Contributions to Zoology, 72 (2-3) 119-130 (2003)

SPB Academic Publishing bv, The Hague

Evolution of reef-associated decapod through time, with

particular reference to the Maastrichtian type area

René+H.B. Fraaije

Oertijdmuseum de Groene Poort, Bosscheweg 80, NL-5283 WB Boxtel, the Netherlands

Keywords: Decapod evolution, K/T boundary, biostratigraphy

Abstract prior to 1987 by various authors (Bosquet, 1854;

van Binkhorst, 1857; Binkhorst van den Binkhorst,

The result of of some twenty years intensive collecting from 1861; Noetling, 1881; Pelseneer, 1886;Forir, 1887a- in the strata Maastrichtian type area is a collection ofmore than c, 1889; Mulder, 1981) suffer from a lack of strati-

1,200 generally small-sized anomuranand brachyuran remains. graphic control. A taxonomic revision of most of The of the stratigraphical ranges thirty-one species known to these was carried out Collins al. date species by et (1995). from the Maastricht Formation (Late Maastrichtian) are

shown Since 1987, new from the Maastrichtian and five successive decapod assemblages are discussed. species

For the first time, crustacean to decapod remains now turn out type area were described and discussed by Fraaye be useful biostratigraphic tools on a local to regional scale. & Collins (1987), Feldmann et al. (1990), Jagt et

al. (1991, 1993), Collins et al. (1995), Fraaye (1996

a-c, 2002), Fraaye & van Bakel (1998) and Jagt et Introduction al. (2000).

Rigid collecting from six key sections (see Collins

Brachyurans utilize a broad of feeding types, array et al. 1995, p. 168, fig. 1) during the past two de- including deposit feeding, filter feeding, seaweed cades has resulted in an extensive, stratigraphically grazing, and have been scavenging predation. They well-documented, decapod, crustacean collection

a major of many marine communities component containing over 1,200 specimens and housed at the from the Late onwards and have probably Oertijdmuseum de .GroenePoort, Boxtel (the Neth-

played an role in the evolution of important ma- erlands). rine ecosystems. Unfortunately, they are only rarely described by paleontologists and are also frequently

overlooked by biologists in Recent marine com- Carapace size and morphology through time munities, especially in the tropics where reach

their & 1978). highest diversity (Zipser Vermeij, size is be related thought to to predation pres- Apart from paleoecological factors and possible sure (Vermeij, 1978). The relatively small size of destruction after death, their in apparent scarcity the of crabs majority Mesozoic was probably con- paleontological collections is often due probably trolled by predation pressure of simultaneously to their small sizes, and therefore the relatively reef teleost fish evolving (Vermeij, 1978) and co- chance of being overlooked in the field (Bishop, occurring cephalopods. Apart from size selection, •986; Plotnick et The size of al., 1990). average predation pressure can also lead to avoidance strat- anomuran and brachyuran crab remains in the fine- egies, i.e., strategies devised to minimize the risks grained sediments of the Maastrichtian area type of For envis- being predated upon. crabs, we may 18 frss than 10 mm. Almost all decapod crustacean age four strategies that play a continuous role dur- species from the Maastricht Formation described ing theirevolution: 1 - to hide and live in crevices;

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- 2 3 - burrow into loose camouflage their carapace; during which a slower and stabilized evolutionary

4 - swim in sediments; open waters. pattern dominated.This model with pulses of rapid

in the first is Carapace morphology group gen- diversification, mixing and more stable periods, in somewhat erally quadratic or elongate. Such crabs roughly 30 million year cycles, perfectly matches have been successful from the Late Jurassic the on- polyphyletic origin of such groups as the swim- wards and have retained crabs. After each consistently a relatively ming period, however, more and

small size. The vast area of shelf seas with an ex- more ecological niches were successfully filled,

of redundant bioherms after the Callovian the pansion leading to recent, most diverse (in size, mor- transgression offered various new ecological niches phology and feeding strategy) decapod crustacean and led to the diversification of rapid prosopids fauna ever seen on earth.

(Forster, 1985). The other three strategies, which

probably evolved during the , are char- acterized by a considerable size increase through Cretaceous/Ccnozoic decapod crustacean time. The strongly elongate and burrowing Raninidae evolution

( crabs) are examples of the third group. They

in appear the Early Cretaceous and show an in- The Late Cretaceous diversification of crabs coin-

in size crease and diversity through time, with max- cides with two similar radiations in other preda-

ima the Late Cretaceous (Feldmann et al., teleost and during tory groups, fish gastropods (Taylor,

and Eocene (Forster & Mundlos, 1982). The These 1996) 1981). groups also seem to have thrived during Raninidae lived in shallow-water predominantly periods of transgression with maximum (semi-iso- environments the Mesozoic but during today are, lated) shallow-water habitats. The rapid evolution due to the Ceno- probably strong competition during of these predatory groups and others probably had zoic, effects mainly deep-water organisms. profound upon the structure of all benthic The of the ancestors Recent swimming crabs communities, and thus also affected crab evolution.

that are wider than long and with To the possess carapaces put evolutionary patterns of decapod crus- in antero-lateral spines. They appeared the latest in the Maastrichtian in taceans type some perspec- Cretaceous the the (Fraaye, 1996a). During Eocene, tive, faunal data for five other well-studied regions crabs their largest swimming quadrupled carapace and ages are summarized in Fig. 1. As the majority

size in their Maastrichtian comparison to probable of the type Maastrichtian decapods is linked with

It is not mere coincidence that ancestors. periods reef-associated carbonates all decapod crustacean of increase in size and of diversity crabs are linked faunas compared are from similar environments. with sea level stands et all global high (Haq ah, 1987). Further, faunas represent the most diverse of

radiations crabs Major explosive adaptive their as known. Studies among age presently used are Bishop occurred the Late the during Jurassic, -Ceno- (1983, 1986: Lower Albian, Texas), Muller& Col- the manian, Campanian-Maastrichtian, and during lins (1991: Upper Eocene, Flungary), Muller (1984: the Eocene and Miocene These transgressions. pe- Middle Miocene, Hungary and Austria), Guinot

riods are characterized stands by high leading to (1985: Recent, French Polynesia) and personal ob-

many shallow, more or less isolated, seas. Conse- servations (Lower Kimmeridgian, southern Ger-

crabs quently, more ecospace and new niches for many). came into existence. In isolated diver- seas, rapid Brachyuran crab diversity increased in time (Fig.

sification of clades was probably very common. 1), implying a strong evolutionary radiation from The clades this of monophyletic produced by type the Late Jurassic onwards. It is also clear that cer- have event recently been referred to as tain ‘species brachyuran groups played a more prominent flocks’ (Yacobucci, 1996). After sea level fall, newly role in the shallow marine ecosystems than they

evolved crab forced populations were to retreat and do currently. For instance, the Dynomenidae and

mixing led to in turn strong competition, possibly Raninidae first appeared during the Late Jurassic

leading to extinction and faunal turnovers (Fraaye, and Early Cretaceous, respectively. After a rapid

1996b). This was followed of by periods stasis, evolutionary radiation during the Late Cretaceous,

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Fig. I. Reef-associated decapod crustacean faunas through time.

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2. crustacean from Fig. Stratigraphic ranges of decapod genera Maastrichtian type area.

they dramatically declined from the Eocene onwards. cessful group, however, is that of the Xanthidae.

hold forthe their The same may true Calappidae, although Having originated during the Early Cretaceous, to a somewhat lesser extent. The decline of the evolutionary adaptations resulted in the by far most

Raninidae possibly corresponds to the synchronous diverse group of brachyurans in reef-associated biotic radiation of clypeasteroid echinoids and sub- faunas.

niche and the sequent competition and replacement. Decapod crustaceans are, probably were,

Other such the food for groups, as Portunidae, Cancridae, most important source cephalopods (Fraaye

Majidae, Leucosiidae, Parthenopidae and Grapsidae, & Jager, 1996; Jager & Fraaye, 1997). The preda-

tion however, display a rapid evolutionary radiation in pressure on crabs by relatively fast swimming post-Eocene times, as do the Ocypodidae and Pilum- and hunting ammonites such as Placenticeras and

in nidae from the Miocene onwards. The most suc- Sphenodiscus the Campanian and Maastrichtian

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era which apparently do not cross the K/T bound-

of them in in ary some reality may occur younger

For deposits. two genera, Stephanometopon and

Aulacopodia, generic classification and subsequent

possible evolutionary offspring is uncertain at the

moment. The extinction of Graptocarcinus and

possibly Glyptodynomene seems to correspond to

a niche-displacement by the apparently better adapt-

ed Dromiopsis spp. (Fraaye, 1996b).

Of the fifteen genera known from the Danian of

Denmark and first in the Sweden, ten appear Cre- 3. Fig. Number of decapod crustacean species and percentages taceous (Collins & Jakobsen, 1994). for Percentages the Middle Danian of Fakse (Denmark). of faunal elements in the Middle Danian of Fakse,

based upon field observations, are shown in Fig. 3. may have led to the introduction of the swimming The successful crossing of the K/T boundary and xanthids (Bishop, 1991; Fraaye, 1996a, 1997). In persistance in Recent faunas of the Calappidae, their turn and these well-adapted swimming crabs found Carpiliidae Xanthidae probably are the result a new and abundant food source in planktonic am- of their predatory feeding adaptations, evolved monites such the Late as Scaphites spp. (Westermann, 1996). during Cretaceous, especially the special- In the ization Maastrichtian of the Vistula River valley of the chelae for breaking and crushing mol-

(central a of luscan shells & Poland) very high percentage speci- (Taylor, 1981; Zipser Vermeij, 1978) mens of constrictus have and the of Such Hoploscaphites paired, development teleplanic larvae. a more or less regular punctures in their body relatively long planktonic larval phase is known in chambers, assumed to be the result of decapod crus- tropical Recent representatives of these families, tacean but is attacks (Radwahski, 1996; Fraaye, 1997). rare or absent in taxa inhabiting cooler seas Identical holes in the same place of the body cham- (Vermeij, 1978). bers are known from the Upper Cretaceous of USA

(N.H. Landman, pers. comm.).

The known of all Substrate stratigraphic ranges genera and decapod diversity in the present in the Maastrichtian type area are presented Maastriehtian type area

m Fig. 2. Of the twenty-three anomuran and bra- chyuran genera known to date from this area, ten Percentages for decapod families are plotted and

first in appear this interval, ten do not extend be- shown separately for the Emael, Nekum and Meers- yond the K/T boundary, whereas thirteen pass this sen members in Fig. 4. From the base of the Emael unscathed, one of survives to Member the which, Paguristes, up section, Paguridae (hermit crabs) Ihis day, and two, Cretachlorodius and Leptoides show a clear increase in number and diversity, di- gave rise forms. to Recent Danian decapod crusta- rectly correlated with an increased availability of cean faunas rather are poorly known, partly because their ‘mobile homes’, empty gastropod shells. In of few accessible for instance the relatively outcrops (as Emael Member, the Raninidae and Calappidae

m the In region studied here) and partly because of dominate. the Nekum and Meerssen members, luck of collection effort. for the fauna of the remain Except Calappidae a more or less constant, Denmark and Sweden (Collins & Jakobsen, 1994), important component whereas the Raninidae be- and to a lesser degree those from Greenland (Collins come proportionally less dominant. The Callianas- & Rasmussen, 1992), sidae is the commonest in the Nekum Argentina (Feldmann et ah, group Member,

1995) and Antarctica the (Feldmann et ah, 1993), but rapidly decline in the overlying member. Tory-

base decapod crustacean data for the Danian is rather nommids reached their acme in the Nekum Mem-

with meager, overall poor biostratigraphic control. ber and rapidly declined in the lower Meerssen

Therefore, it is be the to expected that of ten gen- Member, probably as a result of competition with

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Fig. 4. Percentages for decapod crustacean families in the three highest members of the Maastricht Formation in the type area.

efficient swimmers of fields of the the presumably more amongst currence seagrass during deposition

latter Meerssen the Xanthidae (Fraaye, 1997); some of these uppermost Emael, Nekum and lower mem- represent Tethyan invaders (Fraaye, 1996a). The bers (Liebau, 1978; Felder et ah, 1980; Voigt, 1981).

the Dynomenidae and Flomolidae display the same trend The sea grasses enabled colonisation of small

as pagurids going up-section from the Nekum to epizoans such as bryozoans (Voigt, 1981), fora-

the Meerssen members. The Galatheidae and Car- minifera (Sprechmann, 1981) and calcareous algae

I iidae make their first in the Meerssen Sediments which accumulate around pi appearance (Brasier, 1975).

communities Member (Fig. 5), but are far outnumbered by other sea grass are distinctive and, if they

families. Within the interval studied, five succes- escape channeling and other forms of associated

be erosion within the stand chance sive decapod assemblages may recognised, as biotope, they a high

follows of Sea is (in ascending order): preservation (Brasier, 1975). grass no-

(I) raninid table for its ability to influence the character of

(II) raninid/diogenid/calappid the sediment substrate. The dense plant growth

(III) callianassid/calappid/torynommid probably reduced current velocities, whereas the

(IV) calappid/xanthid/raninid rhizomes stabilised the accumulated sediments. In-

(V) calappid/dynomenid faunal filter feeders, e.g., Protocallianassafaujasi,

The diversity increase between the Emael and Meers- thrived in the nutrient-enriched sediments around

sen members is intimately linked with an increase sea grasses, making use of things such as decaying of substrate diversity (Fig. 6). Accumulations of plant material (Sven et ah, 2001). The accumulated

dead shell material section. The dead hard of the facili- grew markedly up parts sea grass-epibionts

tated other entire spectrum of live/dead interactions (ecologi- the colonization of epizoans such as fungi,

of in cal consequences shell accumulation) benthic sponges, corals, brachiopods, serpulids, lunulitiform

communities was referred to as ‘taphonomic feed- bryozoans, boring and encrusting bivalves, sessile back’ by Kidwell & Jablonski (1983). The change gastropods and cirripedes. In the lower Meerssen

from a predominantly soft-bottom dwelling com- Member(units IVf-1 to -4) this colonization finally

munity in the lower Emael Member (and underly- led to the development of small-sized bioherms,

ing members), to the firm ground and shell-gravel vertically and laterally alternating with hard grounds

dwelling communities in the Nekum and Meerssen (Voigt, 1974; van den Elsen, 1985) and relief in-

in members, was probably triggered by the mass oc- fillings (Zijlstra, 1995). Cavities hardgrounds

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5. Fig. chart in the Maastrichtian Decapod crustacean species range type area.

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Fig. 6. Major palaeoenvironmental factors in the Maastrichtian type area which influenced decapod crustacean communities,

played an important role in Late Maastrichtian eco- the larvae to settle in deeper habitats and migrate

systems as refugia and domiciles (Voigt, 1959). inshore at larger sizes. The infrequent availability

The number and size of holes and crevices in (caused by storm disturbance and occupation by

reefsubstrates is positively correlated with the num- competitors) of holes and adult-juvenile predatory ber and size of decapod crabs and shrimps (Reaka- interactions was probably important in the evolu-

These Kudla, 1990). decapod crustaceans each prey tion of decapod crustaceans in reef communities their on own and each others’ juveniles, forcing over time.

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Abele documented that the num- of (1974, 1982) fragility decapod crustacean carapaces, in con- ber of decapods in a habitat is a function of the trast to some of their claw fragments, would rule structural number of of out and in complexity (= substrates) any reworking. Therefore, crabs, par- that habitat The author ticular same (Abele, 1976, 1982) raninids preserved in situ, are amongst the also concluded that habitats comparable support best groups suited to document evolutionary rela- similar numbers of decapods, regardless of size of tionships within the Emael, Nekum and Meerssen the in overall available species pool. Referring to this members the type area. Naturally, the excellent hypothesis, the habitat of the Middle Danian fauna preservation of remains decapod crustacean may of Fakse that be (Denmark) most closely approximates attributed further to the low organic content and of the Nekum Member in the Maastricht area. The the high pH of the warm-water depositional envi- occurrence of Raniliformis baltica in both settings ronment, which would buffer the effects of acids

(Danian of Den- Limburg [Geulhem Member] and produced by soft-tissue decay (Plotnick et ak, 1988). mark) might suggest a similar depositional envi- A second bloom of Raninoides quadrispinosus in ronment as well. The crab fauna also the lower points to Meerssen Member, without striking dif- environmental similarities between the Nekum and ferences in carapace morphology in comparison to

Geulhem members in the Maastricht an ob- material from the Gronsveld and area, Valkenburg mem- servation backed crinoid by data (J.W.M. Jagt, pers. bers, shows this species to have been the least af-

The difference fected comm.). most important between by changes in substrate and depth within the the environment of the Nekum and Maastrichtian. depositional type Thus far, large-sized and highly

Geulhem a members is substantial drop in tempera- ornamented specimens ofEumorphocorystes sculp- indicated ture, by the drastic decline of bioherms tus are only known from the Nekum and underly- and all the the bottom-dwelling groups, with ing members, whereas in Meerssen Member a

of echinoids possible exception (Felder, 1981; van significantly smaller sized and but partially orna- den Elsen, This of mented 1986). drop temperature already morphotype occurs. More specimens are started in the upper halfof the Meerssen Member needed to determine whether these different mor-

(sections IVf-5 are and -6) where decapod crustacean photypes ontogenetic or phylogenetic (van Bakel remains are extremely rare or absent. et ah, work under way).

Zijlstra (1995) documented storm-induced sedi- mentation cycles in the type Maastrichtian carbon- ates. The of highest degree sedimentary, e.g., trough Acknowledgements cross beds, channels, spillover and tempestite sheets, and orientation I wish to thank to J.W.M. taphonomic features, e.g., parallel Jagt (Maastricht), G. van der Zwaan ot F. Schram the serpulid Pyrgolopon and other elongated (Utrecht), (Amsterdam) and J. Meulenkarap (Utrecht)

for their of this suggestions leading to improvement paper. T. bioclasts, indicating high energy and storm-gener- van Hinte prepared the figures. This is contribution to IGCP ated a deposits, occur in the middle Meerssen Mem- project 362, ‘Tethyan and Boreal Cretaceous’. ber (sections IVf-3 and -4). Norris (1986) showed that amongst crabs the preservation potential pro- duced by storm-induced burial is high. relatively References This is supported by the near-uniform ‘upside-down’

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