From f04d23caa9e45094c17d2c3d912e3e300506d55b Mon Sep 17 00:00:00 2001 From: ggserver Date: Fri, 6 Dec 2024 19:41:25 +0000 Subject: [PATCH] Add updates up until 2024-12-06 19:35:34 --- .../C9/2F1AC921FFC1FFC2F2F0FF7CFCB6599D.xml | 337 ++++++ .../C9/2F1AC921FFC1FFC3F2F0F9CAFC495E81.xml | 175 +-- .../C9/2F1AC921FFC3FFC0F2F0FCAAFB345BA2.xml | 161 +-- .../C9/2F1AC921FFDDFFD8F2F0FC18FBC25D28.xml | 1021 +++++++++++++++++ 4 files changed, 1527 insertions(+), 167 deletions(-) create mode 100644 data/2F/1A/C9/2F1AC921FFC1FFC2F2F0FF7CFCB6599D.xml create mode 100644 data/2F/1A/C9/2F1AC921FFDDFFD8F2F0FC18FBC25D28.xml diff --git a/data/2F/1A/C9/2F1AC921FFC1FFC2F2F0FF7CFCB6599D.xml b/data/2F/1A/C9/2F1AC921FFC1FFC2F2F0FF7CFCB6599D.xml new file mode 100644 index 00000000000..1c3aedb10ba --- /dev/null +++ b/data/2F/1A/C9/2F1AC921FFC1FFC2F2F0FF7CFCB6599D.xml @@ -0,0 +1,337 @@ + + + +New Eocene species of the crinoid genera Holopus and Cyathidium (Cyrtocrinida: Holopodidae) from north-eastern Italy + + + +Author + +Martinez-Soares, Pablo +Institut de Systématique, Evolution, Biodiversité (ISYEB) UMR 7205 CNRS-MNHN-SU-EPHE-UA, Muséum national d’Histoire naturelle, 57 rue Cuvier, 75005 Paris, France. + + + +Author + +Roux, Michel +0000-0003-3119-9609 +Institut de Systématique, Evolution, Biodiversité (ISYEB) UMR 7205 CNRS-MNHN-SU-EPHE-UA, Muséum national d’Histoire naturelle, 57 rue Cuvier, 75005 Paris, France. +michel.roux51@gmail.com + + + +Author + +Giusberti, Luca +0000-0002-4401-5410 +luca.giusberti@unipd.it + + + +Author + +Gatto, Roberto +0000-0001-5849-1242 +roberto.gatto@unipd.it + + + +Author + +Eléaume, Marc +0000-0003-1999-1177 +Institut de Systématique, Evolution, Biodiversité (ISYEB) UMR 7205 CNRS-MNHN-SU-EPHE-UA, Muséum national d’Histoire naturelle, 57 rue Cuvier, 75005 Paris, France. & Institut de Systématique, Evolution, Biodiversité (ISYEB) UMR 7205 CNRS-MNHN-SU-EPHE-UA, Muséum national d’Histoire naturelle, BP 225, 29182 Concarneau Cedex, France. +marc.eleaume@mnhn.fr + + + +Author + +Améziane, Nadia +0000-0002-8546-9961 +Institut de Systématique, Evolution, Biodiversité (ISYEB) UMR 7205 CNRS-MNHN-SU-EPHE-UA, Muséum national d’Histoire naturelle, BP 225, 29182 Concarneau Cedex, France. +nadia.ameziane@mnhn.fr + +text + + +Zootaxa + + +2024 + +2024-11-28 + + +5541 + + +4 + + +401 +437 + + + + +http://dx.doi.org/10.11646/zootaxa.5360.3.8 + +journal article +10.11646/zootaxa.5541.4.1 +1175-5326 +14241160 +4F812F60-9242-4F44-8E25-99381FD7E8B3 + + + + + + +Genus + +Cyathidium +Steenstrup + +in +Michaelis & Scherk, 1847 + + + + + + +Type-species. + +Cyathidium holopus +Steenstrup + +in +Michaelis & Scherk, 1847, by monotypy. + + + + +Synonymy. + +Cyathidium +( +Steenstrup 1847 +) + +in Michaelis & Scherk, 1847: 119, 150; + +Micropocrinus +Michelin, 1851: 93 + +; +Sieverts, 1931:173 +; + +Pseudocupressocrinus +Valette + +in +Lambert & Valette, 1934: 59 +; + +Cyathidium +Sieverts-Doreck, 1938: 29 + +; + +Micropocrinus +Biese & Sieverts-Doreck, 1939: 117 + +; + +Cyathidium +Rasmussen, 1961: 238–239 + +; +1978 +: T839. + + + + +Remarks. + +Žítt +et al. +(2014) + +thoroughly described the wide range of shape variation in a lower Turonian population of 136 aboral cups attributed to + +Cyathidium +aff. +depressum + +. The ratio H/D varies from 0.2 to +c. +1.0 and usually increases during growth ( +Fig. 7 +). The cup curvature is also greatly variable with H/h from 1 to 2.7. Details of characters discriminating the extant species such as interradial morphology and fulcral ridge arrangement ( + +Améziane +et al. +1999 + +) are only preserved in a few specimens, and only parts of the external surface show growth lines and peculiar ornamentation. Therefore, attribution to a new species or to previously described species merely based on aboral cups requires well-preserved specimens and discriminating qualitative characters. Intact fossil specimens with a retracted crown are known exclusively in + +C. holopus + +( +Baumiller & Gaździcki 1996 +; +Donovan & Jakobsen 2004 +; +Hess & Messing 2011 +; +Nielsen 1913 +; +Rasmussen 1961 +). The vault covering the distal arm series is composed solely of IIBr1, their height being substantially greater than their width. IIBr have not yet been described from the large species + +Cyathidium vlieksi +Jagt, 1986 + +, but are known (Jagt pers. comm.). This species has an aboral cup seemingly very similar to some specimens of + +C. holopus + +. Intraspecific variation of the aboral cup in + +C. holopus + +and + +C. vlieksi + +show two extreme morphotypes: one very depressed, like in + +C. depressum +, + +and another as high or taller than wide with conspicuous growth lines. The lower Santonian species, + +C. senessei +, + +differs from all other known congeners in having a stout and tumid curved IBrax, with relatively small distal articular facets facing inwards and subperpendicular to the upper plan of the IBr circlet ( + +Améziane +et al. +1999 + +; + +Heinzeller +et al. +1996 + +; +Rasmussen 1961 +). The central space provided by the IBrax circlet ( +Rasmussen 1961 +, pl. 34 fig. 7c) probably houses IIBr1 keystone plates, but these have not yet been described. + +Žítt +et al. +(2014) + +figured the main discriminating characters of dissociated brachials of + +C. +aff. +depressum + +from the lower Turonian. Unfortunately, IIBr were not available in the original description of + +C. depressum + +from the upper Cenomanian but IBr1 were illustrated ( +Rasmussen 1961 +). The presence of granulation on the aboral cup appears highly variable among fossil assemblages and cannot be used as a specific discriminating character except when there are sufficient ossicles to demonstrate the stability of this trait. Another early Campanian form from southern +England +and north-west +France +will be published soon (Gale & Jagt, in press). + + +Included fossil species and occurrence. + +Cyathidium chiampoensis + + +sp. nov. + +(lower Lutetian of Chiampo valley, north-eastern +Italy +), + +C. depressum + +including + +C. +aff. +depressum + +first described by +Žítt 1996 +(upper Cenomanian–lower Turonian, +Germany +and +Bohemia +), + +C. gastaldii + +(Middle Miocene near Turin, northwestern +Italy +), + +C. holopus + +(upper Maastrichtian of +Netherlands +, Danian of +Denmark +and +Sweden +, lower Eocene of Seymour Island, +Antarctica +), + +C. senessei + +(lower Santonian of Bugarach, southern +France +), + +C. vlieksi + +(Maastrichtian of +Belgium +, +Netherlands +), +C. +sp. nov. +(Gale & Jagt, in press; lower Campanian, north-western +France +and southern +England +). + + + + +Stratigraphic distribution. +Upper Cenomanian–Recent. + + + + \ No newline at end of file diff --git a/data/2F/1A/C9/2F1AC921FFC1FFC3F2F0F9CAFC495E81.xml b/data/2F/1A/C9/2F1AC921FFC1FFC3F2F0F9CAFC495E81.xml index 0a3cedf59e9..62612783a4b 100644 --- a/data/2F/1A/C9/2F1AC921FFC1FFC3F2F0F9CAFC495E81.xml +++ b/data/2F/1A/C9/2F1AC921FFC1FFC3F2F0F9CAFC495E81.xml @@ -1,88 +1,88 @@ - - - -New Eocene species of the crinoid genera Holopus and Cyathidium (Cyrtocrinida: Holopodidae) from north-eastern Italy + + + +New Eocene species of the crinoid genera Holopus and Cyathidium (Cyrtocrinida: Holopodidae) from north-eastern Italy - - -Author + + +Author -Martinez-Soares, Pablo -Institut de Systématique, Evolution, Biodiversité (ISYEB) UMR 7205 CNRS-MNHN-SU-EPHE-UA, Muséum national d’Histoire naturelle, 57 rue Cuvier, 75005 Paris, France. +Martinez-Soares, Pablo +Institut de Systématique, Evolution, Biodiversité (ISYEB) UMR 7205 CNRS-MNHN-SU-EPHE-UA, Muséum national d’Histoire naturelle, 57 rue Cuvier, 75005 Paris, France. - - -Author + + +Author -Roux, Michel -0000-0003-3119-9609 -Institut de Systématique, Evolution, Biodiversité (ISYEB) UMR 7205 CNRS-MNHN-SU-EPHE-UA, Muséum national d’Histoire naturelle, 57 rue Cuvier, 75005 Paris, France. -michel.roux51@gmail.com +Roux, Michel +0000-0003-3119-9609 +Institut de Systématique, Evolution, Biodiversité (ISYEB) UMR 7205 CNRS-MNHN-SU-EPHE-UA, Muséum national d’Histoire naturelle, 57 rue Cuvier, 75005 Paris, France. +michel.roux51@gmail.com - - -Author + + +Author -Giusberti, Luca -0000-0002-4401-5410 -luca.giusberti@unipd.it +Giusberti, Luca +0000-0002-4401-5410 +luca.giusberti@unipd.it - - -Author + + +Author -Gatto, Roberto -0000-0001-5849-1242 -roberto.gatto@unipd.it +Gatto, Roberto +0000-0001-5849-1242 +roberto.gatto@unipd.it - - -Author + + +Author -Eléaume, Marc -0000-0003-1999-1177 -Institut de Systématique, Evolution, Biodiversité (ISYEB) UMR 7205 CNRS-MNHN-SU-EPHE-UA, Muséum national d’Histoire naturelle, 57 rue Cuvier, 75005 Paris, France. & Institut de Systématique, Evolution, Biodiversité (ISYEB) UMR 7205 CNRS-MNHN-SU-EPHE-UA, Muséum national d’Histoire naturelle, BP 225, 29182 Concarneau Cedex, France. -marc.eleaume@mnhn.fr +Eléaume, Marc +0000-0003-1999-1177 +Institut de Systématique, Evolution, Biodiversité (ISYEB) UMR 7205 CNRS-MNHN-SU-EPHE-UA, Muséum national d’Histoire naturelle, 57 rue Cuvier, 75005 Paris, France. & Institut de Systématique, Evolution, Biodiversité (ISYEB) UMR 7205 CNRS-MNHN-SU-EPHE-UA, Muséum national d’Histoire naturelle, BP 225, 29182 Concarneau Cedex, France. +marc.eleaume@mnhn.fr - - -Author + + +Author -Améziane, Nadia -0000-0002-8546-9961 -Institut de Systématique, Evolution, Biodiversité (ISYEB) UMR 7205 CNRS-MNHN-SU-EPHE-UA, Muséum national d’Histoire naturelle, BP 225, 29182 Concarneau Cedex, France. -nadia.ameziane@mnhn.fr +Améziane, Nadia +0000-0002-8546-9961 +Institut de Systématique, Evolution, Biodiversité (ISYEB) UMR 7205 CNRS-MNHN-SU-EPHE-UA, Muséum national d’Histoire naturelle, BP 225, 29182 Concarneau Cedex, France. +nadia.ameziane@mnhn.fr -text - - -Zootaxa +text + + +Zootaxa - -2024 - -2024-11-28 + +2024 + +2024-11-28 - -5541 + +5541 - -4 + +4 - -401 -437 + +401 +437 - -http://dx.doi.org/10.11646/zootaxa.5360.3.8 + +http://dx.doi.org/10.11646/zootaxa.5360.3.8 -journal article -10.11646/zootaxa.5541.4.1 -1175-5326 -14241160 -4F812F60-9242-4F44-8E25-99381FD7E8B3 +journal article +10.11646/zootaxa.5541.4.1 +1175-5326 +14241160 +4F812F60-9242-4F44-8E25-99381FD7E8B3 @@ -101,7 +101,7 @@ Figures 8A–C ; -Table 4 +Table 4 Synonymy. @@ -120,14 +120,24 @@ Frisone Type material. - -Two aboral cups including the + +Two +aboral cups including the holotype ( MGP-PD 33229 , -Fig. 8A–B -), one well-preserved + +Fig. 8 +A +– +B + +), + + +one +well-preserved paratype ( MGP-PD 33230 @@ -156,28 +166,19 @@ Small species (D< - - -Type -stratum. - -Lower Lutetian -tuffite horizon - -. +Type stratum. +Lower Lutetian +tuffite horizon. - - -Type -locality. - -Cengio -dell’Orbo quarry (Cava Boschetto) in the +Type locality. +Cengio dell’Orbo quarry +( +Cava Boschetto +) in the Chiampo Valley , north-eastern Italy - . @@ -187,7 +188,7 @@ dell’Orbo quarry (Cava Boschetto) in the Aboral cups belonging to small specimens with cup diameter up to 2.72 mm , H equal or substantially lower than D (H/D 0.42), usually D greater than Db. Quantitative characters of the type series listed in -Table 4 +Table 4 . The holotype , which is the largest specimen, has a curved aboral cup and an external surface showing 8–9 conspicuous parallel growth lines interrupting numerous perpendicular ridges; base smaller than D straight on one side ( diff --git a/data/2F/1A/C9/2F1AC921FFC3FFC0F2F0FCAAFB345BA2.xml b/data/2F/1A/C9/2F1AC921FFC3FFC0F2F0FCAAFB345BA2.xml index f605211035d..f37e1511506 100644 --- a/data/2F/1A/C9/2F1AC921FFC3FFC0F2F0FCAAFB345BA2.xml +++ b/data/2F/1A/C9/2F1AC921FFC3FFC0F2F0FCAAFB345BA2.xml @@ -1,88 +1,88 @@ - - - -New Eocene species of the crinoid genera Holopus and Cyathidium (Cyrtocrinida: Holopodidae) from north-eastern Italy + + + +New Eocene species of the crinoid genera Holopus and Cyathidium (Cyrtocrinida: Holopodidae) from north-eastern Italy - - -Author + + +Author -Martinez-Soares, Pablo -Institut de Systématique, Evolution, Biodiversité (ISYEB) UMR 7205 CNRS-MNHN-SU-EPHE-UA, Muséum national d’Histoire naturelle, 57 rue Cuvier, 75005 Paris, France. +Martinez-Soares, Pablo +Institut de Systématique, Evolution, Biodiversité (ISYEB) UMR 7205 CNRS-MNHN-SU-EPHE-UA, Muséum national d’Histoire naturelle, 57 rue Cuvier, 75005 Paris, France. - - -Author + + +Author -Roux, Michel -0000-0003-3119-9609 -Institut de Systématique, Evolution, Biodiversité (ISYEB) UMR 7205 CNRS-MNHN-SU-EPHE-UA, Muséum national d’Histoire naturelle, 57 rue Cuvier, 75005 Paris, France. -michel.roux51@gmail.com +Roux, Michel +0000-0003-3119-9609 +Institut de Systématique, Evolution, Biodiversité (ISYEB) UMR 7205 CNRS-MNHN-SU-EPHE-UA, Muséum national d’Histoire naturelle, 57 rue Cuvier, 75005 Paris, France. +michel.roux51@gmail.com - - -Author + + +Author -Giusberti, Luca -0000-0002-4401-5410 -luca.giusberti@unipd.it +Giusberti, Luca +0000-0002-4401-5410 +luca.giusberti@unipd.it - - -Author + + +Author -Gatto, Roberto -0000-0001-5849-1242 -roberto.gatto@unipd.it +Gatto, Roberto +0000-0001-5849-1242 +roberto.gatto@unipd.it - - -Author + + +Author -Eléaume, Marc -0000-0003-1999-1177 -Institut de Systématique, Evolution, Biodiversité (ISYEB) UMR 7205 CNRS-MNHN-SU-EPHE-UA, Muséum national d’Histoire naturelle, 57 rue Cuvier, 75005 Paris, France. & Institut de Systématique, Evolution, Biodiversité (ISYEB) UMR 7205 CNRS-MNHN-SU-EPHE-UA, Muséum national d’Histoire naturelle, BP 225, 29182 Concarneau Cedex, France. -marc.eleaume@mnhn.fr +Eléaume, Marc +0000-0003-1999-1177 +Institut de Systématique, Evolution, Biodiversité (ISYEB) UMR 7205 CNRS-MNHN-SU-EPHE-UA, Muséum national d’Histoire naturelle, 57 rue Cuvier, 75005 Paris, France. & Institut de Systématique, Evolution, Biodiversité (ISYEB) UMR 7205 CNRS-MNHN-SU-EPHE-UA, Muséum national d’Histoire naturelle, BP 225, 29182 Concarneau Cedex, France. +marc.eleaume@mnhn.fr - - -Author + + +Author -Améziane, Nadia -0000-0002-8546-9961 -Institut de Systématique, Evolution, Biodiversité (ISYEB) UMR 7205 CNRS-MNHN-SU-EPHE-UA, Muséum national d’Histoire naturelle, BP 225, 29182 Concarneau Cedex, France. -nadia.ameziane@mnhn.fr +Améziane, Nadia +0000-0002-8546-9961 +Institut de Systématique, Evolution, Biodiversité (ISYEB) UMR 7205 CNRS-MNHN-SU-EPHE-UA, Muséum national d’Histoire naturelle, BP 225, 29182 Concarneau Cedex, France. +nadia.ameziane@mnhn.fr -text - - -Zootaxa +text + + +Zootaxa - -2024 - -2024-11-28 + +2024 + +2024-11-28 - -5541 + +5541 - -4 + +4 - -401 -437 + +401 +437 - -http://dx.doi.org/10.11646/zootaxa.5360.3.8 + +http://dx.doi.org/10.11646/zootaxa.5360.3.8 -journal article -10.11646/zootaxa.5541.4.1 -1175-5326 -14241160 -4F812F60-9242-4F44-8E25-99381FD7E8B3 +journal article +10.11646/zootaxa.5541.4.1 +1175-5326 +14241160 +4F812F60-9242-4F44-8E25-99381FD7E8B3 @@ -103,7 +103,7 @@ Figures 8F, G ; -Table 6 +Table 6 @@ -144,22 +144,18 @@ Non Type material. -A single aboral cup ( +A + +single aboral cup ( holotype -) from the Miocene of the Superga Hill near Turin and two plaster casts are housed in the palaeontological collection of the -MNHN -in Paris (MNHN-F-R68284). Several casts of this -holotype -were sent to collections outside of -France -. Two of them are housed in the Palaeontological Museum of Turin University ( +) from the Miocene of the Superga Hill near Turin and two plaster casts are housed in the palaeontological collection of the MNHN in Paris (MNHN-F-R68284). Several casts of this holotype were sent to collections outside of France. Two of them are housed in the Palaeontological Museum of Turin University ( Manni & Pacioni 2021 ). The cast of an aboral cup ( -MPURLS NS -6/800) having nearly the same size and belonging to the Palaeontological Museum of Rome ( +MPURLS NS 6/800 +) having nearly the same size and belonging to the Palaeontological Museum of Rome ( Manni 2005 -) is also a cast of the -holotype +) is also a cast of the holotype + . @@ -179,7 +175,7 @@ Description of the . Aboral cup rounded subpentagonal, with granulated external surface as indicated in Michelin’s diagnosis. Quantitative characters listed in -Table 6 +Table 6 . Wall relatively thick ( 2.5–2.7 mm ); interradial processes conspicuous but irregularly preserved; synarthrial articulation of radials worn, aboral ligament pit inconspicuous ( @@ -192,7 +188,7 @@ Aboral cup rounded subpentagonal, with granulated external surface as indicated Other specimen. Manni (2005) figured a small aboral cup (quantitative characters listed in -Table 6 +Table 6 ) with a granulated external surface belonging to the Michelotti Collection housed in the Museum of Palaeontology of “La Sapienza” University of Rome (MPURLS 2534). He attributed it to a juvenile of C. gastaldii @@ -246,7 +242,12 @@ Boschetto (= Cengio dell’Orbo quarry, Chiampo Valley, Lutetian), specimens her Occurrence. -Serravallian (Middle Miocene) of Superga Hill near Turin, northwestern +Serravallian (Middle Miocene) +of +Superga Hill +near +Turin +, northwestern Italy . diff --git a/data/2F/1A/C9/2F1AC921FFDDFFD8F2F0FC18FBC25D28.xml b/data/2F/1A/C9/2F1AC921FFDDFFD8F2F0FC18FBC25D28.xml new file mode 100644 index 00000000000..a38c1805d93 --- /dev/null +++ b/data/2F/1A/C9/2F1AC921FFDDFFD8F2F0FC18FBC25D28.xml @@ -0,0 +1,1021 @@ + + + +New Eocene species of the crinoid genera Holopus and Cyathidium (Cyrtocrinida: Holopodidae) from north-eastern Italy + + + +Author + +Martinez-Soares, Pablo +Institut de Systématique, Evolution, Biodiversité (ISYEB) UMR 7205 CNRS-MNHN-SU-EPHE-UA, Muséum national d’Histoire naturelle, 57 rue Cuvier, 75005 Paris, France. + + + +Author + +Roux, Michel +0000-0003-3119-9609 +Institut de Systématique, Evolution, Biodiversité (ISYEB) UMR 7205 CNRS-MNHN-SU-EPHE-UA, Muséum national d’Histoire naturelle, 57 rue Cuvier, 75005 Paris, France. +michel.roux51@gmail.com + + + +Author + +Giusberti, Luca +0000-0002-4401-5410 +luca.giusberti@unipd.it + + + +Author + +Gatto, Roberto +0000-0001-5849-1242 +roberto.gatto@unipd.it + + + +Author + +Eléaume, Marc +0000-0003-1999-1177 +Institut de Systématique, Evolution, Biodiversité (ISYEB) UMR 7205 CNRS-MNHN-SU-EPHE-UA, Muséum national d’Histoire naturelle, 57 rue Cuvier, 75005 Paris, France. & Institut de Systématique, Evolution, Biodiversité (ISYEB) UMR 7205 CNRS-MNHN-SU-EPHE-UA, Muséum national d’Histoire naturelle, BP 225, 29182 Concarneau Cedex, France. +marc.eleaume@mnhn.fr + + + +Author + +Améziane, Nadia +0000-0002-8546-9961 +Institut de Systématique, Evolution, Biodiversité (ISYEB) UMR 7205 CNRS-MNHN-SU-EPHE-UA, Muséum national d’Histoire naturelle, BP 225, 29182 Concarneau Cedex, France. +nadia.ameziane@mnhn.fr + +text + + +Zootaxa + + +2024 + +2024-11-28 + + +5541 + + +4 + + +401 +437 + + + + +http://dx.doi.org/10.11646/zootaxa.5360.3.8 + +journal article +10.11646/zootaxa.5541.4.1 +1175-5326 +14241160 +4F812F60-9242-4F44-8E25-99381FD7E8B3 + + + + + + + +Holopus fabianii + +sp. nov. + + + + + + +Figs. 7 +, +8H–J +, +9–11 +; +Tables 7–10 + + +Synonymy. + +Holopus spileccense +Manni & Pacioni, 2021: 69–73 + +, figs. 4–6; + +Frisone +et al +., 2020: 306 + +( +non +Schlüter 1878 +). + + + + +Type material. + +Eight +aboral cups including the +holotype +( + +Fig. 8 +H +– +I + +), + + +one +figured +paratype +( +paratype 1 +, +Fig. 8J +), + + +six +non-figured +paratypes +( +paratypes 2–7 +); + + +17 +figured brachials including 12 IBrax ( +paratypes 8–19 +, +Fig. 10A–L +) and five IIBr ( +paratypes 20–24 +, +Fig. 11A–H +); + + +14 +non-figured IBrax ( +paratypes 46–59 +), + + +21 +non-figured IIBr ( +paratypes 25–45 +) including three keystone plates and three IIBr from distal series + +. + + +Other material examined. +17 IBrax, 58 IIBr of Wp< +2.8 mm +, 21 IIBr of Wp> +2.8 mm +. + + + + +Etymology. +Species dedicated to Ramiro Fabiani (1879–1954) for his contributions to stratigraphy and palaeontology of the Paleogene from north-eastern +Italy +. + + + + +Diagnosis. +Cup with conspicuous constriction in the lower half; upper part of cup with rounded pentagonal cross-section; pronounced radial ribs and marked interradial ribs; adoral face without external interradial process; calycinal cavity unknown; estimated maximum cup size: H 21.0 mm, D +14.8 mm +; IBrax about as high as wide, with Y-shaped ridge, upper part of ridge (V-shape) irregularly doubled, usually with outer branches reaching angles of maximum brachial width (Wd) and inner branches corresponding to radial axis of brachial ramifications; small IBrax with only a few tubercles or none at all; large IBrax with several tubercles especially in the proximal half; external ornamentation of proximal IIBr consisting of X-shaped positioned tubercles, muscular synarthries with fulcral ridge shorter than maximum brachial width axis, adoral border forming an obtuse angle giving relatively narrow V-shape section; keystone plates with distal muscular synarthry roughly triangular and rounded, proximal synarthry symmetrical with fulcral ridge as long as Wp, external face smooth, shaped like an isosceles triangle, four conspicuous parallel symplexial crenulations on each side; distal IIBr series higher than broad, muscular synarthries with inconspicuous fulcral ridge, marked aboral ligament pit and axial nervous canal, conspicuous adoral muscle pits. + + + + +Type stratum. +Lower Lutetian +, tuffite horizon. + + +Type locality. +Cengio dell’Orbo quarry +in the +Chiampo Valley +, north-eastern +Italy +. + + + + +FIGURE 9. +Main quantitative characters of +Holopodidae +from the Eocene of north-eastern Italy. Cy: + +Cyathidium +, Ho + +: + +Holopus, +Ho. + +a.: juvenile of + +H. alidis + +(MNHN-IE-2014-2185). Straight lines: values of H/D and H/Wd. Other values in mm. + + + + +Description of type series. +Among the 8 aboral cups of the type series (quantitative characters listed in +Table 7 +), the +holotype +was selected as the best preserved aboral cup ( +Fig. 8H, I +). +Holotype +medium sized (H +3.83 mm +, D +2.75 mm +), with a strong constriction (Db/Dcs 1.68) located in the aboral half and separating adoral two-thirds of cup from a broad basal part with flat attachment surface with a greater diameter than the adoral side (D/Db 0.88); adoral part with rounded pentagonal cross-section demarcated by five conspicuous radial ribs with intermediate ribs more pronounced on one side ( +Fig. 8H +) than on the other ( +Fig. 8I +); adoral face without external interradial process, calycinal cavity filled with recrystallized sediment making it impossible to identify any possible internal interradial processes, if present. Other aboral cups of type series are of variable sizes (H +1.72 to 6.55 mm +, D +1.76 to 4.62 mm +), usually height greater than adoral diameter (H/D 1.21 to 1.48) except in small +paratype +7 (H/D 0.98), H/D increasing from nearly 1 to more than 1.4 with size ( +Fig. 9 +), similar characters as the +holotype +, more or less conspicuous, attachment surface often smaller than adoral diameter and very irregular, sometimes appearing as a mold of a subcylindrical substrate ( +Fig. 8J +), cup with moderate curvature (H/h~1.3) only in +paratype +3. + + +Type series of IBrax ( +paratypes +8 to 19) illustrate changes in form and ornamentation with size ( +Fig. 10 +, +Table 8 +). Smallest IBrax nearly as high as wide; external surface ( +Fig. 10A–E +) with a more or less thick Y-shaped ridge, distal part of ridge (V-shape) irregularly doubled with usually outer branches reaching angles of maximum brachial width (Wp) and inner branches corresponding to radial axis of brachial ramifications; proximal external surface without tubercles on either side of axial ridge. Larger IBrax about as wide as high except +paratypes +10 and 18 which show H/ Wd 0.82 and 0.84, respectively; external ornamentation ( +Fig. 10I–L +) showing Y-shaped ridge composed of a short lower segment and predominantly longer V-shaped segments corresponding to radial axis of brachial ramifications, 10 to 20 large tubercles developed irregularly over the entire external surface, sometimes transforming former ridges into rows of tubercles ( +Fig. 10K–L +). Asymmetric +paratype +17 possibly belongs to a lateral trivium plate ( +Fig. 10J +). Aboral interradial process moderately developed. + + +Type series of IIBr, proximal series to keystone plate ( +paratypes +20 to23 and 25 to 39) with external ornamentation predominantly X-shaped, always with a more or less marked axial ridge and few tubercles ( +Fig. 11D–E +). Ratios H/h, H/Wp and Wp/Wd independent of size ( +Table 9 +). Height asymmetry mostly due to oblique articulation with IBrax, 12 cases with H/h<1.3, 4 cases with 1.5<H/h<1.7 ( +Fig. 11E +) and 3 cases with 1.95<H/h<2.05 ( +Fig. 11D +). Lateral faces with 4 to 6 parallel symplexial crenulations, sometimes restricted to the highest one. Muscular synarthries with fulcral ridge shorter than maximum brachial width (W). +Two types +of muscular synarthries: one symmetrical with equal fulcral ridge segments parallel to W ( +Fig. 11A +), another asymmetrical with unequal fulcral ridge segments oblique to W ( +Fig. 11B +); most IIBr with one synarthry asymmetrical and the other symmetrical (IIBr1 or IIBr2?), other IIBr with two articular facets asymmetrical (IIBr1) or symmetrical (IIBr2 or 3?). Four IIBr nearly as high as wide and asymmetrical, assigned to IIBr1 of a bivium ( +Fig. 11E +). Articulations with adoral border forming an obtuse angle, ambulacral groove shallow, giving the whole ossicle a relatively flat V-shaped section. + + +Keystone plates of type series ( +paratypes +24 and 40–42, +Table 10 +) with distal muscular synarthry roughly triangular and rounded ( +Fig. 11F +); proximal synarthry symmetrical, with fulcral ridge as long as Wp ( +Fig. 11H +); external face smooth, shaped as an isosceles triangle ( +Fig. 11G +), four conspicuous parallel symplexial crenulations on each side ( +Fig. 11F +). + + +A few brachials from the distal IIBr series ( +paratypes +43–45, +Table 10 +) higher than broad, muscular synarthries similar to the distal articulation of the keystone plates with inconspicuous fulcral ridge, marked aboral ligament pit and axial nervous canal, conspicuous adoral muscle pits. + + + + + +Complementary data from material not included into the +type +series. + +The additional material allows us to investigate in more detail the range of variation of quantitative characters (Supplementary +Tables 1–2 +) and ratios in the main +types +of brachials. All material examined including the types-series: 8 aboral cups, 43 IBrax, 46 IIBr of the proximal series (including 14 with Wp> +2.8 mm +) and 60 IIBr with Wp< +2.8 mm +(distal series plus a few keystone plates poorly preserved). Ratio of IBrax to aboral cups 5.4, very close to expected ratio of 5; ratio of IBrax to proximal IIBr 1.07, far from the expected ratio of 4 or 6 indicating a high IIBr deficit. Height asymmetry (H/h) of the proximal IIBr is independent of their size. In 50% of the proximal IIBr 1.1<H/h<1.3 (possibly IIBr2 or IIBr3), in 34.6% 1.4<H/ h<1.75 (possibly IIBr1 or IIBr2) and in 15.4% 1.75<H/h<2.3 (possibly IIBr1). Quantitative characters of IBrax attaining maximum values with H +4.78 mm +, Wd +5.30 mm +, Wp +4.15 mm +and Wm +3.25 mm +(Appendix +Table 1 +). + + + + +Remarks. +The high number of isolated ossicles of + +Holopus fabianii + + +sp. nov. + +from the +type +locality provides a unique insight into intraspecific character variation in the genus + +Holopus + +. To date, no comparable data have been published for extant species. However, the absence of a crown preserved with connected brachials makes it impossible to establish the precise number of IIBr in the arm’s proximal series. The combination of H/h asymmetry, which appears to increase proximally in extant species, and the fact that 50% of the IIBr have a H/h close to 1 suggest that there might have been at least 2–3 IIBr preceding the keystone plate. Like in the extant species + +H. alidis + +, the position of the keystone plate could have varied depending on the arm and the developmental stage ( + +Bourseau +et al +. 1991 + +). + + +The recrystallized sediment within the calycinal cavity masks possible inner interradial processes, whose presence and characters would have been useful to describe and compare to extant species. These structures might have been absent as in + +H. spileccense + +(see below). + + +The maximum size of the aboral cup and IBrax of + +H. fabianii + + +sp. nov. + +can be estimated by comparing the greatest width of the facets corresponding to the same articulation joining two adjacent ossicles. The proximal articulation of the largest IIBr ( +paratype +20) is 1.6 times wider (Wp +5.06 mm +) than the largest distal articulation of IBrax (Wm +3.25 mm +). The largest proximal articulation of IBrax is 2 times wider (Wp +4.15 mm +) than the largest distal articulation of the available aboral cups ( +paratype +2, Wr +2.03 mm +). Thus, the estimated size of the individual to which +paratype +20 brachial belonged was 3.2 times larger than +paratype +2. Therefore, the aboral cup of + +H. fabianii + + +sp. nov. + +could possibly be as large as H 21.0 mm and D +14.8 mm +, a size intermediate between the known maximum aboral cup sizes of + +H. alidis + +(H +13 mm +, D +9 mm +) and + +H. rangii + +(H +30 mm +, D +25 mm +). + + + +Holopus fabianii + + +sp. nov. + +displays closer affinities with + +H.alidis + +than with + +H.rangii + +,as both possess a differentiated keystone plate beyond IIBr2 separating proximal and distal arm series ( +Figs. 5C–H +and +Fig. 11F–H +). Both species also have an aboral cup with rather marked radial ribs. Yet, + +H. alidis + +differs in having a distinctly granulated external cup and brachials, including the keystone plates, with a predominantly tuberculated ornamentation unlike the nearly smooth keystone plates found for + +H. fabianii + + +sp. nov. + +( +Fig. 5C +, + +Fig. 5 +G + +, Bourseau +et al +. 1991, pl. 12). However, judging from our observation on + +Cyathidium + +, the relevance of the latter character might be called into question. The IBrax ornamentation with Y- or V-shaped ridges common in + +H. fabianii + + +sp. nov. + +is restricted to juveniles of + +H. alidis + +( +Fig 4B, C +) and tends to disappear under the tuberculate ornamentation in older specimens. Moreover, the adoral side of the fulcral ridge of the proximal-most brachials often conspicuously crenulated in + +H. alidis + +( +Fig. 5A–B +) is not found in more distal synarthries ( +Fig. 5E +). This character appears to be absent in + +H. fabianii + + +sp. nov. + +( +Fig. 11A–B +). Furthermore, the IIBr fulcral ridges in + +H. alidis + +( +Fig. 5B +) and + +H. rangii + +( +Fig. 6D, F +) are as long as the brachial is wide, which contrasts significantly with what is observed in + +H. fabianii + + +sp. nov. + +( +Fig. 11A–C +). This appears to be due to a lateral aboral displacement of the adoral surfaces of the synarthries, “opening” the articulation and thus giving the brachial a rather flat V-shape. The brachials of + +H. alidis + +(and + +H. rangii + +) are thus far thicker than those of + +H. fabianii + + +sp. nov. + +, and, in IBrax, exhibit a more pronounced distal interradial process. + + + + +FIGURE 10. +Variation in primibrachial ornamentation in + +Holopus fabianii + + +sp. nov. + +from Chiampo (MGP-PD 33235). A–G, J: small specimens, A: paratype 8, B: paratype 9, C: paratype 10, D: paratype 11, E: paratype 12, F: paratype 13, G: paratype 14, J: paratype 15, H, I, K, L: large specimens, H: paratype 16, I: paratype 17, K: paratype 18, L: paratype 19. Scale bars: 1 mm. + + + + +FIGURE 11. +Secundibrachials of + +Holopus fabianii + + +sp. nov. + +from Chiampo belonging to proximal series (MGP-PD 33236); A–E: IIBr from proximal part of proximal series, A: symmetrical muscular synarthry from a small specimen (paratype 20), B: asymmetrical muscular synarthry (paratype 21), C–D: proximal-most IIBr of a large specimen (paratype 22), C: oblique distal view, D: external face, E: IIBr of a bivium (paratype 23), F–H: keystone plate, distalmost IIBr of proximal series (paratype 24), F: oblique view of distal muscular synarthry, G: oblique view showing triangular external face and proximal muscular synarthry at left, H: proximal muscular synarthry. Scale bars: 1 mm. + + + + + +Holopus plaziati + +, known exclusively from isolated brachials (7 IBrax + 1 proximal IIBr), comes from the lower Eocene of the Corbières in southern +France +( + +Roux +et al +. 2021 + +). Its IBrax differ from those of + +H. fabianii + + +sp. nov. + +and + +H. spileccense + +in being generally smaller with a smaller H/Wd ( +Fig. 7 +), and in having muscular synarthries with usually conspicuous crenulation on the adoral side of the fulcral ridge and sometimes on the adoral border of the muscular area. While +syntype +2 of + +H. plaziati + +appears similar to the IBrax of + +H. fabianii + + +sp. nov. + +, IBrax of +syntypes +3 and 7 have adoral articulation facets larger than in + +H. fabianii + + +sp. nov. + +In + +H. plaziati + +this increases the size of the distal interradial process and the general thickness of the brachial, resulting in an ambulacral groove that is more deeply carved into the ossicle. The conspicuous lateral symplexial crenulations in + +H. plaziati + +are variable (parallel to undulated or branched). Its aboral surface displays irregular tuberculate ornamentation without formation of interradial or Y-shaped ridges. + + + +TABLE 7. +Quantitative characters of the eight aboral cups in the type-series of + +Holopus fabianii + + +sp. nov. + +Abbreviations: see Fig. 1A. *: estimated value (broken ossicle). D used as growth index. Values in mm except ratios. Holotype: MGP-PD 33232; Paratype 1: MGP-PD 33233; Paratypes 2–7: MGP-PD 33234. + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + +
SpecimenDHhhtDbDcsWrH/DH/hH/htD/Dcs
Paratype 71.81.71.70.81.2–1.71.61.20.981.002.071.13
Paratype 12.53.02.81.61.7–2.31.71.41.211.061.831.49
Holotype2.73.93.72.53.11.81.71.391.021.531.49
Paratype 6>3*4.4-2.13.2–4.13.1-<1.4*-1.351.06
Paratype 33.34.93.82.85.03.2-1.481.291.741.03
Paratype 5>4*5.1-3.7>2.7*2.4-<1.4*-1.35>1.5*
Paratype 4->5*>3*-4.5–5.3---1.6*--
Paratype 24.66.56.54.57.3–8.13.92*1.421.001.441.17
+ + + +TABLE 8. +Quantitative characters of IBrax of + +Holopus fabianii + + +sp. nov. + +belonging to figured paratypes. Abbreviations: see Fig. 1B. Wd used as growth index. Values in mm except ratios. Paratypes 8–19: MGP-PD 33235. + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + +
SpecimenFigureWdHWpWmH/WdH/WpWp/WmWp/Wd
Paratype 810A1.81.81.01.11.021.780.930.57
Paratype 1110D1.91.91.01.01.031.900.970.54
Paratype 1210E2.22.21.51.31.011.461.160.69
Paratype 910B2.22.11.11.30.961.960.850.49
Paratype 1510J2.32.21.21.20.951.820.980.52
Paratype 1010C2.62.11.21.30.821.760.960.46
Paratype 1410G2.72.4-1.80.89---
Paratype 1310F2.82.62.61.70.921.011.510.91
Paratype 1610H3.83.43.42.00.901.021.640.88
Paratype 1710I4.14.02.92.40.971.351.240.72
Paratype 1810K4.43.73.12.40.841.181.300.71
Paratype 1910L4.94.53.92.40.921.171.580.78
+ + +Manni and Pacioni (2021) +attributed 30 brachials from +Cava +Boschetto (= Cengio dell’Orbo quarry, the type-locality of + +H. fabianii + + +sp. nov. + +) to + +H. spileccense +(Schlüter) + +, a taxon known only from cups found in the Ypresian strata of Spilecco Hill (see below). These 30 brachials are housed in the MGP-PD collections. +Manni and Pacioni (2021) +figured the ornamentation of 5 IBrax and 6 IIBr from the proximal series but did not identify keystone plates. The variation of X-shaped ridges on IIBr was well-illustrated and corresponds to the range observed in the material here ascribed to + +H. fabianii + + +sp. nov. + +The quantitative characters of these brachials also fall within the variation range of those of + +H. fabianii + + +sp. nov. + +(except IBrax MGP-PD 31442 which presents the largest Wd, +5.3 mm +) and they should thus be attributed to this species. + + +Occurrence. +Middle Eocene (Lutetian) of Chiampo Valley, north-eastern +Italy +. + + + + \ No newline at end of file