Decapod Diversity in the Mid-Cretaceous of Northern Spain

Decapod diversity in the mid-Cretaceous of northern Spain

Report Molengraaff Fonds – grant to Adiël Klompmaker, 2010

Introduction

During the late Albian – early Cenomanian (102-97 Ma) interval both sea level and global temperatures were high (Haq et al., 1988; Larson, 1991; Scott, 1995), while oceanic spreading rates peaked (Larson, 1991). Hence, epicontinental seas expanded markedly and the number of reefs increased worldwide (Kiessling, 2002). Reefal deposits assigned to the Albeniz Unit of the Eguino Formation (López-Horgue et al., 1996), of late Albian – early Cenomanian age, are found near Alsasua, western Navarra (northern Spain; Fig. 1). This unit overlies upper Albian siliclastic sediments and consists of two stages, documenting firstly the establishment of carbonates on a ramp and, secondly, the formation of patch reefs with marly deposits in between (López-Horgue et al., 1996). Six patch reefs have been observed over an area of some 30 square kilometers (Fig. 2).

Fig. 1. A, Location of Spain in Europe. B, Location of the study area in northern Spain. C, A detailed map of the study area with the location of the Koskobilo and the Monte Orobe quarries, the latter in which a contemporaneous decapod fauna was found in the past.

From these levels, numerous decapod crustaceans have already been described, in particular from the Monte Orobe patch reef (Van Straelen, 1940, 1944; Ruiz de Gaona, 1943; Via Boada, 1981, 1982; Gómez-Alba, 1989; López-Horgue et al., 1996; Fraaije et al., 2008). At a nearby locality in the Aldoirar reef, which is composed mainly of scleractinian corals, algae and a few rudistid bivalves, fieldwork was conducted during three consecutive years (2008-2010) at the disused Koskobilo quarry, from which

Fraaije et al. (2009) had previously recorded a paguroid anomuran. Klompmaker et al. (2011a, see attachment) recorded the first gastrodorid from the Cretaceous Period (Gastrodorus cretahispanicus), recognized in assemblages collected at this locality, and Klompmaker et al. (2011b, see attachment) added Rathbunopon obesum.

Fig. 2. An overview of the patch reefs in the Alsasua area during the late Albian – early Cenomanian. The Koskobilo quarry is located in the southeastern part of the Aldoirar patch reef. Modified after López- Horgue et al. (1996, fig. 3).

The purpose of this report is to provide information on the entire decapod fauna from Koskobilo, and its diversity compared to other localities in the world consisting of Cretaceous sediment yielding a rich decapod fauna. Moreover, sites within the quarry are compared to one another based on the finds of the author in the summer of 2010.

Diversity in Koskobilo

The total number of specimens from the Koskobilo quarry is approximately 1000 (collected in the summers of 2008-2010), of which about 250 were collected by the author during the fieldtrip in 2010. Taken together the decapod fauna of Koskobilo consists of 37 species in this single locality, many of which are new to science (see Table 1). Although not all published yet (Klompmaker et al., in prep.), it appears that the Koskobilo quarry is going to yield about twenty new species in total, and also several new genera. Numerous species have also been discovered in recent coral reefs. Abele (1974) mentioned 55 decapod species in a coral environment with six substrates, and Abele (1976) noted 55 and 37 decapod species for two localities off the Pacific coast of Panama.

Table 1. The list of decapod species known from the Kokskobilo and Monte Orobe quarries. Koskobilo Monte Orobe 1 "Xanthosia" n. sp. Annieporcellana dhondtae Fraaije et al., 2008 2 Annuntidiogenes worfi Fraaije et al., 2009 Annuntidiogenes ruizdegaonai Fraaije et al., 2008 3 Caloxanthus n. sp. Distefania incerta (Bell, 1863) 4 Cretatrizocheles (n. gen.) n. sp. Distefania 'transiens' (Wright and Collins, 1972) 5 Distefania n. sp1 Eodromites grandis (Von Meyer, 1857) 6 Distefania n. sp2 Eomunidopsis navarrensis (Van Straelen, 1940) 7 Distefania incerta (Bell, 1863) Eomunidopsis orobensis (Ruiz de Gaona, 1943)

8 Eodromites grandis (Von Meyer, 1857) Etyxanthosia fossa (Wright & Collins, 1972) 9 Eomunidopsis n. sp. Glyptodynomene alsasuensis Van Straelen, 1944 10 Eomunidopsis navarrensis (Van Straelen, 1940) Goniodromites laevis (Van Straelen, 1940) 11 Eomunidopsis orobensis (Ruiz de Gaona, 1943) Graptocarcinus texanus Roemer, 1887 12 Etyxanthosia fossa (Wright & Collins, 1972) Homolopsis edwardsii Bell, 1863 13 galatheoid Navarradromites (n. gen.) n. sp. 14 Gastrodorus cretahispanicus Klompmaker et al. 2011 Navarrahomola (n. gen.) n. sp. 15 Glyptodynomene alsasuensis Van Straelen, 1944 Necrocarcinus labeschei (Eudes-Deslongchamps, 1835) 16 Goniodromites laevis (Van Straelen, 1940) Paragalathea multisquamata Via Boada, 1981 17 Graptocarcinus texanus Roemer, 1887 Paragalathea ruizi (Van Straelen, 1940) 18 Hispanigalathea (n. gen.) n. sp. Paragalathea straeleni (Ruiz de Gaona, 1943) 19 Hispanigalathea n. sp. Pithonoton bouvieri Van Straelen, 1944 20 Laeviprosopon n. sp1 Rathbunopon obesum (Van Straelen, 1944) 21 Laeviprosopon n. sp2 Sabellidromites scarabaea (Wright and Wright, 1950) 22 Laeviprosopon n. sp3 Viaia (n. gen.) n. sp. 23 Laeviprosopon n. sp4 Xanthosia cf. 'X. similis' (Bell, 1863) (=X. fossa?) 24 Mesotylaspis (n. gen.) n. sp. 25 Navarracaris (n. gen.) n. sp. 26 Navarradromites (n. gen.) n. sp. 27 Navarrahomola (n. gen.) n. sp. 28 Nykteripteryx (n. gen.) n. sp. 29 Paragalathea multisquamata Via Boada, 1981 30 Paragalathea ruizi (Van Straelen, 1940) 31 Paragalathea straeleni (Ruiz de Gaona, 1943) 32 Pithonoton bouvieri Van Straelen, 1944 33 Priscinachus n. sp1 34 Priscinachus n. sp2 35 Rathbunopon obesum (Van Straelen, 1944) 36 Torynomma n. sp. 37 Viaia (n. gen.) n. sp.

Stratigraphic and biogeographic notes

The Koskobilo fauna sheds light on the transition from Late Jurassic decapod faunas to Cretaceous faunas. Gastrodorus has only been found in Upper Jurassic sediments previously, but is now also known from the mid-Cretaceous. Surprisingly, Eodromites grandis was hitherto known only from the Late Jurassic, but is now also found in Koskobilo. Specimens of this species also represent the youngest specimens from this genus. The same applies for the four species of Laeviprosopon. Striking similarities exist between the decapod fauna from Koskobilo and the Paleocene (Danian) Fakse fauna from Denmark, possibly due to a similar depositional environment containing corals. Examples include Caloxanthus, Xanthosia, and some galatheids.

Many genera are new to Iberia (Spain and Portugal) and have not been encountered in Monte Orobe or were misidentified from Monte Orobe. These include (obviously) all new genera, Gastrodorus, Caloxanthus, Priscinachus and Torynomma.

Koskobilo versus Monte Orobe

In comparison to Monte Orobe, a patch reef in a small abandoned quarry some 4 km north of Koskobilo, the diversity in Koskobilo is markedly higher. This may be due to several factors: a) the smaller species were recognized in Koskobilo, b) possibly more collecting time in Koskobilo based on the number of specimens from Monte Orobe in the Museo Geológico del Seminario de Barcelona in comparison with the collection from Koskobilo, although a part of the collection of the Museo Geológico del Seminario de Barcelona was loaned and not returned (P. Artal, pers. comm. 2008) or c) Koskobilo contained several sites within the quarry yielding slightly different decapod faunas, possibly related to a different microenvironment, see below. Another observation is that some of the decapods appear larger in Monte Orobe (e.g. Distefania incerta and Eodromites grandis), and more decapods are preserved with a cuticle instead of internal molds in Koskobilo.

Diversity in the Cretaceous

This decapod diversity is unique for the Cretaceous. In fact, Koskobilo will become the richest decapod locality in the world most likely with 37 species. Table 2 shows an overview of other decapod rich localities in the world yielding at least ten species. As can be seen, the ENCI quarry in the Netherlands is second with 31 species. More species are planned to be described from this quarry though (Fraaije, pers. comm., 2011), so this locality might rival Koskobilo in the future, if no additional species will be discovered from the Koskobilo quarry. Both localities have in common that they were a coral reef. Table 3 shows the richest decapod formations from the Cretaceous with at least ten species. The Eguino Formation which includes the Koskobilo and Monte Orobe localities tops this list as well with 42 species followed by the Maastricht Formation.

An interesting question is whether the peak in diversity is caused by a preservational bias or is due to ecological reasons (i.e. a natural high number of species within fossil coral reefs). A preservational bias may be considered likely as sedimentation rates are generally high in carbonate environments. Tucker (1990: p. 33-34) mentioned that ‘carbonate sediments can accumulate rapidly compared with other sedimentary rock types’. He notes sedimentation rates in the order of 0.1-6 millimeters per year for carbonate systems depending on the position within a reef. Schlager (1999) mentioned sedimentation rates from the same order of magnitude. The preservation of the decapods from Koskobilo is, however, by no means very good, which may be expected with high sedimentation rates. For example, the vast majority of the carapaces are broken and complete carapaces are uncommon. Moreover, not a single complete specimen is found with the venter, abdomen, and appendages still attached. Even though it is expected that the majority of the material exhibits a moldic preservation, at least some complete specimens would be expected. Apparently, there was sufficient time to allow for degradation and disassociation of dead decapods either due to biological or mechanical causes. Thus, sedimentation rates may be high compared to other sedimentary environments, but it was insufficient to cause exquisite preservation of the decapod fauna.

A high number of decapods are also known from recent reefal environments. Abele (1974) found that decapods were especially diverse in a subtidal Pocillopora coral community and in a rocky intertidal habitat in Panama in comparison to numerous other sedimentary environments including several beaches (sand-mud), a marsh, and mangroves. One type of substrate within this rocky intertidal habitat comprised Porites corals. Furthermore, there appears to be a positive correlation between the coral head size and the number of decapod species and individuals that inhabited the coral head (Abele, 1976). In addition, it is noted that the number of decapod species increased with increasing complexity of the habitat; that is, with more substrates in a certain habitat (Abele, 1974). A coral reef exhibits many different microenvironments, and has, in addition, different parts such as the reef front, reef flat and back reef. A reef would be an ideal place for many decapods to live, they use the reef for shelter, as a feeding site, and as a source of nutrition (Abele, 1974). In conclusion, the peak in diversity in Koskobilo is ecological in nature and not so much preservational.

For the stratigraphy of British formation and localities, Benchley & Rawson (2006) is followed. New species that are just listed but have not appeared in the formal literature yet after many years, are not taken into account in the species count in the tables below. A specimen of which the genus can be determined, but not the species (sp.) is only counted when this is the first specimen of the genus from the locality or formation. Otherwise, it might represent a poorly preserved specimen of a species of the same genus that is also present in the same locality or formation. Hence, duplication is avoided here.

Table 2. Decapod rich localities from the Cretaceous. # Localities Formation Age Rock Unit Country species Koskobilo quarry Eguino Formation Albian-Cenomanian Spain 37 ENCI quarry Maastricht Formation Maastrichtian The Netherlands 31 Petréval Craie glauconieuse Formation lower Cenomanian France 26 Monte Orobe quarry Eguino Formation Albian-Cenomanian Spain 23 Hakel Unknown lower-middle Cenomanian Lebanon 20 Hadjoula Unknown lower-middle Cenomanian Lebanon 20 Atherfield, Isle of Wright* Atherfield Clay Formation lower Aptian Great Britain 16 White Hart Pit, Wilmington, Devon West Melbury Marly Chalk Formation lower Cenomanian Great Britain 15 Folkstone, Kent Gault Formation middle-upper Albian Great Britain 15 Cambridge Upper Greensand Formation upper Albian Great Britain 15 Pargny-sur-Saulx Formation des Argiles tégulines de Courcelles lower-middle Albian France 12 Blue Springs Coon Creek Formation Maastrichtian Mississippi, USA 11 Sahel Alma Unknown Santonian Lebanon 11 Bemelen Maastricht Formation Maastrichtian The Netherlands 10 Berg en Terblijt Maastricht Formation Maastrichtian The Netherlands 10 locality 10, James Ross Island Santa Marta Formation ?upper Santonian-Campanian Antarctica 10

Grayson County, 2 mi N of Denison, 0.5 mi E of Kde type locality on S side St. Louis-San Fransisco Railway track Denton Formation Albian Texas, USA 10 South Dakota, Heart Tail Ranch Pierre Shale Formation Campanian USA 10 Bully Gault Formation middle-upper Albian Great Britain 10

* Four new species have been mentioned by Simpson (1985), but they have not been formally described yet, and, thus, have not been included in the count for this formation.

Table 3. Decapod rich formations from the Cretaceous. # Formation Age Rock Unit Location species Eguino Formation* Albian-Cenomanian Spain 42 Maastricht Formation** Maastrichtian The Netherlands 39 Upper Greensand Formation upper Albian Great Britain 33 Hakel & Hadjula & Maifuk (not official fm) lower-middle Cenomanian Lebanon 30 West Melbury Marly Chalk Formation lower Cenomanian Great Britain 27 Craie glauconieuse Formation lower Cenomanian France 26 Pierre Shale Formation Campanian South Dakota, USA 25 Pawpaw Formation upper Albian-lower Cenomanian Texas, USA 20 Merchantville Formation lower Campanian New Jersey, USA 20 Gault Formation middle-upper Albian Great Britain 19 Coon Creek Formation Maastrichtian Mississippi/Tennessee, USA 17 Denton Formation Albian Texas, USA 17 Atherfield Clay Formation*** lower Aptian Great Britain 17 Formation des Argiles tégulines de Courcelles lower-middle Albian France 16 Santa Marta Formation ?upper Santonian-Campanian Antarctica 14 Sahel Alma (not official fm)**** Santonian Lebanon 11 Pender Formation lower Campanian British Columbia, Canada 11 Fossil Bluff “Formation”***** Berriasian-Albian Antarctica 10 Glen Rose Formation upper Aptian- lower Albian Texas, USA 10 Formation Gault lower-middle Albian France 10 Calcaires à Spatangues Formation****** lower Hauterivian France 10 * Obviously, not all the species from Koskobilo have been published yet, but this is planned to be submitted at the end of 2011 or otherwise in 2012 depending on some other publications that need to be published first. ** Jagt et al. (2000) mentioned seven new species from the Maastricht Formation in a conference abstract. These have, however, not been described after eleven years and are not incorporated into the species count. The disassociated claws Aulacopodia riemsdyki Bosquet, 1854 and Pseudomicippe granulosa Pelseneer, 1886 could be of numerous carapaces (see Jagt et al., 2000), and, thus, have not been counted here. *** Four new species have been mentioned by Simpson (1985), but they have not been formally described yet, and, thus, have not been included in the count for this formation. **** Drobna n. sp1 and sp2 Roger, 1946 have not been incorporated in the count. Garassino (1994) citing Ejel & Dubertret (1966) mentioned that the Shel Alma deposits represent the upper part of the Globotruncana concavata zone (Santonian). On the other hand, Garassino & Schweigert (2006: p. 70) mentioned the deposits at Sahel Alma to be Senonian in age (Turonian- Maastrichtian) based on the same source. They (p. 71), however, mentioned Cretasergestes sahelalmaensis from Sahel Alma to be Cenomanian in age. I follow Garassino (1994) for the age assignment as Hallett (2002: p.189) mentioned that Globotruncana concavata is an index microfossil for the Santonian. ***** Sensu Taylor et al. (1979). The Fossil Bluff Formation has later been subdivided into several formations (see Fig2: Miller & MacDonald, 2004). However, the exact position of the decapods within the new formations cannot be determined. Therefore, I retain the Fossil Bluff “Formation” here. ****** A. Milne Edwards (1865: 341-347) reports on two new species from the neocomien of Yonne, one of which from the middle Neocomien, which might be the Calcaires à Spatangues Formation. Since this is not certain, those species have not been tallied here. Additional remark: Gérard Breton (pers. comm. April 2011) suspects that the 30 species (among which many synonymies exist, see Van Straelen, 1936 and Schweitzer et al., 2010) reported by Robineau-Desvoidy (1849) from the Neocomian of Saint- Sauveur-En-Puisaye (Yonne) originate from the Calcaire à Spatagues Formation. Since Robineau-Desvoidy (1849) does not

6 report on a particular formation or stage within the Cretaceous, I refrain from including them in my overview as also the Valanginian Calcaire de Bernouil Fm, the lower Barremian Argiles ostréennes Fm, and the Upper Barremian Sables & Argiles panachées Fm crop out in Puisaye, a district within the Yonne department in France.

Sites within the Koskobilo quarry

Several sites within the quarry were discovered during field work in the summers of 2008 and 2009 (Fig. 3). Sites 1-4 yielded a comparatively rich decapod fauna compared to other places in the quarry. This seemed to be related to the amount of biogenic debris/material that could be discovered as more visible debris would generally mean a higher chance of finding decapods. At site 5, much biogenic debris was found as well, suggesting that a high number of decapod specimens could be present as well. This was, however, not the case at all based on field observations in 2009.

In 2010 I collected decapods for 9 hours at sites 1-4, and 2 hours at site 5 because this site was not yielding many decapods and because of time constraints. Collecting was performed in such a way that there was an about equal hours of collecting in the sunshine at every site (4 hours sun rest cloudy/in shadow for sites 1 and 4 and 3.5 hours in the sun and the rest cloudy/in the shadow for sites 2 and 3). This is because personal observations in 2008 and 2009 suggested that more decapods could be discovered in bright sunlight. After every one or several hours, I changed sites. For comparison among sites, I selected only those specimens with the central portion of the cervical groove (the ‘main’ groove on decapods) present to avoid a bias caused by preferential breakage of certain species. Preferential breakage causes numerous fragments to form. Counting all these fragments would imply that certain species that preferentially break would be overrepresented. By counting only those specimens with the axial point of the cervical groove present, this bias can be avoided. This central part is suitable because a cervical groove is present in virtually all decapods, decapods have only one cervical groove, and the central part is less likely to be degraded in comparison to other more distal parts of the carapace. In paguroids, the carapace often is less calcified toward the posterior carapace. In other words, the choice for the axial part of the cervical groove, usually located somewhere halfway the carapace, is ambiguous. Moreover, carapace terminology differs from other decapods (see Fraaije et al., 2008: Fig. 2). In this particular case, the presence of >50% of the transversal groove on the anterior carapace serves as the criterion whether or not to include a paguroid specimen into the count.

Fig. 3. The Koskobilo quarry with five sites indicated. Modified source: Google Earth

As can be seen from Table 4, the number of specimens and the number of species differs significantly. Most species and specimens are found at site 2, whereas site 4 yields the lowest number of specimens and site 4 yields the lowest number of species based on sites 1-4 only. Two hours were spent collecting at site 5 in comparison to nine at sites 1-4. It is very likely that site 5 would have yielded the lowest number of specimens and species, had I collected more hours there. This was also suggested by a survey of all sites in 2009.

For eveness the Simpson Index is used (formula given below, where n = the total number of organisms of a particular species and N = the total number of organisms of all species). Hence, a larger D means a lower diversity. D = (n / N)2

For site 1 it is 0.18, for site 2 it is 0.16, for site 3 it is 0.14, and for site 4 it is 0.21. This means that eveness is about equal for every site. However, the species that are dominant per site differ.

Table 4. The number of specimens found per species per site, as well as the total number of specimens and species per site. Site 1 Site 2 Site 3 Site 4 Site 5 Annuntidiogenes worfi 1 Caloxanthus n. sp. 3 2 2 3 Edodromites grandis + Navarradromites n. sp. 3 7 1 Eomunidopsis navarrensis 8 23 9 26 Eomunidopsis orobensis 6 1 1 Gastrodorus cretahispanicus 1 Glyptodynomene alsasuensis 1 2 Goniodromites laevis 22 10 1 6 Graptocarcinus texanus 3 2 10 10 Viaia n. sp. 1 Navarrahomola n. sp. 1 Laeviprosopon spp. 2 2 1 2 Indet. 1 2 Navarracaris n. sp. 1 paguroid 1 8 2 Distefania incerta 2 1 3 3 Distefania sp. 4 1 5 Paragalathea ruizi 2 20 4 7 1 Galatheid 1 Pithonoton bouveri 2 2 1 Rathbunopon obesum 1 1 1 "Xanthosia" n. sp. 1

Number of specimens 59 85 40 66 1 Number of species 15 17 14 13 1

The reason for a difference in the number of species and specimens per site may be related to microenvironments within the reef. In exploring the hypothesized microenvironments within the mid- Cretaceous Koskobilo patch reef in Spain, I made five thin sections. The thin sections were described and compared, and photos were taken of the various fossils found within the thin sections. For each hand sample and the accompanying thin section a Folk and Modified Dunham name was determined.

In addition, the geopetal fabrics in the thin section were identified and an attempt was made to determine the microfacies. In both cases Flügel (2004) was used. To determine the microfacies, the following questions were answered (Flügel, 2004: p. 580), with the exception of question 10, which I added.

1) Which fossil groups are represented by the skeletal grains? 2) Which fossil groups are abundant, common, and rare?

3) If calcareous algae or foraminifera occur, which major systematic units (e.g., dasyclad green algae or corallinacean red algae; orbitolinid or nummulitinid foraminifera) are present? 4) Are there differences in preservation of the skeletal grains? 5) Are the sizes of the skeletal grains within comparable ranges or are some conspicuously larger or smaller than the average grains? 6) Do the skeletal grains represent benthic (mobile or sessile) or planktonic organisms? 7) Do the grains exhibit biogenic encrustations? 8) Which fossil groups appear commonly associated (e.g. radiolarians and sponge spicules, or echinoderms and bivalves)? Are there striking grain associations (that might point to specific paleoclimatic conditions?) 9) Are the fossil concentrations allochtonous or autochtonous? 10) What is the general relative size of the fragments?

The results from the Folk (1959, 1962) and modified Dunham (Embry & Clovan, 1971, after Dunham 1962) classifications are listed in Table 5. All carbonate rocks were fossiliferous as the names imply. There was only one definite framework of an in situ coral community (site 2). Since the cement has been recrystallized into (sub)equant, small calcitic crystals, I chose to biosparite in many cases.

Table 5. The Folk and Dunham names for the carbonate rocks. Folk modified Dunham 2 biolithite boundstone (framestone) 5 unsorted biosparite floatstone in TS (rudstone in hand sample) 1 poorly washed biosparite packstone 4 unsorted biosparite rudstone 3 unsorted biosparite packstone

General thin section description per thin section

The fossils immediately identified in the thin section (TS) of site 2 were colonial scleractinian corals (Plate 1) and a cross-sectional view of a gastropod (Plate 2). It was noted that many parts of the hand sample were recrystallized as the corals suddenly faded out halfway the individual polyps. Other fossils that were found are bryozoan, foraminifers (forams), a possible annelid, red algae, and an echinoderm fragment (Plate 2). Corals are by far the largest fossil in this thin section. Even though this, the diversity is not low; the evenness is low because of the coral dominance. The coral are interpreted to be in situ based on field observations. There are several blobs with biolithic hash. Especially the coral structures are recrystallized. Peloids are present, but not as much as in TS 1.

The only way to tell what is up or down is by noting several transverse cross-section through the several coral. This means that you either look at the bottom or at the top (see Plate 1). No other geopetal fabrics were found.

The thin section 5 contained mainly red algae (Plate 3) and few large biolithic structures, so the eveness is fairly low. The red algae, in itself small (mm scale), make up the largest fragments in this thin section with the exception of one large algal blob. Other fossils include forams, echinoids, possible ostracods, and a possible holothurian (Plate 4). The overall diversity can be considered to be moderate/high. Lots

10 of recrystallization had taken place. No peloids were observed indicating that bioturbation was absent to very limited in this environment.

In terms of geopetal fabrics (see p. 180 in Flügel, 2004; Fig. 4 for all thin sections with the exception of the first one), ‘sandfang’ and the deposition type (in this case on red algae) were present. There also was alignment of the larger red algae fragments. However, smaller fragment were not aligned in one part of the thin section. The one large algal fragment (see Figure 3C) showed encrustation by possibly a bryozoan colony. One the other side there is accumulation of debris indicating that the algae had a different orientation in the latter case. The geopetal structures do line up, so up and down could be interpreted.

TS 1 exhibits many small biolithic fragments as well. There is one large coralline fragment. Because the TS consists of small fragments of organisms, identification is hard. The fossils that have been encountered are bivalves, bryozoa, echinoid fragments, forams, algae, and corals (Plate 5). The diversity is fairly high as well as eveness. Recrystallization is limited, possibly because this TS is heavily pelletized (so lots of bioturbation), also compared to the TSs described above.

In this TS no particular geopetal fabric could be found, possibly due to the high degree of reworking due to bioturbation, which is evidenced by the large number of fecal pellets.

TS 4 contained many orbitolinid forams as the largest fossils (up to 4 mm long, Plate 6). In addition, bivalves, forams, echinoid fragments, bryozoa, a possible ostracod, possible brachiopods, and a coral fragment were found (Plate 7). The shell fragment (either brachiopods or bivalves) are relatively large as well. The fossils in the TS are relatively diverse and have an intermediate eveness. Again, lost of recrystallization is observed. Only few peloids were observed, some of which occur in patches.

Also in this case, sandfang and deposition on larger biolithic fragments does occur. Sandfang does occur in bivalve shells, while deposition is observed on orbitolinids. In addition, internal geopetals were found inside some shells that may have been bivalves. There is also one case of possible encrustation. The shells and orbitolinids are oriented in the same general direction, although convex up criterion could not be used in this case. All the geopetal structures point to one direction.

The TS 3 contains very small biolithic fragments compared to the other TSs. There are only two blobs of larger algal fragments. The fossils include green, red and charophytic algae, forams, bryozoa, echinoid fragments, and a bivalve fragment (Plate 8). Its diversity is not extremely high; eveness is high as there is no fossil that is dominant. There are some dark spots in which small biolithic fragments have accumulated. Recrystallization is very prominent again. There are only very few peloids present indicating that bioturbation was present but rare.

The geopetal fabrics were helpful in determining what was up and down. Deposition on biolithic fragments was observed for bryozoa and algae. One case of sandfang was observed. Almost all the large elongated biolithic fragments were oriented in the same way, suggesting a preferred orientation. Some fragments were, however, way off. Possibly some reworking was at work. A pattern in convex up was not found.

Fig. 4. A) Thin section 5: deposition on red algae, B) TS 5: sandfang on a red algae, C) TS 5: possible bryozoan encrustation on algae, D) TS 4: deposition on shells, E) TS 4: possible bryozoan encrustation on recrystallized shell fragment, F) TS 4: sandfang in shell, G) TS 4: sandfang and internal geopetal fabric, H) TS 3: massive deposition on bryozoan fragment, I) TS 3: sandfang in shell.

Microfacies

Hereunder are the answers to the questions for the microfacies analysis per sample (Table 6-10):

Table 6 TS 2 colonial scleractinian corals, gastropod, bryozoan, forams, a 1 possible annelid, red algae, and an echinoderm fragment 2 abundant: corals; rest: common/rare 3 mainly red algae 4 no 5 corals far larger than rest 6 mainly benthic sessile (corals + red algae) 7 no 8 not much evidence 9 corals autochtonous; rest possibly allochtonous 10 very large and small

Table 7 TS 5 red algae, green algae, forams, echinoids, bivalve (rare), and 1 possible ostracods, possible holothurians 2 abundant: red algae; rest: common/rare 3 red algae 4 no 5 variety in sizes red algae 6 benthic sessile 7 one grain only 8 no 9 red algae partly autochtonous; rest likely allochtonous 10 large and small

Table 8 TS 1 bivalves, bryozoa, echinoid fragments, forams, algae, and 1 corals, (pellets) 2 all common/rare 3 hard to tell yes, some shells are 'ghosts', one very well preserved and 4 some are intermediate yes, with with exception of one large corraline fragment and 5 one shell fragment 6 mostly benthic (mobile and sessile) 7 no

8 - 9 shells mostly autochtonous, most allochtonous 10 small mainly

Table 9 TS 4 orbitolinids, bivalves, forams, echinoid fragments, bryozoa, a 1 possible ostracod, possible brachiopods, and a coral fragment 2 abundant: orbitolinids; rest: common/rare 3 orbitolinids 4 yes, within shells 5 difference in shell sizes 6 benthic (mainly sessile if orbitolids are sessile indeed) 7 one encrustation, possibly formed ex-situ 8 - 9 orbitolinids autochtonous; most shells allochtonous 10 fairly large, some smaller

Table 10 TS 3 green, red and charophytic algae, forams, bryozoa, echinoid fragments, 1 and a bivalve 2 all common/rare 3 planktonic forams & red algae 4 yes, shells mostly ghosts, some better preserved 5 lots of small debris and larger shells 6 more benthic (both mobile and sessile) than fair number of planktonic 7 no 8 - 9 shells mostly autochtonous; rest possibly allochtonous 10 small

Microfacies analysis – Depositional environment

The five samples studied might represent five different microfacies/environments. The most obvious ones are sites 2 and 5. They differ in the main fossil (corals versus red algae). Also, these corals are found to be in situ in the quarry while a part of the red algae might be ex situ. The diversity was also higher in the case of site 2, possibly because site 5 may have inhibited a higher diversity because of its structure having less cavities compared to corals. The Folk and Dunham names differ as well. The distinction of these two microfacies is in line with the decapod fauna (see Table 4). Possibly the

14 decapods were not able to cope with an environment consisting of mainly red algae. They certainly would have had fewer places to hide.

Site 2 would have been more likely a part of the actual reef itself, possibly part of the upper part of the reef front, where many branching corals can be found usually. Alternatively, it might have been a local patch of corals near the reef talus. The algae level might be more part of the reef flat, with lower energy, although more research is needed in this case.

The other three thin sections (and thus sites) are more alike in terms of Folk and Dunham names. They are also closely spaced geographically (within 50 meters in map view), while site 2 is hundreds of meters away. They also have in common that they all seem to have at least two types of preservation based on the shells. This suggests some transport. They do, however, differ markedly in major aspects. Site 4 is the only thin section that exhibits many orbitolinid forams, while the TS from site 3 yields more planktonic forams as compare to all the other TSs and, moreover, exhibits the smallest biolithic grains overall. Site 1 differs from the other ones by the presence of many fecal pellets, suggesting an actively bioturbating community, which is corroborated by the fact that no geopetal fabric could be found within this thin section. In terms of decapod fauna, sites 1, 3, and 4 are to a certain extend comparable in terms of number of specimens and specimens. They are also geographically closest. The decapod fauna differs from site 2 in that Paragalathea ruizi, the group of Edodromites grandis + Navarradromites n. sp. and paguroids are more abundant at site 2. In general, an overall pattern appears where decapods may be smaller on average at site 2. More research is necessary to confirm this. Sites 1, 3, and 4 probably represent different parts of the reef talus, because of the overall smaller, broken grains in comparison with site 2, suggesting transport. The constant supply of food (detrital material) might have provoked an active benthic life community leaving behind fecal pellets at site 1. Except for the orbitolinids at site 4, sites 1 and 4 are alike in terms of their microfacies.

Future research

Subsequent research will focus on the completion of a huge manuscript on the decapods from Koskobilo. The hope is that this can be submitted at the end of 2011 or in the beginning of 2012. (This depends on the speed with which other manuscripts, critical for this manuscript, are going to be submitted and accepted. Furthermore, I intend to make more thin sections of the several sites within the quarry to increase the sample size. Lastly, new taxa from Koskobilo will be integrated into the existing database of Drs. Feldmann and Schweitzer (my advisors at Kent State University) to generate the Mesozoic decapod diversity curves. I hypothesize that the diversity of Mesozoic decapod crustaceans (crabs, lobsters, shrimp etc.) is related to Mesozoic reef diversity. This is in line with data on the preferred habitats of extant decapods, which are coral reefs with their many microhabitats (Abele, 1974, 1976).

Attachments

Attachment 1: KLOMPMAKER, A.A., P. ARTAL, R.H.B. FRAAIJE, AND J.W.M. JAGT. 2011. Revision of the family Gastrodoridae (Crustacea, Decapoda), with description of the first species from the Cretaceous. Journal of Paleontology, 85 (2):226-233.

Attachment 2: KLOMPMAKER, A.A., P. ARTAL, AND G. GULISANO. 2011. The Cretaceous crab Rathbunopon: revision, a new species and new localities. Neues Jahrbuch für Geologie and Paläontologie Abhandlungen, 260 (2):191-202.

Attachment 3: Accepted article for Palaeontology. KLOMPMAKER, A.A., P. ARTAL, B.W.M. VAN BAKEL, R.H.B. FRAAIJE, AND J.W.M. JAGT. Etyid crabs (Crustacea, Decapoda) from mid-Cretaceous reefal strata of Navarra, northern Spain.

Furthermore, I presented a poster at a regional meeting of the Geological Society of America: http://gsa.confex.com/gsa/2011NE/finalprogram/abstract_186213.htm The poster can be downloaded at: http://www.personal.kent.edu/~aklompma/pittsburghklompmaker.pdf

References:

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Plate 1. Corals from site 2. Note that B is the same as A, but then under crossed polarized light. It shows that recrystallization was prominent in this TS.

Plate 2. Several fossils from site 2. A) foram, B) echinoderm fragment, C) gastropod, D) possible annelid, E) red algae, and F) possible algae.

Plate 3. Several examples of red algae from the site 5.

Plate 4. Several fossils from site 5. A) green algae, B) foram, C) echinoid, D) bivalve, E) possible ostracod, F) possible holothurian.

Plate 5. Several fossils from site 1. A) bivalve, B) bryozoans, C) echinoderm, D) possible foram, E) red algae encusting an unrecognizable, recrystallized fossil fragment, F) coral fragment. Note the abundance of fecal pellets in the pictures.

Plate 6. Orbitolinid forams from site 4.

Plate 7. Several fossils from site 4. A) bivalve and possible ostracod hash, B) foram, C) echinoid fragment, D) bryozoan, E) possible brachiopod, F) coral fragment.

Plate 8. Several fossils from site 3. A) red algae and possibly green algae, B) charophytic algae, C) possible planktonic forams, D) bryozoa (encrusted), E) echinoid, F) bivalve.