associated with deformation of the ecological void left by Mesozoic marine reptiles echoid ossicles where the parasite housed itself. (Table 4). In fact, tooth dentition in marine mammals Fish bite marks are well preserved on complete closely parallels that of the Mesozoic marine reptiles asterioids and asteroid ossicles from the Late (Massare, 1987, 1997). Cretaceous White Chalk of northwestern Germany Conversely, many benthic invertebrate (Neumann, 2000). On some specimens, serrated predators, mcH as naticid and muricid gastropods tooth marks may be related to galeoid shark and various decapod crustaceans, were seemingly predation. Regurgitate pellets are also common in little affected by the terminal Cretaceous extinctions. the White Challr, and indicate that predation by Several groups of shell-drilling predators evolved these durophagous fish may have been size or diverszed within the Cenozoic (for review, see selective (Neumann, 2000). Vermeij, 1987); some groups, such as the Vertebrates.-As with late Paleozoic fish, neogastropods, evolved in the Late Cretaceous. armor does not appear to have been a significant Prosobranch gastropod predators, the dominant part of the response to escalation among Mesozoic drilIers, were much more common in the Cenozoic marine vertebrates. W~ththe exception of relatively than at any other time, though the Mesozoic record slow-moving placodonts and marine turtles, none needs to be more throughly examined (Kowalewski of the marine vertebrates developed my unusual et al., 1998). The record of octopod shell drilling is armor during the Mesozoic. Indeed, within chiefly Cenozoic, with the soft-bodied octopod actinopterygian fish there is a distinct trend toward fossil record primarily within the Cretaceous to reduction of ganoid scales in favor of lighter and Paleogme engeser 1988; Harper, 2002). less protective cycloid and ctenoid types (Patterson, Several major groups of vertebrate shell- 1994; Benton, 1997j. Presumably, this scale crushing and shucking predators that may have reduction reflects the ineffectiveness of dermal seriously impacted benthic and pelagic marine armor against large predators, which demonstrably biotas evolved or diversified during the Cenozoic: swallowed prey whole (Voitil, 1990). This further shell-crushing sea turtles (i.e., the single genus reduction in armament is clearly coordmated with Caretta), the coral reef telcost fishes and other the development of improved swimming speed, teleosts, rays and skates, dwing marine and shore buoyancy control, and maneuverabiiity in the birds, pinnipeds, sea otters, gray whales, and Cretaceous teleosts. Jnhun, this increased mobility humans. Among mammals, the origination of may well have triggered adaptations for impraved pinnipeds (seals and walruses), the cetaceans speed, maneuverability, andlor stealth among larger (especially the gray whale) in the Eocene, and sea predators, such as the neoselachian sharks, otters (Carnivora; Famrly Mustelidae) in the Iate plesiosaurs, ichthyosaurs, and mosasaurs. Miocene also potentially impacted Cenozoic WmRAND BREirT-POST-PALEOZOIC PATTERNS IN ibMRlNE PREDATION

TABLE 4--Cenozoic marine vertebrates and their functional feeding groups.

Taxonomic Functional Prey Forensic Evidence Group Feeding (potentially traceable in the Group fossil record)

Mustelidae Generalist Abalone(Ha1iotis);sea Shell damage consisted of: (Sea otters) carnivores urchins (Strongylocenfrotus fractured middle sections of franciscanus, S. purpura fus); shells as a result of being hit with kelp crabs (Pugetfia ); rock stones by sea otter; larger shells crabs (Cancer); turban snails may have fractured middle (Tegula ); octopus (Octopus ); sections; edge damage may be bivalves (Tivela, Saxidomus, due to otter gnawing on the Tressus ); sea stars ( Pisaster ) edges of the shell or chipping urith a stone Cetaceans Generalists Krill, whales, dolphin, squid, Gray whales suck pits in the callianassids, small bivalves; benthos to gather food; pits may no specialists be ephemeral, but may be preserved; no other information available Pinnipeds Generalist Two genera are speciaiists on Walruses may leave marks on molluscs and crustacea shells, but no record of this as yet Sea Turties Generalist One genus (Caretta) specialist Care tta may leave marks on on molluscs molluscan prey, but no record of this as yet Sea Snakes Generalists Crush prey, but no record of their predatory prowess as yet Diving Marine Birds Generalists Eat crustacea, molluscs, fish; Extensive literature on birds and one genus appears to how they forensically alter prey specialize on molluscs Marine Crocodiles Generalist Birds, Rsh, turtles, humans, No forensic information available golf bails, etc. benthic invertebrate prey. Vermeij (1987) reviewed today: the squilIoids and the gonodactyloids,which the molluscivorous habits of some of these groups, have very different means of feeding. The squilloids and here we discuss their more generalist feeding either attack prey with their dactylar spines, or grasp behavior, add or update several other groups, and prey between the toothed margins of their propodus suggest possible alternative scenarios to his and dactylus (Kunze, 1983). Squilloids typically escalatory hypothesis. prey on hh, polychaetes, and very smallcmstacems Stomtopods.-Stomatopod crustaceans are (Schram, 1986).h gonodactyloids, the propodus is obligate carnivores and vicious predators. swung fiom an anterioventrd position, and prey is Stomatopods that crush the shells of heir prey by "smashed" on the lower part of the dactylus (Kunze, pounding them with blunt expanded segments of 1983). Gonodactyloids feed typically on hard- their maxillipeds (e.g., Burrows, 1969) did not shelled prey like molluscs and Iarge crustaceans evolve until the Cenozoic (Hof and Briggs, 1997; (Schram, 1986).Both types, however, can also scoop Hof, 1948). Two major groups of stomatopods exist up prey from the benthos with their maxillipeds. PALEOWOLO GICM SOCIETY PAPERS, V 8,2002

The gonodactyloids can smash small to large puffer fish or non-molluscan invertebrates holes in molluscan prey (Hof, 1998; Ahyong and (Table I). Clearly more work needs to be done on Harling, 2000). Stomatopods can also shear the parthenopid crabs. Durophag ous cancrid crabs gastropod shells in half and break the outer lip (Cancer spp.) eat a diversity of prey, such as (Geaq et al., 1991). Additionally, small puncture polychaetes, squid, crustaceans, fish, and wounds in molluscan prey, cded ballistic traces echinoderms, following the dominant macro- (e.g., the trace fossil Belichnm), are atbibuted to invertebrates in the habitat; whereas Ovalipid crabs stomatopods (Pether, 1995). Most of the extant may predominantly eat molluscs (Stem 1993). groups have an actual fossil record extending back Lobsters also crush shells, but usually only only to the Eocene, with the shell-smashing fmgments are left (Cox et al., 1997). In modern seas, gonodactylid group originating in the Miocene rock lobsters are known to prey extensively on (Hof and Briggs, 1997; Hof, 1998). Approximately moHuscs, such as abalones and turban snails in some 400 living stomatopod species are recognized localities (Van Zy3 et al., 1998; Branch 2000), and (Manning, 1995). echinoderms in others (Mayfield et al., 200 1). Very Despite their long history, only afew examples little is known about lobster foraging and how it would of gonodactylid shell-breaking predation are affect the fossil record of invertebrate hd-shelled known from the Neogene fossil record. Geary et prey (reviewed in part by JYalker et al., 2002). al. (1991) described a few cases of putative Gastropods.-Gastropod predators that chip stomatopod shell damage from Pliocene localities and wedge open molluscan prey (e.g., Buccinidae, in Florida. Baluk and Radwanski (1996) also FascioIaridae, and Melongenidac) originated in the documented stomatopod damage on diverse Late Cretaceous, but diversified in the Cenozoic; gastropods from Miocene localities in Europe. however, the shell-chipping record in prey shells Stomatopod predatory damage should be easily is known. only from the Pliocene (Vermeij, 1987). recognized, and documentation of this damage in Buccinid gastropods chip their outer lips in the more assemblages would enhance the process of preying on other mollusks, and then paleoecologicd picture of these creatures. subsequently .yep& their self-inflicted breakage Brachytkran crabs and lobsters.-The second (Carder, 1951; Nielsen, 1975).Diet1 and Alexander major wave of crustacean adaptive radiation (1998) noted that this type of lip damage occurs in occurred in the Paleogene. Brachyuran crabs had buccinids as old as Mocene. Older buccinids, which appeared in the Mesozoic, but new families of crabs range back to the late Cretaceous, have not yielded such as the Portunidae, Cancridae, Grapsidae, and evidence of this distinctive lip damage. Hence, the Ocypodidae arose in the Eocene (Table 1). Most shell-prying habit of buccjnids may have evolved brachyuran crabs are generalist and opportunistic within-the Neogene. feeders, and few are durophagous (Table I). The best evidence for predation in the Cenozoic Brachyurans with heavily toothed claws apparently fossil record comes from predatory driIlholes evolved in the Paleocene (Vermeij, 1983), but this preserved in prey ranging from protists, such as has not been studied in detail. Crushing daws are foraminifera, to many phyla of invertebrates, such a foxmidable too1 far peeling and crushing as bryozoans, molluscs, brachiopods, and molluscan shells and also for crushing other echinoderms (Can-iker and Yochelson, 1968; Sohl, crustacea or hard-shelled prey. The distinctive 1969; Taylor, 1970; Sliter, 1971; Bishop, 1975; peeling of calappid crabs has been documented in Boucot, 1981,1990; Bromley, 1981; Vermeij, 1987; fossil and recent shells (Bishop, 1975; Vermeij, Kabat, 1990; Kowdewski, 1993; Kowalewski and 1982, 1987). The parthenopid crabs, which Flessa, 1997). Prosobranch gastmpods are the originated in the Late Cretaceous, are known to primary shell drillers in marine environments, eat molluscs only in the laboratory (Vermeij, 1978), although nudibrauchs (Vayssiereidaej, flatworms, and the few reports available show them eating nematodes, and the protist forademhave also WALKERAND BRETT-POST-PALEOZOIC PATTERNS INkMRINE PREDATION evolved drilling apparatuses (Woelke, 1957; SGter, Cassids use sulfuric acid from their proboscis gland 1971 ; Carriker, 1981 ; HalIock and Talge, 1994). and the radda to cut out (rather than drill) an Eight families or superfadies of molluscs irregular hole in echinoderm tests (Kabat, 1990); have evolved drilling behavjor. Naticids are the best however, most workers use the term "drillhole" for studied, but the other groups of shell drillers were their predatory traces. Although the earliest Wed reviewed by Kabat (1990). Gastropods, in echinoderm dates back to the Early Cambrian, most particular, have evolved a variety of means to prey drilling predation on echinoids is hewn only from upon hard-shelled prey, and one major family the Cretaceous and Cenozoic (Sohl, 1969; Bw et (Cassidae) and two little known groups al., 1972; Nebelsick and Kowalewsh, 1999). The (Marginellidae and Nassasiidae) of shell drillers earliest drilholes attributed to cassids were originated in the Cenozoic; the naticids and described hmthe Early Cretaceous (Albian) of muricids persisted from the Mesozoic into the Texas, but the cassid drilling record is much more Cenozoic (Fig. 7). In drilling gastropods (e-g., extensive in the Cenozoic, especially from the muricids, naticids), the small rasping organ (the Eocene to present (Hughes and Hughes, 1981; radula) and the accessory boring organ are used to McClintock and Marion, 1993; Nebelsick and drill holes in prey shells (Carriker, 1969).Predatory Kowalewski, 1999). cephalopods, such as the octopus, also use the Muricids (Neogaskopoda) diversified greatly radula to bore into mollusc shells. in the Paleogene, occurring primarily in tropical Capulids (Capulidae, Mesogastropoda) are to subtropical waters, although they are found in specialized ectoparasites on molluscs and temperate and cooler regions as well (Vokes, 1971, echinoderms (Kabat, 1990). Capulids drill their prey 1990; Vemeij, 1996; Vermeij and Carlson, 2000). to extract nutrients fiom the host's feeding currents. During times of reduced food, muricids may drill They drill sharp-sided cylindrical holes and leave conspecifics (Spanier, 1986,1987). An increase in an attachment scar on their host's shell (Matsukama, such cannibalistic boring in muricids has been 1978; &bat, 1990). Capulid-host associations date associated with sea Ievel changes in the Red Sea back to the Late Cretaceous, where capulids are (Spanier, 1987). Rarely, some muricids drill their known to associate with inoceramid bivalves own opercula or bore into dead empty shells (Hayami and &e, 1980). Drilled inoceramids, (Prezant, 1983). While increasing drillhole size can however, are not reported for this assemblage. be correlated with increasing size of muricid Capulid attachment scars and she11 morphology that predator for some species, this does not hold for codom to their host are rePo& in modem and others (Urrutia and Navarro, 200 1).Muricids may middle Pleistocene assemblages (On, 1962; Chant- also change their ckilling behavior and preferred Mackie and Chapman-Smith, 1971). Such &Zing location.on the prey with ontogeny (Umtia associations should be highly reliable, and could and Navarro, 20013. Drillholes in inarticulate potentially be found in the fossil record (Boucot, brachiopods are rare but reported in Recent 1990).Actual evidence of capulid drilling, however, communities, and may be due to muricid predation is known only from one report from the late (Paine, 1963; Kowalewski and Flessa, 1997). Pleistcxene of Japan (Matsha, 1478). Thus, very Similar drillholes in fossil inarticulate brachiopods little is known about this intriguing parasitic drilling reported horn the Tertiary of Seymour Island, behavior in the fossil record. Antarctica, and the eastern United States (Cooper, CaTsid (Tonnoidea, Mesogastropoda) predatory 1988; Wiedman et al., 1988; Bitner, 1996) may be holes (not true moles,but rather rasped areas) in attributed to a muricid predator. echhodems date back to the Early Cretaceous, but The record of naticid predatory drillholes has have been Little studied despite their ubiquity in been. used extensively to examine the evolution of Cenozoic and modern echinoids (Hughes and predatory behavior, escalation hypotheses, and Hughes, 1981; Nebelsick and Kowalewski, 1999). cost-benefit analyses in modern and fossil PALEOATTOLOGICAL SOCIETY PAPERS, V 8,2002 assemblages (Taylor et al., 1980; Kitchell et al., toxins to relax the prey, rather than feedjng through 1981; Ktchell, 1986; Kelley, 1988; Kabat, 1990; the hole. Under SEh4, the calcareous microsmcture Anderson et al., 1991; Kelley and Hansen, 2001; is seen to be greatly etched, suggesting a dominant see below). The naticid subfamily Polinicinae solutional mechanism for chilling. diversified greatly in the Cenozoic, and the Cephalopods.lSheU-mshing md crustacean- polinicid body fossil and predatory trace fossil crushing nautiloids diversified after the record is extensive, especialIp after the Oligocene Cretaceous-Tertiary extinction, and remained (Sohl, 1969; Taylor et al., 1983). relatively abundant into the Miocene, when The Nassariids (Nmgasbopods) are carnivores nautiloids were quite diverse and abundant in or scavengers, and until recently, thek predatory continental shelf habitats across the globe (Ward, drilling habits were in question (Kabat, 1990; 1987). The earliest Nautilus is known from the Kowalewski, 1993). Kabat (1990), in fact, Eocene-early Oligocene, but no fossils are known suggested that driVing did not occur in this group, from the upper Oligocene to Pleistocene (Teichert although the possibility of nasariid drilling was and Matsumota, 1987). In modern seas, nautiloids mentioned by Fischer (1963). Recently, Morton extend from Fiji in the east to the Indian Ocean in and Chan (1997) have shown. unequivocally that the west, and from New Caledonia to Japan (Ward, some nassarids can drill prey. A few (8 of 30 1987). Natiloids forage for prey or Mirstaceanmolts individuals) laboratory-reared juveniles of across great depth ranges. There may be up to seven NassaPius festivus were found with stereotypically- species of nautiloids in modern oceans, but several sited boreholes on the ventral surface of their main of the species designations are debated (Saunders body whorl (Morton and Chan, 1997, their fig. 1). and Ward, 1987; Ward, 1987). The boreholes varied in morphology, from The prey of Nutilm is seemingly quite different elongate, irregular drillholes to spheiical from that of the Mesozoic ammonoids. countersunk borings that were clearly rasped with Unfortunately, the feeding ecology of Nadlus is the radda and aided by chemical dissolution. It is poorly known, but it is thought to be both a predator thought that drilling may be a juvenile behavior and a scavenger (Ward, 1987). While direct that is lost in the adults, as no adult ;assariids have observations of predation are Iacking, evidence ~orn unequivocally been found to drilI prey. crop dissections suggests that nautifoids cat To date, two species (Amtroginella johnsoni crustaceans, especially crabs (Saunders and Ward, and Aakstroginekla muscaria) of marginellid 1987; Ward, 1987; Nixon, 1988). The crop of gastropods from southeasternAustralia are known N~xutilmmromphalus, for example, has often been to ddl into molluscan prey (Ponder and Taylor, found to contain many hermit crabs of one species 1992). Parabolic in sectional shape and circular to (Ward, 1987j. However, this dietary findjng may subcircular in outline, the studied drillholes range be biased in that nearly all Nautilzm studied are in length from 1.13 mm to 3.1 mm. AdditionaIly, caught in traps, which also attract crustaceans. marginellid drillholes are countersunk with a very Additionally, mutiloids have been directly observed smallinner diameter relative to the outer diameter. by divers to eat molts from lobsters and slipper This heropening may have an irregular shape lobsters. With their large, chitinous jaws tipped with that can be used to distinguish these borings from calcium carbonate, nautiloids shred their prey or those of other predatory gastropods such as scwenged items into very ~rnall-~iecesof about 5 naticids. Naticids make larger drillholes (see mm3 mixon, 1988). Predators of Nautilus include Kowalewski, 1993). However, marginellid sharks, triggerfish, hums, octopods, and prhaps ~llholesare similar to octopus drillings (Ponder other nautiloids (Ward, 1987). . and Taylor, 19921, and thus may be dEcult to Ammonites and nearly all belemnoids became distinguish in the fossil record. Like octopods, extinct during the Cretaceous-Tertiary crisis. marginelids may only use the ddhole for injecting However, other coleoid cephalopods, such as the WMKERMD BRETi-POST-PALEOZOIC PATTERNS IN MARINE PREDATION cuttlefish Sepia, the squids (hligo), and Nautiltas, predatory group. Using the jaws of their Aristotle's are common in the Cenozoic (Nixon, 1988). The lanterns, echinoids are able to graze corals and even Sepiida, in particular, diversfid greatly in the nibble on the tests of distantly related clypeasteroid Cenozoic. Benthic sepiids and cuttlefish mainly feed "sand dollars" Wier, 1977). An unusual modem on small crustacea, such as prawns (Nixon, 1988). predatory interaction between deep-water cidaroids Drillholes in molluscan prey from Recent and crinoids was documented by Bauder et al. octopods are well known (Fugita, 1916; Pilson and (1999): the cidaroid, Calcocidaris micm, devours Taylor, 1961; Wodinsky, 1969; Nixon, 1980; the stalked isocrinoid, Endoxocrinss parrae. Bromley, 1981; Kowalewski, 1993). Octopuses use Another cidaroid, Histocidaris nuttiagi, also secretions and abrasion from an accessory salivary contained crinoids in. its gut (BaumiUer et d.,1 999). papilla in the Mingprocess (Nixon, 1979,1980). Choadrichthyes.-Most sharks are Their drillholes are distinctly irregular or oval, with opportunistic predators, with Limited exceptions a very small inner borehole diameter (Kabat 1990). such as the planktivorous whale sharks (Cortks, Despite their ubiquity in modem habitats, their 1999). Intriguingly, both sharks and bony fishes ability to select particular prey, and their shell- have evolved similar suites of prey capture collecting habits, few of their borulgs (trace fossils strategies, including suction, grasping, biting, of Oichnm spp.) have been reported in the fossil gouging, and fdter feeding (Motta et al., 2002). record (Brornley, 1993; Harper, 2002). Robba and Inertial suction feeding is thought to be ancestral Ostinelli (1975) first reported octopod drillings in bony fishes, while the ancestxal condition of from the Pliocene of Italy. Bromley (1993) reports sharks most likely involved grasping the prey and octopus drillings from the Pliocene of Greece. dismembering it with Little upper jaw protrusion Walker (199 1,2001) reports octopod drillings f5om (Lauder, 1985; Motta et al., 2002). Some sharks, the late Pleistocene of the GaTApagos TsIands, and especially durophagous forms, use an inertial for the late Pliocene of Ecuador. Harper (2002) suction prey capture method similar to the bony rccords octopus drillings from the Plio-Pleistocene fish. Suction feeding has arisen many times within of Florida. Octopods may also drill nautiloids, the shark group, chiefly in relation feedhg on many shells of which have multiple drillholes be&c prey (Motta et al., 2002). Specializations (Saunders et- al., 1991). Octopods also make for suction feeding include rapid jaw opening, a "kitchen middens" of their favored prey type which round terminaI mouth, reduced dentition, and the can be found outside their den; the shells are ability to produce large suction pressures (Motta commonly drilled (Walker, 1990). et al., 2002). Whale sharks (Rhincodoa) possess Echinodemzs.-Evidence for the rise of asteroid these features, but are planktivorous. Thus, predation in the Cenozoic is reviewed by Vermeij durophagy may be an exaptation from a primady (1987). Gastropods were found within the oral disc adaptive form of suction feeding in sharks. of the sea star, Ctenophoraster in Eocene- Few shark groups are known to have evol~ed Oligocene deposits from Antarctica (Blake and durophasous members, and thus durophagy is Zinsmeister, 3979). This type of in siht predation considered a rare form of feeding. Seven species has a long fossil history dating back to the of chimaerids (Holocephdi), one species of horn Paleozoic, but is rareIy reported fmm Cenozoic shark weterodontidaej, one species of nurse shark localities. It is important to note that many (Orectolobifomes), two species of the classically Cenozoic predators that ingest their prey whole, predatory Carcharhiformes, and seven species of such as sea stars, don? leave an imprint on their rays (Rhobatoidea, Rajoidea, My liobatioidea) are prey (see Vermeij, 1987). These predators are still known to be durophagous. Stout, flattened teeth abundant in the Cenozoic, and some have evolved and robust jaws are the hallmarks of durophagy. to prey on reef corals {such as Acanthmterplanci~. Durophagy in sharks, however, does not Regular echinoids have emerged as a major necessarily mean that they eat molluscs; many PALEONTOLOGICAL SOCIETY PAPERS, V 8,2002

Figure 11 WALKER AhD BRETl-POST-PALEOZOIC PATTERNS LNihNRliVE PREDATION durophagous sharks eat crustaceans and/or fish with teeth up to 17 cm long. This giant shark may (Wilga and Motta, 2000). For example, the have been specialized for fcedrng upon whales chimaenids have pavement tooth structures like (Gottfried et al., 1996). Shark teeth were found the myliobatids, and feed primarily on molluscs embedded in a whale jaw preserved in Pliocene and crabs piGaincomo and Perier, 1996). Horn sediments, suggesting a potential shark attack on shahfeed primarily on limpets, bivalves, and blue the whale (J3ernere and Cerutti, 1982). crabs (Smith, 1942; Wilga and Motta, 2000). The The radiation of deep-water Neoselachian- nurse shark, Ginglyrnostom cirrat~nn,eats fish and Squaliformes sharks, to which most of the modern crustaceans (Motta et al., 2002). The bonnethead forms belong (e-g., Somniosinae, Cenbophorinae, (Carcharhiniformes) shark, Sphyma tibuvo, has most Etmopterinae, Oxynotinae), began in deep molarifom teeth and modified jaw structures that waters with demersd forms originating after the function in crushing hard-shelled crustaceans, but Cenomanim-Turonian anoxic event; the second it can also eat fish (Wilga and Motta, 2000). radiation of Squalifomes sharks (most of the Another member of Carcharhiniformes, the Dalatiidae) began in the Early Tertiary after the smoothhound or dogfish, Mustelus, has cusped Cretaceous mass extinction, and these epipelagk teeth and also feeds primarily on crustaceans sharks radiated into shallow waters (Fig. 9; see dso (Russo, 1975; Yamaguchi and Taniuchi, 2000). Adnet and Cappetta, 2001, their fig. 4, p. 241). The Some skates (Rajoidea), guitdsh (Rhinoboidei), dentition of most of these groups is quite varied, and says (Myliobatoidea) have crushing and but most are heterodont. Most squalifomes dme grinding dentition for crunching crustacean or on fish and cephalopods (Cort6,1999). molluscan prey but they canalso feed on polychates All major living families of durophagous rays (Gregory et al., 1979; Wilga and Motta, 2000). were established by the middle Eocene (Vermeij, Neoselachan sharks became the top predators 1987 after Maisey, 1982), but their effect on the in the Cenozoic seas. During the Paleogenc Pcriod resultant fossil record of molluscs and other prey the successful galeomorphs (Galea) radiated into is not known. It is clear that despite their pavement several clades, including the dogfish and gray type dentition, rays eat a wide variety of food that sharks that feed on crustacear~sand molIuscs, the is not necessarily hard-shelled prey. Some prey basking and whale sharks that strain krill from sea items ingested, however, may be incidental to their water, and the white sharks that eat fish, seals, and foraging for larger prey items. Modern bat rays, cetaceans (Benton, 1997j. Members of the latter such as My1iobdis califomicq feed on bivalves, group attained large size, culminating with crustaceans, and polychaetes; bivalves are the Carcharodun megalodon (Fig. 11.2-11.4) in the dominant food item for most size cIasses, except Miocene and Pliocene, a lS20-meter-long shark for the adults (Gray et al., 1997). Prey items within

FlGURE I l4enozoic marine vertebrate predators. 1,Teleost paracanthopterygian fish, Mmnichythes. 2, Silhouettes of modern great white shark Carcharodon carcharias and Neogene C. megalodon. 3, Outline and skeleton of C. megalodon. 4, Carcharodon megalodon; reconstructed jaws of C.megalodon (perhaps overestimated). 5, Batoid sting ray Raja. 6, Reconstruction of flightless marine, wing-propelled swimming birds drawn at same scale in swimming posture: lower figure is modern Emperor penguin; upper shows reconstruction of extinct pelecaniforrne plotopterid. 7, 8, Early whale with limbs, Ambulocetus in two postures; Middle Eocene. 9, Reconstruction of oldest known whale Pakicetus (Early Eocene). 10, Early large whale Basilosaurus; note tiny head with distinctive, rnufi-cusped teeth; Late Eocene. 11, Early (Oligocene-Miocene) pinniped Enalioarcfos. 12, Desmatophocid seal Allodesmus, Miocene. Figures adapted from Benfon (19973. PALEONTOLOGICAL SOCIETY PAPERS, V 8,2002

Myliobatis stomach contents vary ontogenetically sediment away and to enlarge the burrow (Gray et and with the sex of the ray (Gray et al., 1997). In al., 1997). Similar shallow pits are present in the study by Gray et d. (1997), juvenile bat rays Pleistocene Iocalities, and are indicative of ray fed on small clams & 5 mm clam siphon diameter), feeding activities (Fig. 12) (Howard et al., 1977). benthic shrimp, and polychaetes . Adult rays These pits could be correlated with associated predominately fed on polychaetes, large clams fragmented mollusc deposits, but, to date, this has (> 5 mm clam siphon diameter), and Cancer crabs. not been exarnined The largest rays preferred large clams and Cancer Pods of deep-water gast~opodsattributed to crabs. Large clams and crabs were eaten by adult either fecal masses or regurgitated remains from female rays, whereas subadult and adult male rays shell-eating sharks or other predators, were fed primariiy on polychaetes and burrowing shrimp. described from bathyal Pliocene deposits from In other studies, large females also predominantly Ecuador (Hasson and Fischer, 1986, p. 35). fed on echiuran worms (Karl and Obrebski, 1976). However, a recent analysis of these shell "nests" Rays and skates excavate shallow pits in soft revealed that they are not related to predation sediments in search of prey (Fig. 12) (Gregory et (Walker, 200 1). al., 1979). The cownose ray (Rhinoptem bonasus) 0steicthyes.-The diversscation of teleosts and possibly other batoids repeatedly dale (Fig. 11.1) in the Cenozoic is unprecedented among sediments and water through the mouth and vent vertebrates: presently some 23,670 species are it out the gill slits; the pectoral fins act to move the assigned to 38 orders and 425 families (Patterson, 1994). This is largely the result of development of two clades during the Cenozoic, the Os&ophysi in fresh water and the very successful Acanthomorpha (over 21,000 extant species) in all environments [Maisey, 1996). Teleosts, ranging from tarpons to tunas, became the most common piscivorous open-water predators during the Cenozoic. Certain fast-swimming large predatory teleosts, such as swordfish, seemingly filled a part of the fast-swimming piscivorous predator guild held by ichthyosaurs and primitive teleosts (e.g. Xiphac~nm)during much of the Mesozoic. Ray-finned teleosts with molluscivorous habits originated and diversxied in the Eocene, a few otlier groups in the Oligocene and Miocene (Vermeij, 1987). Additionally, a mjor evolutionary radiation occurred in the tmpical reef fish fauna of the Eocene. Most of the fossil. record of reef fish comes from Late Cretaceous to Miocene Tethyan reef deposits of southern Europe (Rosen, 1988; Cboate and Bellwood, 1991). The best reef fish fossils, however, are from the Eocene of Monte Bolca, Italy (Blottc, Fl G U RE 12-Benthic feeding by batoid rays. 1980; Choate and Bellwood, 1991; Bellwood, 1996). upper figure, diagram of eagle ray WiobatisjeHing mae fossils are exceUently preserved-some retain water through gill slits to excavate circular feeding pigmentatiOR--and mass mo~gv depression. Lower figure, drawing of ray in feeding probably related to poisonous algal blooms (Choate position on excavated pit. From Gregory et al. (1 979) and Howard et al. (1 977). and Bellwood, 199i). WALKERAND BhTYT-POST-PALEOZOIC PATTERNS IN h4AHNE PREDATION

FIGURE I +Recent shells dredged from Ria de Arosa, Galicia, Spain, showing healed (2,23,24) and unhealed fracturing attributable to crustaceans. 1,2, Fragmented bivalve Ghlamys. 3-14, Fragments of bivalve Venus. 15-23, Gastropod Nassarius. 24-28, Turriteita. From Cad& (1 968).

This extraordinary group of reef teleosts of the dumphagous Sparidae, which evolved in the evolved rapidly, coinciding with the evoIution of Miocene (moate and Bellwood, 1991). Thus, with the coral taxa that dominate reefs today (Rosen, the evolution of the scleractinian coral species in 1988; Choate and Bellwood, 1991). Within a 20- the Eocene (Acropom, Porites, and Pocillopora), million-year period, most teleost families that occur the reef fishes evolved as well. Since that time, in modem reefs had appeared, with the exception reef fish morphology has remained relatively stable FIGURE I ?-Recent shells dredged from Ria de Arosa, Galicia, Spain, showing healed (2,23,24) and unhealed fracturing attributable to crustaceans. 1,2, Fragmented bivalve Chlamys. 3-14, Fragments of bivalve Venus. 15-23, Gastropod Nassarius. 24-28, Turriteila. From Cadde (1958).

This extraordinary group of reef teleosts of the durophagous Spxidae, which evolved in the evolved rapidly, coinciding with the evolution of Mocene (Choate and Bellwood, 1991). Thus, with the coral taxa that dominate reefs today (Rosen, the evolution of the scleractinim coral species in 1988; Choate and Bellwood, 1991). WI~a 20- the Eocene (Acropora, Porites, and PocilZopom), million-year peri od, most teleost families that occur the reef fishes ev~lvedas well. Since that time, in modem reefs had appeared, with the exception reef fish morphology has remained reiatively stable PALEONTOLOGICAL SOCIETY PAPERS, V 8,2002

though the duration of the Cenozoic. thought that most sea snakes are piscivorous, with a Today, predatory and grazing reef fish in the few species that are "generalists"-that is, that feed hdo-Pacific alone comprise over 4000 species of on both fish and invertebrates such as crustaceans fish, representing I8 % of all living fishes (Choate and molluscs (McCosker, 1975; Glodek and Voris, and Bellwood, 1991). The triggerfish 1982; Voris and Voris, 1983; Heatwole, 1987). (Tetraodontidae, Balistidae) are known to crush Saltmarsh snakes (natricines) eat small fish and prey in their jaws. Triggerfish durophagous fiddler crabs; the granulated file snake (Acmchordus specializations include the loss of jaw protrusion, granulatus) eats fish, crustaceans, and snails enlarged jaw adductor muscles, and stout teeth. CfTeatwole, 1987). Sea snakes swallow their prey Sparidae (parrot fish) also crush corals and other whole, but it would be very useful to ho~vwhat the hard prey. taphonomic quality of the invertebrates are once they Fish other than kopical reef fish can fragment pass through the gut of the sea snake. That is, what shell mated (Cadie, 1968 Cate and Evans, 1994; size hgments? Is there any indication of gastric Norton, 1995j; although they may be responsible acids on the fragments? How much do they eat of for a majority of fragmented shelly remains in the varying prey items? fossil record, direct evidence linking them to the Sea ibrtIes4enozoicfossil MIes are known scene of the crime is lacking. Alternatively, fish from a number of localities dating from the that puncture shelly hardpaas are known (Norton, Paleocene (Weem, 1988). The Chelonoidea first 1988), and it would be possible to trace this specific appeared in the late Early Cretaceous and the type of shell damage in the fossil record, although Cenozoic fauna includes the survivors of the this has not been zttempted. Cretaceous-Tertiary mass extinctioi: Dermo- Sea smkes.4ea snakes evoIved from varanid chelyidae and Cheloniidea Wirayama, 1997j. Of ancestors (as did mosasaursj in the mid-to-Late these, the Demochelyidae, with their thin shells Cretaceous, and diversifred greatly in the Cenozoic and fontanellization, are poorly preserved; (Caldwell and Lee, 1997; heand CaldweU, 2000). chelonids are better preserved, and are slightly better From the Late Cretaceous to Eocene, there were known (Weems, 1985). Despite th~staphonomic several genera of marine snakes representative of problem, no catastrophicterminal Cretaceous event the booidean fady Palaeophidae (Hecht eet al., is mident in the record of sea turtles (Weemq, I98 8). 1974; Heatwole, 1987). Early Tertiary fossils of Sea turtles had declined in diversity by the late sea snakes are very abund.ant and globally Campanian, and were low in diversity during the widespread. However, the Palaeophidae are not the Maastrichtian and Danian, but recovered in the direct ancestors of modern sea snakes; rather, the Thanetian and Ypresian stages of the early Cenomic Family EIapideae (terreshal, venomous snakes of (Weem, 1988). The cheloniids underwent a major the cobra family) is thought to have given rise to diversification in the late Paleocene (Weems, 1988). the extant sea snake fauna between 35 and 25 This pattern of diversity matches the global pattern million years ago, in the Oligocene to Miocene. of oceanic cooling and warming in Late Cretaceous However, the modern genera are not well hown to early Tertiary time (Weems, 1988, p. 143, his as fossils Weatwole, 1987). The modern fauna of fig. 27). The later Tertiary sea turtles are too poorly sea snakes, Laticaudinae and Hydrophiinae, are known to allow us to extrapolate diversity at this comprised of 12 genera and approximately 448 time, but jn general, dversity dcclined f3om he species (Hecht et al., 1974) distributed chiefly in Eocene to the five cosmopolitan species remainig subtropical to tropical oceans. A few saltmarsh and today (Weems, 1988). estuarine snakes also occur in temperate North Although modern turtles are morphoTagically America. Most sea snakes are nearshore creams similar, their feeding preferences differ: the (within the 100 m isobath; Hecht et al., 1974). Cheloniidae are omnivorous, and herbivorous Although they not particularly well studied, it is (Hkayama, 1997) adult geen turtles (Cheloaia WMKER AhD BRETT-POST-PALEOZOIC PmERNS INhKANNE PREDATION mydas) eat sea gasses and algae; the hawksbills included among the oldest fossils of the Division (Eretmochelys imbricata) eat sponges, and Kemp's Neognathae, with Jite Cretaceous records for the ridley (hpidochelyskernpii) eats crustaceans. The transitional shore birds (Feduccia, 1995). Several Dermochelyidae are rather fond of jellyfish. orders of marine birds have fossil records extendhg Caretta has massive jaws with a b3xtrating at least to the early Paleogene; these include surface; this genus is most common in temperate Anseriformes (ducks), Gawiformes (loons), and waters, and the present distribution of this sea turtle Charadsiformes (shore birds). dates back to the early Pliocene {Dodd and Morgan, Foot-propelled loons (0. Gaviifermes; Late 1992; Panis et d.,2000). Caretta eats sea pens, Cretaceous(?) to Recent) and grebes (0. crustaceans, and molluscs among many other prey Podicepifonnes ; Miocene to Re cent) appear highIy items (Hendrickson, 1980; Plotkin et d.,1993; convergent on the Cretaceous ichthyomithines and Nicholls, 1997). One species, Caredta caretta, the hesperornithines, but are not closely related loggerhead turtle, is hokvn to eat a variety of prey (Chiappe, 1995). Con~astingly,gliding albatrosses (Mortimer, 1982; Bjorndal, 1 985). Camtta off the (0.Procellarifomes; Eocene to Recent), some with Texas shelf feed primarily on sea pens inthe spring, wingspans exceeding 3.5 m, gulls (0. and benthic crabs in the summer and fall (Ploclrin Charadriformes; Eocene to Recent), and pelicans et al., 1993). They can also eat molluscs, and cormorants (0.Pelicaniformes; Eocene to antbxopogenic debris (e.g., fishing line, plastic mash Recent) seemingly fill a guild similar to the bags), Diopatm tube worms, barnacles, fish, Mesozoic sea-going, piscivorous pterosaurs. seaweed, whip coral, sea pansies, sea anemones, Finally, the penguins (0.Sphenisciformes; Eocene stomatopods, shrimp, and jellyfish (Plotkin et at., to Recent) (Fig. 114, includmg some 25 genera, 1993). The small bivalve molluscs present in their have an excellent fossil record, primarily in the stomachs may come from the digested tubes of Southern Hemisphere (Simpson, 1975); &verse Diopatra, and perhaps may not have been directly fossils are especially common in the Eocene to fed upon. Scavenging gastropods that feed on dead Miocene of New Zealand and Seymour Island. fish or crabs, such as Nassarius acutlks, may have These amphibious birds have become specikd been eaten accidently as the turtle went after for rapid underwater fight even as they have lost decaying fish fPloth et al., 1993). aerial flying abilrty. Interestingly, an extinct clade Caretta populations from Merent geographic of flightless pelicanifom birds, the Plotopteridae areas feed on different types of prey. In the western (Eocene to Miocene), convergently evolved Mediterranean, Caretta carem eat fish and hrnicate elongate paddle-like wings for underwater flight. salps, although they can also eat benthic Some of the Pacific plotopterids attained lengths crustaceans and molluscs (Toms et al., 2001). of 2 m (Olson and Hasegawa, 1979). All of these However, the fish may be from scavenged by-catch birds are primarily piscivores and their abundance that is thrown overboard by fisherman. Caretta can attests to the proliferation of small teleost fish in also forage on jellfish at the ocean surface (Plotkin near-surface seawater. et al., 1993). The variety of prey that these The diversification of diving and other coastal loggerheads eat is impressive, denoting a generalist marine birds also may have greatly impacted the (Plotkin et d.,1993). Of allthe sea turtles that exist fossil record of crustaceans and molluscs (Vermcij, today, Care-etta is the only generalist. 1977,1987).Although several groups origmated jn Sea and Shore Birds.-The extinction of the Late Cretaceous, the diversification of shore pterosaurs and early toothed divi~gbirds in the Late birds and diving marine birds took place chiefly in Cretaceous left open another important niche for the Paleogene (Vermeij, 1987). Oyster catchers, marine piscivorous predators. It seems that this however, originated in the Neogene (Olson and void was filled rapidly by the evolution of Steadman, 1978). Diving marine birds catch fish neognathan sea birds (Fig. 11 -6).Aquatic birds are that had previously preyed on molluscs; the PMEONTOLOGICM SOCIETET PAPERS, V8,2002 molluscs are then transported to nests often far produced by foraging rays and flatfish (Cadee, from their original habitat (Teichert and Serventy, 1990). However, rays excavate sand around the 1947; Smith, 1952; Lindberg and Carlton, 1969; circumference of the foraging pits, whereas Lindberg and Kellogg, 1982). Coastal birds such shelducks excavate to only one side (Cad&, 1990). as the oyster catchers and eiderducks can prey These distinguishing characteristics could be directly on large numbers of molluscs and other obliterated by the tides, and therefore it may be invertebrates (SchSer, 1972; Cadke, 1989,1995). difficult to distinguish bird foraging pits from those Oyster catchers penetrate molluscan prey by of benthic fish predators. Foraging pits of aquatic stabbing slighdy gaping valves with their beaks, birds are known only from the Holocene. resulting in broken mollusc shells (Drinnan, 1957; Pinraiped Muwzmccks.-Pllmipeds-seals, sea Carter, 1968; Cadee, 1995). Oyster catcher lions, and walruses (Fig. 11.1 1)-in modern seas predation often produces distinctive shell damage, have a global distribution, occur in enormous with one valve Iragmented and the other numbers in some regions of the world, and are able untouched (Drinnan, 1957). However, at least half to dive to great depths in the ocean in search of of the prey shells may be left intact as these birds food (Table 4). Therefore, some forms must have also insert their beak between the valves, had a significant impact on hardshelled supposedly without harming the shells. communities, although many eat fish. Some of the The diet of hemng gulls may consist of up to pimipeds evolved molar crowns with hyper- 70% marine molluscs. Prey are diectly scooped mineraked cutting edges for crushing and piercing up by the gull from the marine environment and the hard exoskeletons of crustaceans and molluscs transported to the shore where the shells are (Haley, 1986). Pmnipeds evolved in the ~ocene,and dropped over tidal flats or other hard surfaces, thus have had over 40 &on years to affect the which fragments the shells so that the meat can be evolutionary history of molluscs and arthropods; extracted (Cadke, 1995). Often, birds select however, there is no indication that they did so. particular sizes of prey, which can affect the Walruses (Family Odobenidaej feed mainly on resultant fossil record of-coastal molluscan benthic invertebrates, and have a peculiar feeding assemblages (Cadke, 1989). Cadke (1995) has style: they suck out the siphon or foot of bivalves estimated for the Dutch Wadden Sea that shell- using their piston-like tongue while their mouth crushing shore birds may fragment np to 35% of works as a vacuum pump (Muizon, 1993). the annual shell carbonate production. Because the Wahses have large and deep palates, a wide, blunt Dutch Wadden Sea benthos has up to 75% shell snout with strong muscular insertions, and a fragments, myof these fragments may be of rediction of maxillary dentition (Muizon, 1993). biological rather than physical origin (Cade'e, The fmks are thought to have a primarily social, 1995). Thus, in shallow Cenozoic seas, coastal and rather than foraging fmction. Walruses also leave sea birds would have been important agents of shell long, narrow feeding tracks or small excavated pits frag mentaion (CadGe, 1995). These fragments must that can be seen in side-scan sonar' (Oliver et al., occur in the fossil record (Trewin and Welsh, 1972), 1983; Nerini, 1984, her fig. 3). but it is generally impossible to pinpokt exactly AUknonm odobenine odobenids (walruses) are who fragmented the shells. bottom feeders and are first known fiom the Miocene Shore birds may also leave benthic feeding (Repehg, 1976). Several.extinct species from the traces in soft sediment. For example, gulls such as Pliocene are known to have been rnolluscivorous Lam ridibundm may make koughs up to 3 mlong, (Repenning,1976) and were widespread at that time 15 cm wide, and 3 cm deep jn soft shore sediments in the northern hemisphere (Muizon, 1993). as they forage for food (Cadke, 1990). Shelducks However, the modern walrus {Odoberzus}has a fossil make smaller pits (60 cm in diameter and 10 cm record only fiom the Pleistocene (Repenning, 1976). deep). These feeding traces are similar to those There is no direct fossil record of pinniped predatory WALKER AND BRE7T-POST-PALEOZOIC PAT7ENS IN MARINE PREDATION behavior, with the possible exception of putative structure (Estes and Palmisano, 1974; Estes et al., coprolites packed with crab parts, attributedto seals 1982). It is also known that where humans have (Glaessner, 1960; Boucot, 1990). preyed on sea otters for their furry pelts, the Sea otters.--Sea otters (Table 4) evolved resultant fossil record is skewed toward during the cooling period of the late Miocene, and herbivorous limpets and sea urcb; where sea are restricted to temperate regions (Van Blaricom otters are not preyed upon, the stratigraphic record and Estes, 1988). Otters occur in shallow coastal shows abundant kelp beds and fish populations areas, where they eat a variety of invertebrate prey (Simenstad et al., 1978). (Haley, 1986; Van Blaricom and Estes, 1988). In Cetaceam.-Cetaceans (whales and dolphins) soft-sediment habitats, they are known to prey upon originated from land-dwelling artiodactyls in the endobenthic bivalves (Kvitek et al., 1992, 1993). early to middle Eocene (hgerich et a]., 2001; Sea otters ingest copious amounts of echoderm Thewissen et al., 2001j. Early forms such as and molluscan prey-taking in up to 35% of their Ambtklocet~s(Fig. 11.7-1 1.9) probably were body mass in invertebrate prey every day (Hines amphibious and may have behaved like seals and Pearse, 1982)-but their predatory effects in (Thewissen et d.,2002). It is not known what these the fossil record remain unknown. Hines and ancient toothed whales fed on. By the late Eocene Pearse (1982) used the size, structure, and breakage the gigantic (20 m) and Mymarine Bmilosaurus characteristics of empty abalone shells to document seems to have occupied the gddof large Mesozoic the selectivity of the predator and the source of marine rept3es, such as mosasaurs (Fig. 11.lo). Its abalone mortality in a rocky subtidal habitat off of sharp, multiply cusped, undifferentiated teeth were central California. Gormand sea otters prefer apparently adapted to fish capture, although a abalones in California, and can consume about ten relatively small head limited prey size (Benton, abalones a day (Costa, 1978). Similarly, cracked 1997). The toothed whales (Suborder Odontoceti) shells were used to infer otter predation on bivalve diverged in the Oligocene and radrated during the prey in southeastern Alaska {Kvitek et al., 1992). Miocene into a large number of smaller, dolphin- In this area, sea otters substantially impacted the like lineages (Barnes, 1984). These whales evolved population of endobenthic bivalves and epibenthic highly sensitive echolocation and fast-s wimming urchins (Kvitek et al,, 3992, 1993). Additionally, behaviors. They are well adapted for chasing down foraging pits dug by otters atwacted predatory sea and capturing fish, sharks, and, in some cases, other stars, which then ate any exposed molluscs. Otters whales. Apparently, these odontocetes re-evohed digging for cIams also exhumed buried shells many of the adaptations of Mesozoic pursuit (Kvitek et al., 1982), suggesting that biological predators, specifically the ichthyosaurs (see remeis common in these areas. The reworked Massare, 1987, 1997). The largest toothed whales, shells then become settling sites for epibenthic sperm whales, are of uncertain origin, but molecular invertebrates. Curiously, sea otter populations may studies of Milinkovitch (1995) suggest that they may be controIled by paralytic shellfish poisoning in actually have been derived, in the Oligocene Epoch, these areas CKyitek et al., 1993). from the baleen whales rather than the oodntrxete The predatory record of these creatures should whales. Sperm whales are well adapted for deep be discernible because sea otters have peculiar diving in pursuit of squid prey and perhaps occupy carnassial teeth that are flat and rounded for the guild of some Cretaceous mosasaurs. crushing prey, and their lower incisors are used to An unprecedented find of a walrus-like whale scoop meat out of shells. One thing is certain, skull from the Pliocene of Peru indicates that one however: where sea otters occur, their effects on rare form of whale may have been durophagous populations of their favored food items should be on molluscs andlor crustaceans (Muizon, 1993). great. Sea otter predation on sea urchins has a Odobenocetops peruvianus did not have an considerable effect on nearshore community elongated rostrum, but had large ventrally chected PALEO~VTOLOGICALSOCIETY PAPERS, Y 8,2002 premaxillary tusks, a deep-vaulted palette without population of gray whales (estimated in 1984 at teeth, and strong muscle scars on the premaxillae, 15,500 whales) could turn over 3,565 km2/yr of sea which indicate durophagy (Muizon, 1993). Its bottom while feeding, considerably impacting the morphology is similar to the Beluga and narwhal benthic communities where they feed Werini, 1984). whales (Monodontidae). Gray whale fossils, however, are ody known The Suborder Mysticeti (baleen whales) from the late Pleistocene, although several cIosely originated in the Oligocene (Whitmore and Sanders, related groups are known from the Miocene of North 1976) and developed sheets of horn- or hair-like America (J3ames and Mehod, 1984). The obligate baleen for sieving water to collect pelagic organisms, barnacle parasite of gray whales, Ciyptukpw, is also especially krill-a form of predation previously only known from the late Pleistocene (Barnes and evolved by certain bony fish (e.g., Mesozoic McLeod, 1984). It is known that there were two pachyconnids, and whale sharks) and perhaps by a allopaaic populations of the gray whale in the early Triassic marine reptile, the placodont Henodus. All Holocene, one in the Noah Pacfic and one in the of these organism attained large size, and mysticete North Atlantic, which is now extinct. whales include the largest known organisms. Order Sit-enia (sea cawsJ.--Sea cows date from While a number of cetaceans may eat some the Eocene, aad are a very small group of mammals benthic fma,it is only the gray whde (Mysteceti, that feed chiefly on sea grasses, algae, or water Eschrichtiidae, Eschrichtius robustus) that hyacinths (Damning, 1976; Savage, 1976). One consistently raids the benthos in search of particular fossil Sirenian, however, may have fed invertebrates (e.g., tubiculous amphip ods and on benthic molluscs. Miosiren from the late callianassid shrimp) to- complement its fare of Miocene of Belgium displays thickened tooth pelagic prey such as squids, mysid shrimp, and fish enamel and cusp modifications, which indicate that (Norris et al., 1983; Nerini, 1984). Gray whales are it may have fed on rnoIluscs (Savage, 1976). also known to skim eelgrass mats for both Other mammals that forage far marine crustaceans and sea grasslalgae, and sandy muddy invertebrates.-Raccoons (Procyon) forage for habitats for gaskopods, bivalves, and tube-building cnzstaceans in temperate to subtropical tidepools polychaetes (e.g ., Diopat~aand Onuphis; Nerini, and saIt marshes (Ricketts et al., 1985;Walker, pers. 1984,her table 2). Buccinids, neptunids, thaids, and obs., 1997). The first known Procyon is horn the naticids are just a few of the gastropods that have upper Pliocene; there are several Pleistocene fossil been found among gray whale stomach contents; species as well (Arata and Hutchison, 1964). Macow Mya,and Mytilm are some of the ingested Fossils of Procyon are known from all over the bivalves. The gray whale is able to sieve sediments continental United States, as well as Baja California through its hckbaleen plates, which have coarser and Canada (Arata and Hutchson, 1964). Coyotes hairs than other baleen whales (Nemoto, 1970). and other mammals also can feed in the intertidal Gray whales leave very large feeding pits in zone of temperate regions (Ricketts et d.,1985). shallow, nearshore to intertidal mudflats that are Rats, in particular, can prey on over 40 different often the only record of their feeding behavior types of intertidal organisms, especially key hole (Nerini, 1984, her fig. I). On one benthic foraging limpets, porcellanid crabs, and cancrid crabs dive, it is possible -Ear one whale to make a series of (Navarrete and Castilla, 1993). shallow pits that are usually arrayed in a slight curve Humrzpts.-Lastly, the origmation of humans and range from 1 to 3 rn long and from 0.5 to 1.5 m in the late Pleistocene added to the potential for wide. Gray whales are known to commonly feed coastal foraging and selection of particular in Baja California lagoons, along their migratory invertebrate food items as evidenced by abundant range from the Bering Sea to Baja California (a kitchen midden sites around the world, as well as 6000-km range), and in the northern Bering, tools embedded in late Pleistocene coral reefs (see Chukchi, and Beaufort Seas @mini, 1984).An entire Walter et al., 2000). Humans have been using sea WALKER AND BRETT-POST-PALEOZOIC PATTERNS IN AURlNE PREDATION creatures for food and ornamentation for many remains, predation preserved in situ, and prey thousands of years based on archaeological shell organisms in stomach contents are rare in midde~ls(e.g., Speed, 1969; Avery and Siegfried, Cenozoic deposits just as they are rare in 1980; Jerardino et al., 1992). For example, hChile, Mesozoic and Paleozoic assemblages (Hitzsehel the rocky coast has been exploited by humans for et al., 1968; Boucot, 1990; Brett, 1990). While food for at least 8,500 years (Moreno, 2001). This there is an extensive literature on coprolites, most foraging was tied closely to settlement of the studies focus on terrestriaI and vertebrate remains; Pacific region of South America, and has only few if any coprolites in marine environments can recently been recognized as a force that affects the be tied with reliability to a specific predator resultant ecological community structure ofan area (Bishop, 1975; Boucot, 1990). (Moreno et al., 1984). Ecological shifts in seafood Echinoderms.-In many localities, not least in biota directly or indirectly caused by humans are the Danish basin, the Cretaceous-Tertiary extinction known from thc present day (Castilla and Duran, greatly affected the invertebrate biota. However, 1985; Castilla, 1999) and ffom the stratigraphic several echinoderm groups do not appear to have record (Simenstad et al., 1978; Kirch, 1983). been greatly affected by this extinction event, and show an increase in diversity directly above the EVOLUTIONARY VIGNETTES: boundary in the Danish basin (Kjaer and Thomen, SELECTED PATTERNS 1999).There are several examples of shallow-water OF PREDATION stalked crinoids from the early Cenozoic (Oji, 1996); FROM TEE CENOZOIC and further movement oflshore of isdd(stalked) crinoids occurred in the Miocene in the Caribbean The Mesozoic Marine Revolution hypothesis region (Bottjer and Jablonslu, 1988; Donovan, (Vermeij, 1977, 1987) has been subjected to many 2001). Deeper-water crinoids have a relatively tests, £rom several sources of evidence, chiefly to constant genefic composition hrnthe Miocene to determine: (I) if shell armor increases through time; the Recent; the Plio-Pleistocene regional extinction (2) if shell predators increase through he;and (3) had little effect on this group (Donovan, 2001). d lethal shell injuries increase through time (see also Shallow water areas remain populated by stern-less Vmeij, 19831. In the following sections we review comahlid crinoids (Donovan, 2001). This suggests and critique some of the primary lines of argument. that mobility and cryptic habitats may have enabled Most of the putative durophagous functiond groups this group to swivein the face of high predation in re-evolved in the Cenozoic, and, one could argue, shallow water. became more common during this time than in the Arm autotomy is common jn stalkless crinoids, Mesozoic. However, some of this apparent increase but has not been well documented in stalked may represent biases such as the Raupian "pull of crinoids (Oji, 1986). The ability to autotomize the Recent" and the better record of well-preserved crinoid arms dates back at least to the Triassic (Oji fossils. It is also well known that aragonitic and OJsamoto, 1994). It is thought that autotomy Mesozoic invertebrates, especially molluscs, are not acts as a "lizard-tail'yefense (after Baumiller et as well preserved as calcitic forms (except for al., 1999): arms can be dropped quickly into the ammonoids in black shales); whereas in the mouths of predators, while the rnajn body of the Cenozoic more aragonitic forms are preserved, crinoid is Ieft to regenerate new arms. It is possible giving us a more detailed picture of the potential that isocrinids exploited this ability and that this is predatory panorama. She11 repair, drilkg, and other what allowed them to survive the putative increased features can be distinguished on Cenozoic hardparts fish predation in the late Mesozoic (Oji and much more easily than on older ones. This is not a Okamoto, 1994). Modem crinoids from bathyal gloom-and-doom scenario, just a realistic one. depths have more regenerated arms than crinoids Examples of prey in coprolites or regurgitated from deeper depths (Oji, 1996). P~OIVTOLOGICALSOClETY PAPERS, V 8,2002

Stalk shedding is also a common occurrence surface (Nebelsick, 1999). Fish bite marks on in isocrinoids, and may be a deterrent to predators. clypeasteroid echinoids are also reported horn the Baumiller et al. (1999) hypothesize that crinoids upper Miocene of Argentina (Zinsmeister, 1980). have evolved various antipredatory strategies since h the modern Atlantic and Gulf region, there the Devonian: a pIanktonic (e.g., Uintacrinus) or are 95 asteroid species in 56 genera, with a pseudoplanktonic (e.g., Seimcnnus) Westyle, stalk- depauperate (because of lack of work) record in shedding abilities (e.g ., in isocrinids, camatuIids), the Cenozoic Caribbean region dating to the early short-bursts of swimming (e-g., comahllids), and Paleocene (Donovan, 2001). Asteroids are known life in cryptic habitats (e.g.,comatulids) . from the Eocene to Pleistocene in Florida, and in Kier (1977) plotted global diversity of some horizons thek fragments are very abundant echinoids through the Cenozoic, and showed (Oyen and Poaell, 2001). Amazing preservation of limited diversity jn the Paleocene and Oligocene complete specimens of He1 imteu microbranchiusis Epochs, with peaks in echinoid diversity in the horn from the Pliocene of Florida (Oyen and Eocene and Miocene-Pliocene Epochs; the record Portell, 2001). of regular echinoids was not as good as that of Ophiuroids are one of the most &verse extant irregular echinoids. Regular echinoids are echinoderm groups in the Caribbean region, but commonly fragmented, and their fragments usually have a "poor" fossil record because of their easily are not studied by taxonomists (Greenstein, 1993) disarticulated skeletons and a lack of work on these or are not collected (Oyen and Portell, 2001). creatwes @onovan, 200 1; Oyen and Portell, 2001). Clypeasteroids evolved in the Paleocene and Nonetheless, a number of dense stalked, crinoid- diversified rapidly, aided by the evolutionary ophiuroid associations are known from before the innovation of numerous small tube feet and spiue- Jurassic; a near absence of these dense assemblages free branching food grooves. Flattening of the test after the Jurassic was postulated to be due to meant that only the top fraction of the sediment predation pressure (Aronson, 1987, 1991). could be sieved for food particles mer, 1982). Intri,ouingly, however, the Tertiary La Meseta Records of predation on Cenozoic echinoids Formation, Antarctic Peninsula, contains localized are rare, even though in modem seas predation on dense assemblages of autochthonous ophiuroids echinoids is well documented (Nebelsick, 1995, and crinoids representing shallow-water facies 1999). Drilled echinoid tests arc known from the (Aronson et al., 1997). The incidence of sublethal Eocene Upper Ocala Formation in North Central arm injuries was low in this assemblage, suggesting Florida (Gibson and Watson, 1989). Some of these that predation was rare; possibly in high latitude drillholes were predatory; others were parasitic. cool-water areas predation is suppressed. Parasitic eulimids are known to drill the aboral MoUuscs.-Molluscs provide the most sides of echinoids; commonly an ecboid displays important Cenozoic database for examining multiple drillholes made by parasitic gastropods evolutionary questions regarding the fossil record (Berry, 1956). Cassid drillings on irregular of predation because they ari globally widespread, echinoids are known from the Eocene of the very abundant, well preserved, and present in many Atlantic Coastal Plain (Woodcock and Kelley, different facies. Therefore, most studies have 2001) and elsewhere (see Cassid review, th~s focused primarily on escalationin marine molluscs. paper). Sand dollars (Pamscutella hobarthi) from Shell repair and shell drilling in mollvscs have the lower Miocene of the Austrian Molasse Zone provided the database by which to examine displayed repaired scallop-shaped areas on their Fhanerozoic predatory kends. Shell repair data has tests resulting from predation, possibly by reguiar not been applied with as much success as drilling echoids (Nebelsick, 1999). Lethal predation was predation, most likely because shell repair can be indicated by large round holes cutting though the a consequence of a variety of physical and echinoid test or by bite marks penetrating the oral biological destructive factors. Shell repair may WALKER RRTD BRETT-POST-PALEOZOIC PAlTERilTS IN kMRlNE PREDATION show an increase in the Cretaceous and Cenozoic, (Vermeij et al., 1980, 1982; Vermeij, 1982). This or it may not, and a closer examination of shell method does not take into account the fact that older repair d~ringthis time is warranted (Table 3). Little shells may have more shell repair than younger work has been done for Cenozoic localities to shells, and thus can result in an overestimate of shell examine shell repair with respect to habitat, species, repair for an assemblage (although Vermeij has and stratigraphic interval. recognized this problem). Further, more instances Do lethal shell injuries (or shell repair) of shell repair than actual sample size are commonly increase through late Mesozoic-Cenozoic time?- reporkd which makes the data dXficuIt to interpret. Traces of non-fatal peeling in moUuscs are evident Therefore, it is important to determine which method as scars on the shell (Figs. 8.2, 13; Table 4) that is most useful in examining the fossil record of shell result from repair of the outer shell lip by the mantle repair and to be consistent with that method. edge (Robba and Ostinek, 1975; RaffaeVi, 2978; Comparing papers that use Merent methods is Elner and Raffaelli, 1980; Vermeij et d.,1982; difficult and tenuous at best. Vermeij, 1982; Allmon et d.,1990; Cadee et al., Interpretations of shell repair must be carefully 1997). Frequency of shell repair is often cited in evaluated especially In regard to equating frequency order to compare temperate with tropical and deep- of shell repair with intensity of predation (Cadie et sea with shallow-sea habitats, as well as to examine al., 1997; Cadke, 1999). There are several factors within- and between-habitat predation, and the that complicate the interpretation of shell repair. temporal dynamics of shell repair. It appears that First, it is =cult to distinguish repair that may shell repair may increase through the Phanerozoic, have been provoked by physical factors, such as with higher incidence of shell repair in the burial or crushing between stones (e.g., Raffaelli, Cenozoic-indicating that durophagous predators 1978; Cadie, 1999). Seff-inflicted wounds resulting become more of a threat to moIluscam prey (e.g., from the process of predation that are then Vermeij et al., 1980, 1982; Vermeij, 1983, 1987; subsequently repaired can also inflate estimates of Diet1 et al., 2000). But analysis of the data on shell shell repair For example, buccjnid gasf~opodschip repair (Table 3) dustrates fiat there are no real their outer lips in the process of preying on other differences in shell repair frequency between the molluscs and then rep* their self-inflictedbreakage Mesozoic and-Cenozoic, despite 'the better record mielsen, 1975).Second, shell repair frequencies do of marine durophagous predators at this time. not directly correlate with the intensity of predation, Shell repair must be interpreted with caution, as a total absence of scars may mean either that as researchers use different methods and predation did not occur or that predators were 100% interpretations in analyses of shell repair data. Two efficient (Schoener, 1979). Third, the incidence of methods are used to estimate shell repair repair on a shell needs to be tied to the age of the frequencies. First, shell repair frequencies can be organism, as older snails may exhibit more shell estimated by dividing the number of shells with repair than. younger ones. Th~smay be especially one major repair (jagged scar) by the total number true for deep-sea snails that may exhibit slower of shells in the sample (after Robba and OstinelJi, growth rates and increased longevity with depth 1975; Raffaelli, 1978; Elner and Maelli, 1980; (Vale and Rex, 1988). Fourth, certain life history GeBer, 1983; Vale and Rex, 1988, 1989; Cad& et traits (slow growing vs. fast growing, particular al., 1997; Walker, 2001). This is the more behavior) and feeding mode may affect whether and conservative estimate for shell repajr, as snails can when a shell is exposed to predation. Fifth, some survive injury more than once. If the snail is older, species may be more prone to predation than others it may display more instances of shell repair. Second, in an assemblage (Hoffmeister and Kowalewski, shell repair frequency has also been calculated as 200 1; Kelley and Hamen, 2001 ;Walker, 2001); and, the total number of scars in all shells divided by the using the metric of only one species' repair total number of shells in the sample (Table 3) frequencies can bias the results for an entire PMEONTOLOGICAL SOCIETY PAPERS, V8,2002 assemblage. Lastly, for a time-averaged assemblage, overabundance of drilled shells in some localities. shell repair hquencies might be higher than what Additionally, drilled shells are more prone to would be found in the living population at any one taphonomic breakage than undrilled shells, and time because of the patchiness of predation (and such breakage may be more common Ln some associated physical factors). localities than others (Roy et al., 1994). Left vs. The consensus is, however, that conspicuous right val~esof bivalves and pedical vs. brachial shell repair (i.e., conspicuously peeled shells with valves of brachiopods are also differentially subsequent repair) is most likely the result of transported andlor preserved (Brett and Allison, predation. That is, only deeply peeled injuries that 1998). Thus, it would be important to know the are subsequently repaired can reliably be used in valve kequencies of an assemblage, and whether the analysis of shell repair, whereas repaired nips they are biased. It would also be important to know or edge chippings may not be indicative of ifddhg predators were actually found in the same predation (Walker and Voight, 1994; Walker, 2001). assemblage as the drillholes (e-g.,Hansen and Consequently, although shell rep& is not a good KeUey, 19951, but not all papers that examine indicator of predation intensity, it is instrumental drilling predation discuss this issue. in providing a record of predators within a habitat It is also important to examine more than one when body fossils of the predators are missing. locality within a time period, as the record of Shell Drilling through Time.--Shell drilling predation is seongly controlled by habitat (Vermeij frequency is less ambiguous in interpretation: a et al., 1981; Geller, 1983; Hansenand Kelley, 1995; completed drillhole signifies prey mortality. Also, Cadie et al., 1997; Hoffmeister and Kowdewski, particufar borehole horphologies may be 2001). Location within. a sequence may al& affect associated with specific gastropod or octopod the density of drilled shells, as trtransgressive lag predators (Carriker and Yochelson, 1968; Kabat, deposits formed after a major sequence boundary 1990; Kowalewski, 1993; KowaEewski et al,, (e.g., extinction?) commonly contain more biotic 1998). Nonetheless, certain caveats also apply to information as a result of longer time averaging the study of drilling predation, (Brett, 1995; Holland, 2000). Therefore, one must Escalation striclies of drilling predators and be careful in interpreting the pattern and process their prey have not generally taken into account of drilling through the Phanerozoic, as it is not as the particular facies and associated biota of simple as merely counting drilled maper temporal analyzed assemblages (with the exception of stratigraphic sequence. As Boucot said, "Nature Hoffmeister and Kowalewski, 200 1). Essentially, does not take place wihan ecoiogical vacuum"; all assemblages are Weated a? if they were the same nor should evolutionary interpretations using the facies (e.g., onshore and offshore assemblages are fossil record be decougled from facies studies. grouped). Environmental differences between even these caveats, based on an analysis of assemblages, however, can aRect the morphology over 150,000 gastropod and bivalve shells from of the tax&-some species are larger in nearshore the Gulf and Atlantic Coastal Plain (GACP), KelIey environments than they are in offshore environments and Hansen (2001) suggested that the interaction (or vice versa). This gradient in morphology may between naticid drilling predators and their prey not be related to predation. does not necessdy show escalation from the Sedimentary facies couId also have Cretaceous to Oligocene. After examination of a taphonomic effects. For example, assemblages number of localities, they found that there is an deposited above storm wave base may sort drilled episodic pattern to drilling frequency, with mass and andrilled shells differently compared to extinctions resetting the "arms race" for faunas. offshore assemblages. Drilled and lmdrilled shells Drilling within the most of their Cretaceous can be differentially transported in nearshore localities was greater than several of their late settings and thus there may be a bias toward an Eocene localities and similar to early Oligocene WaLKERAhrD BREm-POST-PALEOZOIC PATTEWS IN hMRlNE PREDATION localities (their table 8.1, p. 653). This is a various assemblages examined. significant finding given that previously Vermiej Escalated species are thought to be more (1987) had used naticid drilling as one line of sensitive to changes in primary productivity because evidence illustrating escalation in Cretaceous to maintaining heavy armor or high speeds to avoid Eocene faunas. In the Cretaceous Vermeij (1 987) predators requires high metabolic rates and thus an found limited drillmg, but by the Eocene, drilling uninterrupted food source (Vermeij, 1987). had reached modern levels. Therefore, Hansen et al. (1999) tested whether Kelley and Hansen (19931, in contrast, did not purported escalated species (those with find an ever-increasing trend in naticid drillkg antipredatory adaptations such as heavy armor) were frequencies from the Cretaceous to Eocene for more vulnerable to extinctions caused by climate GACP molluscs. Escalation could also mean that a change and associated environmental changes. Ten predator gets better at selecting prey; however, she11 characters deemed important for predator Kelley and Hansen (1993) did not flnd mytemporal resistance were evaluated for GAPC molluscs across trends toward increased drillhole site stereotypy in various mass extinction events associated with naticids. Molluscan prey were found to have more climatic cooling andlor a decline in primary incomplete drillholes and multiply drilled shells, productivity (e.g., Cretaceous-Paleocene; Eocene- indicating that prey effectiveness may have Oligocene; middle Miocene; Pliocene-Pleistocene). escalated, but Kelley and Hansen's (2001) data did Importantly, all these assemblages were deposited not show a trend for most of the periods examined. in relatively shallow shelf environments with Keiley and Hansen (2001) also examined roughLy similar grain sizes; all but one assemblage diEerences in morphology within molluscan genera was a bulk collection. Hansen et al. (1999) found that may be related to escalation. Although many that escalated species, overall, were not more genera were examined, four particularly Iong- vulnerable to climate-related mass extinction. Only ranging Miocene genera from the GAFP-two ornamented Pliocene gasbopod species were more gastropod predators (Empira hems and Never& susceptible to extinction than their weakly duplicicatu) and two frequently ddled naticid prey ornamented counterparts. In another shrdy, Kelley (Bicorbula idoma and Stewartia anockoptta)--were et al. (20013 found that recovery faunas after a mass analyzed for different morphological charateristics extinction event were not more vulnerable tu (thelr table 8.3, p. 159). In this case, however, the enhanced drilling pmsure, contrary to hypothesized gastropod predators are also cannibalistic. Results predictions. Additionally, no overall trend in showed that shell size (height) did not change for unsuccessful drilling was seen from the late either E. heros, B. idonea, or S. anodoata (no data Cretaceous to Pleistocene. are reported for N. duplicata), indicating that these Spatial trends in drilling predation vary by prey species found no sjze refuge from predation environment in fossil studes. Hansen and Kelley over time. Shell thickness (which wodd make aprey (1995) used 27,554 specimens of GACP molluscs item more difficult to M}did not change for E. from the Eocene and found a statisticdy si&cant heros, decreased for N. duplicata, slightly increased difference in drilling frequency between the inner- for 3.idowa and increased greatly for S. mdonta. to middle- shelf Moodys Branch Formation and Internal volume (an indicator of the amount of food the outer-shelf Yazoo Formation, the deeper site a predator can take in) did not change within the having a higher frequency of drilling predation. Wocene. Thus, it appears that most prey characters However, for the five other assemblages examined deemed to be directly related to predatory escalation from the Moody's Branch, there was no significant did not demonstrably change within the Miocene bathymetric trend. Drilling frequency was also (except for shell thickness in S. anodunta). It would highly correlated with the percentage of naticids be interesting to how whether drilling frequency and their preferred prey within each assemblage. increased or stayed relatively the same across the Hoffmeister and Kowalewski (2001) examined PAL,EOATOLOGIC1QL SOCIETY PAPERSJ V 8,2002 spatial and environmental variation in drilling Molluscan conchiolin layers: Are they predation in the middle Miocene of Cenbal Europe. antipredatory ?-Conchiolin is the organic The sampling methodology allowed for component of molluscan shells composed of comparisons within provinces, between provinces proteins, polysaccharids, and glycosaminoglycans (Boreal vs. Paratethys), and between facies (fme- (Table 6) (Wilber and Simkiss, 1968; Gsegoire, grained vs. coarse-grained siliciclastics). They 1972;Wainwright et al., 1982). The periostracum found that unsuccessful and multiple drillholes and non-calcareous operculae are composed occurred more frequently in the Boreal province chiefly of conchiolin, while the nacreous layers than the Paratethys province; the same ftcies dso and other shell microstructures contain various included molluscs with different drilling quantities of conchioh. Thus, conchiolin has a very frequencies-with as much as a three-fold old history, putatively stemming from the oldest difference between sampIes collected in adjacent shelled mollusc in the Cambrian Period. What is sites from the same facies! They concluded puzzling, however, is that only a few goups of unequivocally that spatial variation should be molluscs+hiefly the freshwater bivalves (e-g., evaluated independently before any large-scale Margaritiferidae, Unionidae, Mutelidae), estuarine temporal bends are inferred for predation. Clearly to marine bivalves (Corbiculidae and Solenidae), multiple collections with emphasis on facies need and a few marine species-have conchiolin to be included in the temporal analysis of escalatory represented as separate sheets within their shells predation hypotheses. vaylor et al., 1969;Anderson, 1992;Harper, 1994). Shell ornamentation {spines).-Spine Conchioh, as a protein, is thought to form at a high development requires extra amounts of calcium metabolic cost to the organism-and, perhaps carbonate in seawater, and shallow tropical marine because of this, there appears to be an evolutionary waters meet this requirement (Nichol, 1945; tendency to lose conchiolin layers (references in Stanley, 1970). Spines have inspired varied Kardon, 1998). Thus, there must be some hypotheses concerning their function as evolutionaq reason for majntaining conchiolin in antipredatory architecture (Table 5). Few studies, molluscan shells despite its high metabolic cost of however, have focused on alternative hypotheses production (Table 6).It has long been hypothesized such as whether spines are phylogenetic legacies that the conchioltn sheets deterred predation by of shell building (as in cardiid bivalves), non-aptive ddhgmolIuscan predators, such naticid (Lewy consmctional artifacts, exaptations, or adaptations. and Samtleben, 1979) or muricid gastropods paylor, Few workers have endeavored to apply such 1970,1981), and most of the work done to test this philosophical rigor, and many have created adaptive hypothesis has focused on the corbulid bivalves scenarios. Another important possible function of (Fischer,, 1963; Lewy and Sanitleben, 1979; De spines could be to increase surface area for the Cauwer, 1985; Anderson et al., 1991; Anderson, settlement of epibionts, and for trapping debris that 1992; Harper, 1994). The corbulids (Family camoUnages the sheUs (Vance, 1978;Feifarek, 1987; Corbulidae) are small, inequivalved bivalves wid a Stone, 1998). Stone+(1998j has shown that spines globose shell form, a single byssus thread, and on epifaunal bivdves deter the attack of muricid shallow burrowing habits (e.g., Stanley, 1970).They shell-drilling predators, but muricids can still bore first appeared in the Middle Jurassic, with the in areas where spines are absent. h the same study, greatest diversification taking place in the spines were found not to deter predatoxy attacks by Cretaceous and Eocene (e.g., Hallam, 1976). sea stars that engulf their prey. The rise of spinose There appear to be three contrasting temporal ornamentation in bivalves predates the radiafion of "trends" related to whether conchroIin reduces the predatory Muricidae in the Albian, and actually predation. The first is that conchiolin does extends back to the late Paleozoic in the effectively reduce predation on corbulids through superfamily Pectinoidea (Stone, 1998). their evolutionary history. Fischer (1963), for WALKER AND BRETT-POST-PMEOZOIC PATTERNS IN MEPREDATION

TABLE 5-Hypotheses for the origin of skeletal spines in marine invertebrates.

Hypothesis Evidence Examples Reference

Spines deve7op in calcium Less energetically costly to make Spondyhs arnerkanus Stanley, 1970 carbonate supersaturated sea spines in tropical waters waters; most common in tropics

Spines are antipredatory Primary spines that project Spondylus americanus Logan, 1 974 outward may protect mantle edge

HoIIow spines and keels are for Economy of mass Ammonoids Birkelund, pelagidplanktonic existences 1981

Spines have no function Constructional artifact? Alternative hypothesis Carter, 1967; for any invertebrate Kauffrnan, -1 aca Spines function in filter-feeding Spines cover opening to animal Poricthophenid Rudwick, 1970 brachiopods

Spines are an ancestral Spines form in various ways, even Gardii-dbivalves; Schneider and condition; phylogenetic within closely related farniries anornalodesrnatans Carter, 2001 constraint

Spines vary with Various spine types depending on Spondylus arnericanus Logan, 1974 environmental conditions substrate the larvae zitach to (Jurassi+Recent) of the substrate Attachment to substrate Spines act as attachment Cemented bivalves like Logan, 1974 mechanisms Spondylus whose right valve is attached to substrate; this hypothesis does not function for the left vaive Spines discourage epibionts Spines and pedicel!aria in some Sea urchins; the bivalve Logan, 1974 echinoderms discourage biant Spondylus americanus settlement; perhaps barbed secondary spines of Spondylus americanus reduce biont settlement Spines acts as suppotts for - Brachiopods (Jurassic Rudwick, 1965; sensory mantle tissue; "mantle Acanthothirus ) Logan, 1974 outposts" to give early warning signals of danger

Spines serve a camouflagic Hair-like barbed spines typical of Late Paleozoic Grant, 1966; function, breaking up the neanic stage of the left valve of productoid brachiopod, Logan, 1974 distinctive outIine of shell Spondyius which get covered with Waagenoconcha; aLgae and sediment; spines are Spondylus americanus thickly encrusted with epiboints

Spines stabilize the shell on a The Cretaceous Logan, 1974; shifting substrate Spondy[us spinosr~s Carter, 1972 Spines keep the feeding Logan, 1974 margins of the shell above the substrate PALE0iVTOLOGIC.M SOCIETY PAPERS, V 8,2002

TABLE 6-Evolutionary findings concerning whether conchiolin serves an anti-predatory function against drilling predators for corbulid bivalves, Alternative hypotheses are discussed in text.

Evolutionary Evidence Reference conclusion

Adaljtation Conchiolin arises at the same time as Harper (1994) drilling predators in the Cretaceous Exaptation* Conchiolin arises in the Middle Jurassic, Kardon (1998) well before origin of drilling naticids; *however, no temporal trend in drilling predation

Not Anti-predatory No temporal trend in drilling predation Anderson et a!. (1 991); Anderson (1992) Phylogenetic Conclusions drawn from synthesis of This paper Constructional Artifact the literature -

example, suggested that Recent corbuLid species (1994) found that there was no ,significant were less Irkely to be completely drilled than fossil difference in possession of conchiolin sheets species. Kardon (1998, p. 73) also suggests that in between temperate and tropical localities. Given temporally and spatially separated fossil samples these contrasting findings, it is important to of corbulid bivalves, conchiolin layers are effective examine some of the salient evolutionary deterrents of naticid predation. The second is that hypotheses regarding conchiolin as an drilling predation actually increased in corbulids antipredatory deterrent, such as cost-benefit over their evolutionary history. For example, analyses and whether conchiolin is an adaptation although using a limited data set, Taylor et d. or exaptation (or neither) against predation. (1983) suggested that corbtilids showed enhanced Corbulids have small size and effective valve predation from the Late Cretaceous to Eocene. And armor (i.e., relatively thick valves with conchiolin lastly, others suggest that there is no spatial or sheaths), and thus, according to Kitchell et al.'s tempord trend in ddhgpredation In corbulids. (1981) cost-benefit model, would represent a high For example, in Late Cretaceous to Pliocene drilling investment with low benefits weCauwer, corbulid fossils from Europe and N'orth America, 1985). For example, Kelly (1988) found that De Cauwer (1985) found no trend toward increased predation on corbulids was lower than would be complete drilling was found. Similarly, for predicted by the cost-benefit model (but see Miocene to Pleistocene fossil corbulids from the Anderson, 1992). Yet corbulids are heavily drilled Dominican Republic and Florida, there appears to in many localities, and this may be aresult of their be no spatid or temporal pattern in complete or tendency to cluster, their shallow burrowing depths, incomplete drilling, strongly indicating that and their slug$shness @e Cauwer, 1985). Perhaps conchiolin layers are not effective deterrents to drillkg predators may mistakenly drill empty shells naticid predation (Anderson, 1992). Likewise, in the presence of chemical attractants in the Harper (1994) reported that there was no exhalant water of the corbulid associations sign5ciant Merence in drilling frequency among (Carriker, 1981; De Cauwer, 1985); or there may almost all geological samples examined Erom the be hydrodynamic and taphonomic reasons for the Cretaceous to Plio-Pleisotocene. Further, Harper preponderence of drilled. corbulids in some WALKER RRrD BRETT-POST-PALEOZOIC PATTERNS IiVhJAHNE PREDATION localities (De Cauwer, 1985). calcium carbonate-saturated regions, and have Anderson et al. (1991) tested the cost-benefit done so for most of their geologic history. model cf Kttchell et al. (1981), aEd showed that a The most promising line of research concerning corbulid bivalve (Varicorbula caloosae) was no conchiolin, however, stems from the finding of more likely to be drilled by a naticid predator than mechanical tests that the conchiolin in corbdids may by a venerid bivalve (Chione cancellata), among function to inhibit crack propagation, which in turn Pleistocene fossils £corn Florida. Anderson (1992) may be a deterrent to shell-cmshng predation examined many species of corbuiids from the (Kardon, 1998). It remains to be tested whether Miocene and Pliocene of the Dominican Republic conchioh layers do inhibit shell-crushing predators. and from the Pliocene and Fleistocene of Florida Biornechanical tests using corbulid bivalves, in and found that the incidence of drilled, addition to feeding experiments with live incompletely drilled, and multiply Medvalves dumphasous crustaceans, are needed to address chis was highly variable in space and time. This result hypothesis. An historical analysis of shell repair in was similar to other studies on drilled bivalves, and corbulids through time is warranted. therefore indicates that conchioh was generally Lastly, Kardon (1988) found that naticid not part of the antipredatory arsenal. Rather, drilling rates were not significantly slowed by alternative evolutionary hypotheses, such as conchiolin layers. Further, although Kardon (1998) conchiolin as a retardant of shell dissolution or a states that conchiolin has acted as an effective deterrent to crab-cmshing predation (Anderson, deterrent against &Xing predation in the corbulid 3992; Kardon, 19981, for example, need to be fossil record (p. 73, but see her p. 761, her data do advanced and tested. The main evolutionary not support this claim (p. 75, her table 2). Her results, question, as Harper (1994) pointed out in fact, support the findings of Anderson et al. (1991) (paraphrasing Gould and Vrba, 1982), is whether and Anderson (1992) that there is temporal variation conchiolin layers are a beneficial trait that is in drilling through time in corbulids, with no enhanced by natural selection (adaptation), or apparent trend. It would dso be important to how whether conchiolin layers are an exaptation, a from which facies these corbulids came, and whether beneficial trait that is secondarily co-opted for taphonomic (hydrodynamic or biotic) conditions another function. Experimental tisting is required affected their preservation. to detemhe if a trait is tmly beneficial; and there Although Kardon (1998 j suggests that should also be a temporal correspondencebetween conchioh is an exaptation, and Harper (1994) the evolution of the trait and the proposed selective suggests that concfiiolin is an adaptation, a review agent (Harper, 1994). of the data to date indicates that conchiolin may Accordingly, Kardpn (1998 j tested three be an artifact of construction. Of course, this hypotheses concerning the evolutionary importance statement needs to be refuted by scientific tests. of conchiolin: 1) it retards shell dissolution; 2) it That is, without further tests with shell-crushing increases shell strengh and thus deters crushing predators, we cannot know 5 conchiolin is indeed predation; and 3) it inhibites shell drilhg by naticid a beneficial trait, either co-opted or evolved by the gastropods. She also examined the fossil record of organism against predation. Other hypotheses were naticid drilling predators, and compared it to that discussed by Harper (1994), such as protection of conchiolin-bearing corbuiids (which hail from against nonpredatory borers or assistance with the Middle Jurassic; but see Harper, 1994) to hermetic sealing, and these could be rigorously examine the evolution of the trait in association tested as well (see Table 6). The oldest corbulids with its putative selective agent (the naticids). Her (Jurassic Covbulomima) were marine organisms, experimental results show that conchiolin did and had conehialin before the evolution of naticid retard shell dissolution, although-as she clearly drilling during the Early Cretaceous (Kardon, pointed out-the majority of corbulids live jn. 19981, further suggesting that conchiolin was not PUEONTOLOGICA L SOCIETY PAPERS, V 8,2002 evolved as a deterrent to drilling predation. marine crocodilians, ichthyosaurs, and plesiosauts. However, it would be very important to howthe Following a setback in the Late Triassic, predators environmental conditions of the origin and made a major re-advance in the mid-Mesozoic with diversification of corbulids; also, for those the evolution of new groups of decapods, corbulids with more than one conchiolin layer, ammonites, neogastropods, and teleost fishes, as whether they are from "physiologica1ly" more well as neoselachian sharks and marine reptiles. stressful environments, such as brackish water or Some of these groups are thought to have been anoxic environments. It would also be important durophagous, but that does not mean they ate ro how their cladistic relationships with respect exclusively molluscan prey. Limited data from ths to their environment and conchiolin form. time indicates that drilling predation existed, but occurred at low very frequencies. SUMMARY DISCUSSION: The Late Cretaceous saw unprecedented levels AN EPISODIC HISTORY of diversity of marine predaceous vertebrates OF PREDATfON including pliosaurs, plesiosaurs, and mosasaurs. The patCretaceous-Tertiary extinction decimated Predation in marine communities evolved marine reptiles. Drilling and shell peeling through several phases of intensification with frequencies pick up in the Late Cretaceous minor setbacks following mass extinctions correspondingto the evolution of new durophagous (Fig. 14). The Perm-Triassic extinctian crisis and shell-drilling groups. The drilling frequencies formed a major setback for all marine communities. from this time are no d-ifferent from those reported This certainly included &my predatory taxa (e.g., from Cenozoic localities; indeed, drilling an'd shell many ammonoids, nautiloids, phyllocarids, repair data from the later Cretaceous and Cenozoic predatory archeogastropods). However, certain show no apparent trends. marine predators, notably bony fishes and sharks, The Cretaceous-Tertiary mass extinction seem to have been less strongly affected by this eliminated all large marine predators, including the major extinction than were many benthic mosasaurs, plesiosaurs, and many sharks and fish. invertebrates (Knoll et al., 1996). Thus, predators Additionally, pterosaurs and early marine birds seem to have rebounded rather rapidly and by the were eliminated. However, many benthic Middle Triassic a variety of new predator guiIds invertebrate and fish predatoxy groups sunived; had appeared, including decapod crustaceans with and during the Paleogene, predatory benthic crushing claws, and shell-crushing sharks and bony invertebrates showed a spurt of evolution with fish. However, data from the Triassic regarding neogastropoda and new groups of decapods, while shell repair and drilling predation are almost non- the teleosts and neoselachian sharks both..underwent existent. New groups of carnivorous marine reptiles parallel rapid evolutioaary radiations; these were also appeared in the Triassic, including joined by new predatory guilds of sea birds and durophagous placodonts, and piscivorous and marine mamals. Ultimately, many of the large perhaps cephalopod-eating pachypleurosaurs, vembrate predator gdds were refdIed by newly nothosaurs, ichthyosaurs, and the fist plesiosaurs, evolved groups of marine mammals (cetaceans, Ceratite ammonoids and some marine reptiles pimipeds) and birds (gulls, albatrosses, penguins). (c.g., placodonts, nothosaurs j became extinct Despite the fact that a new suite of predators during Late Triassic crises. However, other Lineages evolved in the Cenozoic, there are no apparent (e.g., ammanites, ichthyosaurs, plesiosaurs) escalatory trends in durophagous predation. survived to form the stem groups for new Jurassic All of this would seem to suggest episodic, but radiations. The Jurassic to Early Cretaceous saw generally increasing predation pressure on marine the rise of malacostracan crustaceans with crushing organisms through the Mesozoic-Cenozoic chelae and predatory vertebrates-in particular, the interval. Theoretically, there should have been a WMKERAhTD BRETl-POST-PALEOZOIC PATTERNS IN MARWE PREDATION

1 Phases Extinctions 1 Mid Miocene

Late Eocene

Cret-Ted (K-T)

Cenomanian (c-T) Aptian

Tthonian

Plieins-Toarcian I L-Carnian (Late Trias}

Changxingian (P-T) Maokouan

Mid - Carb.

Hangenberg (end Dev.) Kellwasswer (Fr,- Fm.) Taghanic (Giv.- ~r.)

End Ordovician Ashgilt

Biomere Events

FIGURE 14-Summary diagram, showing phases of escalation in marine predator-prey systems and major extinction events. P~EONTOLOGICALSOCIETY PMERS, V 8,2002

Paleogene marine revolution in the molluscan crinoids in modern oceans; other clades may realm, because of the increased abundance of continuously evolve new predators, as Vermeij drilling neogastropods, the -first seal records of (1987) has argued based on the gastropod fossil durophagous stomatopods and decapod crustaceans, record. Schneider and Carter (2001) show that and the evolution of specializd bird and mammalian cardiid spine forms in Mesozoic and Cenozoic predators. However, most of these durophagous groups appear to be a Paleozoic ancestral condition, groups are generalists, and it may be that they had and appear not to be related to the putative a diffuse effect on thejx invertebrate prey. Mesozoic Marine Revolution. Thus, a clade-by- Finally, several new groups of carnivorous clade analysis of predation would be most useful, - marine mammals and birds originated in the as the different groups each have their own Miocene. Walruses, gray whales, and humans arose evolutionary histories and ecological constraints. in the Neogene and affected the coastal hard-shelled This review shows that not all morphology in biota in the areas whe.re they foraged or settled. Thus, benthic organisms need be directly related to a Neogene phase of further predator intensification predation. Additionally, matdurophagous predators is also suggested. However, there is no direct do not prey specifically on molluscs. They also prey evidence that prey were selectively affected (except on hard-shelled crustaceans, a major group of for the widespread decimation of species by humans organisms deemed to have caused selective pressure and their alteration of marine habitats). toward escalated armor in gastropods (Vermeij, The Cenozoic record seems to provide an 1987). We also must strive to examine predation in excellent window into predation and its effects, but assemblages spatially across different envixonments, few have examined the tehporal trends in predation mindful of taphonomic bias, if we are to' derive during this time (except for naticid rnoIluscan evolutionarily and paleoccologcaIly meaningful drillers; e-g.,Hansen et al., 1999; Kelley et al., 2001). interpretations. The Phanerozoic record of Given that many predators leave their signature on predation is there, but it has not been fully explored; shells and other prey, it is just a matter of re- it is especially important to consider multiple examining the fossil record with the specific intent worhg hypotheses about Phanerozoic predation as we seek to interpret this record. to look for predation. More work needs to be done Vermeij reviewed in this area, especially on drilling records from (1987) the record of rnolluscivorous predators, and their multifarious othcr gastropod groups, and on putative shcLl repair methods of predation in the Phanerozoic. He made records that allow a comparison of Paleogene with a plea for more data on the responses of prey species Neogene localities. Additionally, in the Cenozoic, in Mesozoic and Cenozoic assemblages (Vermeij, vast deep-sea (bathyd and deeper) fossil deposits 1987, p. 239).Fifteen years later, his plea still stands. of mollrtscs are well preserved in u~liftcdterraces in tectonically active regions of the world, allowing for comparisons of predation (shell repair, shell drillhg) between shallow benthic and deep sea We dedicate this second entree to Richard fossil assemblages (Walker and Voight, 1994; Bambach for his pioneefing work concerning Walker et d.,2002). seafood through time. We greatly appreciate Although escalation is sometimes cast as an M. Kowalewski and P. Kelley for commissioning ongoing "arms race,"' in actuality the predatory us to produce this synthesis. We thank L. Anderson, record shows episodes of abrupt biotic R. Feldmann, C. Hickman, A. Hoffmeister, reorgetion during and after mass extinctions, G. Storrs, H. Greene, 0. Keppel, J. S. Pearse, punctuating longer interludes of relative stability R. Portell, and J. Voight for their thoughtful (Brett et al., 1996). Some clades may retain the comments concerning numerous predation-related historical legacy of the Paleozoic predatory queries. A. Hofieister, W. Miller 111, and two revolutions, as could be argued for the stalked anonymous reviewers greatly improved the first WALKER AND BRETT-POST-PALEOZOIC PATTERNS IN AfARfNE PREDATION draft of this paper. E. Reitz graciously allowed for facilitating the international editing of this access to her zooarchaeology collections at the manuscript. This is not an exhaustive treatment, Georgia Museum of Natural History, Athens, and any predatory comments regarding this paper Georgia. We also wish to thank Ruth Mawson and need to be addressed to the fust author who will Peter Cockle, of Macquarie University, ~ustralia, bristle with delight!

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