MENATHYRIS WILSONI (BRACHIOPODA), NEW GE NUS
AND SPE CIES FROM THE MID DLE TRI AS SIC (LADINIAN)
OF SOUTH ERN IS RAEL
Howard R. FELDMAN
Di vi sion of Pa le on tol ogy (Invertebrates), Amer i can Mu seum of Nat u ral His tory, New York, NY 10024-5192, USA;
e-mail: feldspar4@optonline.net
Feldman, H. R., 2013. Menathyris wilsoni (Brachiopoda), new ge nus and spe cies from the Mid dle Tri as sic (Ladinian) of south ern Is rael. Annales Societatis Geologorum Poloniae, 83: 81–85.
Ab stract: Menathyris wilsoni gen. et sp. nov. is de scribed from the Lime stone and Marl Mem ber (Early Ladinian)
of the Saharonim For ma tion (early Anisian–late Carnian), at Har Gevanim, Makhtesh Ramon, south ern Is rael. The Saharonim For ma tion rep re sents the main transgressive phase of the Mid dle Tri as sic. The sub unit, from which the shells were col lected, con tains the molluscs Neoschizodus laevigatus (Ziethen), Myophoriopsis cf. subundata (Schauroth), Lima cf. tellei Bittner, Pecten discites (Schlotheim), Ostrea montiscaprilis Klipstein, Pseudo-placunopsis fissistriata (Winkler), Gevanites awadi (Parnes), and the cono dont Pseudofurnishius murcianus (Van den Boogaard), the last-men tioned in dic a tive of the Fassanian (Early Ladinian).
Key words: brachi o pod, Tri as sic, Ladinian, Is rael, Negev, Si nai Pen in sula.
Manu script re ceived 18 June 2013, ac cepted 14 August 2013
IN TRO DUC TION
This pa per re cords part of a longterm study of the Ju
-ras sic and Tri as sic fau nas of the Mid dle East, with em pha sis
on the Si nai Pen in sula, Is rael and Jor dan. The pres ent study
rep re sents a con tin u a tion of work on the Tri as sic rocks and
fau nas of the Sephardic Prov ince in Makhtesh Ramon,
sou-thern Is rael. Feldman (2002, 2005) de scribed a new
brachiopod spe cies, Coenothyris oweni, as well as a brachi o
-pod-dom i nated com mu nity from the Saharonim For ma tion.
The data pre sented herein per mit a palaeo eco logi cal anal y
-sis of the Saharonim For ma tion that will re sult in a better
un der stand ing of the palaeobiogeographical con text of the
fau nas within the Sephardic Prov ince and elu ci da tion of the
evo lu tion ary his tory of the brachi o pod spe cies within the
province.
There have been nu
mer
ous stud
ies on Muschelkalk
(Tri as sic) brachi o pod fau nas (Awad, 1946; Popiel-Barczyk
and Senkowiczowa, 1989; Pálfy and Török, 1992;
Senko-wiczowa and Popiel-Barczyk, 1993), al though Tri as sic
bra-chiopods from Is rael have been ne glected in the lit er a ture
and the Terebratulida, in gen eral, have been poorly rep re
-sented in the Tri as sic (Hagdorn and Sandy, 1998).
The Tri as sic se quence in Is rael con sists of mostly ma
-rine sed i ments with ev i dence of oc ca sional progradations of
ter res trial en vi ron ments and ranges in thick ness from 500–
1100 m (Druckman, 1966; Parnes, 1986). Shaw (1947) first
de scribed the Tri as sic se quence of Ramon, fol lowed by Zak
(1964, 1968) and Druckman (1969, 1974, 1976). The ex po
-sure at Har ‘Arif (Negev) was first de scribed by Zemel et al.
(1956). Eicher (1947) dis cov ered the first un ques tion able
conodonts of clearly Tri as sic age from these Si nai and Negev
out crops. Al though a de tailed de scrip tion of this fauna
(Bro-tzen, 1956) was first pub lished on the palaeontological and
biostratigraphical fea tures of the out crop, Zak (1957) made
the lithostratigraphic sub di vi sion fol lowed in this pa per.
In the pres ent pa per, a new brachi o pod ge nus and spe
-cies, Menathyris wilsoni, is de scribed from Makhtesh
Ra-mon in south ern Is rael.
GEO LOG I CAL SET TING
The brachi o pods de scribed herein were col lected from
the Saharonim For ma tion at Makhtesh Ramon, south ern Is
-rael (Fig. 1). Makhtesh Ramon is a large (40 km long, 8 km
wide), north
east-trending, ero
sional cirque in the Negev
that rep re sents the cen tral breached part of the Ramon
anticlinorium, in which sed i men tary rocks of Cre ta ceous to Tri
-as sic age are ex posed (Sohn, 1968; Feldman, 2002).
Accor-ding to Sohn (1968), Makhtesh Ramon is the larg est and sou-
thernmost of three breached domes in the Negev, the other
two be ing Hamakhtesh Hagadol (Makhtesh Hathira/Kurnub
Anticline) and Hamakhtesh Haqatan (Makhtesh Hazera).
The Saharonim For ma tion (Fig. 2) con sists mostly of
car bon ates with shale, marl and evaporite beds that rep re
-sent the main transgressive phase of the Mid dle Tri as sic
(Benjamini et al., 2005). Zak (1957) sub di vided the for ma
-tion at Makhtesh Ramon, south ern Is rael, into the fol low ing
mem bers: (1) the fossiliferous Lime stone Mem ber, con sist
-ing of me ter-thick clay and marly shale ho ri zons, topped by
Awad’s (1946) “Mot tled Scar”; (2) the Lime stone and Marl
Mem ber, con sist ing at the base of 9 m of dark, nod u lar
bio-turbated micrites, over lain by fossiliferous lime stones and
marls that rep re sent off shore to intertidal en vi ron ments of
de po si tion (Parnes, 1975, 1986; Benjamini et al., 2005); (3)
the Lime stone and Gyp sum Mem ber that con sists of var i ous
lithofacies, such as fine-grained do lo mite, foraminiferal and
oncoidal lime stones, al gal stromatolites, and shale,
marl-stone and gyp sum in ter ca la tions (Parnes, 1986; Benjamini
et al., 2005); and (4) the Lime stone Mem ber, seen in sev eral
cuestas with gray, bioturbated lime stones form ing led- ges,
al ter nat ing with gypsiferous shales, dolomicrite and al gal
stromatolites (Benjamini et al., 2005).
Ac cord ing to Benjamini et al. (2005) the lithofacies of
the Saharonim For ma tion in di cates, at the end of the
Ani-sian, rapid deep en ing, fol lowed by an os cil lat ing pat tern of
fa cies rang ing from deeper ma rine en vi ron ments and
shallow ing pro gres sively through shelf and mar ginal ma rine fa
-cies.
As so ci ated fauna: The ho ri zon, in which the ma te rial
was found, also con tains the molluscs Neoschizodus
laevi-gatus (Ziethen), Myophoriopsis cf. subundata (Schauroth),
Lima cf. tellei Bitner, Pecten discites (Schlotheim), Ostrea
montiscaprilis Klipstein, Pseudoplacunopsis fissistriata
(Winkler),
Pseudoplacunopsis fissistriata (Winkler), Geva-
nites awadi (Parnes), and the cono dont Pseudofurnishius
murcianus (Van den Boogaard), the lastmen tioned in dic a
-tive of the Fassanian (early Ladinian).
MA TE RI ALS AND METH ODS
The shells are all ar tic u lated. Trans verse se rial sec tions
were pre pared with a WOKO 50P grind ing ma chine with
slice-spac ing of 100 to 200 µm. Draw ings of the spec i mens
were made with a cam era lucida and mea sure ments were
taken with a dig i tal cal i per and rounded off to 0.1 mm. Prior
to pho to graph ing, the shells were coated with am mo nium
chlo ride.
Ab bre vi a tions: AMNH, Amer i can Mu seum of Nat u ral His
tory, New York; USNM and NMNH, United States Na tional Mu -seum and Na tional Mu seum of Nat u ral His tory, Smith so nian In sti tu tion, Wash ing ton, D.C.
Mea sure ments: All mea sure ments are in mil li me ters (mm). Fig. 1. Lo ca tion map of sam ple lo cal ity in Makhtesh Ramon,
south ern Is rael (Is rael grid co or di nates 1370/9998-1379/9994). The Menathyris wilsoni gen. et sp. nov. ho ri zon is de noted by an “X” (mod i fied from Feldman, 2002)
Fig. 2. Strati graphic sec tion of the Tri as sic Saharonim For ma -tion at Har Gevanim, Makhtesh Ramon, south ern Is rael. Menathyris wilsoni gen. et sp. nov. was col lected from Sub unit 44. Sub -units 12–15, 31 and 42 rep re sent Coenothyris oweni beds (see Feldman, 2002). Sub unit des ig na tions are af ter Zak (1964) and Parnes (1975)
SYS TEM ATIC PALAE ON TOL OGY
Subphylum RHYNCHONELLIFORMEA Wil liams,
Carlson, Brunton, Holmer and Popov, 1996
Class RHYNCHONELLATA Wil liams, Carlson, Brunton,
Holmer and Popov, 1996
Or der TEREBRATULIDA Waagen, 1883
Suborder TEREBRATELLIDINA Muir-Wood, 1955
Superfamily ZEILLERIOIDEA Allan, 1940
Fam ily ZEILLERIIDAE Allan, 1940
Menathyris ge nus novum
Type spe cies: Menathyris wilsoni gen. et sp. nov. by monotypy. Other spe cies: Ge no type spe cies only.
Et y mol ogy: In mem ory of Mena SchemmGreg ory for her im por
tant con tri bu tions to Palaeozoic and Me so zoic brachi o pod palae -on tol ogy. Her un timely death has dealt a blow to our pro fes si-on.
Di ag no sis: Small terebratellids subpentagonal to subpyriform in
out line, com monly ventribiconvex; shells wid est at midlength, an -te rior commissure ranges from rectimargina-te to incipiently uni-plicate. Ven tral beak low, rounded, fo ra men small, permesothyrid; sulcus weak, shal low and nar row. Sur face smooth with nu mer ous, very fine growth lines, an te rior half of shell costate. Car di nal pro -cess lack ing, small teeth ar tic u late with shal low sock ets, hinge plates sup ported by me dian sep tum nar rowly tri an gu lar in cross sec tion; den tal plates not pre served, dis tal ends of hinge plates give rise to de scend ing branches of brachial loop that broad ens to form lat er ally curv ing crural pro cesses.
Re marks: Menathyris gen. nov. is sim i lar to Aulacothyris
Dou-villé in over all ap pear ance and ex ter nal mor phol ogy but dif fers in its smaller size, much shal lower and nar rower sulcus, con vex ity and costation. The shells are sim i lar to Zeilleria arethusa Dubar (see Dubar, 1942, pl. I, figs. 1–6), but dif fer in out line and costa-tion; Z. arethusa is more oval in out line, rather than subpentagonal or subpyriform. The costae of Z. arethusa orig i nate at the umbos and are more nu mer ous whereas those of Menathyris gen. nov. orig i nate no more than half the dis tance from the umbo. There is no in di ca tion of in cip i ent uniplication in Z. arethusa, whereas it is very ev i dent in the new ge nus. Se rial sec tions sug gest a re la tion -ship to the Angustothyrididae, but Menathyris gen. nov. dif fers from all of these forms in its smaller size, more dis tinct costation, and shape of the hinge teeth in cross sec tion. The clos est morpho-type to Menathyris gen. nov. among the angustothyrids is Angustothyris Dagys but this ge nus dif fers in its ro bust and al most bul -bous hinge teeth, lack of costation, larger size and deeper, U-sha-ped septalium.
Menathyris gen. nov. is placed in the Zeillerinae, since it most closely re sem bles the gen era therein and is dis tinctly dif fer ent from known taxa within the subfamily.
Menathyris wilsoni gen. et sp. nov.
Figs 3, 4
In cluded spe cies: Type spe cies only by monotypy. Di ag no sis: As for ge nus be cause of monotypy.
Et y mol ogy: In hon our of Mark A. Wil son for his sub stan tial con tri
-Fig. 3. Menathyris wilsoni gen. et sp. nov. A1–4. AMNH 85660, dor sal, ven tral, lat eral, an te rior views; B1–4. AMNH FI-85648, holotype, dor sal, ven tral, lat eral, an te rior views; C1–4. AMNH FI- 85651, dor sal, ven tral, lat eral, an te rior views; D1–4. AMNH FI85643, dor sal, ven tral, lat eral, an te rior views. All spec -i mens were col lected from a hard l-ime stone bed, over ly -ing the Gevanites beds, Sub unit 44, equiv a lent to the lower half of the Lime stone Marl Mem ber, Saharonim For ma tion in Makhtesh Ra-mon, south ern Is rael (Is rael grid co or di nates 13725/99954)
Fig. 4. Se rial sec tions (A–P) of Menathyris wilsoni gen. et sp. nov. AMNH FI85653. Orig i nal length of shell was 12.7 mm. Dis tances are those from the ven tral beak and num bers de note dis -tances be tween sec tions in mm. The dor sal valve is be low. Abbre-viations: c – crura, cp – crural pro cess, ds – den tal socket, ms – me dian sep tum
bu tions to palae on tol ogy and ge ol ogy, par tic u larly the evo lu tion and palaeoecology of en crust ing and bioeroding in ver te brates, as well as the or i gin and diagenesis of car bon ate rocks (es pe cially hard-grounds), cal cite sea dy nam ics, and Pleis to cene sea-level change.
Ma te rial: The col lec tion con sists of 14 spec i mens; the holotype
(AMNH FI-85648) and paratypes (AMNH FI-85643-85647; 85649-85655; 85660) have been placed in re pos i tory in the palaeontological col lec tion of the Amer i can Mu seum of Nat u ral His -tory, New York.
Type Lo cal ity: The shells were col lected from a hard lime stone
bed over ly ing the Gevanites beds, Sub unit 44, Mid dle Mem ber (of Druckman, 1974) equiv a lent to the lower half of the Lime stone Marl Mem ber (of Parnes et al., 1985; Parnes, 1986), Saharonim For ma tion in Makhtesh Ramon, south ern Is rael (GPS co or di nates: N30.58587497°, E34.87624599°; Is rael grid co or di nates 13725/ 99954). The age of the sub unit, ac cord ing to Parnes et al. (1985), is early Ladinian, ly ing 10 m above the beds con tain ing Gevanites awadi Parnes (Fassanian) and 20 m be low the beds with Gevanites epigonus Parnes and G. cornutus Parnes (early Longobardian).
De scrip tion: Shells small (Ta ble 1), subpentagonal to pyriform in
out line, inequivalve, but rang ing from dorsibiconvex to ventri-biconvex, an te rior commissure mostly rectimarginate, but some valves are incipiently to slightly uniplicate; lat eral commissure rectimarginate; umbo sub stan tial, rounded, beak low, suberect to erect, beak ridges rounded, pedicle fo ra men small, permesothyrid, an te rior half of shell costate with about 12–15 costae orig i nat ing about half to two-thirds of the dis tance from umbo, sides rounded; sur face with nu mer ous, fine con cen tric growth lines that are ero-ded on even slightly worn shells.
Pedicle col lar pres ent within the first 13 mm of the ven tral umbo. No car di nal pro cess de vel oped. Poorly de fined peg-like teeth from the ven tral valve are in serted into shal low sock ets in the dor sal valve. Short, ven trally con vex hinge-plates are sup ported by a low me dian sep tum, which per sists to well over half the shell length, di min ish ing near the an te rior commissure. Septalium wide and shal low, den tal plates not pre served; dis tal ends of hinge
plates flat ten and give rise to de scend ing branches of the brachial loop which broaden to form lat er ally curv ing crural pro cesses. A cross-sec tion of the trans verse band of the brachial loop was not ob tain able.
Re marks: Terebratula semiplicata Klipstein (1843, pl. 15, fig. 3)
from the Tri as sic Keuper Marls, St. Cassian, Tyrol, dif fers from Menathyris wilsoni gen. et sp. nov. in its lack of a sulcus (no sulcus on ei ther valve), subelliptical out line, and six costae at the an te rior commissure (3 large, 3 small). Terebratula praemarginata Klip-stein (1843, pl. 15, fig. 6) also from the Tri as sic Keuper Marls, St. Cassian, Tyrol, dif fers in its lack of costae at the an te rior com-missure, erect beak, broad in cip i ent sulcus, very weak, fine growth lines and semispherical gen eral out line. T. semiplicata Klipstein is sim i lar to T. praemarginata in that both shells are punctate, ven-tribiconvex and permesothyrid. Zeilleria itoensis (see Dubar, 1942, pl. II, figs. 1–8) from the Li assic of Mo rocco is sim i lar in size (L, 11.9; W, 11.9; T, 10.7) and costation to Menathyris wil-soni gen. et sp. nov., but dif fers in its more biconvex and elon gate shells.
Zeilleria sp. from the Lower Pliensbachian of Sterlange, France (NMNH 195551) is larger (L, 25.6; W, 18.7; T, 12.7) than Menathyris wilsoni gen. et sp. nov. subpentagonal in out line with sharply de fined beak ridges and lacks costation. Zeilleria cf. quadrifida Lamarck from the mid dle Lias of Somerset, United King -dom (USNM 104824) is con sid er ably larger than Menathyris gen. et sp. nov. (L, 28.0; W, 24.0; T, 14.0), has well de fined beak ridges, a bilobate out line and lacks costation. Zeilleria sartha-censis d’Orbigny from the Hettangian of the United King dom (USNM 75593) dif fers from Menathyris wilsoni gen. et sp. nov. in its larger size (L, 18.6; W, 15.1; T, 14.5), well de fined beak ridges, a slightly bilobate out line, a telate permesothyrid fo ra men, and lack of costation. Somalitela re sem bles Menathyris wilsoni gen. et sp. nov. in costation, but dif fers in size (L, 26.9; W, 25.1; T, 17.8), elon gate-oval out line, flat tened umbo and more an gu lar beak ridges.
Shells of Parathyridina semiarata Dubar from the Ju ras sic (Lotharingian) of Jebel Anrhemeur, south west of Marrakech, Mo -rocco (USNM Acc. 198144) are con sid er ably larger and more globose than Menathyris gen. et sp. nov. (L, 26.2; W, 25.2; T, 19.4). One of the shells has 20–21 costae that orig i nate about one-third of the dis tance from the beak. Two (uncataloged) spec i mens from the Ju ras sic (Bajocian) of the Si nai Pen in sula, Egypt, have costae that orig i nate half the dis tance from the beak, but are oth er -wise sim i lar to the shells from Mo rocco. Both spec i mens of Parat- hyridina have a uniplicate an te rior commissure with su per im -posed, al ter nat ing costation.
Ac knowl edge ments
I thank the fol low ing for com ments, sug ges tions and dis cus -sions: Ar thur J. Boucot (Or e gon State Uni ver sity), Fran cis Hirsch (Naruto Uni ver sity of Ed u ca tion), Mena SchemmGreg ory (Uni -ver sity of Coimbra, de ceased), Mark A. Wil son (the Col lege of Wooster). I am grate ful to the Geo log i cal Sur vey of Is rael for pro -vid ing of fice space and ac cess to the li brary dur ing my ten ure as a Vis it ing Sci en tist. Thanks go to re view ers Maria Aleksandra Bit-ner (Pol ish Acad emy of Sci ences) and Shuzhong Shen (Nanjing In sti tute of Pa lae on tol ogy) for their crit i cal com ments; they greatly im proved the manu script. Steve Thurston (Amer i can Mu seum of Nat u ral His tory) de serves thanks for his ex cel lent pho to -graphic work. I thank the Touro Col lege Fac ulty Re search Fund for sup port ing this pro ject and thank the mem bers of its peer re -view panel.
Ta ble 1
Mea sure ments (in mm) of Menathyris wilsoni gen. et sp.
nov. from the Ladinian of south ern Is rael
Specimen Length [mm] Width [mm] Thickness [mm] Apical Angle [°] AMNH FI-85643 14.5 11.6 8.9 85 AMNH FI-85644 14.2 11.2 9.1 86 AMNH FI-85645 14.6 13.2 x 74 AMNH FI-85646 11.2 9.2 6.0 84 AMNH FI-85647 11.7 9.0 8.6 82 AMNH FI-85648 holotype 14.0 12.6 8.5 95 AMNH FI-85649 11.9 10.7 6.0 90 AMNH FI-85650 12.6 11.4 8.4 92 AMNH FI-85651 14.9 12.5 9.8 88 AMNH FI-85652 14.4 15.4 x 80 AMNH FI-85653* 12.7 10.5 8.1 78 AMNH FI-85654 17.0 14.0 est 10.9 86 AMNH FI-85655 15.2 12.5 x 90 AMNH FI-85660 13.1 11.5 9.0 97 * – sec tioned spec i men; x – dam aged spec i men; est, es ti mated mea sure -ment. Fig ured spec i mens: holotype AMNH FI-85648: paratypes AMNH FI-85643, FI-85651, FI-85660
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