The Rugose Corals Tabulate Corals A Devonian coral garden Tabulae Rugose corals get their name because the exterior of Tabulate corals are always colonial. There are 212 species of corals many of their forms has a wrinkly appearance. They are Their name is derived from a flat shelf known from the fossil beds at the often called “horn corals” because their form may resemble (tabula) that separates individuals Falls of the Ohio. Not all occur in CORALS the horn of a cow or goat. In fact, the largest horn coral vertically in the colony (figure 6). The the coral beds; some may be found (Siphonophrentis elongata, figure 1) was referred to as a individual corallites that make up the in the slightly higher (younger) at “petrified buffalo horn” by colony are typically 0.5 - 8 mm wide. rock layers. Additional studies may Figure 11. Underside Falls of the Ohio settlers due to its shape. Figure 6. Despite the small individuals, these reveal more or fewer species. It takes of Alveolites coral Horn corals show a wide corals can be quite large. The most experience to identify corals with State Park variation in form, although common coral is a form called Favosites (Emmonsia). accuracy. The random orientation of many of the solitary external form is not a Emmonsia was given its own genus name, but some rugose corals make identification even more difficult. Alan Goldstein, Interpretive Naturalist distinguishing feature for specialists consider it to be a sub-genus of Favosites. These Look for evidence of Devonian hurricanes. Can you find identification. Most rugose corals vary from a few inches (or centimeters) to 15 feet inverted colonies? Many corals radiate toward a center corals have septa radiating (4.5 m) across. The larger colonies probably had hundreds point, indicating they are upside-down (figure 11). Some from the center (like bicycle of thousands (or millions) of small of these colonies were more than 1,000 pounds! Others spokes) when observed in polyps living at one time. are on their side. Many colonial corals show rhythmic cross-section (figure 2). Figure 2. Coral Septa The colonial form of tabulate banding, whether these are daily seasonal or lunar cycles Cystiphylloides is characterized by corals reflects the environment. As are unknown. Horn corals that are “J” or “L” shaped are dissepimentaria, bubble-like structures in a modern reef, colonial corals were also jostled by storms, and bent back towards sunlight forming layers. Some have a both as can grow in mounds, ‘bushes,’ (see figure 1). internal structures (figure 3). Figure 7. Favosites sheets and plates. The coral beds are best exposed late summer through the (Emmonsia). detail Looking closely at the overall shape The first two fall. Visitors have the rare opportunity to walk on an ancient The fossil beds at the Falls of the Ohio State Park, and internal structure, one can appreciate forms are dominant, typical of high energy sea floor to get a good idea how corals were distributed if located between Louisville, Kentucky and Clarksville, the enormous variety of the 212 species conditions, where there is strong wave one could go back in time with snorkeling gear. Indiana, are world-renowned. First described by that have been recorded at the Falls of Figure 3. action and currents. Favosites (Emmonsia) the Ohio. Cystiphylloides grow in mounds (figure 7) and very thick European settlers paddling down the Ohio River It is easy to understand why branching forms. The largest [F. (E.) References & Suggested Reading over 250 years ago, the abundance of corals is still identifying new species is so ramosa] form colonies over 50 feet (15 m) striking today. Are these fossil corals related those difficult. across with other corals filling in the space *Davis, W. J., 1885, Kentucky Fossil Corals, living in today’s oceans? In a word, “no.” Modern Further “complicating” factors between branches as the colony was buried. Kentucky Geological Survey. (scleractinian) corals first appear in the fossil record - many rugose corals are not Alveolites and Thamnopora grow in bushes Figure 8. Greb, Stephen F., et al, 1993, Fossil Beds of the Falls in the early Triassic period, about 241 million years Thamnopor solitary, but colonial - and some with branches from 1/4” to one inch thick of the Ohio, Kentucky Geological Survey. ago. The corals from the Falls of the Ohio are much may be either form! Individual (figure 8). The latter occurs in colonies as large as 8 feet Oliver, W. A., Jr., 1976, Devonian Noncystimorph older, about 390 million years old! Some varieties corallites in a colonial coral (2.4 m) across. Platyaxum is an uncommon coral, growing Colonial Rugose Corals in the New York Area, superficially resemble living corals, because they lived may grow nearly parallel to in a plate-like form. U. S. Geological Survey, Professional Paper 869. Figure 4. Eridophyllum in a similar habitat - shallow, warm, tropical seas. neighbors, but occasionally Two other corals deserve particular mention because *Stumm, E.C., 1964, Silurian and Devonian corals contact with them, such as Eridophyllum (figure 4). they are unlike other tabulate corals. Syringopora grows in of the Falls of the Ohio: Geological Society of If the rugose coral is massive and the individuals are in mound or bush-like forms consisting of curved, straight or America Memoir 93. Two orders of coral dominated the Devonian seas: full contact with one another, it forms beautiful geometric gently undulating tubes (figure 9). Aulocystis is a dendroid rugosa and tabulata. Both became extinct at or near patterns, like Prismatophyllum prisma (figure 5). The (branching) coral with a tight bush-like form * Out of print, check public or university library or on-line. the end of the Permian period, about 240 million years largest colonial rugose coral (figure 10). ago. Both orders appeared in the early Ordovician on the Indiana shore (directly period and their diversity peaked during the Devonian. below the Interpretive Center) is a Prismatophyllum LEFT: The mission of the Interpretive Services is to provide informa- colony 11 feet (3.3 m) Figure 9. Syringopora tion and offer interpretive experiences with Indiana’s natural across. A 30–foot (10 m) and cultural resources to visitors, staff and a diverse public. colony is reported on the RIGHT: Kentucky side. Figure 10. Aulocystis Figure 5. Prismatophyllum Cover photo: Figure 1. Siphonophrentis elongata Fossil Coral Names in Current Use

Rugose Corals Edaphophyllum bifurcatum (Hall) Schlotheimophyllum versiforme (Hall) Favosites “Emmonsia” convexa (Davis) Enallophrentis concava (Hall) Siphonophrentis elongata (Rafinesque & Clifford)( figure )2 Favosites “Emmonsia” cymosa (Davis) Acinophyllum mclareni Fagerstrom Enallophrentis ? curvata (Hall) Siphonophrentis yandelli (Edwards & Haime) Favosites “Emmonsia” emmonsi (Rominger) Acinophyllum stokesi (Milne-Edwards & Haime) Enallophrentis ? cyathiformis (Hall) “Skoliophyllum” squamosum (Nicholson) [Probably new genus] Favosites “Emmonsia” eximia (Davis) Acrophyllum clarki Davis Enallophrentis duplicata (Hall) Stauromatidium trigemma (Davis) Favosites “Emmonsia” epidermata (Rominger) Acrophyllum conigerum (Greene) Enallophrentis ? foliata (Hall) Stereolasma (?) exile (Davis) Favosites “Emmonsia” ocellata (Davis) Acrophyllum ellipticum Davis Enallophrentis inflata (Hall) Stereolasma gallicalcar (Davis) Favosites “Emmonsia” radiciformis (Rominger) Acrophyllum oneidaense (Billings) Enallophrentis simplex (Hall) Stereolasma parvulum (Davis) Favosites “Emmonsia” ramosa (Rominger) Aemulophyllum exiguum (Billings) Enallophrentis trisutura (Hall) Stereolasma rectum (Hall) Favosites “Emmonsia” tuberosa (Rominger) Aemulophyllum exiguum elongatum (Davis) Eridophyllum apertum (Hall) (figure 4) Tabulophyllum? bellicinctum Greene Favosites argus Hall Amplexiphyllum cruciforme (Hall) Eridophyllum archaici (Billings) Tabulophyllum? greeni (Davis) Favosites biloculi Hall Amplexiphyllum tenue (Hall) Eridophyllum coagulatum (Davis) Tabulophyllum? perplicatum (Hall) Favosites clausus Rominger Aulacophyllum conigerum Davis Eridophyllum seriale Edwards & Haime Tabulophyllum? sinuosum Hall Favosites clelandi Davis Aulacophyllum mutabile Davis Eridophyllum tumidulum (Hall) Tabulophyllum? tripinnatum (Hall) Favosites goldfussi d’Orbigny Aulacophyllum perlamellosum (Hall) Hadrophyllum orbignyi Edwards & Haime Tabulophyllum zaphrentiforme Davis Favosites hamiltoniae Hall Aulacophyllum pinnatum Hall Hallia strigata (Greene) Triplophyllum terebrata (Hall) Favosites mundus Davis Aulacophyllum sulcatum (d’Orbigny) Heliophyllum agassizi Greene Zaphrentis phrygia Rafinesque & Clifford Favosites patellatus Stumm Bethanyphyllum arctifossa (Hall) Heliophyllum alternatum Hall Favosites pirum Davis Bethanyphyllum depressum (Hall) ? Heliophyllum coalitum (Rominger) Tabulate Corals Favosites placentus Rominger Bethanyphyllum nanum (Davis) Heliophyllum denticulatum Hall Favosites proximatus Stumm Bethanyphyllum pocillum (Davis) Heliophyllum ethelanum (Davis) Alveolites asperus (Rominger) Favosites quercus Davis Bethanyphyllum prateriforme (Hall) Heliophyllum gurleyi Greene Alveolites constans Davis Favosites ramulosus Davis Bethanyphyllum robustum (Hall) Heliophyllum halli Edwards & Haime Alveolites expatiatus (Rominger) Favosites rotundituba Davis Bethanyphyllum validum (Hall) Heliophyllum incrassatum Hall Alveolites goldfussi Billings Favosites turbinatus Billings Bethanyphyllum vesiculatum (Hall) Heliophyllum infundibulum Hall Alveolites minimus Davis Lecfedites canadensis (Billings) Blothrophyllum romingeri Stumm Heliophyllum ingens (Davis) Alveolites mordax Davis Platyaxum foliatum Davis “Blothrophyllum” trisulcatum (Hall) Heliophyllum insigne (Davis) Alveolites squamosus Billings Platyaxum frondosum (Nicholson) (Needs to be studied. Neither Blothrophyllum or Tabulophyllum) Heliophyllum latericrescens Hall Alveolites winchellana (Miller) Platyaxum orthosoleniskum (Werner) Bordenia knappi Hall Heliophyllum pocillum (Davis)? Antholites speciosus Davis Platyaxum undosum Davis “Breviphrentis halli” (Edwards & Haime) Heliophyllum tenuiseptatum Billings Aulocystis auloporoidea (Davis) Pleurodictyum cornu Stumm “Breviphrentis” nitida (Hall) Heliophyllum venatum Hall Aulocystis fascicularis (Davis) Pleurodictyum cylindricum (Michelin) “Breviphrentis” ovalis (Hall) Heliophyllum verticale Hall Aulocystis frutectosa (Davis) Pleurodictyum insigne (Rominger) Breviphrentis (?) planima (Hall) [further study needed] Heliophyllum yandelli (Rominger) Aulocystis (?) incrustans (Davis) Pleurodictyum maximum (Troost) Bucanophyllum ohioense (Nicholson) “Heterophrentis” annulata (Hall) Aulocystis jacksoni (Grabau) Pleurodictyum michelinoides (Davis) Cayugaea (?) subcylindricum Stumm Heterophrentis is not a valid genus. Aulocystis nobilis (Billings) Pleurodictyum papillosa (Davis) Cladionophyllum cicatriciferum (Davis) “Heterophrentis” colletti (Hall) Aulocystis (?) procumbens Davis Pleurodictyum planum (Davis) Coleophyllum romingeri (Hall) “Heterophrentis” irregularis (Hall) Aulocystis transitorius Stumm Pleurodictyum spiculata (Greene) Compressiphyllum davisana (Miller) “Heterophrentis” rafinesqui Aulopora culmula Davis Pleurodictyum wardi Greene Craterophyllum adnascens (Greene) (Edwards & Haime) Aulopora edithana Davis Romingeria commutata Beecher Craterophyllum (?) latiradium (Hall) “Heterophrentis” subcompressa (Hall) Aulopora tubiporoides (Yandell & Shumard) Romingeria fasciculata Davis “Craterophyllum” magnificum (Billings) Homalophyllum fusiformis (Hall) Bractea arbor (Davis) Romingeria umbellifera (Billings) Cyathocylindricum gemmatum (Hall) Homalophyllum herzeri (Hall) Bractea frutex (Davis) Romingeria uva Davis Cyathocylindricum opulens Oliver Homalophyllum ungulum (Rominger) Bractea impedita (Davis) Romingeria vannula Davis Cylindrophyllum gradatum (Greene) Iowaphyllum knotti (Davis) Chonostegites clappi Edwards & Haime Striatopora (?) alba Davis Cystiphylloides americanum (Edwards & Haime) Kionelasma coarticum (Hall) Chonostegites tabulatus (Edwards & Haime) Striatopora bellistriata Greene Cystiphylloides crassatum (Greene) Kionelasma? conspicuum (Hall) Cladopora acupicta Davis Striatopora cavernosa Rominger Cystiphylloides hispidum (Davis) Kionelasma mammiferum (Hall) Cladopora bifurca Davis Syringopora hisingeri Billings Cystiphylloides infundibuliformis (Greene) (figure )3 Odontophyllum convergens (Hall) Cladopora (?) gracilis Davis Syringopora perelegans Billings Cystiphylloides limbatum (Davis) Prismatophyllum bella (Davis) Cladopora gulielmi Davis Thamnopora distans (Nicholson) Cystiphylloides nanum (Hall) Prismatophyllum conjunctum (Davis) Cladopora (?) imbricata Rominger Thamnopora limitaris (Rominger) (figure )8 Cystiphylloides plicatum (Davis) Prismatophyllum ovoideum (Davis) Cladopora labiosa (Billings) [often confused with Alveolites Thamnoptychia alternans (Rominger) Cystiphylloides pustulatum (Hall) Prismatophyllum prisma Lang & Smith (figure )5 winchellana (Miller)] Thamnoptychia tuberculata (Stumm) Cystiphylloides quadrangulare (Hall) Prismatophyllum truncata Stewart Cladopora (?) robusta Rominger Thamnoptychia vermiculosa (Leseur) Cystiphylloides tenuiradium (Hall) Scenophyllum (?) coniferum (Greene) Favosites “Emmonsia” amplissima (Davis) Diplochone greenei (Miller) Scenophyllum conigerum (Rominger) Favosites “Emmonsia” arbuscula (Hall) Acknowledgement: The writer would like to thank the late Dr. William A. “Disphyllum” synaptophylloides Stumm Schlotheimophyllum typicum (Davis) Favosites “Emmonsia” bacula (Davis) Oliver, Jr. for critical review of the faunal list with updated names