OCCURRENCE OF THE REGULAR URCHIN EUCIDARIS TRIBULOIDES FROM THE TAMIAMI FORMATION (PLIOCENE) OF FLORIDA
ROGER W. PORTELL FLORIDA MUSEUM OF NATURAL HISTORY GAINESVILLE, FLORIDA and CRAIG W. OYEN GEORGIA SOUTHERN UNNERSITY STATESBORO, GEORGIA The echinoids of the Pliocene Tamiami and the echinoids, Echinocardium goth Formation of Florida are typically abun icum (Ravenel, 1848) and Encope tami dant, well-preserved, and reasonably well amiensis Mansfield, 1932. Numerous known. In a 1963 monograph, Kier listed pectens and oysters were also collected. All nine species of echinoids from the specimens from UF locality CH026 were Tamiami Formation. These were Arbacia collected in a gray, medium- to fine crenulata Kier, 1963, Lytechinus variega grained quartz sand with a modest tus plurituberculatus Kier, 1963, Clype amount of heavy mineral content. No arag aster.crassus Kier, 1963, C. sunnilanden onitic-shelled taxa were preserved, only sis Kier, 1963, Encope tamiamiensis calcitic-shelled taxa typically associated Mansfield, 1932, E. michelini imperforata with the Tamiami Formation were present Kier, 1963, Mellita aclinensis Kier, 1963, (see taxa above). At UF locality CH026, Rhyncholampas evergladensis (Mansfield, the Tamiami Formation is overlain by a 2 1932), and Echinocardium gothicum meter thick shell bed attributable to the (Ravenel, 1848). Phelan (1972) eliminated Fort Thompson Formation. previous confusion between E. michelini L. Other fossil occurrences of E. tribuloides Agassiz, 1841, and E. aberrans Martens, are known from outside Florida. Donovan 1867, and re-identified Kier's E. michelini and Embden (1996) summarized t h e imperforata as E. aberrans. Kier (1992) Jamaican occurrences, including speci agreed with Phelan's specific assessment mens from the Plio-Pleistocene August but still considered his earlier (1963) sub Town Formation, early Pleistocene specific designation to be valid. Since Kier Manchioneal Formation and Old Pera (1963), no new additions to the echinoid Beds, and late Pleistocene Port Morant fauna of the Tamiami Formation have and Falmouth formations. Elsewher e, been reported, until now. Lewis and Donovan (1991) reported radi Between 1989 and 1996, fossil collec oles from the Pliocene and late Pleistocene tions were made by the Invertebrate of Tobago. Cutress (1980) confirmed sever Paleontology staff and volunteers of the al published reports from the Plio Florida Museum of Natural History Pleistocene Playa Grande Formation, and (FLMNH) at University of Florida (UF) Pleistocene Abisinia, Cerro Gato, an d locality CH026 (also known as the Tortuga formations of Venezuela. Cutress HandyPhil Pit) in western Charlotte also discounted reports by Jackson (1922) County, Florida (text-figure 1). These col and Sanchez Roig (1949) of E. tribuloides lections yielded hundreds of isolated regu from the Miocene (?) of Cuba. lar urchin test plates and radioles and Extant E. tribuloides occurs from Cape three partial, but flattened, regular urchin Hatteras, North Carolina, east to tests (UF 39528, UF 60203, and UF Bermuda, throughout the Caribbean, the 72022). Recent analysis of these urchin Gulf of Mexico, and south to Rio de remains ind.icates that they belong to Janeiro, Brazil (Serafy, 1979). Its known Eucidaris tribuloides (Lamarck, 1816), a depth range has been recorded between 0 genus and species not previously recorded and 800 meters. However, it is most com from the Florida fossil record. Other com monly found at depths of less than 50 mon taxa collected at this locality included meters (Serafy, 1979). Eucidaris tribu the brachiopod, Glottidia inexpectans loides inhabit rocky areas, typically under Olsson, 1914; the gastropods, Dicathais rocks and in small crevices. They are also handgenae Portell and Vokes, 1992, and known to inhabit seagrass beds (Hendler Ecphora quadricostata (Say, 1824); the et al., 1995). barnacle, Tamiosoma advena Zullo, 1992; 99 100 Tulane Studies in Geology and Paleontology Vol. 30
cules only modestly larger than surrounding N secondary tubercles. Ambulacra are narrow, slightly sinuous, and widest at ambitus. Poriferous zones in minor, sunken grooves. Pores round to slightly ellipti t cal, separated by a low, interporal partition. Ambulacra approximately one-fourth as wide as interambulacra areas. Interambulacra plates up to two times wider HANDYPHIL PIT CHARLOTIE than tall. Interradial tract about two times (CH026) COUNTY wider than adradial tract. Secondary tubercles densely distributed throughout plate surface. Interradial sutur~ simple, non-serrate. · Primary radioles have limited shape varia tion, but include both cylindrical and truncate 20 40 km forms. Shaft may be slightly inflated in proxi mal to medial portion. Acetabulum diameter · Text Figure 1. Map showing locality of approximately 44-61 % of radiole width. the· first reported occurrence of the fossil Radioles up to 42.9 mm long, and 3.5 to 4.5 mm Eucidaris tribuloides (Lamarck, 1816) wide. Milled ring is located 2.0 to 2.7 mm above from Florida. b-ase, and collar is 1.6 to 2.4 = long with coni cal shape widest at milled ring. Neck is narrow ACKNOWLEDGMENTS and located 3.5 to 4.6 mm above base. Shaft has numerous longitudinal series of low nodules, We are indebted to Phillip Whisler of which may not be visible on more poorly pre Gainesville, Florida for his assistance in served or altered samples. Distal end ofradioles the field, and particularly, for collecting marked by low-ribbed crown, with a central and donating fossil E. tribuloides speci prominence. mens UF 72022 and UF 60203. Jimmy The three flattened tests (UF 72022, UF and Pat Philman, Englewood, Florida, 60203, 39528) are incomplete and compacted; kindly allowed access to their shell pit (UF therefore, not all diagnostic features are well locality CH026). Burchard Carter, Georgia preserved. More detailed species descriptions Southwestern State University and are available in Weisbord (1969), Phelan (1970), Stephen Donovan, University of the West Cutress (1980), Donovan (1993), and Hendler et Indies reviewed earlier drafts of the man al. (1995). uscript and provided helpful comments. This is University of Florida Contribution Discussion: Comparison of the fossil to Paleontology 486. specimens from the Tamiami Formation with several modern E. tribuloides speci mens from the Florida Keys show most SYSTEMATIC PALEONTOLOGY morphologic· characteristics to be similar among the samples. The Florida fossils Class ECHINOIDEA Leske, 1778 also are comparable with E. tribuloides Order CIDAROIDA Claus, 1880 described by Donovan (1993) from the Family CIDARIDAE Gray, 1825 Falmouth Formation (Pleistocene) in Genus EUCIDARIS Doderlein, 1887 Jamaica, as well as samples of this species EuCIDARIS TRIBULOIDES (Lamarck, 1816) reported by Weisbord (1969) from the Plate 1, figures la-b, 2, 3 Playa Grande Formation of Venezuela (Plio-Pleistocene). Many of Weisbord's Description: Test typically flattened both Venezuelan samples are now reposited in adapically and adorally. Primary tubercles non the Florida Museum of Natural History crenulate, perforate, having a conical boss near and, therefore, were available for compari ly circular in outline, and grading into scro son by the authors. bicule without a basal terrace. Areoles circular Cutress (1980) compared samples of fos at ambitus, becoming transversely elongate sil and modern E. tribuloides collected near apical and oral regions. Scrobicular tuber- from sites in the Caribbean and Gulf of
; No.2 Tamiami Formation Eucidaris 101 Eucidaris tribuloides Biometrics Summary
I. Modem Eucidaris tribuioides
Number of Plates Mean of Medial Area Mean of Areole Measured Width (MAW) MAW/ArW Width CArW) Ratio Mean n=lS X=2.48mm ir-6.18 mm X=0.40
II. Fossil Eucldaris tribuloides (famiami Formation, Pliocene)
Number of Plates Mean of Medial Area Mean of Areole MAW/ArW Measured Width (MAW) Width CArW> Ratio Mean n=20 x=4.32mm x=7.32 mm X=0.58
ArW MAW
Text Figure 2. Summary statistics for single-plate measurements of medial area width (MAW), areole width (ArW), and medial area width I areole width ratio (MAW I ArW). Measurements are in mm, and were obtained from locations indicated on the test plate illustration and discussed in the text. (Illustration modified from Cutress, 1980). Mexico region with several fossil and/or ratio of 0.51 for E. clavata and 0.97 for E. modem species of Eucidaris, including E. tribuloid es, while the ApD/P erD mean madrugensis (Sanchez Roig, 1949), E. ratio was 0. 77 for E. clavata and 1.02 for thouarsii (Valenciennes, 1846), and E. E. tribuloides. clauata Mortensen, 1928. Cutress reported Eucidaris thouarsii, a Recent species several key differences between E . tribu from the eastern Pacific Ocean, has wider loides and each of these species using both median areas on interambulacral plates qualitative and quantitative (biometric) and wider associated areoles .. Data accu characteristics, and we have summarized mulated by Chesher (1972) for the ratio of these differences here. Eucidaris clavata, MAW/ArW of E. tribuloides shows a ratio a modern species known only from range of 0.6-0.8, while on equivalent-sized Ascension and St. Helena Islands in the specimens of E. thouarsii, Cutress (1980) South Atlantic, has fewer interambulacral calculated the mean ratio to be slightly plates per column, contains more conflu lower, 0.58, over a range of 0.4-0. 78. An ent areoles, and lacks fan-shaped septal additional biometric ratio, areole width bundles. Biometric analysis of the two (ArW) to the width of a single interambu species completed by Pawson (1978) lacral plate (IApiPlW) also supports a sep revealed significant differences in the aration of the species, with the ratio of the median area width (MAW) of ArW/IAmPlW mean ratio for E. thouarsii the interambulacral plates to the associat equal to 0.68 whereas for E. tribuloides ed areole width (ArW), as well as in the the mean ratio is equal to 0.58, ratio of the apical system diameter (ApD) Finally, comparisons with E. madrugen to the peristome diameter (PerD) for the sis are important, though the characters of species. He reported a mean MAW/ArW this species are somewhat difficult to deci- 102 Tulane Studies in Geology and Paleontology Vol.30
pher from the literature. Jackson (1922) only one of the two plates and its width is and Sanchez Roig (1949) reported .cidaroid considered. Since the sum of two single. specimens from the Miocene of Cuba, plate median area widths is not equal to which Cutress subsequently reassigned to the "entire" median area width deter E. madrugensis. Sanchez Roig (1949) mined from two plates, one cannot simply caused confusion in the process of naming extrapolate data from one set to the other. the species, by using both the names A second problem in examining the data is Dorocidaris madrugensis (in the text) and distinguishing from which region on the Leiocidaris madrugensis (in the figure plates a median area width was mea caption) for the same series of figures in sured. Each interambulacral plate is pen his plates. Furthermore, Cutress (1980) tagonal in outline, but the apex of the pen· determined through examination of the tagon is asymmetrical, so measurements figured specimens that not all of the speci will vary unless specific endpoints are mens were actually eucidarids, but were identified for the median area width posi Cretaceous phyllacanthid species instead. tion. No specific diagrams were provided Cutress determined that E. madrugensis in Pawson (1978) or Cutress (1980) to is closely related (likely the ancestral illustrate exactly where their measure· form) to E . tribuloides, but differs by hav ments are located on the test plates, mak ing whorled radioles in adults, a larger ing replication of equivalent measure areole width, and having radiole collars ments on the Tamiami Formation speci· that are only half as long as in E. tribu mens illustrated in Plate I less certain. loides. In addition, Donovan and Paul (in Text-figure 2 shows summary data and press) described the presence of low spin calculations for MAW, ArW, and ules 9n the primary radioles of E. madru MAW/ArW ratios (using single-plate mea· ge ns is whereas primary radioles of E. surements) for a specimen of E. tribu· tribuloides do not possess such spinules. loides from the Florida Keys and for sever· A cautionary note must be made regard al FLMNH samples of the fossil E. tribu· ing the biometric data gathered and their loides reported herein. In addition, text· use in the ·diagnostic ratios described figure 2 illustrates the exact location mea· above. Cutress (1980) pointec;l out that in sured on each plate, for each of the bio most of her measurements of median area metric variables, to help alleviate any con width (MAW) and calculations of the fusion in how our ratios were determined. MAW/ArW ratios, she used only single It is interesting to note that the plate measurements. Since most of her MAW/ArW mean ratio for the Florida fos work dealt with fossil cidaroids rather sils is 0.58 while the modern specimen has than modern specimens, she recognized a mean ratio of 0.40. The implications of that imperfect preservation of the fossils this may be that the fossil material is not often resulted in incomplete skeletal from E . tribuloides, but rather a new remains to identify and measure. This is species. Pawson (1978) believed such bio· in contrast to Pawson (1978) and metric ratios important enough. to justify Mortensen (1928), where they determined specific differentiation between E. clavata the usual, "entire" median area measure and E. tribuloides, so such variation ments from the pair of associated and between the modern and fossil samples articulated interambulacral plates. The used in this study may warrant further difference between these methods will examination and interpretation if more cause a different ratio . to result, because complete specimens are recovered in the
PLATE 1
Figures 1-3. Eucidaris tribuloides (Lamarck, 1816) . la-b. UF 72022. a. Oral surface of flattened test showing partially exposed lantern. b. Aboral surface of flattened test. Both XL ,2. UF 60203. Lateral view of flattened test. XL .3. UF'80300. Radiole. X2. No. 2 Tamiami Formation Eucidaris 103
1a
2
3
PLATE 1 102 Tulane Studies in Geology and Paleontology Vol. 30
pher from the literature. Jackson (1922) only one of the two plates and its width is and Sanchez Roig (1949) reported cidaroid considered. Since the sum of two single specimens from the. Miocene of Cuba, plate median area widths is not equal to which Cutress subsequently reassigned to the "entire" median area width deter E. madrugensis. Sanchez Roig (1949) mined from two plates, one cannot simply caused confusion in the process of naming extrapolate data from one set to the other. the species, by using both the names A second problem in examining the data is Dorocidaris madrugensis (in the text) and distinguishing from which region on the Leiocidaris madrugensis (in the figure plates a median area width was mea caption) for the same series of figures in sured. Each interambulacral plate is pen his plates. Furthermore, Cutress (1980) tagonal in outline, but the apex of the pen determined through examination of the tagon is asymmetrical, so measurements figured specimens that not all of the speci will vary unless specific endpoints are mens were actually eucidarids, but were identified for the median area width posi Cretaceous phyllacanthid species instead. tion. No specific diagrams were provided Cutress determined that E. madrugensis in Pawson (1978) or Cutress (1980) to is closely related (likely the ancestral illustrate exactly where their measure form) to E. tribuloides, but differs by hav ments are located on the test plates, mak ing whorled radioles in adults, a larger ing r.eplication of equivalent measure areole width, and having radiole collars mep.ts on the Tamiami Formation speci that are only half as long as in E. tribu mens illustrated in Plate 1 less certain. loides. In addition, Donovan and Paul (in Text-figure 2 shows summary data and press) described the presence of low spin calculations for MAW, ArW, and ules 9n the primary radioles of E. madru MAW/ArW ratios (using single-plate mea gensis whereas primary radioles of E. surements) for a specimen of E. tribu tribuloides do not possess such spinules. loides from the Florida Keys and for sever A cautionary note must be made regard al FLMNH samples of the fossil E. tribu ing the biometric data gathered and their loides reported herein. In addition, text use in the -diagnostic ratios described figure 2 illustrates the exact location mea above. Cutress (1980) pointe<;l out that in sured on each plate, for each of the bio most of her measurements of median area metric variables, to help alleviate any con width (MAW) and calculations of the fusion in how our ratios were determined. MAW/ArW ratios, she used only single It is interesting to note that the plate measurements. Since most of her MAW/ArW mean ratio for the Florida fos work dealt with fossil cidaroids rather sils is 0.58 while the modern specimen has than modern specimens, she recognized a mean ratio of 0.40. The implications of that imperfect preservation of the fossils this may be that the fossil material is not often resulted in incomplete skeletal from E. tribuloides, but rather a new remains to identify and measure. This is species. Pawson (1978) believed such bio in contrast to Pawson (1978) and metric ratios important enough. to justify Mortensen (1928), where they determin'ed specific differentiation between E. clavata ·the usual, "entire" median area measure and E. tribuloides, so such variation ments from the pair of associated and between the modern and fossil samples articulated interambulacral plates. The used in this study may warrant- further difference between these methods will examination and interpretation if more cause a different ratio. to result, because complete specimens are recovered in the
PLATE 1
Figures 1-3. Eucidaris tribuloides (Lamarck, 1816) . la-b. VF 72022. a. Oral surface of flattened test showing partially exposed lantern. b. Aboral surface of flattened test. Both Xl. .2. VF 60203. Lateral view of flattened test. Xl. -3. VF"80300. Radiole. X2. Tamiami Formation Eucidaris No. 2 103
11
2
3
PLATE 1 104 Tulane Studies in Geology and Paleontology Vol. 30
future. Unfortunately, the fossils currently Paleontology, v. 70, no. 3, pp. 485-493. available are not articulated and have DONOVAN, S.K., and C.R.C. PAUL, in press, been flattened during fossilization, pre Echinoderms of the Pliocene Bowden shell venting a more thorough qualitative and bed, southeast Jamaica: Contributions to quantitative analysis of the specimens Tertiary and Quaternary Geology, Leiden. with a broader group of biometric fea HENDLER, G., J.E. MILLER, D.L. PAWSON, tures. It appears as though at least two of and P.M. KIER, 1995, Sea stars, sea urchins, the three best fossil specimens are near and allies: Echinoderms of Florida and the the largest e~d of the size range reported Caribbean: Smithsonian Institution Press, for E. tribuloides. However, without better 390 p. preserved fossil specimens, it is impossible JACKSON, R.T., 1922, Fossil echini of the West to justify a new species description. Based Indies: Carnegie Institute of Washington, on the fossils currently available, we Publication no. 306, 102 p. believe the best interpretation is to treat LEWIS, D.N., and S.K. DONOVAN, 1991, The them as E. tribuloides. This is the first fos Pliocene Echinoidea of Tubago, West Indies: sil occurrence of this species from the Tertiary Research, v. 12, pp. 139-146. United States. KIER, P.M., 1963, Tertiary echinoids from the Caloosahatchee and Tamiami formations of LOCALITY DATA Florida: Smithsonian Miscellaneous Collections, v. 145, no. 5,. 63 p. The following is a collecting locality of the KIER, P.M., 1992, Neogene paleontology in the Invertebrate Paleontology Division, northern Dominican Republic: The class Florida Museum of Natural History, Echinoidea (Echinodermata): Bulletins of University of Florida (UF). AID.erican Paleontology, v. 102, no. 339, pp. 13-27. CH026 Shell pit approximately 1.2 kilometers MORTENSEN, T., 1928, A monograph of the east of Grove City, Charlotte County, Florida Echinoidea: vol. 1, Cidaroida, C.A. Reitzel, (NE 1/4, NE 1/4, Sec. 16, T41S, R20E; Copenhagen, 551 p. Englewood Quadrangle, USGS 7.5 minute FAWSON, D.L., 1978, The echinoderm fauna of series, 1987). Ascension Island, South Atlantic Ocean: Smithsonian Contributions to the Marine Sciences, no. 2, 31 p. LITERATURE CITED PHELAN, T.F., 1970, A field guide to the cidaroid echinoids of the northwestern CHESHER, R.H., 1972, The status of knowl Atlantic Ocean, Gulf of Mexico, and edge of Panamanian echinoids, 1971, with Caribbean Sea: Smithsonian Contributions to comments on other echinoderms: Biological Zoology, no. 40, 67 p. Society of Washington Bulletin, v. 2, pp.· 139_ PHELAN, T.F., 1972, Comments on the echi 158. noid genus Encope, and a new subgenus:· CUTRESS, B.M., 1980, Cretaceous and Tertiary Proceedings of the Biological Society of Cidaroida (Echinodermata: Echinoidea) of Washington, v. 85, no. 8, pp. 109-130. the Caribbean area: Bulletins of American SANCHEZ ROIG, M., 1949, Los equinodermos Paleontology, v. 77, no. 309, 221 p. f6siles de Cuba: Paleontologia Cubana, v. 1, DONOVAN, S.K., 1993, Jamaican Cenozoic 302 p. echinoidea, in Biostratigraphy of Jamaica SERAFY, D.K., 1979, Echinoids (Echinoder (R.M. WRIGHT, and E. ROBINSON, eds.): mata: Echinoidea): Memoirs of the Hourglass Geological Society of America Memoir 182, Cruises, v. 5, part 3, 120 p. pp. 371-412. WEISBORD, N.E., 1969, Some Late Cenozoic DONOVAN, S.K., and B.J. EMBDEN, 1996, echinoidea from Caho Blanco, Venezuela: Early Pleistocene echinoids of the Bulletin.s of American Paleontology, v. 56; no. Manchioneal Formation, Jamaica: Journal of 252, pp. 273-372.
August 27, 1997