Frederick Hotchkiss, PhD

annotations to published articles

Hotchkiss, F.H.C. 1977. Ophiuroid Ophiocanops (Echinodermata) not a living fossil. Journal of natural History 11: 377-380. Additional comments in Hotchkiss (1995): Ophiocanops does not have the 'auluroid' vertebrae of the Oegophiurida (p. 423); the extraordinary gonadal and stomachal characters perhaps are retained from the Oegophiurida; hence perhaps a surviving member of the stem group of the Order Phrynophiurida (p. 415, 423 and Fig. 6). Am aware of the following 2001 paper but have not seen it: Fujita, T., S. Irimura & W.W. Kastoro. Biology of a rare ophiuroid Ophiocanops fugiens Koehler, 1922 (Echinodermata) associated with black corals, with notes on the specimens collected from Lembeh Strait, Bitung, Indonesia. In: Terazaki, M., A. Taira, M. Uematsu, Y. Michida and T. Kaneko, eds., Proceedings of the 11th JSPS Joint Seminar on Marine Science, Center for International Cooperation, Ocean Research Institute, University of Tokyo, Tokyo, 2001, pp. 326-333. For more on this topic see 2008 paper by Sabine Stöhr, Chantal Conand and Emilie Boissin: Brittle stars (Echinodermata: Ophiuroidea) from La Réunion and the systsematic position of Ophiocanops Koehler, 1922. Zoological Journal of the Linnean Society, 2008, 153:545-560 [they describe Ophiocanops multispina n.sp., transfer Renetheo felli to Ophiocanops, and present many new observations on morphology including upper arm plates, a second under arm plate, side arm plates that meet ventrally so that there is no resemblance to the ambulacral groove of Paleoozoic Oegophiurida, and place Ophiocanops in the Ophiomyxidae].

Hotchkiss, F. H. C. 1979. Case studies in the teratology of starfish. Proceedings of The Academy of Natural Sciences of Philadelphia 131:139-157. The historical review on pp. 139-143 is only a sampling of what could have been included. Some additional references, in historical order, follow: 1882. Bell, F.J. Note on the echinoderm fauna of the Island of Ceylon, together with some obversations on heteractinism. Ann. Mag. Nat. Hist., ser. 5, 10:218-225; 1887. Bell, F.J. The echinoderm fauna of the island of Ceylon. The Scientific Transactions of the Royal Dublin Society, series 2, 3:643-657, pls. 39-40. [an oblique cut regenerates obliquely to the arm axis]; 1912. Richters, C. Zur Kenntnis der Regenerationsvorgange bei Linckia. Zeitschrift fur Wissenschaftliche Zoologie 100:116-175; 1914. Schapiro, J. Uber die Regenerationserscheinungen verscheidener Seesternarten. Arch. f. Entwmeck. 38:210-251; 1915. Nusbaum, J. & M. Oxner. Zur Restitution bei dem Seestern Echinaster sepositus Lam. Zool. Anzeiger 46(6):161-167; 1917. Zirpolo, G. Notizia di alcuni Asteroidi anomali pescati nel Golfo di Napoli, Echinaster sepositus Gray e Asterias glacialis O.F. Muller. Bollettino della Societa di Naturalisti in Napoli 30:20-29; 1918. Zirpolo, G. Un caso di rigenerazione parziale delle braccia in un Astropecten aurantiacus L. Pubblicacazioni della Stazione Zoologica di Napoli 2(2):169-175, pl. 10 [a 5-rayed individual loses two adjacent rays and some disk; only a single ray regenerates, resulting in a 4-rayed individual]; 1924. Zirpolo, G. Ulteriori notizie di Asteroidi anomali. Bollettino della Societa di Naturalisti in Napoli 36:305-346, pls. 6-8; Zirpolo, G. Notizia di un Echinaster sepositus Gray con sei braccia nel Golfo di Napoli. Atti Pontif. Accad. Nuovi Lincei, Anno 77:161-163. [No external or internal irregularities, providing his argument for hexamery from the first moment of life]; 1926. O'Donohue, C.H. On the summer migration of certain starfish in Departure Bay, B.C., Canada. Fisheries Research Board of Canada. Contributions to Canadian Biology and Fisheries 1:455-472; Zirpolo, G. Gemmazioni, rigenerazioni ipertipiche ed ipotipie studiate nell'Astropecten aurantiacus L. Bollettino della Societa di Naturalisti in Napoli 38:167-221 [includes examples of tetramery arising when two arms regenerated in place of three, and when one arm regenerated in place of two]; 1929. Zirpolo, G. Le forme cometoidi dell'Asterias tenuispina Lmk. Bollettino della Societa di Naturalisti in Napoli 40:25-35; Zirpolo, G. Nuovo caso di gemmazione in un Astropecten aurantiacus L. Bollettino della Societa di Naturalisti in Napoli 40:83-87; Zirpolo, G. Notizia di Asteroidi irregolari. Bollettino della Societa di Naturalisti in Napoli 40:221-231; 1931. Abeloos, M. Les potentialites regeneretrices de la face dorsal des bras des Asteries, Trifurcation dorsal d'un bras chez Solaster papposus (Linck). Bull. Biologique de la France et de la Belgique fonde par Alfred Giard, 65(3):394-405. 1933. Mortensen, Th. The Godthaab Expedition, 1928, Echinoderms. Meddelelser om Gronland 79(2):1-62 + plate I. [on p. 17 arm number in Solaster papposus: nearly all have 10 arms, one has 12, a few have 9, one has 8 arms]; 1938. Domantay, J.S. An unusual bud due to heteromorphosis in Echinaster luzonicus (Gray). Phillipp. J. Sci. Manila 64:281-283; 1941. Fisher, W.K. A new genus of sea stars (Plazaster) from Japan, with a note on the genus Parasterina. Proc. USNM 90(3114):447-456, pls. 66-70; 1945. Fisher, W.K. Unusual abnormalities in sea-stars. J. Wash. Acad. Sci. 35:296-298; 1951. Alfano, B. Commemorazione del Prof. Giuseppe Zirpolo. Bollettino della Societa di Naturalisti in Napoli 59:39-64 [on the life and publications of G. Zirpolo]; 1958. Dollfus, R.Ph. Courbe interbrachiale chez une asterie du genre Stellasteropsis. Bull. Soc. Zool. Fr. 83:294-297 [compares 4-armed and 5-armed individuals]; 1967. Raup, D.M. & E.F. Swan. Crystal orientation in the apical plates of abberant echinoids. The Biological Bulletin 133(3):618-629 [ocular IV and genital 4 are judged missing in two four-part urchins; of special interest because interradius IV/V is the location of closure of the hydrocoele crescent to form the ring canal; ray IV is on the ventral or posterior horn of the hydrocoele crescent]; 1969. Kenny, R. Growth and asexual reproduction in the starfish Nepanthia belcheri (Perrier). Pacific Science 23(1):51-55. 1975. Hinegardner, R.T. Morphology and genetics of sea urchin development. American Zoologist 15:679-689 [breeding experiments and loss of symmetry control]; 1986. Horowitz, A., S. Able, & H.L. Strimple. Abnormalities in Pentremites Say (Blastoidea) from the Pella Formation (Upper Mississippian) of Iowa. Journ. Paleontology 60(2):390-399; 1989. Ausich, W.I. & T.W. Kammer. Teratological specimen of Agaricocrinus americanus (Roemer) (Lower Mississippian, Crinoidea). J. Paleont 63(6):945-946; 1991. Boursin, J. Un curieux (b)oursin ou coussin? Xenophora, number 53:25 [square 4-armed Halityle regularis from New Caledonia]; 1992. Ramalingam, K. & A. J. Antony. A note on the tetramerous starfish (Oreaster sp.). Comparative Physiology and Ecology 17(3):121-122; 1999. James, D.B. Abnormal asteroids from the seas around India. Mar. Fish. Infor. Ser., T. & E. Ser., number 158:21-22; 2000. Hotchkiss, F.H.C. On the number of rays in starfish. American Zoologist 40;340-354 [this paper inludes additional references not repeated here, as well as a new synthesis for the topic]; 2002. Dupont, S. & J. Mallefet. Abnormal forms in the brittle-star Amphipholis squamata: a field study. J. Mar. Biol. Ass. U.K. 82:491-493 [deviations from pentamerism are not a heritable character but are a consequence of environmental perturbations on the metamorphosis of larvae and/or abnormal regeneration of arms]; 2007. Ceranka, T. Symmetry disorders of the test of the Miocene echinoid Echinocyamus from Poland. Acta Palaeontologica Polonica 52(3):503-518.

Behavioral evidence for a physiological anterior in locomotion in starfish was examined on pp. 153-154. Concerning behavioral evidence for nonrandom use of arms in ophiuroids see: Welte, N.T. & L.M. Lutton. 2003. Appendage selection during brittlestar locomotion. Journal of the Pennsylvania Academy of Science 77(1):15-19. This paper revealed a pattern of two adjacent arms being used least for leading and most for propulsion. The identity of the arms with respect to the location of the madreporite was not examined and deserves further study.

Hotchkiss, F.H.C. 1980. The early growth stage of a Devonian ophiuroid and its bearing on echinoderm phylogeny. Journal of Natural History 14:91 96. The paper documented primary rosette plates in an oegophiurid with alternating ambulacral plates (Lysophiurina). The Lysophiurina and Zeugophiurina lineages diverged no later than Early Ordovician time. The Zeugophiurina are the presumed stem group of the Silurian-Recent Ophiurida. Documentation of a primary rosette in Paleozoic Ophiurida can be mentioned: in Ophiaulax decheni (Devonian, Ophiurinidae) [R. Haude & E.T. Thomas. 1983. Ophiuren (Echinodermata) des hohen Oberdevons im nördlichen Rheinischen Schiefergebirge. Paläont. Z. 57(1/2):121-142]; in Aganaster gregarius (Mississippian, Ophiolepididae) [F.H.C. Hotchkiss & R. Haude. 2004. Observations on Aganaster gregarius and Stephanoura belgica. Pp. 425-431 in Heinzeller & Nebelsick (eds.), Echinoderms: München]. Additional references on postlarval growth stages in Recent ophiuroids include: 1998. Sumida, P.G. and P.A. Tyler. postlarval development in shallow and deep-sea ophiuroids (Echinodermata: Ophiuroidea) of the NE Atlantic Ocean. Zoological Journal of the Linnean Society 124:267-300. 2005. Stöhr, S. Who's who among baby brittle stars (Echinodermata: Ophiuroidea): postmetamorphic development of some North Atlantic forms. Zoological Journal of the Linnean Society 143:543-576 [includes information on the euryalid Asteronyx loveni: the plates of the primary rosette are conspicuous at first, but do not endure: at 3.2 mm disk diameter they are small fragments at the center of the disk].

Hotchkiss, F. H. C. 1982. Ophiuroidea (Echinodermata) from Carrie Bow Cay, Belize. Pp. 387-412 in The Atlantic Barrier Reef Ecosystem at Carrie Bow Cay, Belize, I. Structure and Communities. K. Rutzler and I.G. Macintyre, editors. Smithsonian Contributions to the Marine Sciences, No. 12. Corrections to identifications in this paper were published by Hendler & Peck (1998. Ophiuroids off the deep end: Fauna of the Belizean fore-reef slope. Pp. 411-419 in Echinoderm Biology, Burke et al. [eds.], Balkema, Rotterdam), as follows: Ophiomitrella glabra (H.L. Clark) is regarded as a junior synonym of Ophioblenna antillensis, Axiognathus squamatus (Delle Chiaje) as a junior synonym of Amphipholis squamata, and the six-rayed fissiparous Ophiostigma sp. was nominally referred to Ophiostigma isocanthum [but see below]. The Ophioderma sp. juvenile is Ophioderma rubicundum, and the Sigsbeia murrhina Lyman sensu Hotchkiss specimens are Sigsbeia conifera Koehler. The Ophiurochaeta littoralis Koehler sensu Hotchkiss specimens were reidentified as an undescribed species of Ophiurochaeta [four undescribed Ophiurochaeta species provisionally designated A, B, C and D are mentioned by Hendler and Peck]. In a subsequent paper the six-rayed fissiparous Ophiostigma sp. was described as the new species Ophiostigma siva by Hendler [1995. New species of brittle stars from the Western Atlantic, Ophionereis vittata, Amphioplus sepultus, and Ophiostigma siva, and the designation of a neotype for Ophiostigma isocanthum (Say) (Echinodermata: Ophiuroidea). Contributions in Science, Natural History Museum of Los Angeles County No. 458].

Hotchkiss, F.H.C., S.E. Churchill, R.G. Gelormini, W.R. Hepp, R.J. Rentler & M.T. Tummarello. 1990. Events of autotomy in the starfish Asterias forbesi and A. vulgaris. Presented at 7th International Echinoderms Conference, Sept. 9/14, 1990, Atami, Japan. Published 1991, pp. 537-541 in T. Yanagisawa et al. (eds.), Biology of Echinodermata, A.A. Balkema, Rotterdam, 590 pp.

Hotchkiss, F. H. C. 1993. A new Devonian ophiuroid (Echinodermata: Oegophiurida) from New York State and its bearing on the origin of ophiuroid upper arm plates. Proceedings of the Biological Society of Washington, vol. 106, no. 1, pp. 63 84. To the comment on p. 73 regarding the very straight arms of the holotype and the existence of a stiffening reaction in ophiuroids, see remarks of W.K. Spencer (Monograph p. 246 citing von Uexkull, and p. 316 on Furcaster with arms stretched out very stiffly) that a very strong stimulus to the arm produces a rigor ... and the arms feel like small sticks. On the subject p. 75 that upper arm plates do not stay with the arm segments that become included in the disk, Ishida & Inoue (1993:104 Ophiura sarsii) noted that "about 4 dorsal arm plates at the base of the arm are incorporated into the disk". Further to the evidence p. 76 against the idea that upper arm plates get incorporated into the radial shields, note A.M. Clark 1966 Echinodermata [Port Phillip Survey 1957-1963], Memoirs of the National Museum, Melbourne No. 27, pp. 289-384, on p. 337 regarding Amphiura (Ophiopeltis) parviuscutata that basal arm segments have their dorsal arm plates reduced or even absent inside the disk area. Regarding the possible origins and homologies of the radial shields, these were not treated comprehensively on pp. 75-76 except where there was some suggestion that related radial shields to upper arm plates. For other origins of radial shields see for example Spencer (Monograph p. 244) and Matsumoto (Monograph p. 369). The statement of a reviewer that "it is incorrect to infer from Lyman's illustrations that radial shields grow by adding platelets" is true only for ophiurid ophiuroids in which radial shields grow from a single tiny scale which initially formed at the edge of the disk. Brian Stewart showed that the radial shields of Astrobrachion constrictum grow by proximal sequential addition of scales of stereom to the proximal ends (1966. Growth dynamics of the radial shields of the euryalid snake star Astrobrachion constrictum. Invertebrate Biology 115(4):321-330). On the subject of retaining for the moment the two suborders Lysophiurina and Zeugophiurina p. 80, note W.K. Spencer (Monograph p. 9) on " remarkable analogous (homoplastic) course of development" of two series, and that "For the present both series are included in the Ophiuroidea". I did not mean to suggest that Strataster might in some way be ancestral to crown group ophiuroids. The evidence seen in Strataster helps us understand the origin of upper arm plates in all lineages of ophiuroids. A paper by Turner & Heyman, 1995, Proc. Biol. Soc. Wash 108(2):292-297 has important observations on the presence of radial shields and dorsal arm plates in Ophiosyzygus and many other ophiomyxids.

Hotchkiss, F.H.C. 1995. Lovén's law and adult ray homologies in echinoids, ophiuroids, edrioasteroids and an ophiocistioid (Echinodermata: Eleutherozoa). Proceedings of the Biological Society of Washington [DC], vol. 108, no. 3, pp. 401 435. Lovén’s Rule in the echinoid order Cidaroida: Lovén’s Rule is evident in the pattern of alternation of the ambulacral plates that begins at the edge of the peristome in a 7 mm diameter specimen of Histocidaris elegans (A. Agassiz) figured by Mortensen (1927: Fig. 6). Reference: Mortensen, Th. 1927. Report on the Echinoidea collected by the United States Fisheries Steamer “Albatross” during the Philippine Expedition, 1907-1910. Part 1 The Cidaridae. USNM Bulletin 100, vol. 6, part 4, pp. 243-312, pls. 48-80. Lovén’s Rule in the echinoid order Temnopleuroida: in Temnopleurus apodus only the buccal plates Ib, IIb, IIIa, IVb, and Va carry tubefeet, the other buccal plates being quite rudimentary (Mrtsn 1943 Monograph III.2 p. 101); in Paratrema doederleini with only five buccal plates, the five buccal plates that are present are the plates Ib, IIb, IIIa, IVb, Va (R. T. Jackson, 1927, p. 451, fig. 8; confirmed by Mrtsn 1943 Monograph III.2 p. 274; a specimen with only four buccal plates has the A. II buccal plates missing); however in Prionechinus saggitiger with only five buccal tube feet the pattern is bbbba (Agassiz, 1881, Challanger Report, pl. VIa, fig. 12; numbering of ambulacra not stated). Lovén’s Rule in the echinoid order Echinoida: In Strongylocentrotus droebachiensis and in Psammechinus miliaris the buccal tube-feet first appear in the plates Ib, IIb, IIIa, IVb, Va (Jackson 1927, pp. 451-452; David & Mooi, 1995, The ontogenetic basis of Lovén’s Rule clarifies homologies of the echinoid peristome, pp. 155-164, in Echinoderm Research 1995, Emson, Smith & Campbell (eds), Balkema, Rotterdam; key reference on Lovén’s Rule in echinoids and other Echinodermata). Lovén’s Rule in the echinoid order Holasteroida: the largest ambulacral basicoronals in pourtalesiids definitely exhibit the aabab pattern early in ontogeny (David & Mooi 1995). To the list of references concerning Fell's identification of Kirk's ophiuroid as Ophiomyxa (p. 415), add that his first mention was in his 1962 key to the genera of ophiuroids. Additional and more recent sources of information relating to Table 2 include: Emlet, R.B. 2006. Direct development of the brittle star Amphiodia occidentalis (Echinodermata,Ophiuroidea, Amphiuridae) from the northeastern Pacific Ocean. Invertebrate Biology 125(2): 154-171 (place of hydrocoele closure and location of the stone canal). Further to evidence on changes of symmetry axes (Note 5): In the Order Echinoida, the oval test of Parasalenia gratiosa var. boninensis is elongate in the IIIb-5b direction, that of Echinometra mathaei in the 3a-Ia direction, that of Hetrocentrotus mamillatus and of H. trigonarius in the IVb-1b direction, and the test of Colobocentrotus mertensii is elongate in the 4a-1b direction using Lovén's nomenclature (Hayato Ikeda, 1939, The shape of the test of Colobocentrotus mertensii Brandt (Echinoidea, Echinometridae), Annotationes Zoologicae Japonenses 18(3):194-201, pls. 9-10). A specimen of C. mertensii with its long and short axes apparently interchanged, as judged by the location of the madreporite, may be a case of situs inversus (unpublished suggestion of FHCH; specimen described by Hiroshi Ohshima, 1939, Some rare abnormalities in asteroids and echinoids,Zool. Mag. Tokyo, 51:158-162). The ovoid test of Tiarechinus can also be mentioned (Treatise on Invertebrate Paleontology 1966 Part U pp. U436-U437). The test is circular but the apical system is elongate in the III-5 direction in Orthopsis ruppellii, Order Orthopsida (Treatise fig. 326,1h), likewise in Glyphocyphus (G.) radiatus, Order Temnopleuroida (Treatise pp. U414-415). The details for SET I specimen numbers 106-112 are listed below: 106. Protasteridae, Eugasterella africana Jell & Theron, South African Museum, Cape Town, paratype, SAM K1015, figured Jell & Theron (1999:177, Fig. 48A), madreporite in III/IV interradius. 107. Protasteridae, Eugasterella africana Jell & Theron, South African Museum, Cape Town, paratype, SAM K1014, figured Jell & Theron (1999:177, Fig. 48C) , madreporite in III/IV interradius. 108. Protasteridae, Eugasterella africana Jell & Theron, Geological Survey of South Africa, Bellville in Capetown No. B4561B, paratype, figured Jell & Theron (1999:177, Fig. 48B), madreporite in III/IV interradius. 109. Protasteridae, Eugasterella africana Jell & Theron, Geological Survey of South Africa, Bellville in Capetown No. B4561D, paratype, figured Jell & Theron (1999:176, Fig. 47C), madreporite in III/IV interradius. 110. Protasteridae, Eugasterella africana Jell & Theron, Geological Survey of South Africa, Bellville in Capetown No. B4561a, holotype, figured Jell & Theron (1999:175, Fig. 46A), madreporite in III/IV interradius. 111. Protasteridae, Strataster ohioensis Kesling & LeVasseur, , Geological Survey of South Africa, Bellville in Capetown No. B0196, figured Jell & Theron (1999:181, Fig. 51C), madreporite in III/IV interradius. 112. Cheiropterasteridae, Hexuraster weitzi (Spencer), Roy Oosthuizen Collection, Zwartskraal, Prince Allbert No. RO 45, figured Jell & Theron (1999:163, Fig. 39) , madreporite in III/IV interradius. The details for SET II specimen numbers 215-219 are listed below: 215. Protasteridae sp. Specimen not figured by Jell & Theron. Specimen details still not known. Arms score ?AB*A? 216. Eophiuridae, Haughtonaster reedi Rilett, Geological Survey of South Africa, Bellville in Capetown No. B4567, figured Jell & Theron (1999:158, Fig. 36B). Arms score AA?*AB 217. Protasteridae, Strataster ohioensis Kesling & LeVasseur, , Geological Survey of South Africa, Bellville in Capetown No. B4512, figured Jell & Theron (1999:183, Fig. 53). Arms score AAB*?? 218. Protasteridae, Strataster ohioensis Kesling & LeVasseur, , Geological Survey of South Africa, Bellville in Capetown No. B4513, figured Jell & Theron (1999:182, Fig. 52D). Arms score ?AB*?B 219. Protasteridae, Strataster ohioensis Kesling & LeVasseur, , Geological Survey of South Africa, Bellville in Capetown No. B4513, figured Jell & Theron (1999:179, Fig. 49b). Arms score AAB*A? NEW OBSERVATION: Specimen SUG299 of Encrinaster tischbeinianus (Roemer), Family Encrinasteridae, from De Doorns, Ceres District, Lower Devonian Bokkeveld Group, property of the Geological Collections, Stellenbosch University, South Africa, oral surface, figured Jell & Theron (1999:164, Fig. 40B-D): The specimen has a madreporite (enlarged in their Fig. 40D; madreporite not recognized by FHCH in the contact print in 1994). Using a magnifier to examine their Fig. 40C, the arms score AAB*AB. The five arms conform with Lovén’s AABAB rule. The madreporite is located in the III/IV interradius. The specimen fulfills SET I criteria and is assigned Set I reference number 113. Including the new observation, Lovén’s rule applies to the ophiuroid families Eophiuridae, Cheiropterasteridae, Encrinasteridae and Protasteridae (not just the Protasteridae as originally reported).

Hotchkiss, F.H.C., D. M. Rudkin, and S. Anderson. 1997. A case for rearmament -- the oldest known evidence of regeneration in sea stars (Abstract). 7th Canadian Paleontology Conference, University of Saskatchewan, Saskatoon, September 26-30, 1997. [Also: Rudkin, D. M., F. H. C. Hotchkiss and S. Anderson. A case for rearmament -- the oldest known evidence of regeneration in sea stars. Royal Ontario Museum Nineteenth Annual Research Colloquium, 20 November 1997, Toronto, Abstracts of Papers, p. 1 (abbreviated abstract).] We reported a specimen of Promopalaeaster wilsonae (Raymond 1912) with two regenerating arm tips, collected by S. Anderson from the Upper Member of the Bobcaygeon Formation (Middle Ordovician: Kirkfieldian) of south-central Ontario, and now housed in the Royal Ontario Museum (ROM 53320). The Anderson fossil is the oldest [earliest] example of an asteroid with a regenerating ray, supplanting a regenerating specimen of Cnemidactis girvanensis from the Upper Ordovician of Girvan, Scotland reported by Spencer (1918 pt. III, p. 161, pl. 12 fig. 5). Arm stumps that had healed without regenerating the lost arm were described in the Ordovician starfish Urasterella ulrichi by Schuchert (1915:37).

Hotchkiss, F.H.C. 1998. Discussion on pentamerism: The five-part pattern of Stromatocystites, Asterozoa, and Echinozoa. Pp. 37-42 In R. Mooi and M. Telford (eds.) Echinoderms: San Francisco. A.A. Balkema, Rotterdam. Postscript notes: Nichols (1967b) noted that the crystallographic alignment hypothesis of Cockbain (1966) is not consistent with c-axis orientations in echinoid genital plates; clarified and expanded arguments that support the suture-line hypothesis. There is much additional literature relevant to this topic that could not be fitted into the original paper; also there is literature that has come out since the paper. 1889. Bather, F.A. Trigonocrinus, a new genus of Crinoidea, from the "Weisser Jura" of Bavaria; with the description of a new species, T. liratus.--Appendix. Sudden deviations from normal symmetry in Neocrinoidea. Quarterly Journal of the Geological Society for February 1889, pp. 149-171, pl. 6 [p. 166 pentamery persists through natural selection based on mechanical principles; in pentamerous calyx every line of weakness is met halfway by a solid plate]. 1915. Clark, A.H. A monograph of the existing crinoids. The comatulids. USNM Bull. 82, vol. 1, part 1, 406 pages [p. 148: sutures are lines of weakness; with an odd number of radial divisions the sutures do not line up and the construction is stronger; the relation between the bilateral crustacean type and the pentaradiate echinoderm type is explained and provides explanation for the establishment of pentaradial symmetry]. 1928. Bather, F.A. The fossil and its environment. Quarterly Journal of the Geological Society of London 84(2):lxi-xcviii [pp. lxxi-lxxii present his ideas on the origin in Pelmatozoa of a primitive trisactiny followed by a forking of the lateral rays; a pentactiny superimposed upon a trisactiny; states the advantage of tripartite and quinquepartite over quadripartite: each line of weakness abuts on a mass of strength; calling it the principle of bonding] 1975. Strathman, R.B. Limitations on diversity of forms: branching of ambulacral systems of echinooderms. The American Naturalist 109:177-190. 1987. Anderson, D.T. Developmental pathways and evolutionary rates. In: Rates of Evolution, K.S.W. Campbell & M.F. Day (eds.), Allen & Unwin, Sydney [p. 146 pentamerous symmetry determined during development by a counting process]. 1998. Tolić, Iva Marija and Nenad Trinajstić. The number five in biology. Periodicum Biologorum 100(3):259-266 [Fivefold symmetry in animals, plants, viruses, biomolecules, crystals (quasicrystals)]. 2001. Changizi, M.A. The economy of the shape of limbed animals. Biological Cybernetics 84:23-29 [wire-minimization principles, animal limb number, and body-to-limb proportion; predicts six or more rays for Asteroidea]. 2002. Barrio, R.A., P.K. Maini, J.L.Aragón and M. Torres. Size-dependent symmetry breaking in models for morphogenesis. Elsevier Physica D 168-169(2002):61-72 [discusses de novo appearance and control of pentagonal symmetry of echinoids]. 2006. Parsley, R.D. and Y. Zhao. Long stalked eocrinoids in the basal Middle Cambrian Kaili Biota, Taijiang County, Guizhou Province, China. J. Pallaeont. 80(6):1058-1071 [pp. 1069-1070 discuss the ontogeny and phylogeny of ambulacral symmetry, from a 1-1 pattern to a 2-2 pattern to a 2-1-2 pattern].

Hotchkiss, F.H.C. 1998. A "rays-as-appendages" model for the origin of pentamerism in echinoderms. Paleobiology 24:200-214. A gentle criticism of the idea that there are analogies between appendages of bilateral forms and the rays of echinoderms is thoughtfully presented by R. Mooi, B. David & G. Wray, 2005, Arrays in rays: terminal addition in echinoderms and its correlation with gene expression, Evolution and Development 7(6):542-555. I accept their comments and claim only the analogy; the phrase rays-as-appendages seemed apt because Baterson's rule of duplication was discovered from analysis of duplicated appendages, and does not apply to axial structures; however Bateson's rule is not limited to appendages (it operates also in ciliate protozoa). Further to the mention on p. 203 of the paper by Hinegardner (1975): The report of loss of symmetry control and the rearing of four-part "square" Lytechinus pictus challenges the working hypothesis of locked-in pentamerism. Therefore it is important to mention that these are highly inbred lines with high mortality as larvae and at metamorphosis, facts which suggest that the square imagos are the result of teratological incomplete development. I do not agree with D.G. Stephenson's (Proc. European Colloq. Echinoderms,Brussels, 1979) interpretation that Hinegardner's square sea urchins are the result of loss of symmetry as soon as domestication relaxes the constraints of natural selection; nor Stephenson's corollary that pentamerism must be extrtemely adaptive for echinoids. Stephenson also presented the idea that "pentamerism is an ancestral character retained because any disadvantages of pentamerism are less than the disadvantages of changing it", and claimed that "this possibility has been ruled out by Hinegardner's (1975) breeding experiments." I propose instead that the loss of symmetry reported by Hinegardner be interpreted along the lines described by John M. Opitz (1985. The developmental field concept. American Journal of Medical Genetics 21:1-11, see Final generalizations on pp. 9-10): "all primary malformations qua developmental field defects are causally nonspecific"; "most will be shown to be causally heterogeneous"; "the causes are many, but the final common developmental pathways are few"; "most primary malformations are anomalies of incomplete formation"; "most field defects are multifactorially determined (not 'inherited'), hence have a low empiric recurrence risk"; "because oligogenes of major effect do play a role in the developmental history of all fields, sooner or later Mendelian inheritance of a primary malformation should be expected and observed"; "most field defects occur per se but can be and frequently are components of complex syndromes, including aneuploidy syndromes"; "the epigenetically hierarchichical nature of field development is responsible for the fact that, regardless of how small or limited the embryonic primordium, so long as it is still a field, its development can be altered genetically and environmentally." A further reference to diatoms with pentagonal symmetry: P.A. Simms & R. Ross. 1990. Triceratium pulvinal and T. unguicularum, two confused species. Diatom Research 5(1):155-169 [figs. 13-16 of T. unguiculatum forma quinquangulum]. Further to the matrix of ray homologies and especially concerning the entry for Luidia: My primary source on the location of hydrocoel closure and the madreporite in Luidia is Bury 1895 Fig. 21 and text pp. 67-68 [Bury 1895 The metamorphosis of echinoderms. Q. J. Microsc. Sci. n.s. 38:45-136 + pls. 3-9] [this source is mentioned in the caption to my matrix of ray homologies, but change "Fig. 22" to "Fig. 21"]. The observations of Bury were latched onto by F.A. Bather [1915 Studies in Edrioasteroidea IX. The genetic relations to other echinoderms. Geol. Mag., n.s., dec. VI, vol. 2, pp. 393-403]. On page 401 “It is important to notice here that in an Asteroid larva (Bipinnaria asterigera) described by Bury, the [hydrocoel] closure does still take place in the M plane” and mentioned further on p. 402. Mentioned again by Horstadius [1939 Uber die Entwicklung von Astropecten aranciacus L. Pubb. Staz. Zool. Napoli 17(2):221-312]. On p. 276 he says that in all known starfish (including Astropecten) the madreporite is between hydrocoel lobes I-II, except in Luidia it is between lobes V and I. On p. 278 he appears to give his own confirmatory observations on this point in recounting his work on Luidia ciliaris. NOTE NOTE NOTE – his system of Roman numerals is customary numbering of the linear series of hydrocoel lobes as seen in developing imago of metamorphosing larva, so is different than in my matrix. His Roman I is my edrio and asteroid D; his Roman V is my edrio and asteroid C; his Roman II is my edrio and asteroid E. Some newer papers on the development of the echinoderm body plan: 2006: Swalla, B.J. Building divergent body plans with similar genetic pathways. Heredity 97:235-243.

Hotchkiss, F.H.C., Rudolf J. Prokop and Václav Petr. 1999. Isolated skeletal ossicles of a new brittlestar of the Family Cheiropterasteridae Spencer, 1934 (Echinodermata: Ophiuroidea) in the Lower Devonian of Bohemia (Czech Republic). Journal of the Czech Geological Society 44:189-193. To the occurrences of the family Cheiropterasteridae, add the Silurian and Lower Devonian of Bolivia (Branisa, Leonardo. 1965. Los fosiles guis de Bolivia I. Paleozoico. Index Fossils of Bolivia I. Paleozoic. Servicio Geologico de Bolivia, Boletin No. 6, 282 pp., 80 pls. [?Loriolaster sp. p. 108 pl. xxii fig. 1] [?L. cf. L. mirabilis p. 128 pl. xxxii fig. 9]). The replacement name for Hexura Spencer, 1950, is Hexuraster Jell & Theron, 1999.

Hotchkiss, F.H.C., Rudolf J. Prokop and Václav Petr. 1999. Isolated vertebrae of brittlestars of the Family Klasmuridae Spencer, 1925 (Echinodermata: Ophiuroidea) in the Devonian of Bohemia (Czech Republic). Journal of the Czech Geological Society 44:329-333. To the occurrences of the family Klasmuridae, add isolated ossicles in the Devonian of Poland by Bockzarowski (2001, especially pp. 13-20) with descriptions of new species of Pectenura: P. excubitor, P. formosa, P. pecten, and P. senta. Also isolated lateral element from a borehole, Devonian of Germany, by Haude (2004, on p. 244) [Haude, R. 2004. Cour. Forsch.-Inst. Senckenberg 251:237-251]. Regarding Antiquaster magrumi add: Kesling & Chilman (1975, see pp. 168, 176, and plate 39) [UMMP Papers on Paleontology No. 8]. For redescription of Protasteracanthion and P. primus, and Antiquaster made a synonym of Protasteracanthion, see Alexander Glass (2006. Ph.D. Thesis. The brittle star fauna of the Hunsruck Slate and a phylogeny of the Paleozoic Ophiuroidea. Univ. Illinois at Urbana-Champaign. For abstract see Dissertation Abstracts International 67(11B):6270].

Hotchkiss, F.H.C. 2000. On the number of rays in starfish. American Zoologist 40:340-354. Correction to the statement concerning holothurians on p. 340: the tentacles of holothurians correspond to the rays of other echinoderms, and holothurians have FIVE-PLUS organization of rays/tentacles around the mouth (Hotchkiss 1998:rays as appendages, p.208). The 8-rayed specimen of Luidia senegalensis on p. 345 is deposited in the Smithsonian National Museum of Natural Histsory as USNM 1112625. Further evidence that tetramery at metamorphosis is the result of teratological incomplete development (p. 349) comes from an Arbacia punctulata that developed 4-part symmetry while its genetically identical twin developed 5-part symmetry (the parents had typical 5-part symmetry; Marcus, N.H. 1979. Developmental aberrations associated with twinning in laboratory-reared sea urchins. Developmental Biology 70:274-277). Likewise in the ophiuroid Amphipholis squamata, deviations from pentamerism are not heritable but are a consequence of environmental perturbations on the metamorphosis of larvae and/or abnormal regeneration of arms [Dupont, S. & J. Mallefet. 2002. Abnormal forms in the brittle-star Amphipholis squamata: a field study. J. Mar. Biol. Ass. U.K. 82:491-493]. Some additional references: 2000. Sanchez, P. The sequence of origin of the postmetamorphic rays in Heliaster and Labidiaster (Echinodermata: Asteroidea). Revista Chilena de Historia Natural 73:573-578. 2007. Herringshaw, L.G., M.P. Smith and A.T. Thomas. Evolutionary and ecological significance of Lepidaster grayi, the earliest multiradiate starfish. Zoological Journal of the Linnean Society 150:743-754.

Hotchkiss, F.H.C. 2000. Inferring the developmental basis of the sea star abnormality “double ambulacral groove” (Echinodermata: Asteroidea). Revista Chilena de Historia Natural 73:579-583. The specimen (p. 582) from Marblehead, MA of Asterias vulgaris with double ambulacral groove on one of only four rays is deposited in the collections of the California Academy of Sciences, CASIZ 173969. The specimen (p. 581) from a student at Lehigh University of Asterias forbesi with double ambulacral groove on one of its five rays is deposited in the US National Museum, Smithsonian Institution, echinoderm collection, USNM 1082952. The Zoological Record for 1904, p. 34, mentions fusion of rays in Ctenodiscus, referencing Michaailovskij, M. 1904. Die Echinodermen der zoologischen Ausbeute des Eisbrechers “Jermaks” von Sommer 1901. Annuaire Mus. St. Petersb. ix, pp. 157-188. The research collection of sea stars of H. Barraclough Fell contained a Pentagonaster pulchellus with double ambulacral groove. This collection is now at the US National Museum.

Hotchkiss, F.H.C. and R. Haude. 2004. Observations on Aganaster gregarius and Stephanoura belgica (Ophiuroidea: Ophiolepididae) (Early Carboniferous and Late Devonian age). Pp. 425-431 In Heinzeller & Nebelsick (eds.), Echinoderms:München. Taylor & Francis Group, London. The purpose of the research was to elucidate the structure of the oral frame, the structure of the distal parts of the rays, and the status of radial shields. It was noted that this was partly accomplished with new material. More specimens need to be examined to substantiate and expand the findings, especially any observations on mouth papillae. MPRI 0077 is a slab that has five fairly intact disks and many isolated portions of arms of A. gregarius. One specimen, in oral view, preserves one arm out to the 18th arm segment. This arm confirms that the under arm plates of Aganaster abruptly cease after the 15th arm segment, and that podial pores perforate the lateral arm plates beginning on the 16th arm segment. This arm increases the number of observations of this question to N = 3. To be noted in passing: Pearse (1947. Zoological names - a list of phyla, classes, and orders) listed Order Aganasterida H.L. Clark 1939. I have not found the 1939 H.L. Clark paper, in spite of earnest searching. The Order Aganasterida is listed also by Boettger (1952) and by Blackweldeer (1963).

Hotchkiss, F.H.C., Rudolf J. Prokop and Václav Petr. 2007. Isolated ossicles of the Family Eospondylidae Spencer et Wright, 1966, in the Lower Devonian of Bohemia (Czech Republic) and correction of the systematic position of eospondylid brittlestars (Echinodermata: Ophiuroidea: Oegophiurida). Acta Musei Nationalis Pragae, series B Historia Naturalis 63(1):3-18. MPRI contribution No. 1. For additional mention of Eospondylus in the Hunsrück Slate see Südkamp (1985, and especially 1995 documenting Eospondylus on crinoids). Further to the postscript on an eospondylid ophiuroid of eyelet-type in the Arkona Shale, note similarities of preservation and fauna between the Arkona Shale and the Hunsrück Slate [see e.g.: Úna C. Farrell and Derek E. G. Briggs, 2007, A pyritized polychaete from the Devonian of Ontario; Proc. R. Soc. B 274:499–504. the polychaete in this paper was collected by Michael and John Topor and donated by them to UMMP].

Hotchkiss, F.H.C. 2008. Pattern formation in starfish: arm stumps, regeneration models, and evolution. MPRI contribution No. 3. Poster presentation prepared for the Fifth North American Echinoderm Conference, Melbourne, Florida, July 2008, and also the 4th Workshop of German and Austrian Echinoderm Research, Vienna, October 2008. Other published mentions of ray stumps that do not regenerate: Ruedemann (1916:61) described specimens of Hallaster forbesi with “relatively short, somewhat abruptly terminating rays” as probably arm stumps that failed to regenerate. However Spencer (Monograph p. 298) showed that the arms are flexed aborally giving only a false appearance of being stumps. The abruptly terminating arms of Cholaster peculiaris, which is known from a single specimen, are also the product of arms bent 180 degrees onto themselves (FHCH unpublished); so these are not arm stumps that did not regenerate. Additional examples of regenerating Ordovician starfish: Urasterella sp., ROM 53376, Middle Ordovician, Bobcaygeon Formation, near Lake Simcoe, Ontario, Canada (personal communication, David Rudkin, 4/15/2008). Cnemidactis girvanensis (Schuchert), counterpart specimens BMNH E 52484a,b; Upper Ordovician (Ashgillian), Thraive Glen, Ayreshire, Scotland; figured by W.K. Spencer (1918:161 and pl. XII fig. 5); Owen (1965:565 curatorial notes). Modes of wound closure: FHCH has observed dorso-ventral downward movement like closing a roll-top desk (D-V wound closure) in Astropecten following autotomy. Previous research/comments on symmetric versus asymmetric positional information in the starfish arm: Lovén’s rule and Batesonian duplications of appendages imply asymmetrical positional information. David, Mooi & Telford (Echinoderm Research 1995: 163): “In groups such as Asteroidea, in which the paired plates of the ambulacral columns are directly opposite and do not alternate, it should still be possible to see from early ontogenetic stages in which columns the first plate is laid down, thereby testing the existence of Lovén’s rule even in this group.” Hotchkiss (1998: 207) emphasized that no case of Batesonian duplication of asteroid rays has ever been reported, and he rejected the implications of this negative finding. The implied expectation of these researchers was that positional information in the starfish ray is asymmetric. Impact on the rays-as-appendages hypothesis: Symmetrical positional information in the starfish ray is not consistent with the rays-as-appendages model which explicitly invokes asymmetrical positional information. Either the rays-as-appendages hypothesis should be rejected, or we need some sort of hypothesis-saving explanation. I opt for the latter because the fossil record indicates that alternating ambulacral plates and Loven’s law are plesiomorphic, and these features imply asymmetric positional information. Somehow, the starfish arm has changed from an original asymmetric condition to a symmetric condition. Other examples of failure to regenerate following L-R bilaterally symmetrical wound closure and stump formation occur in planarians (headless planarians): Child, C.M. 1911. Experimental control of morphogenesis in the regulation of Planaria. Biological Bulletin 20:310-331. Della Valle, P. 1914. L’inibizione della rigenerazione del capo nelle planarie mediante la cicatrizzazione. Archivio Zoologico Italiano 7:275-311.

some references to birds/turtles drumming for worms

American woodcock. " Later, at 10:30 a.m., this bird and another fed in bright sunshine on the lawn, nonchalantly pulling worms from the sod. They had found the only square footage in town that was frost free. Their side-to-side- shifting, accompanied by deliberate weight shifts from foot to foot, must cause enough vibration to stir the worms below." Joe Choiniere, director of Mass Audobon's Wachusett Meaadow Wildlife Sanctuary. Seasons of the woodcock: the secret life of a woodland shorebird. Sanctuary, The Journal of the Massachusetts Audubon Soociety, Summer 2006, 45(4):3-5. [see p. 5]

Gull in grass park beside Lake Merritt, Oakland, California, spring 1991. Observations by F. Hotchkiss communicated to Ned Newton. "... on the grass, to the front and left of me, only eight feet from the sidewalk, was a gull. The gull seemed to be stepping in place. I stopped to watch it on the grass. It stepped again... it did it clearly with an alternate, rapid step -- stepping in place. In between this stepping it would stop -- it would peer at the ground just in front of itself, but never walking -- just with its feet stationary, planted -- and I would then see it grab an earthworm with its beak -- and quickly swallow it -- and look for another -- and then resume the stepping in place -- without moving an inch. This gull was a talented earthworm drummer. How on earth did it ever learn to do this!!" Ned Newton added the following comments: "Well, it seems that I have heard of Robins and Woodcock drumming up worms. But it must have been fun to see a relatively large bird like a gull doing this. I was telling a friend from Connecticut about the drumming for worms. She was not at all surprised and added that she has seen turtles doing this same thing -- hopping from one foot to another. Somehow they don't have the speed that I would think is required. Anyone who has been out at night catching nightcrawlers knows that you have to be fast. Well, maybe the next time the kids are dancing it would be interesting to look around near by or on the flood and see if worms are raised to the surface." Newton's Notes, April 26, 1991 [Notes from the President], Nashoba Valley Bird Club, The Meadow Plover 2(2) Spring 1991.

For information on turtles stomping for earthworms see:

Ernst, C. H., J. E. Lovich, and R. W. Barbour. 1994. Turtles of the United States and Canada. Washington, D.C.: Smithsonian Institution Press. [summary of information]

Kaufmann, J. H. 1986. Stomping for earthworms by wood turtles, Clemmys insculpta: a newly discovered foraging technique. Copeia 1986(4):1001-1004.

Kaufmann, J. H., J. H. Harding and K. N. Brewster. 1989. Worm stomping by wood turtles revisited. Bull. Chicago Herpetol. Soc. 24: 125-126.

Kirkpatrick, David T. and Catherine Kirkpatrick. 1996. Stomping for earthworms by Clemmys insculpta in captivity. Bulletin of the Chicago Herpetological Society 31(2):21-22.

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Frederick Hotchkiss, PhD annotations to published articles Hotchkiss, F.H.C. 1977. Ophiuroid Ophiocanops (Echinodermata) not a living fossil. Journal of natural History 11: 377-380. Additional comments in Hotchkiss (1995): Ophiocanops does not have the 'auluroid' vertebrae of the Oegophiurida (p. 423); the extraordinary gonadal and stomachal characters perhaps are retained from the Oegophiurida; hence perhaps a surviving member of the stem group of the Order Phrynophiurida (p. 415, 423 and Fig. 6). Am aware of the following 2001 paper but have not seen it: Fujita, T., S. Irimura & W.W. Kastoro. Biology of a rare ophiuroid Ophiocanops fugiens Koehler, 1922 (Echinodermata) associated with black corals, with notes on the specimens collected from Lembeh Strait, Bitung, Indonesia. In: Terazaki, M., A. Taira, M. Uematsu, Y. Michida and T. Kaneko, eds., Proceedings of the 11th JSPS Joint Seminar on Marine Science, Center for International Cooperation, Ocean Research Institute, University of Tokyo, Tokyo, 2001, pp. 326-333. For more on this topic see 2008 paper by Sabine Stöhr, Chantal Conand and Emilie Boissin: Brittle stars (Echinodermata: Ophiuroidea) from La Réunion and the systsematic position of Ophiocanops Koehler, 1922. Zoological Journal of the Linnean Society, 2008, 153:545-560 [they describe Ophiocanops multispina n.sp., transfer Renetheo felli to Ophiocanops, and present many new observations on morphology including upper arm plates, a second under arm plate, side arm plates that meet ventrally so that there is no resemblance to the ambulacral groove of Paleoozoic Oegophiurida, and place Ophiocanops in the Ophiomyxidae]. Hotchkiss, F. H. C. 1979. Case studies in the teratology of starfish. Proceedings of The Academy of Natural Sciences of Philadelphia 131:139-157. The historical review on pp. 139-143 is only a sampling of what could have been included. Some additional references, in historical order, follow: 1882. Bell, F.J. Note on the echinoderm fauna of the Island of Ceylon, together with some obversations on heteractinism. Ann. Mag. Nat. Hist., ser. 5, 10:218-225; 1887. Bell, F.J. The echinoderm fauna of the island of Ceylon. The Scientific Transactions of the Royal Dublin Society, series 2, 3:643-657, pls. 39-40. [an oblique cut regenerates obliquely to the arm axis]; 1912. Richters, C. Zur Kenntnis der Regenerationsvorgange bei Linckia. Zeitschrift fur Wissenschaftliche Zoologie 100:116-175; 1914. Schapiro, J. Uber die Regenerationserscheinungen verscheidener Seesternarten. Arch. f. Entwmeck. 38:210-251; 1915. Nusbaum, J. & M. Oxner. Zur Restitution bei dem Seestern Echinaster sepositus Lam. Zool. Anzeiger 46(6):161-167; 1917. Zirpolo, G. Notizia di alcuni Asteroidi anomali pescati nel Golfo di Napoli, Echinaster sepositus Gray e Asterias glacialis O.F. Muller. Bollettino della Societa di Naturalisti in Napoli 30:20-29; 1918. Zirpolo, G. Un caso di rigenerazione parziale delle braccia in un Astropecten aurantiacus L. Pubblicacazioni della Stazione Zoologica di Napoli 2(2):169-175, pl. 10 [a 5-rayed individual loses two adjacent rays and some disk; only a single ray regenerates, resulting in a 4-rayed individual]; 1924. Zirpolo, G. Ulteriori notizie di Asteroidi anomali. Bollettino della Societa di Naturalisti in Napoli 36:305-346, pls. 6-8; Zirpolo, G. Notizia di un Echinaster sepositus Gray con sei braccia nel Golfo di Napoli. Atti Pontif. Accad. Nuovi Lincei, Anno 77:161-163. [No external or internal irregularities, providing his argument for hexamery from the first moment of life]; 1926. O'Donohue, C.H. On the summer migration of certain starfish in Departure Bay, B.C., Canada. Fisheries Research Board of Canada. Contributions to Canadian Biology and Fisheries 1:455-472; Zirpolo, G. Gemmazioni, rigenerazioni ipertipiche ed ipotipie studiate nell'Astropecten aurantiacus L. Bollettino della Societa di Naturalisti in Napoli 38:167-221 [includes examples of tetramery arising when two arms regenerated in place of three, and when one arm regenerated in place of two]; 1929. Zirpolo, G. Le forme cometoidi dell'Asterias tenuispina Lmk. Bollettino della Societa di Naturalisti in Napoli 40:25-35; Zirpolo, G. Nuovo caso di gemmazione in un Astropecten aurantiacus L. Bollettino della Societa di Naturalisti in Napoli 40:83-87; Zirpolo, G. Notizia di Asteroidi irregolari. Bollettino della Societa di Naturalisti in Napoli 40:221-231; 1931. Abeloos, M. Les potentialites regeneretrices de la face dorsal des bras des Asteries, Trifurcation dorsal d'un bras chez Solaster papposus (Linck). Bull. Biologique de la France et de la Belgique fonde par Alfred Giard, 65(3):394-405. 1933. Mortensen, Th. The Godthaab Expedition, 1928, Echinoderms. Meddelelser om Gronland 79(2):1-62 + plate I. [on p. 17 arm number in Solaster papposus: nearly all have 10 arms, one has 12, a few have 9, one has 8 arms]; 1938. Domantay, J.S. An unusual bud due to heteromorphosis in Echinaster luzonicus (Gray). Phillipp. J. Sci. Manila 64:281-283; 1941. Fisher, W.K. A new genus of sea stars (Plazaster) from Japan, with a note on the genus Parasterina. Proc. USNM 90(3114):447-456, pls. 66-70; 1945. Fisher, W.K. Unusual abnormalities in sea-stars. J. Wash. Acad. Sci. 35:296-298; 1951. Alfano, B. Commemorazione del Prof. Giuseppe Zirpolo. Bollettino della Societa di Naturalisti in Napoli 59:39-64 [on the life and publications of G. Zirpolo]; 1958. Dollfus, R.Ph. Courbe interbrachiale chez une asterie du genre Stellasteropsis. Bull. Soc. Zool. Fr. 83:294-297 [compares 4-armed and 5-armed individuals]; 1967. Raup, D.M. & E.F. Swan. Crystal orientation in the apical plates of abberant echinoids. The Biological Bulletin 133(3):618-629 [ocular IV and genital 4 are judged missing in two four-part urchins; of special interest because interradius IV/V is the location of closure of the hydrocoele crescent to form the ring canal; ray IV is on the ventral or posterior horn of the hydrocoele crescent]; 1969. Kenny, R. Growth and asexual reproduction in the starfish Nepanthia belcheri (Perrier). Pacific Science 23(1):51-55. 1975. Hinegardner, R.T. Morphology and genetics of sea urchin development. American Zoologist 15:679-689 [breeding experiments and loss of symmetry control]; 1986. Horowitz, A., S. Able, & H.L. Strimple. Abnormalities in Pentremites Say (Blastoidea) from the Pella Formation (Upper Mississippian) of Iowa. Journ. Paleontology 60(2):390-399; 1989. Ausich, W.I. & T.W. Kammer. Teratological specimen of Agaricocrinus americanus (Roemer) (Lower Mississippian, Crinoidea). J. Paleont 63(6):945-946; 1991. Boursin, J. Un curieux (b)oursin ou coussin? Xenophora, number 53:25 [square 4-armed Halityle regularis from New Caledonia]; 1992. Ramalingam, K. & A. J. Antony. A note on the tetramerous starfish (Oreaster sp.). Comparative Physiology and Ecology 17(3):121-122; 1999. James, D.B. Abnormal asteroids from the seas around India. Mar. Fish. Infor. Ser., T. & E. Ser., number 158:21-22; 2000. Hotchkiss, F.H.C. On the number of rays in starfish. American Zoologist 40;340-354 [this paper inludes additional references not repeated here, as well as a new synthesis for the topic]; 2002. Dupont, S. & J. Mallefet. Abnormal forms in the brittle-star Amphipholis squamata: a field study. J. Mar. Biol. Ass. U.K. 82:491-493 [deviations from pentamerism are not a heritable character but are a consequence of environmental perturbations on the metamorphosis of larvae and/or abnormal regeneration of arms]; 2007. Ceranka, T. Symmetry disorders of the test of the Miocene echinoid Echinocyamus from Poland. Acta Palaeontologica Polonica 52(3):503-518. Behavioral evidence for a physiological anterior in locomotion in starfish was examined on pp. 153-154. Concerning behavioral evidence for nonrandom use of arms in ophiuroids see: Welte, N.T. & L.M. Lutton. 2003. Appendage selection during brittlestar locomotion. Journal of the Pennsylvania Academy of Science 77(1):15-19. This paper revealed a pattern of two adjacent arms being used least for leading and most for propulsion. The identity of the arms with respect to the location of the madreporite was not examined and deserves further study. Hotchkiss, F.H.C. 1980. The early growth stage of a Devonian ophiuroid and its bearing on echinoderm phylogeny. Journal of Natural History 14:91 96. The paper documented primary rosette plates in an oegophiurid with alternating ambulacral plates (Lysophiurina). The Lysophiurina and Zeugophiurina lineages diverged no later than Early Ordovician time. The Zeugophiurina are the presumed stem group of the Silurian-Recent Ophiurida. Documentation of a primary rosette in Paleozoic Ophiurida can be mentioned: in Ophiaulax decheni (Devonian, Ophiurinidae) [R. Haude & E.T. Thomas. 1983. Ophiuren (Echinodermata) des hohen Oberdevons im nördlichen Rheinischen Schiefergebirge. Paläont. Z. 57(1/2):121-142]; in Aganaster gregarius (Mississippian, Ophiolepididae) [F.H.C. Hotchkiss & R. Haude. 2004. Observations on Aganaster gregarius and Stephanoura belgica. Pp. 425-431 in Heinzeller & Nebelsick (eds.), Echinoderms: München]. Additional references on postlarval growth stages in Recent ophiuroids include: 1998. Sumida, P.G. and P.A. Tyler. postlarval development in shallow and deep-sea ophiuroids (Echinodermata: Ophiuroidea) of the NE Atlantic Ocean. Zoological Journal of the Linnean Society 124:267-300. 2005. Stöhr, S. Who's who among baby brittle stars (Echinodermata: Ophiuroidea): postmetamorphic development of some North Atlantic forms. Zoological Journal of the Linnean Society 143:543-576 [includes information on the euryalid Asteronyx loveni: the plates of the primary rosette are conspicuous at first, but do not endure: at 3.2 mm disk diameter they are small fragments at the center of the disk]. Hotchkiss, F. H. C. 1982. Ophiuroidea (Echinodermata) from Carrie Bow Cay, Belize. Pp. 387-412 in The Atlantic Barrier Reef Ecosystem at Carrie Bow Cay, Belize, I. Structure and Communities. K. Rutzler and I.G. Macintyre, editors. Smithsonian Contributions to the Marine Sciences, No. 12. Corrections to identifications in this paper were published by Hendler & Peck (1998. Ophiuroids off the deep end: Fauna of the Belizean fore-reef slope. Pp. 411-419 in Echinoderm Biology, Burke et al. [eds.], Balkema, Rotterdam), as follows: Ophiomitrella glabra (H.L. Clark) is regarded as a junior synonym of Ophioblenna antillensis, Axiognathus squamatus (Delle Chiaje) as a junior synonym of Amphipholis squamata, and the six-rayed fissiparous Ophiostigma sp. was nominally referred to Ophiostigma isocanthum [but see below]. The Ophioderma sp. juvenile is Ophioderma rubicundum, and the Sigsbeia murrhina Lyman sensu Hotchkiss specimens are Sigsbeia conifera Koehler. The Ophiurochaeta littoralis Koehler sensu Hotchkiss specimens were reidentified as an undescribed species of Ophiurochaeta [four undescribed Ophiurochaeta species provisionally designated A, B, C and D are mentioned by Hendler and Peck]. In a subsequent paper the six-rayed fissiparous Ophiostigma sp. was described as the new species Ophiostigma siva by Hendler [1995. New species of brittle stars from the Western Atlantic, Ophionereis vittata, Amphioplus sepultus, and Ophiostigma siva, and the designation of a neotype for Ophiostigma isocanthum (Say) (Echinodermata: Ophiuroidea). Contributions in Science, Natural History Museum of Los Angeles County No. 458]. Hotchkiss, F.H.C., S.E. Churchill, R.G. Gelormini, W.R. Hepp, R.J. Rentler & M.T. Tummarello. 1990. Events of autotomy in the starfish Asterias forbesi and A. vulgaris. Presented at 7th International Echinoderms Conference, Sept. 9/14, 1990, Atami, Japan. Published 1991, pp. 537-541 in T. Yanagisawa et al. (eds.), Biology of Echinodermata, A.A. Balkema, Rotterdam, 590 pp. Hotchkiss, F. H. C. 1993. A new Devonian ophiuroid (Echinodermata: Oegophiurida) from New York State and its bearing on the origin of ophiuroid upper arm plates. Proceedings of the Biological Society of Washington, vol. 106, no. 1, pp. 63 84. To the comment on p. 73 regarding the very straight arms of the holotype and the existence of a stiffening reaction in ophiuroids, see remarks of W.K. Spencer (Monograph p. 246 citing von Uexkull, and p. 316 on Furcaster with arms stretched out very stiffly) that a very strong stimulus to the arm produces a rigor ... and the arms feel like small sticks. On the subject p. 75 that upper arm plates do not stay with the arm segments that become included in the disk, Ishida & Inoue (1993:104 Ophiura sarsii) noted that "about 4 dorsal arm plates at the base of the arm are incorporated into the disk". Further to the evidence p. 76 against the idea that upper arm plates get incorporated into the radial shields, note A.M. Clark 1966 Echinodermata [Port Phillip Survey 1957-1963], Memoirs of the National Museum, Melbourne No. 27, pp. 289-384, on p. 337 regarding Amphiura (Ophiopeltis) parviuscutata that basal arm segments have their dorsal arm plates reduced or even absent inside the disk area. Regarding the possible origins and homologies of the radial shields, these were not treated comprehensively on pp. 75-76 except where there was some suggestion that related radial shields to upper arm plates. For other origins of radial shields see for example Spencer (Monograph p. 244) and Matsumoto (Monograph p. 369). The statement of a reviewer that "it is incorrect to infer from Lyman's illustrations that radial shields grow by adding platelets" is true only for ophiurid ophiuroids in which radial shields grow from a single tiny scale which initially formed at the edge of the disk. Brian Stewart showed that the radial shields of Astrobrachion constrictum grow by proximal sequential addition of scales of stereom to the proximal ends (1966. Growth dynamics of the radial shields of the euryalid snake star Astrobrachion constrictum. Invertebrate Biology 115(4):321-330). On the subject of retaining for the moment the two suborders Lysophiurina and Zeugophiurina p. 80, note W.K. Spencer (Monograph p. 9) on " remarkable analogous (homoplastic) course of development" of two series, and that "For the present both series are included in the Ophiuroidea". I did not mean to suggest that Strataster might in some way be ancestral to crown group ophiuroids. The evidence seen in Strataster helps us understand the origin of upper arm plates in all lineages of ophiuroids. A paper by Turner & Heyman, 1995, Proc. Biol. Soc. Wash 108(2):292-297 has important observations on the presence of radial shields and dorsal arm plates in Ophiosyzygus and many other ophiomyxids. Hotchkiss, F.H.C. 1995. Lovén's law and adult ray homologies in echinoids, ophiuroids, edrioasteroids and an ophiocistioid (Echinodermata: Eleutherozoa). Proceedings of the Biological Society of Washington [DC], vol. 108, no. 3, pp. 401 435. Lovén’s Rule in the echinoid order Cidaroida: Lovén’s Rule is evident in the pattern of alternation of the ambulacral plates that begins at the edge of the peristome in a 7 mm diameter specimen of Histocidaris elegans (A. Agassiz) figured by Mortensen (1927: Fig. 6). Reference: Mortensen, Th. 1927. Report on the Echinoidea collected by the United States Fisheries Steamer “Albatross” during the Philippine Expedition, 1907-1910. Part 1 The Cidaridae. USNM Bulletin 100, vol. 6, part 4, pp. 243-312, pls. 48-80. Lovén’s Rule in the echinoid order Temnopleuroida: in Temnopleurus apodus only the buccal plates Ib, IIb, IIIa, IVb, and Va carry tubefeet, the other buccal plates being quite rudimentary (Mrtsn 1943 Monograph III.2 p. 101); in Paratrema doederleini with only five buccal plates, the five buccal plates that are present are the plates Ib, IIb, IIIa, IVb, Va (R. T. Jackson, 1927, p. 451, fig. 8; confirmed by Mrtsn 1943 Monograph III.2 p. 274; a specimen with only four buccal plates has the A. II buccal plates missing); however in Prionechinus saggitiger with only five buccal tube feet the pattern is bbbba (Agassiz, 1881, Challanger Report, pl. VIa, fig. 12; numbering of ambulacra not stated). Lovén’s Rule in the echinoid order Echinoida: In Strongylocentrotus droebachiensis and in Psammechinus miliaris the buccal tube-feet first appear in the plates Ib, IIb, IIIa, IVb, Va (Jackson 1927, pp. 451-452; David & Mooi, 1995, The ontogenetic basis of Lovén’s Rule clarifies homologies of the echinoid peristome, pp. 155-164, in Echinoderm Research 1995, Emson, Smith & Campbell (eds), Balkema, Rotterdam; key reference on Lovén’s Rule in echinoids and other Echinodermata). Lovén’s Rule in the echinoid order Holasteroida: the largest ambulacral basicoronals in pourtalesiids definitely exhibit the aabab pattern early in ontogeny (David & Mooi 1995). To the list of references concerning Fell's identification of Kirk's ophiuroid as Ophiomyxa (p. 415), add that his first mention was in his 1962 key to the genera of ophiuroids. Additional and more recent sources of information relating to Table 2 include: Emlet, R.B. 2006. Direct development of the brittle star Amphiodia occidentalis (Echinodermata,Ophiuroidea, Amphiuridae) from the northeastern Pacific Ocean. Invertebrate Biology 125(2): 154-171 (place of hydrocoele closure and location of the stone canal). Further to evidence on changes of symmetry axes (Note 5): In the Order Echinoida, the oval test of Parasalenia gratiosa var. boninensis is elongate in the IIIb-5b direction, that of Echinometra mathaei in the 3a-Ia direction, that of Hetrocentrotus mamillatus and of H. trigonarius in the IVb-1b direction, and the test of Colobocentrotus mertensii is elongate in the 4a-1b direction using Lovén's nomenclature (Hayato Ikeda, 1939, The shape of the test of Colobocentrotus mertensii Brandt (Echinoidea, Echinometridae), Annotationes Zoologicae Japonenses 18(3):194-201, pls. 9-10). A specimen of C. mertensii with its long and short axes apparently interchanged, as judged by the location of the madreporite, may be a case of situs inversus (unpublished suggestion of FHCH; specimen described by Hiroshi Ohshima, 1939, Some rare abnormalities in asteroids and echinoids,Zool. Mag. Tokyo, 51:158-162). The ovoid test of Tiarechinus can also be mentioned (Treatise on Invertebrate Paleontology 1966 Part U pp. U436-U437). The test is circular but the apical system is elongate in the III-5 direction in Orthopsis ruppellii, Order Orthopsida (Treatise fig. 326,1h), likewise in Glyphocyphus (G.) radiatus, Order Temnopleuroida (Treatise pp. U414-415). The details for SET I specimen numbers 106-112 are listed below: 106. Protasteridae, Eugasterella africana Jell & Theron, South African Museum, Cape Town, paratype, SAM K1015, figured Jell & Theron (1999:177, Fig. 48A), madreporite in III/IV interradius. 107. Protasteridae, Eugasterella africana Jell & Theron, South African Museum, Cape Town, paratype, SAM K1014, figured Jell & Theron (1999:177, Fig. 48C) , madreporite in III/IV interradius. 108. Protasteridae, Eugasterella africana Jell & Theron, Geological Survey of South Africa, Bellville in Capetown No. B4561B, paratype, figured Jell & Theron (1999:177, Fig. 48B), madreporite in III/IV interradius. 109. Protasteridae, Eugasterella africana Jell & Theron, Geological Survey of South Africa, Bellville in Capetown No. B4561D, paratype, figured Jell & Theron (1999:176, Fig. 47C), madreporite in III/IV interradius. 110. Protasteridae, Eugasterella africana Jell & Theron, Geological Survey of South Africa, Bellville in Capetown No. B4561a, holotype, figured Jell & Theron (1999:175, Fig. 46A), madreporite in III/IV interradius. 111. Protasteridae, Strataster ohioensis Kesling & LeVasseur, , Geological Survey of South Africa, Bellville in Capetown No. B0196, figured Jell & Theron (1999:181, Fig. 51C), madreporite in III/IV interradius. 112. Cheiropterasteridae, Hexuraster weitzi (Spencer), Roy Oosthuizen Collection, Zwartskraal, Prince Allbert No. RO 45, figured Jell & Theron (1999:163, Fig. 39) , madreporite in III/IV interradius. The details for SET II specimen numbers 215-219 are listed below: 215. Protasteridae sp. Specimen not figured by Jell & Theron. Specimen details still not known. Arms score ?ABA? 216. Eophiuridae, Haughtonaster reedi Rilett, Geological Survey of South Africa, Bellville in Capetown No. B4567, figured Jell & Theron (1999:158, Fig. 36B). Arms score AA?AB 217. Protasteridae, Strataster ohioensis Kesling & LeVasseur, , Geological Survey of South Africa, Bellville in Capetown No. B4512, figured Jell & Theron (1999:183, Fig. 53). Arms score AAB?? 218. Protasteridae, Strataster ohioensis Kesling & LeVasseur, , Geological Survey of South Africa, Bellville in Capetown No. B4513, figured Jell & Theron (1999:182, Fig. 52D). Arms score ?AB?B 219. Protasteridae, Strataster ohioensis Kesling & LeVasseur, , Geological Survey of South Africa, Bellville in Capetown No. B4513, figured Jell & Theron (1999:179, Fig. 49b). Arms score AABA? NEW OBSERVATION: Specimen SUG299 of Encrinaster tischbeinianus (Roemer), Family Encrinasteridae, from De Doorns, Ceres District, Lower Devonian Bokkeveld Group, property of the Geological Collections, Stellenbosch University, South Africa, oral surface, figured Jell & Theron (1999:164, Fig. 40B-D): The specimen has a madreporite (enlarged in their Fig. 40D; madreporite not recognized by FHCH in the contact print in 1994). Using a magnifier to examine their Fig. 40C, the arms score AABAB. The five arms conform with Lovén’s AABAB rule. The madreporite is located in the III/IV interradius. The specimen fulfills SET I criteria and is assigned Set I reference number 113. Including the new observation, Lovén’s rule applies to the ophiuroid families Eophiuridae, Cheiropterasteridae, Encrinasteridae and Protasteridae (not just the Protasteridae as originally reported). Hotchkiss, F.H.C., D. M. Rudkin, and S. Anderson. 1997. A case for rearmament -- the oldest known evidence of regeneration in sea stars (Abstract). 7th Canadian Paleontology Conference, University of Saskatchewan, Saskatoon, September 26-30, 1997. [Also: Rudkin, D. M., F. H. C. Hotchkiss and S. Anderson. A case for rearmament -- the oldest known evidence of regeneration in sea stars. Royal Ontario Museum Nineteenth Annual Research Colloquium, 20 November 1997, Toronto, Abstracts of Papers, p. 1 (abbreviated abstract).] We reported a specimen of Promopalaeaster wilsonae (Raymond 1912) with two regenerating arm tips, collected by S. Anderson from the Upper Member of the Bobcaygeon Formation (Middle Ordovician: Kirkfieldian) of south-central Ontario, and now housed in the Royal Ontario Museum (ROM 53320). The Anderson fossil is the oldest [earliest] example of an asteroid with a regenerating ray, supplanting a regenerating specimen of Cnemidactis girvanensis from the Upper Ordovician of Girvan, Scotland reported by Spencer (1918 pt. III, p. 161, pl. 12 fig. 5). Arm stumps that had healed without regenerating the lost arm were described in the Ordovician starfish Urasterella ulrichi by Schuchert (1915:37). Hotchkiss, F.H.C. 1998. *Discussion on pentamerism: The five-part pattern of Stromatocystites, Asterozoa, and Echinozoa. Pp. 37-42 In R. Mooi and M. Telford (eds.) Echinoderms: San Francisco. A.A. Balkema, Rotterdam. Postscript notes: Nichols (1967b) noted that the crystallographic alignment hypothesis of Cockbain (1966) is not consistent with c-axis orientations in echinoid genital plates; clarified and expanded arguments that support the suture-line hypothesis. There is much additional literature* relevant to this topic that could not be fitted into the original paper; also there is literature that has come out since the paper. 1889. Bather, F.A. Trigonocrinus, a new genus of Crinoidea, from the "Weisser Jura" of Bavaria; with the description of a new species, T. liratus.--Appendix. Sudden deviations from normal symmetry in Neocrinoidea. Quarterly Journal of the Geological Society for February 1889, pp. 149-171, pl. 6 [p. 166 pentamery persists through natural selection based on mechanical principles; in pentamerous calyx every line of weakness is met halfway by a solid plate]. 1915. Clark, A.H. A monograph of the existing crinoids. The comatulids. USNM Bull. 82, vol. 1, part 1, 406 pages [p. 148: sutures are lines of weakness; with an odd number of radial divisions the sutures do not line up and the construction is stronger; the relation between the bilateral crustacean type and the pentaradiate echinoderm type is explained and provides explanation for the establishment of pentaradial symmetry]. 1928. Bather, F.A. The fossil and its environment. Quarterly Journal of the Geological Society of London 84(2):lxi-xcviii [pp. lxxi-lxxii present his ideas on the origin in Pelmatozoa of a primitive trisactiny followed by a forking of the lateral rays; a pentactiny superimposed upon a trisactiny; states the advantage of tripartite and quinquepartite over quadripartite: each line of weakness abuts on a mass of strength; calling it the principle of bonding] 1975. Strathman, R.B. Limitations on diversity of forms: branching of ambulacral systems of echinooderms. The American Naturalist 109:177-190. 1987. Anderson, D.T. Developmental pathways and evolutionary rates. In: Rates of Evolution, K.S.W. Campbell & M.F. Day (eds.), Allen & Unwin, Sydney [p. 146 pentamerous symmetry determined during development by a counting process]. 1998. Tolić, Iva Marija and Nenad Trinajstić. The number five in biology. Periodicum Biologorum 100(3):259-266 [Fivefold symmetry in animals, plants, viruses, biomolecules, crystals (quasicrystals)]. 2001. Changizi, M.A. The economy of the shape of limbed animals. Biological Cybernetics 84:23-29 [wire-minimization principles, animal limb number, and body-to-limb proportion; predicts six or more rays for Asteroidea]. 2002. Barrio, R.A., P.K. Maini, J.L.Aragón and M. Torres. Size-dependent symmetry breaking in models for morphogenesis. Elsevier Physica D 168-169(2002):61-72 [discusses de novo appearance and control of pentagonal symmetry of echinoids]. 2006. Parsley, R.D. and Y. Zhao. Long stalked eocrinoids in the basal Middle Cambrian Kaili Biota, Taijiang County, Guizhou Province, China. J. Pallaeont. 80(6):1058-1071 [pp. 1069-1070 discuss the ontogeny and phylogeny of ambulacral symmetry, from a 1-1 pattern to a 2-2 pattern to a 2-1-2 pattern]. Hotchkiss, F.H.C. 1998. A "rays-as-appendages" model for the origin of pentamerism in echinoderms. Paleobiology 24:200-214. A gentle criticism of the idea that there are analogies between appendages of bilateral forms and the rays of echinoderms is thoughtfully presented by R. Mooi, B. David & G. Wray, 2005, Arrays in rays: terminal addition in echinoderms and its correlation with gene expression, Evolution and Development 7(6):542-555. I accept their comments and claim only the analogy; the phrase rays-as-appendages seemed apt because Baterson's rule of duplication was discovered from analysis of duplicated appendages, and does not apply to axial structures; however Bateson's rule is not limited to appendages (it operates also in ciliate protozoa). Further to the mention on p. 203 of the paper by Hinegardner (1975): The report of loss of symmetry control and the rearing of four-part "square" Lytechinus pictus challenges the working hypothesis of locked-in pentamerism. Therefore it is important to mention that these are highly inbred lines with high mortality as larvae and at metamorphosis, facts which suggest that the square imagos are the result of teratological incomplete development. I do not agree with D.G. Stephenson's (Proc. European Colloq. Echinoderms,Brussels, 1979) interpretation that Hinegardner's square sea urchins are the result of loss of symmetry as soon as domestication relaxes the constraints of natural selection; nor Stephenson's corollary that pentamerism must be extrtemely adaptive for echinoids. Stephenson also presented the idea that "pentamerism is an ancestral character retained because any disadvantages of pentamerism are less than the disadvantages of changing it", and claimed that "this possibility has been ruled out by Hinegardner's (1975) breeding experiments." I propose instead that the loss of symmetry reported by Hinegardner be interpreted along the lines described by John M. Opitz (1985. The developmental field concept. American Journal of Medical Genetics 21:1-11, see Final generalizations on pp. 9-10): "all primary malformations qua developmental field defects are causally nonspecific"; "most will be shown to be causally heterogeneous"; "the causes are many, but the final common developmental pathways are few"; "most primary malformations are anomalies of incomplete formation"; "most field defects are multifactorially determined (not 'inherited'), hence have a low empiric recurrence risk"; "because oligogenes of major effect do play a role in the developmental history of all fields, sooner or later Mendelian inheritance of a primary malformation should be expected and observed"; "most field defects occur per se but can be and frequently are components of complex syndromes, including aneuploidy syndromes"; "the epigenetically hierarchichical nature of field development is responsible for the fact that, regardless of how small or limited the embryonic primordium, so long as it is still a field, its development can be altered genetically and environmentally." A further reference to diatoms with pentagonal symmetry: P.A. Simms & R. Ross. 1990. Triceratium pulvinal and T. unguicularum, two confused species. Diatom Research 5(1):155-169 [figs. 13-16 of T. unguiculatum forma quinquangulum]. Further to the matrix of ray homologies and especially concerning the entry for Luidia: My primary source on the location of hydrocoel closure and the madreporite in Luidia is Bury 1895 Fig. 21 and text pp. 67-68 [Bury 1895 The metamorphosis of echinoderms. Q. J. Microsc. Sci. n.s. 38:45-136 + pls. 3-9] [this source is mentioned in the caption to my matrix of ray homologies, but change "Fig. 22" to "Fig. 21"]. The observations of Bury were latched onto by F.A. Bather [1915 Studies in Edrioasteroidea IX. The genetic relations to other echinoderms. Geol. Mag., n.s., dec. VI, vol. 2, pp. 393-403]. On page 401 “It is important to notice here that in an Asteroid larva (Bipinnaria asterigera) described by Bury, the [hydrocoel] closure does still take place in the M plane” and mentioned further on p. 402. Mentioned again by Horstadius [1939 Uber die Entwicklung von Astropecten aranciacus L. Pubb. Staz. Zool. Napoli 17(2):221-312]. On p. 276 he says that in all known starfish (including Astropecten) the madreporite is between hydrocoel lobes I-II, except in Luidia it is between lobes V and I. On p. 278 he appears to give his own confirmatory observations on this point in recounting his work on Luidia ciliaris. NOTE NOTE NOTE – his system of Roman numerals is customary numbering of the linear series of hydrocoel lobes as seen in developing imago of metamorphosing larva, so is different than in my matrix. His Roman I is my edrio and asteroid D; his Roman V is my edrio and asteroid C; his Roman II is my edrio and asteroid E. Some newer papers on the development of the echinoderm body plan: 2006: Swalla, B.J. Building divergent body plans with similar genetic pathways. Heredity 97:235-243. Hotchkiss, F.H.C., Rudolf J. Prokop and Václav Petr. 1999. Isolated skeletal ossicles of a new brittlestar of the Family Cheiropterasteridae Spencer, 1934 (Echinodermata: Ophiuroidea) in the Lower Devonian of Bohemia (Czech Republic). Journal of the Czech Geological Society 44:189-193. To the occurrences of the family Cheiropterasteridae, add the Silurian and Lower Devonian of Bolivia (Branisa, Leonardo. 1965. Los fosiles guis de Bolivia I. Paleozoico. Index Fossils of Bolivia I. Paleozoic. Servicio Geologico de Bolivia, Boletin No. 6, 282 pp., 80 pls. [?Loriolaster sp. p. 108 pl. xxii fig. 1] [?L. cf. L. mirabilis p. 128 pl. xxxii fig. 9]). The replacement name for Hexura Spencer, 1950, is Hexuraster Jell & Theron, 1999. Hotchkiss, F.H.C., Rudolf J. Prokop and Václav Petr. 1999. Isolated vertebrae of brittlestars of the Family Klasmuridae Spencer, 1925 (Echinodermata: Ophiuroidea) in the Devonian of Bohemia (Czech Republic). Journal of the Czech Geological Society 44:329-333. To the occurrences of the family Klasmuridae, add isolated ossicles in the Devonian of Poland by Bockzarowski (2001, especially pp. 13-20) with descriptions of new species of Pectenura: P. excubitor, P. formosa, P. pecten, and P. senta. Also isolated lateral element from a borehole, Devonian of Germany, by Haude (2004, on p. 244) [Haude, R. 2004. Cour. Forsch.-Inst. Senckenberg 251:237-251]. Regarding Antiquaster magrumi add: Kesling & Chilman (1975, see pp. 168, 176, and plate 39) [UMMP Papers on Paleontology No. 8]. For redescription of Protasteracanthion and P. primus, and Antiquaster made a synonym of Protasteracanthion, see Alexander Glass (2006. Ph.D. Thesis. The brittle star fauna of the Hunsruck Slate and a phylogeny of the Paleozoic Ophiuroidea. Univ. Illinois at Urbana-Champaign. For abstract see Dissertation Abstracts International 67(11B):6270]. Hotchkiss, F.H.C. 2000. On the number of rays in starfish. American Zoologist 40:340-354. Correction to the statement concerning holothurians on p. 340: the tentacles of holothurians correspond to the rays of other echinoderms, and holothurians have FIVE-PLUS organization of rays/tentacles around the mouth (Hotchkiss 1998:rays as appendages, p.208). The 8-rayed specimen of Luidia senegalensis on p. 345 is deposited in the Smithsonian National Museum of Natural Histsory as USNM 1112625. Further evidence that tetramery at metamorphosis is the result of teratological incomplete development (p. 349) comes from an Arbacia punctulata that developed 4-part symmetry while its genetically identical twin developed 5-part symmetry (the parents had typical 5-part symmetry; Marcus, N.H. 1979. Developmental aberrations associated with twinning in laboratory-reared sea urchins. Developmental Biology 70:274-277). Likewise in the ophiuroid Amphipholis squamata, deviations from pentamerism are not heritable but are a consequence of environmental perturbations on the metamorphosis of larvae and/or abnormal regeneration of arms [Dupont, S. & J. Mallefet. 2002. Abnormal forms in the brittle-star Amphipholis squamata: a field study. J. Mar. Biol. Ass. U.K. 82:491-493]. Some additional references: 2000. Sanchez, P. The sequence of origin of the postmetamorphic rays in Heliaster and Labidiaster (Echinodermata: Asteroidea). Revista Chilena de Historia Natural 73:573-578. 2007. Herringshaw, L.G., M.P. Smith and A.T. Thomas. Evolutionary and ecological significance of Lepidaster grayi, the earliest multiradiate starfish. Zoological Journal of the Linnean Society 150:743-754. Hotchkiss, F.H.C. 2000. Inferring the developmental basis of the sea star abnormality “double ambulacral groove” (Echinodermata: Asteroidea). Revista Chilena de Historia Natural 73:579-583. The specimen (p. 582) from Marblehead, MA of Asterias vulgaris with double ambulacral groove on one of only four rays is deposited in the collections of the California Academy of Sciences, CASIZ 173969. The specimen (p. 581) from a student at Lehigh University of Asterias forbesi with double ambulacral groove on one of its five rays is deposited in the US National Museum, Smithsonian Institution, echinoderm collection, USNM 1082952. The Zoological Record for 1904, p. 34, mentions fusion of rays in Ctenodiscus, referencing Michaailovskij, M. 1904. Die Echinodermen der zoologischen Ausbeute des Eisbrechers “Jermaks” von Sommer 1901. Annuaire Mus. St. Petersb. ix, pp. 157-188. The research collection of sea stars of H. Barraclough Fell contained a Pentagonaster pulchellus with double ambulacral groove. This collection is now at the US National Museum. Hotchkiss, F.H.C. and R. Haude. 2004. *Observations on Aganaster gregarius* and Stephanoura belgica (Ophiuroidea: Ophiolepididae) (Early Carboniferous and Late Devonian age)*. Pp. 425-431 In Heinzeller & Nebelsick (eds.), Echinoderms:München. Taylor & Francis Group, London. The purpose of the research was to elucidate the structure of the oral frame, the structure of the distal parts of the rays, and the status of radial shields. It was noted that this was partly accomplished with new material. More specimens need to be examined to substantiate and expand the findings, especially any observations on mouth papillae. MPRI 0077 is a slab that has five fairly intact disks and many isolated portions of arms of A. gregarius. One specimen, in oral view, preserves one arm out to the 18th arm segment. This arm confirms that the under arm plates of Aganaster* abruptly cease after the 15th arm segment, and that podial pores perforate the lateral arm plates beginning on the 16th arm segment. This arm increases the number of observations of this question to N = 3. To be noted in passing: Pearse (1947. Zoological names - a list of phyla, classes, and orders) listed Order Aganasterida H.L. Clark 1939. I have not found the 1939 H.L. Clark paper, in spite of earnest searching. The Order Aganasterida is listed also by Boettger (1952) and by Blackweldeer (1963). Hotchkiss, F.H.C., Rudolf J. Prokop and Václav Petr. 2007. Isolated ossicles of the Family Eospondylidae Spencer et Wright, 1966, in the Lower Devonian of Bohemia (Czech Republic) and correction of the systematic position of eospondylid brittlestars (Echinodermata: Ophiuroidea: Oegophiurida). Acta Musei Nationalis Pragae, series B Historia Naturalis 63(1):3-18. MPRI contribution No. 1. For additional mention of Eospondylus in the Hunsrück Slate see Südkamp (1985, and especially 1995 documenting Eospondylus on crinoids). Further to the postscript on an eospondylid ophiuroid of eyelet-type in the Arkona Shale, note similarities of preservation and fauna between the Arkona Shale and the Hunsrück Slate [see e.g.: Úna C. Farrell and Derek E. G. Briggs, 2007, A pyritized polychaete from the Devonian of Ontario; Proc. R. Soc. B 274:499–504. the polychaete in this paper was collected by Michael and John Topor and donated by them to UMMP]. Hotchkiss, F.H.C. 2008. Pattern formation in starfish: arm stumps, regeneration models, and evolution. MPRI contribution No. 3. Poster presentation prepared for the Fifth North American Echinoderm Conference, Melbourne, Florida, July 2008, and also the 4th Workshop of German and Austrian Echinoderm Research, Vienna, October 2008. Other published mentions of ray stumps that do not regenerate: Ruedemann (1916:61) described specimens of Hallaster forbesi with “relatively short, somewhat abruptly terminating rays” as probably arm stumps that failed to regenerate. However Spencer (Monograph p. 298) showed that the arms are flexed aborally giving only a false appearance of being stumps. The abruptly terminating arms of Cholaster peculiaris, which is known from a single specimen, are also the product of arms bent 180 degrees onto themselves (FHCH unpublished); so these are not arm stumps that did not regenerate. Additional examples of regenerating Ordovician starfish: Urasterella sp., ROM 53376, Middle Ordovician, Bobcaygeon Formation, near Lake Simcoe, Ontario, Canada (personal communication, David Rudkin, 4/15/2008). Cnemidactis girvanensis (Schuchert), counterpart specimens BMNH E 52484a,b; Upper Ordovician (Ashgillian), Thraive Glen, Ayreshire, Scotland; figured by W.K. Spencer (1918:161 and pl. XII fig. 5); Owen (1965:565 curatorial notes). Modes of wound closure: FHCH has observed dorso-ventral downward movement like closing a roll-top desk (D-V wound closure) in Astropecten following autotomy. Previous research/comments on symmetric versus asymmetric positional information in the starfish arm: Lovén’s rule and Batesonian duplications of appendages imply asymmetrical positional information. David, Mooi & Telford (Echinoderm Research 1995: 163): “In groups such as Asteroidea, in which the paired plates of the ambulacral columns are directly opposite and do not alternate, it should still be possible to see from early ontogenetic stages in which columns the first plate is laid down, thereby testing the existence of Lovén’s rule even in this group.” Hotchkiss (1998: 207) emphasized that no case of Batesonian duplication of asteroid rays has ever been reported, and he rejected the implications of this negative finding. The implied expectation of these researchers was that positional information in the starfish ray is asymmetric. Impact on the rays-as-appendages hypothesis: Symmetrical positional information in the starfish ray is not consistent with the rays-as-appendages model which explicitly invokes asymmetrical positional information. Either the rays-as-appendages hypothesis should be rejected, or we need some sort of hypothesis-saving explanation. I opt for the latter because the fossil record indicates that alternating ambulacral plates and Loven’s law are plesiomorphic, and these features imply asymmetric positional information. Somehow, the starfish arm has changed from an original asymmetric condition to a symmetric condition. Other examples of failure to regenerate following L-R bilaterally symmetrical wound closure and stump formation occur in planarians (headless planarians): Child, C.M. 1911. Experimental control of morphogenesis in the regulation of Planaria. Biological Bulletin 20:310-331. Della Valle, P. 1914. L’inibizione della rigenerazione del capo nelle planarie mediante la cicatrizzazione. Archivio Zoologico Italiano 7:275-311. some references to birds/turtles drumming for worms American woodcock. " Later, at 10:30 a.m., this bird and another fed in bright sunshine on the lawn, nonchalantly pulling worms from the sod. They had found the only square footage in town that was frost free. Their side-to-side- shifting, accompanied by deliberate weight shifts from foot to foot, must cause enough vibration to stir the worms below." Joe Choiniere, director of Mass Audobon's Wachusett Meaadow Wildlife Sanctuary. Seasons of the woodcock: the secret life of a woodland shorebird. Sanctuary, The Journal of the Massachusetts Audubon Soociety, Summer 2006, 45(4):3-5. [see p. 5] Gull in grass park beside Lake Merritt, Oakland, California, spring 1991. Observations by F. Hotchkiss communicated to Ned Newton. "... on the grass, to the front and left of me, only eight feet from the sidewalk, was a gull. The gull seemed to be stepping in place. I stopped to watch it on the grass. It stepped again... it did it clearly with an alternate, rapid step -- stepping in place. In between this stepping it would stop -- it would peer at the ground just in front of itself, but never walking -- just with its feet stationary, planted -- and I would then see it grab an earthworm with its beak -- and quickly swallow it -- and look for another -- and then resume the stepping in place -- without moving an inch. This gull was a talented earthworm drummer. How on earth did it ever learn to do this!!" Ned Newton added the following comments: "Well, it seems that I have heard of Robins and Woodcock drumming up worms. But it must have been fun to see a relatively large bird like a gull doing this. I was telling a friend from Connecticut about the drumming for worms. She was not at all surprised and added that she has seen turtles doing this same thing -- hopping from one foot to another. Somehow they don't have the speed that I would think is required. Anyone who has been out at night catching nightcrawlers knows that you have to be fast. Well, maybe the next time the kids are dancing it would be interesting to look around near by or on the flood and see if worms are raised to the surface." Newton's Notes, April 26, 1991 [Notes from the President], Nashoba Valley Bird Club, The Meadow Plover 2(2) Spring 1991. For information on turtles stomping for earthworms see: Ernst, C. H., J. E. Lovich, and R. W. Barbour. 1994. Turtles of the United States and Canada. Washington, D.C.: Smithsonian Institution Press. [summary of information] Kaufmann, J. H. 1986. Stomping for earthworms by wood turtles, Clemmys insculpta: a newly discovered foraging technique. Copeia 1986(4):1001-1004. Kaufmann, J. H., J. H. Harding and K. N. Brewster. 1989. Worm stomping by wood turtles revisited. Bull. Chicago Herpetol. Soc. 24: 125-126. Kirkpatrick, David T. and Catherine Kirkpatrick. 1996. Stomping for earthworms by Clemmys insculpta in captivity. Bulletin of the Chicago Herpetological Society 31(2):21-22.				latest news 5/20/2008 added to the Make it happen listings You Make It Possible! Our latest newsletter describes how our supporters further our mission. Site Update! Fossil Heritage Collection Specimens purchased with the help of our generous donors.
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