PART F, REVISED, VOLUME 2, CHAPTER 11: SYSTEMATIC DESCRIPTIONS of the SCLERACTINIA FAMILY MICRABACIIDAE Rosemarie C
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Micrabaciidae 1 PART F, REVISED, VOLUME 2, CHAPTER 11: SYSTEMATIC DESCRIPTIONS OF THE SCLERACTINIA FAMILY MICRABACIIDAE ROSEMARIE C. BARON-SZABO1,2 and STEPHEN D. CAIRNS1 [1Department of Invertebrate Zoology, Smithsonian Institution, Washington, DC, USA, [email protected] and [email protected]; 2Senckenberg Research Institute, Frankfurt/Main, Germany, [email protected]] INTRODUCTION tion and was even called pseudo-bifurcate by JANISZEWSKA, JAROSZEWICZ, and STOLARSKI The Micrabaciidae is one of the smallest (2013). The unique microstructural skeletal and least diverse scleractinian families, composition was first described by JANISZEW- consisting of the five genera: Micrabacia SKA and others (2011) and later elaborated MILNE EDWArdS & HAIME (20 fossil species), by JANISZEWSKA and others (2015). The MOSELEY (four extant species), Leptopenus molecular monophyly, based exclusively on YABE & EGUCHI (four extant Letepsammia the mitochondrial gene CO1, was discussed species), Rhombopsammia OWENS (two extant by KITAHARA and others (2010), and based species), and MICHELIN Stephanophyllia on four genes, both mitochondrial and (three extant and ten fossil species). Thus, nuclear, by STOLARSKI and others (2011). nearly 40 valid species pertain to this family According to KITAHARA and others (2010) (CAIRNS, 1989, p. 13–24; BARON-SZABO, and STOLARSKI and others (2011), the Micra- 2002, p. 130–131; 2008, p. 147–153), most baciidae, along with the Gardineriidae, form of which (30) are known exclusively as fossils. a deeply diverging basal clade, equivalent to Only the two genera, MOSELEY, Leptopenus the robust and complex clades that encom- 1881 and OWENS, 1986 are Rhombopsammia pass all other Scleractinia. as yet known exclusively from the Holocene. This chapter gives an overview of the The monophyly of the family is supported taxonomy, stratigraphy, and geography of by macromorphological, microstructural, the genera currently included in the family and molecular evidence. The macromorpho- Micrabaciidae. The diagnosis of the family is logical evidence includes a unique pattern followed by the description of its nominate of septal and costal insertion, alternating genus, and then descriptions of the other septa and costae, and the frequent presence genera in alphabetical order. Table 1 (p. 7) of a marginal shelf. These characters are best provides a synopsis of micrabaciid genera, illustrated and discussed by CAIRNS (1989), including key characteristics and stratigraphic who also supplied a key to four (those with ranges. living species) of the five genera. The unique pattern of septal insertion, once thought to Family MICRABACIIDAE be a repeated bifurcation of the third cycle Vaughan, 1905, p. 386 septa (CAIRNS, 1989), was shown by JANISZE- [Micrabaciidae VAUGHAN, 1905, p. 386] [=Stephanophylliidae YABE & EGUCHI, 1934, p. 281] WSKA, JAROSZEWICZ, and STOLARSKI (2013) to be accomplished by unequal-length invagi- Simple, free, cupolate, ahermatypic, azoo- nations of the wall between septa at the xanthellate, non-epithecate sensu lato; wall peripheral edge of the calice, and thus could solid in earliest juvenile stages, becoming not originate from bifurcation. Nonetheless, porous in later ontogenetical stages; porosity the result in adult corals resembles bifurca- of wall achieved by development of concentric © 2017, The University of Kansas, Paleontological Institute, ISSN (online) 2153-4012 Baron-Szabo, R. C., & S. D. Cairns. 2017. Part F, Revised, Volume 2, Chapter 11: Systematic descriptions of the Scleractinia family Micrabaciidae. Treatise Online 98:1–8, 5 fig., 1 table. 2 Treatise Online, number 98 1 mm a b Micrabacia 1 mm c d FIG. 1. Micrabaciidae (p. 2–3). rows of pillars that are part of invaginated septa; septa perforate or compact, margins wall (sensu JANISZEWSKA, JAROSZEWICZ, & highly or slightly dentate; dissepiments STOLARSKI, 2013), which occurs between absent; synapticulae present or absent; wall septa of preceding cycles; septa represent probably marginotheca; columella composed vertical continuations of wall invaginations; of processes of thickening deposits, loose, septa of all cycles fused with nearest ones of spongy, or compact, often elongate; pali former cycle, resulting in pseudo-bifurcation absent; microstructure in thickening deposits pattern; septa of fourth cycle and higher do composed of thin (around 100–300 nm) and not appear simultaneously but are inserted short (1–2 µm) fibers arranged in chip-like in pairs whereby the septum adjacent to bundles; polyp completely invests corallum. older septa arises first; septa composed of Lower Cretaceous (Aptian)–Holocene. thickening deposits unique to this group; Micrabacia MILNE EDWArdS & HAIME, 1849, p. 71 not in crystallographic continuation with [*Fungia coronula GOLDFUSS, 1826–1827, p. 50; SD Rapid Accretion Deposits (RAD) or tradi- MILNE EDWArdS & HAIME, 1849, p. 71] [=Cycla- tional calcification centers; costae beaded or bacia BÖLSCHE, 1866, p. 474 (type, C. stellifera; SD VAUGHAN, 1905, p. 409); =Discopsammia smooth, alternate in position with septa by D’OrbIGNY, 1850, p. 180 (type, Stephanophyllia successive bifurcation, and match number of bowerbankii MILNE EDWArdS & HAIME, 1848, p. Micrabaciidae 3 a c 4.5 mm d 3.5 mm 4.5 mm Micrabacia e b 3.5 mm FIG. 2. Micrabaciidae (p. 2–3). 94; M)]. Solitary, cupolate to discoidal with flat missing, presumably housed at IPB), base (d) and to slightly concave base; marginal shelf narrow; lateral view (e) of corallum. (Milne Edwards & corallum free, small, usually less than 15 mm Haime, 1850, pl. 10,4a,c). in diameter; septa lamellar and perforate; septal Leptopenus MOSELEY, 1881, p. 204 [*L. discus margins nearly vertical peripherally, finely granu- MOSELEY, 1881, p. 205; SD WELLS, 1936, p. 115, lated laterally; synapticulae present; columella syntypes, NHMUK 1880.11.25.158–159]. Solitary, trabecular, papillose, or composed of lamellar discoidal with flat base; corallum extremely fragile segments; thecal pillars often not aligned in contin- and highly porous; costae composed of thin dentate uous concentric rings but in an alternating fashion. ridges that extend far beyond calice to form a [The holotype is assumed to be lost; type material delicate marginal shelf; septa rudimentary, porous, is presumably housed at the IPB (Bonn), but composed primarily of a series of tall spines; synap- repeated attempts by BARON-SZABO to trace the ticulae absent; columella large, spongy or variable specimens have failed. Genus is well described and in shape, composed of loose mass of spines. [Genus discussed in WELLS, 1933, p. 153–171; CAIRNS, is well described and discussed by CAIRNS, 1989, p. 1989, p. 13; BARON-SZABO, 2008, p. 147–153; 13–15.] Holocene: worldwide, including Antarctic, and JANISZEWSKA, JAROSZEWICZ, & STOLARSKI, 2013, 221–5000 m depth.——FIG. 3,1a–b. *L. discus, p. 244–256.] Lower Cretaceous (Aptian)–Eocene: syntype, NHMUK 1880.11.25.159, Holocene, western Europe, Aptian–Paleocene; South Asia, southwestern Atlantic Ocean, Challenger Station Albian; Central Asia, Cenomanian; West Asia, 323, 3473 m depth; a, skeleton surface, cross-section Turonian; eastern Europe, Upper Cretaceous; Central view; b, close-up view of a (new, images courtesy of America, Santonian; northern Europe, Campanian; Ken Johnson). North America, Campanian–Maastrichtian; North Letepsammia YABE & EGUCHI, 1932, p. 58 [*Stepha- Africa, Paleocene–Eocene.——FIG. 1a–d. *M. coro- nophyllia formosissima MOSELEY, 1876, p 561; nula (GOLDFUSS, 1826–1827), Cenomanian (Essen OD; syntype, NHMUK 1880.11.25.156]. Solitary, Greensand), Germany (North-Rhine Westphalia); cupolate with flat, highly porous base; corallum a–b, topotype, GPIH–4842, upper surface (a) and porous but sturdy; costae consist of thin, delicate base (b) of corallum; c–d, topotype, GPIH–4843, ridges; intercostal spaces broader than costae and upper surface (c) and base (d) of corallum (new, penetrated by large pores; thin marginal shelf images by Baron-Szabo).——FIG. 2a–e. *M. coro- present; septa lamellar and highly porous, with nula (GOLDFUSS, 1826–1827), Cenomanian (Essen complex dentition; small synapticulae present, Greensand), Germany (North-Rhine Westphalia); uniting lower, outer edges of all septa; columella a–c, topotype, MB K.2990–1, upper surface (a) and spongy, well developed. [Type and original material base (b) of corallum; c, topotype, MB K.2990–3, are well described by CAIRNS, 1989, p. 15–18, pl. upper surface of corallum (new, from Münster 6,j, 7,g–i, and 8,a–d]. Pliocene–Holocene: East Asia, collection, possibly original type collection, images Pliocene–Pleistocene; Indian and Pacific Oceans, by Baron-Szabo); d–e, sketch of holotype (specimen 97–710 m depth, Holocene.——FIG. 3,2a–b. L. 4 Treatise Online, number 98 1a 6.5 mm 2a 5 mm Letepsammia 2.5 mm 7 mm 1b 3a Leptopenus Rhombopsammia 5 mm 2b 3b 6.5 mm Letepsammia FIG. 3. Micrabaciidae (p. 3–5). Micrabaciidae 5 4.5 mm 1 10 mm 2 Letepsammia Rhombopsammia FIG. 4. Micrabaciidae (p. 3–6). formosissima (MOSELEY, 1876), syntype, NHMUK but sturdy; costae have two rows of aligned gran- 1880.11.25.156, Holocene, Indo-Pacific Ocean, ules, granules equal to or thicker than intercostal 170 m depth, Challenger Station 209; upper spaces; marginal shelf narrow, developed regularly surface (a) and lateral view (b) of corallum (new, or irregularly, absent in some places; septa lamellar images courtesy of Ken Johnson).——FIG. 4,1. L. and perforate only near base; synapticulae present; formosissima (MOSELEY, 1876), syntype,