report on decapod crustaceans from the eocene of zagros basin, iran

Report on decapod crustaceans from the Eocene of Zagros Basin Iran

Alessandro Garassino Ali Bahrami Medhi Yazdi and Francisco J Vega

With 5 figures

Abstract We report on decapod crustaceans from the Eocene of Zagros Basin (Baba-Heydar sec-tion Shahr-e-kurd area Iran) The studied sample includes crustacean penaeids and scyllarids that have been assigned as follows Eopabdehus nov gen with E babaheydariensis n sp (Penaeidae Rafinesque 1815) Eogordonella nov gen with E iranianiensis n sp (Solenoceridae Wood-Mason 1891) and Parsacus nov gen with P eocenicus n sp (Scyllaridae LatReiLLe 1825) Eogordonella iranianiensis n sp is the second report of Solenoceridae in the fossil record The description of Par-sacus nov gen allowed to review the extant genus Parribacus dana 1852 attesting that the fossil species P cristatus foumlRsteR 1984 previously included therein must be assigned to the new genus

Key words Crustacea Decapoda Penaeoidea Solenoceridae Achelata Eocene Iran

1 Introduction and geological setting

The Baba-Heydar section in the Shahr-e-kurd area of Zagros Basin Iran is here investigated in order to study the crustacean horizons of the Pabdeh Forma-tion The section is accessible by the Shahr-e-kurd ndash Kuhrang main road at the vicinity of Qanbarciny village with coordinates 32deg23rsquo1rdquo N 50deg19rsquo66rdquo E (Fig 1) The Pabdeh Formation was deposited in the north-western and southern parts of the Zagros Basin (Fig 2) Two fossiliferous horizons consisting of thin-bedded marly limestones with diverse fishes crusta-ceans insects and plant remains crop out between the marly units in the lower part of the Pabdeh Formation (PRieM 1908 aRaMbouRg 1967 HagHiPouR amp bRants 1971 aLa et al 1980 MuRRis 1980 JafaRian et al

1999 MoHseni et al 2011 asfaRi et al 2014) (Fig 3) The studied specimens subject of this study were col-lected from these two horizons (Fig 3) In the studied area the Pabdeh Formation is about 800 meters thick and includes marls and gray shales with intercalations of thin-bedded limestones (Fig 2) The formation con-sists of thin- to thick-bedded limestones alternating with dark to light gray shales rich in planktonic micro-fauna (MoHseni amp aL-asaM 2004) Many authors (eg boRdenave amp Huc 1995) described the Pabdeh Forma-tion along with its underlying Late Cretaceous Gurpi Formation as a fine-grained matrix-rich carbonate-siliciclastic succession with some hydrocarbon source rock characteristics Petrographic and field studies of the Pabdeh Formation indicate that its sedimentary en-vironment represents a deep sea environment which

copy2014 E Schweizerbartrsquosche Verlagsbuchhandlung Stuttgart Germany wwwschweizerbartde

DOI 101127njgpa20140442 0077-774920140442 $ 300

N Jb Geol Palaumlont Abh 2741 (2014) 43ndash54 ArticleStuttgart October 2014E

44 A Garassino et al

includes shale pelagic lime and calciturbidite facies The existence of planktonic bioclasts related to deep sea such as Globigerina drsquooRbigny 1826 Globrotalia

cusHMan 1927 and abundant micrite indicates the de-position of this group in a deep sea environment The existence of glauconite facies (as authigenic and inter-

Fig 1 A ndash General location map of the studied area B ndash Geological map of the Shahr-e-kurd area the rectangle at the upper left of the map shows the studied locality (modified after ZaHedi 1993)

Report on decapod crustaceans from the Eocene of Zagros Basin Iran 45

bedded in foraminifera) is another indication for the deposition of this facies Evidence from trace fossils microfacies and analyses of turbidities suggests that the depositional environment of the Pabdeh Formation was a distally steepened ramp directed basinward to-wards the intra-shelf basin (MiRZaee-MaHMoodabadi et al 2010)

The age of this horizon is middle to late Eocene based upon the pelagic and benthic foraminifers Hantkenina compressa PaRR 1947 Hantkenina ala-bamensis cusHMan 1925 Marginulina wetherelli Jones 1854 Bolivinoides draco (MaRsson 1878) Laevidentalina communis (drsquooRbigny 1826) Tritax-ia aspera (cusHMan 1926) Lagena spp Uvigerina spp Operculina sp and Nodosaria spp belong to the middle to late Eocene (coxaLL amp PeaRson 2006 213) whereas Morozovelloides bandyi fLeisHeR 1974 indi-cates an early late to middle late Eocene age (PeaRson et al 2006 327) A sample taken close to the Asmari Formation (about 150 m above the second fossiliferous horizon) yielded all assemblages reported by cusHMan amp Mc MasteRs (1936) especially Spiroplectammina sp

2 Material studiedThe studied sample includes 15 specimens assigned as fol-lows Eopabdehus nov gen with E babaheydariensis n sp (2 specimens) (Penaeidae Rafinesque 1815) Eogor-donella nov gen with E iranianiensis n sp (5 speci-mens) (Solenoceridae Wood-Mason 1891) Parsacus nov gen with P eocenicus n sp (8 specimens) (Scyllaridae LatReiLLe 1825) The studied specimens are housed in the palaeontological collection of the University of Isfahan IR Iran (EUIC)

Abbreviations a1 antennula a2 antenna lcxp carapace length (excluding rostrum) lt body total length MBA Museum fuumlr Naturkunde of Humboldt University Berlin P1-P5 pereiopods 1 to 5 s1-s6 pleonal somites 1 to 6 wcxp carapace width

3 Systematic palaeontology

Order Decapoda LatReiLLe 1802Suborder Dendrobranchiata bate 1888

Superfamily Penaeoidea Rafinesque 1815Family Penaeidae Rafinesque 1815

Fig 2 A ndash General view of the first crustacean bearing beds of the Pabdeh Formation in the studied profile Baba-Heydar (Shahr-e-kurd area) B E ndash General view of the second crustacean bearing beds of the Pabdeh Formation in Baba-Heydar (Shahr-e-kurd area) C ndash Close-up view of interbedded opalized nummulitic limestone with crustacean beds D ndash Close-up view of platy cream marls bearing crustacean fossils Photographs by A baHRaMi (Department of Geology University of Isfahan)


Included fossil genera See scHWeitZeR et al (2010) and gaRassino et al (2013)

Genus Eopabdehus nov

Etymology After the Eocene the geological age of the studied specimens and the Pabdeh Formation including the crustacean horizons subject of this study Gender is mas-culine

Type species Eopabdehus babaheydariensis n sp by monotypy

Included fossil species E babaheydariensis n sp (this study)

Diagnosis Elongate rostrum bearing six dorsal proximal teeth and one epigastric tooth carapace with antennal spine hepatic spine well developed short cervical groove

Description As for the type species

Discussion According to scHWeitZeR et al (2010) and ga-Rassino et al (2013) Penaeidae includes presently 25 gen-era from the Early Triassic to the Oligocene Among them only Penaeus fabRicius 1798 has species from the Eocene We justify the erection of Eopabdehus n gen since it does not share morphological characters with Penaeus Indeed according to PeacuteReZ faRfante amp KensLey (1997) the type species P monodon fabRicius 1798 has a rostrum bear-ing dorsal teeth and 2-5 ventral teeth completely different than the rostrum of the new genus which has dorsal teeth limited to the rostrum base According to PeacuteReZ faRfante amp KensLey (1997) only two extant genera Artemesia bate 1888 and Protrachypene buRKenRoad 1934 share an elon-gate rostrum bearing dorsal teeth limited to the rostrum base with the new fossil genus Even though the fossil and extant genera share this character Eopabdehus n gen dif-fers from Artemesia and Protrachypene in having a rostrum with 6 dorsal proximal teeth (vs rostrum with 7 to 14 basal-ly teeth in Artemesia vs rostrum with 7 to 9 basally teeth in Protrachypene)

Fig 3 Stratigraphic section of the Pabdeh Formation in the studied area with position of fossiliferous horizons (arrows) Section by A baHRaMi (Department of Geology University of Isfahan)

Fig 4 A-C ndash Eopabdehus babaheydariensis n gen n sp A EUIC 101400 holoype (r rostrum rt rostral teeth prt postrostral tooth) B EUIC 101401 paratype C Re-construction (a1 unknown a2 P1-P5 and pleopods par-tially reconstructed) D-I ndash Eogordonella iranianiensis n gen n sp D EUIC 101402 holotype (rt rostral teeth s scaphocerite P1-P5 pereiopods dmr dorsal median ridge) E EUIC 101403 paratype (br branchiocadiac ridge hr hepatic ridge sr submargial ridge) F EUIC 101404 para-type G EUIC 101405 paratype H EUIC 101406 para-type I Reconstruction (a1 unknown a2 and P1-P5 partially reconstructed) Scale bar equals 10 mm Photographs by FJ vega (Instituto de Geologiacutea UNAM)


Fig 4 A-C D-I


Eopabdehus babaheydariensis n spFig 4A-C

Etymology The trivial name alludes to the fossiliferous locality of Baba-Heydar Shahr-e-kurd area Zagros Basin IR Iran

Types Holotype EUIC 101400 paratype EUIC 101401

Type locality and age Baba-Heydar Shahr-e-kurd area Iran Pabdeh Formation Eocene

Material and measurements two specimens in lateral view (EUIC 101400 ndash lcxp 43 mm lt 48 mm wcxp 7 mm EUIC 101401 ndash lcxp 32 mm lt 41 mm wcxp 7 mm)

Description Carapace ndash Carapace dorso-laterally flat-tened elongate rostrum bearing six dorsal proximal teeth and one postrostral tooth short cervical groove antennal and hepatic spines well developed Pleon ndash s1 smaller than the others s2-s5 equal in size s6 longer than the previ-ous ones s1-s6 dorsally rounded s1-s6 terga and pleura smooth subtriangular telson distally acute Cephalic ap-pendages ndash Antennulae not preserved antennae partially preserved scaphocerite well developed with apical spine Thoracic appendages ndash P1-P5 partially preserved lacking distal extremity P1-P5 elements elongate and thin Pleonal appendages ndash Pleopods partially preserving subrectangular basipodite uropodal exopod and endopod 13 longer than the telson

Discussion According to PeacuteReZ faRfante amp KensLey (1997) and de gRave et al (2009) the superfamily Penae-oidea includes seven fossil and extant families Aegeridae buRKenRoad 1963 Aristeidae Wood-Mason 1891 Benthe-sicymidae Wood-Mason 1891 Carpopenaeidae gaRassino 1994 Penaeidae Rafinesque 1815 Sicyoniidae oRtMann 1898 and Solenoceridae Wood-Mason 1891 Based upon the morphological characters of these families we assign the studied specimens to Penaeidae because they have an elongate rostrum with slender bearing dorsal teeth that are limited to the rostrum base which can be found in two extant genera Artemesia bate 1888 and Protrachypene buRKen-Road 1934 (vs rostrum short or elongate with ventral or dorsal teeth in Aegeridae vs rostrum usually elongate in females and juvenile males short in adult males with dorsal teeth along its entire length in Aristeidae vs rostrum short triangular with one dorsal tooth in Benthysicymidae vs rostrum elongate with seven-ten dorsal teeth and one ventral tooth in Carpopenaeidae vs rostrum short with both dorsal and apical teeth in Sicyoniidae vs rostrum short with only dorsal teeth in Solenoceridae)

Family Solenoceridae Wood-Mason 1891

Included fossil genera Archeosolenocera caRRioL amp Riou 1991 Eogordonella gaRassino baHRaMi yaZdi amp vega (this study)

Genus Eogordonella nov

Etymology After the Eocene the geological age of the studied specimens and Gordonella tiRMiZi 1960 that shares some morphological characters with the new genus Gender is feminine

Type species Eogordonella iranianiensis n sp by mono-typy

Diagnosis Carapace high distinctly convex posterior to cervical groove short rostrum not overreaching second a1 segment with dorsal teeth only hepatic branchiocardiac and submarginal ridges present slender pleon with dorso-median ridge along entire length on each somite

Included fossil species E iranianiensis n sp (this study)


Discussion According to PeacuteReZ faRfante amp KensLey (1997) Solenoceridae hitherto includes nine extant genera Among them only Gordonella shares some morphological characters with Eogordonella n gen such as having a short rostrum not overreaching second a1 segment with dorsal teeth only epigastric tooth separated from first rostral by more or less conspicuous interval and the carapace and pleon reinforced by numerous ridges Even though the two genera share some morphological characters we justify the erection of the new genus in having weak cervical groove not reinforced by cervical ridge (vs well-marked cervical groove reinforced by cervical ridge in Gordonella) hepatic branchiocardiac and submarginal ridges present (vs anten-nal postantennal cervical hepatic branchiocardiac sub-lateral subhepatic-branchiostegal and submarginal ridges present in Gordonella) telson with smooth lateral margins (vs telson with subapical pair of fixed spines preceded by three pairs of widely spaced spinules in Gordonella) Ac-cording to caRRioL amp Riou (1991) and cHaRbonnieR et al (2010) Solenoceridae is represented in the fossil record by one sole genus Archeosolenocera caRRioL amp Riou 1991 with A straeleni caRRioL amp Riou 1991 from the Late Jurassic (Callovian) of La Voulte-sur-Rhocircne Lagerstaumltte (Ardegraveche France) that is the oldest fossil representative of this family Eogordonella n gen differs from Archeosole-nocera in having well-developed hepatic branchiocardiac and submarginal carapace ridges and a slender pleon with a dorsomedian ridge along entire length on each somite (vs well-developed cervical and hepatic grooves carapace ridges absent and smooth pleonal somites in Archeosole-nocera) Eogordonella n gen is the second report of Sole-noceridae in the fossil record

Eogordonella iranianiensis n spFig 4D-I

Etymology The trivial name alludes to Iran the country where the studied specimens were discovered


Types Holotype EUIC 101402 four paratypes EUIC 101403 101404 101405 101406


Material and measurements Five specimens in lateral and ventral views of which three are exuviae (EUIC 101402 ndash lcxp 61 mm lt 65 mm wcxp 10 mm EUIC 101403 ndash in-complete EUIC 101404 ndash lcxp 36 mm lt 39 mm wcxp 4 mm EUIC 101405 ndash lcxp 47 mm lt 50 mm wcxp 9 mm EUIC 101406 ndash lcxp 38 mm lt 44 mm wcxp 6 mm)

Description Carapace ndash Carapace high distinctly convex posterior to cervical groove short rostrum not overreaching second a1 segment with threefour dorsal teeth only weak cervical groove hepatic branchiocardiac and submarginal ridges well marked Pleon ndash Slender pleon with dorsome-dian ridge along entire length on each somite s1-s5 equal in size s6 slightly longer than the previous ones s1-s6 tergum and pleuron smooth subtriangular telson distally acute Cephalic appendages ndash a1 not preserved well-developed scaphocerite with apical spine Thoracic appendages ndash P1-P3 partially preserved P2-P3 with small thin chelae P4-P5 partially preserved lacking distal extremities P1-P5 flat laterally and smooth Pleonal appendages ndash Pleopods bi-ramous uropodal exopod and endopod 13 longer than the telson

Discussion According to PeacuteReZ faRfante amp KensLey (1997) and de gRave et al (2009) the superfamily Penae-oidea includes seven fossil and extant families Aegeridae buRKenRoad 1963 Aristeidae Wood-Mason 1891 Benthe-sicymidae Wood-Mason 1891 Carpopenaeidae gaRassino 1994 Penaeidae Rafinesque 1815 Sicyoniidae oRtMann 1898 and Solenoceridae Wood-Mason 1891 Based upon the morphological characters of these families we assign the studied specimens to Solenoceridae because they have a short rostrum not overreaching second a1 segment with dorsal teeth only as the extant genus Gordonella tiRMiZi 1960 (vs rostrum short or elongate with dorsal and ventral teeth in Aegeridae vs rostrum usually elongate in females and juvenile males short in adult males with dorsal teeth along its entire length in Aristeidae vs rostrum short tri-angular with one dorsal tooth in Benthysicymidae vs ros-trum elongate with seven-ten dorsal teeth and one ventral tooth in Carpopenaeidae vs rostrum elongate with dorsal and ventral teeth in Penaeidae vs rostrum short with dorsal and apical teeth in Sicyoniidae)

Infraorder Achelata scHoLtZ amp RicHteR 1995Family Scyllaridae LatReiLLe 1825

Included fossil genera Acanthophoenicides audo amp cHaRbonnieR 2012 Biarctus HoLtHuis 2002 Palibacus foumlRsteR 1984 Parribacus dana 1852 Scyllarella RatH-bun 1935 Scyllarides giLL 1898 Scyllarus fabRicius 1775

Subfamily Arctidinae HoLtHuis 1985

Included fossil genera Acanthophoenicides audo amp cHaRbonnieR 2012 Parsacus gaRassino baHRaMi yaZdi amp vega (this study) Scyllarides giLL 1898

Remarks We revise Parribacus dana 1852 among the above-mentioned fossil genera based upon recent results and this study (see discussion below)

Discussion According to scHWeitZeR et al (2010) and audo amp cHaRbonnieR (2012) four fossil species belong to the extant genus Parribacus dana 1852 as follows P cae-sius squiRes 2001 P cottreaui (RogeR 1946) P cristatus foumlRsteR 1984 P japonicus HoLtHuis 1960

Recently feLdMann amp cHaRbonnieR (2011) reviewed the holotype of P cottreaui assigning it to the isopod genus Cirolana LeacH 1818 (Cirolanidae dana 1852)

According to Hu amp tao (1996) the extant P japonicus widespread in the Japanese Sea has fossil representatives from the Pleistocene of Ryukyu Group (Taiwan) unlike HoLtHuis (1960 1991) reported

squiRes (2001) described P caesius from the Eocene of California ascribing it to this genus in dubitative form Indeed as reported by squiRes (2001 21) some morphologi-cal characters such as the general shape of the carapace cervical groove perpendicular to the median ridge and di-rected anterolaterally beyond the median ridge the tuber-culate median ridge extending cross both the gastric and cardiac regions and the raised tuberculate branchial areas are closer to Scyllarides giLL 1898 However even though the outer margin of the 4th antennal segment is serrate in P caesius whereas it is always smooth in Scyllarides we can attest based on the main diagnostic characters of Parriba-cus reported by HoLtHuis (1985 70-71) that P caesius does not belong to this genus for the above-mentioned characters Hence P caesius would be best placed in Scyllarides even though additional study on this species is needed in order to establish its exact systematic placement

foumlRsteR (1984) reported P cristatus (Fig 5H) from the Eocene of Monte Bolca (Verona NE Italy) According to HoLtHuis (1985 70-71) however the main diagnostic char-acters of Parribacus are as follows deep open cervical incision in the anterior part of anterolateral margin antero- and posterolateral margins of carapace provided over their whole length with large sharply pointed teeth of about equal size dorsal surface of carapace without ridges cervical postcervical and branchiocardiac grooves absent 4th-6th an-tennal segment provided over their whole length with large sharply pointed teeth of about equal size Based upon the main morphological characters of P cristatus such as the smooth antero- and posterolateral margins of the carapace well-developed postrostral and posterior postrostral ridges 4th petaloid antennal segment with smooth inner margin slightly serrate outer margin and well-developed median ridge and 6th petaloid antennal segment with smooth inner and outer margins we can attest that this species does not belong to Parribacus for the lack of the main morphological characters of this genus

The species from Italy and the new one from Iran de-scribed below share some morphological characters such


as the well-developed postrostral and posterior postrostral ridges 4th petaloid antennal segment with smooth inner margin slightly serrate outer margin and well-developed median ridge 6th petaloid antennal segment with smooth in-ner and outer margins enough to confirm their ascription to the same genus (see discussion below)

Genus Parsacus nov

Remarks According to HoLtHuis (1991) the short shallow cervical incision and the petaloid antennal segments lacking strong teeth suggest the assignation of Parsacus n gen to Arctidinae Parsacus n gen differs mostly from the fos-sil and extant genera of the subfamily in having postros-tral and posterior postrostral ridges with one row of aligned tubercles and rounded smooth s2-s5 pleura (vs postrostral and posterior postrostral ridges with two rows of aligned tubercles and sharp s2-s5 pleura in Acanthophoenicides vs indistinct postrostral and posterior postrostral ridges and sharp s2-s5 pleura in Arctides HoLtHuis 1960 vs indistinct postrostral and posterior postrostral ridges and sharp or ser-rate rounded s2-s5 pleurae in Scyllarides) Based upon the above-mentioned observations Parsacus n gen represent the fifth report of Arctidinae in the fossil record after Ac-anthophoenicides peterpani audo amp cHaRbonnieR 2012 from the Late Cretaceous (Cenomanian) of Hadjoula (Leba-non) Scyllarides bolcensis de angeLi amp gaRassino 2008 from the early Eocene (Ypresian) of Monte Postale (Italy) S punctatus Woods 1925 from the Early Cretaceous (Al-bian) of Folkestone (United Kingdom) and S tuberculatus (Koumlnig 1825) [= S koenigii (beLL 1858) see quayLe 1987] from the early Eocene (Ypresian) of London Clay Isle of Sheppey (United Kingdom)

Etymology The name alludes to the ancient Greek name of Iran Persis which derives from the name of the main clan of Cirus the Great Parsa Gender is masculine

Type species Parsacus eocenicus n sp by monotypy

Included fossil species P cristatus (foumlRsteR 1984) nov comb P eocenicus gaRassino baHRaMi yaZdi amp vega (this study)

Diagnosis Subcircular carapace dorso-ventrally flattened with short shallow cervical and postcervical incisions ser-

rate anterolateral margin with four spines smooth medio- posterolateral margins postrostral ridge with three aligned tubercles posterior postrostral ridge with seveneight aligned tubercles weak cervical groove sinuous extend-ing until the pregastric tubercle branchiocardiac groove indistinct branchial ridge absent antennular somite with a longitudinal groove 4th petaloid antennal segment with smooth inner margin slightly serrate outer margin and well-developed median ridge 6th petaloid antennal segment with smooth inner and outer margins s1 without median tooth s2-s5 with raised tuberculate median ridge


Parsacus eocenicus n spFig 5A-G I

Etymology The trivial name alludes to the Eocene the geological age of the studied specimens

Types Holotype EUIC 101407 seven paratypes EUIC 101408 101409 101410 101411 101412 101413 101414


Material and measurements Eight specimens almost completely dorso-ventrally preserved (EUIC 101407 ndash lt 42 mm wcxp 19 mm EUIC 101408 ndash lt 38 mm wcxp 19 mm EUIC 101409 ndash lt 37 mm wcxp 18 mm EUIC 101410 ndash lcxp 15 mm wcxp 18 mm EUIC 101411 ndash lcxp 17 mm wcxp 20 mm EUIC 101412 ndash lcxp 13 mm wcxp 18 mm EUIC 101413 ndash lcxp 11 mm wcxp 16 mm EUIC 101414 ndash lcxp 11 mm wcxp 14 mm)

Description Carapace ndash Subcircular carapace dorso-ventrally flattened slightly concave frontal margin with very reduced rostrum and two deep smooth orbital notches orbital notches with curved inner and outer margins dis-tance between orbital notches twice than distance between orbital notches and nearest anterolateral angle lateral mar-gins curved with short shallow cervical and postcervical incisions anterolateral margin with four spines smooth medio- posterolateral margins straight tuberculate poste-rior margin median dorsal ridge divided by weak cervical groove in two parts short postrostral ridge and elongate

Fig 5 A-G I ndash Parsacus eocenicus n gen n sp A EUIC 101407 holotype (ci cervical incision pci postcervical inci-sion ee1 cervical groove s1-s6 pleonal somites mr median ridge 6th as 6th antennal segment 4th as 4th antennal segment) B EUIC 101408 paratype (am anterolateral margin ci cervical incision mm mediolateral margin pci postcervical incision pm posterolateral margin) C EUIC 101411 paratype D EUIC 101410 paratype E EUIC 101412 paratype (pr postrostral ridge ppr posterior postrostral ridge) F EUIC 101409 paratype (am anterolateral margin ci cervical incision pci postcervical incision 4th as 4th antennal segment mr 4th as median ridge of 4th antennal segment) G EUIC 101414 paratype I Parsacus eocenicus n sp reconstruction (P1-P5 pleopods and tail fan unknown) H ndash Parsacus cristatus (foumlRsteR 1984) MAA88 holotype Scale bar equals 10 mm Photographs by FJ vega (Instituto de Geologiacutea UNAM)


Fig 5 A-I


posterior postrostral ridge postrostral ridge restricted to the gastric region with aligned pregastric gastric and car-diac tubercles posterior postrostral ridge with seveneight aligned tubercles cervical groove sinuous extending until the pregastric tubercle branchiocardiac groove indistinct posterior branchial ridge absent dorsal surface of the cara-pace covered with small and large tubercles more or less uniformly arranged Pleon ndash s1 reduced smaller than the others without median tubercle s2-s5 equal in size with a transverse groove and raised tuberculate median ridge rounded smooth s2-s5 pleura s6 and tail fan not preserved Cephalic appendages ndash Antennular somite with a longitu-dinal groove 6th petaloid antennal segment with smooth in-ner and outer margins 4th petaloid antennal segment with smooth inner margin slightly serrate outer margin and well-developed median ridge

Discussion The type species P eocenicus n sp and P cristatus reflect the diagnostic characters of Parsacus n gen Even though the two species share the same geologi-cal age we justify the erection of P eocenicus n sp for (1) palaeogeographical barriers and (2) for some different morphological characters serrate anterolateral margin (vs smooth anterolateral margin in P cristatus) and branchial ridge absent (vs branchial ridge present in P cristatus)

Acknowledgements

We wish to thank cHRistian neuMann (Department of In-vertebrate Palaeontology Museum fuumlr Naturkunde Berlin Germany) for the photographs of the holotype of Parriba-cus cristatus Moreover we wish to thank toRRey nyboRg (Department of Earth and Biological Sciences Loma Linda University California USA) and syLvain cHaRbonnieR (Museacuteum national drsquoHistoire naturelle Paris France) for their helpful reviews and criticism

References

aLa Ma KingHoRn RRf amp RaHMan M (1980) Or-ganic geochemistry and source rock characteristics of the Zagros petroleum province South West Iran ndash Jour-nal of Petroleum Geology 3 61-89

aRaMbouRg C (1967) Reacutesultats Scientifiques de la Mis-sion C Arambourg en Syrie et en Iran (1938-1939) II Les Poissons Oligocegravenes de lrsquoIran ndash Notes et Meacutemoires sur le Moyen-Orient 8 1-210

asfaRi s yaZdi M baHRaMi a amp caRnevaLe g (2014) A new deep-sea hatchetfish (Teleostei Stomiiformes Sternoptychidae) from the Eocene of Ilam Zagros Ba-sin Iran ndash Bollettino della Societagrave Paleontologica Ital-iana 53 (1) 27-37

audo D amp cHaRbonnieR S (2012) New nisto of slipper lobster (Decapoda Scyllaridae) from the Hadjoula La-gerstaumltte (Late Cretaceous Lebanon) ndash Journal of Crus-tacean Biology 32 (4) 583-590

bate CS (1888) Report on the Crustacea Macrura col-lected by HMS Challenger during the years 1873-76 ndash In Report on the scientific results of the voyage of

HMS Challenger during the years 1873-76 under the command of Captain George S Nares RN FRS and the late Captain Frank Tourle Thomson RN 24 1-942

beLL T (1858) A monograph of the fossil malacostracous Crustacea of Great Britain Part 1 Crustacea of the Lon-don Clay ndash Palaeontographical Society Monographs 10 1-44

boRdenave ML amp Huc ay (1995) The Cretaceous source rocks in the Zagros foothills of Iran ndash Revue de lrsquoInstitut Franccedilais du Peacutetrole 50 (6) 121-154

buRKenRoad MD (1963) The evolution of the Eucarida (Crustacea Eumalacostraca) in relation to the fossil record ndash Tulane Studies in Geology 2 (1) 2-17

buRKenRoad MD (1934) Littoral Penaeidea chiefly from the Bingham Oceanographic Collection with a revision of Penaeopsis and description of two new genera and eleven new American species ndash Bulletins of the Bing-ham Oceanographic Collection 4 (7) 1-109

cHaRbonnieR S vannieR J HantZPeRgue P amp gaiLLaRd C (2010) Ecological significance of the arthropod fau-na from the Jurassic (Callovian) La Voulte Lagerstaumltte ndash Acta Palaeontologica Polonica 55 (1) 111-132

caRRioL R-P amp Riou B (1991) Les Dendrobranchiata (Crustacea Decapoda) du Callovien de la Voulte-sur-Rhocircne ndash Annales de Paleacuteontologie 77 (3) 143-160

coxaLL HK amp PeaRson PN (2006) Taxonomy Biostra-tigraphy and Phylogeny of the Hantkeninidae (Clavi-gerinella Hantkenina and Cribrohantkeneina) ndash In PeaRson PN oLsson RK HubeR BT HeMLeben C amp beRggRen WA (eds) Atlas of Eocene Planktonic Foraminifera ndash Cushman Foundation for Foraminiferal Research 213-256 Greenville (East Carolina Univer-sity)

cusHMan Ja (1925) New foraminifera from the Upper Eocene of Mexico ndash Contributions from the Cushman Foundation for Foraminiferal Research 1 4-9

cusHMan Ja (1926) The Foraminifera of the Velasco Shale of the Tampico Embayment ndash Bulletins of the American Association of Petroleum Geologists 10 (6) 1-588

cusHMan Ja (1927) New and interesting foraminifera from Mexico and Texas ndash Contributions from the Cush-man Foundation for Foraminiferal Research 3 111-119

cusHMan Ja amp Mc MasteRs JH (1936) Middle Eocene Foraminifera from the Llajas Formation Ventura Coun-ty California ndash Journal of Paleontology 10 (6) 497-517

dana JD (1852) Crustacea Parts I and II ndash US Explor-ing expedition during the Years 18381839 1840 1841 1842 under the Command of Charles Wilkes USN 13 1-1618

de angeLi A amp gaRassino A (2008) Pseudosquilla les-sinea n sp (Crustacea Stomatopoda Pseudosquillidae) and Scyllarides bolcensis n sp (Crustacea Decapoda Scyllaridae) from the lower Eocene (Ypresian) of Monte Postale (Altissimo Vicenza NE Italy) ndash Atti della So-cietagrave italiana di Scienze naturali e del Museo civico di Storia naturale in Milano 149 (2) 167-178

de gRave S PentcHeff ND aHyong ST cHan T-Y cRandaLL KA dWoRscHaK PC feLdeR DL feLdMann RM fRansen CHJM gouLding LYD LeMaitRe R LoW MEY MaRtin JW ng PKL


scHWeitZeR CE tan SH tsHudy D amp WetZeR R (2009) A classification of living and fossil genera of de-capod crustaceans ndash The Ruffles Bulletin of Zoology Supplements 21 1-109

drsquooRbigny ad (1826) Tableau meacutethodique de la classe des ceacutephalopodes ndash Annales des Sciences Naturelles seacuteries 1 7 96-314

fabRicius JC (1775) Systema entomologiae sistens insec-torum classes ordines genera species adiectis syno-nymis locis descriptionibus observationibus ndash 832 pp Hafriae (Proft)

fabRicius JC (1798) Supplementum Entomologiae Sys-tematicae ndash 572 pp Hafniae (Proft amp Storch)

feLdMann RM amp cHaRbonnieR S (2011) Ibacus cot-treaui RogeR 1946 reassigned to the isopod genus Ci-rolana (Cymothoida Cirolanidae) ndash Journal of Crusta-cean Biology 31 (2) 317-319

fLeisHeR RL (1974) Cenozoic planktonic foraminifera and biostratigraphy Arabian Sea Deep Sea Drilling Project Leg 23A ndash Initial Reports of the Deep Sea Drilling Project 23 1001-1072

foumlRsteR R (1984) Baumlrenkrebse (Crustacea Decapoda) aus dem Cenoman des Libanon und dem Eozaumln Italiens ndash Mitteilungen der Bayerischen Staatssammlung fuumlr Palauml-ontologie und historische Geologie 24 57-66

gaRassino A (1994) The macruran decapod crustaceans of the Upper Cretaceous of Lebanon ndash Paleontologia Lombarda Nuova Serie 3 3-27

gaRassino A vega FJ caLviLLo-canadeLL L cevaL-Los-feRRiZ SRS amp coutintildeo MA (2013) New deca-pod crustacean assemblage from the Upper Cretaceous (Cenomanian) of Chiapas Mexico ndash Neues Jahrbuch fuumlr Geologie und Palaumlontologie Abhandlungen 269 (3) 261-270

giLL T (1898) The crustacean genus Scyllarides ndash Sci-ence new series 7 (160) 98-99

HagHiPouR a amp bRants a (1971) Eocene remains from the Pabdeh Formation north of Ilam ndash Reports of the Geological Survey of Iran 19 81-107

HoLtHuis LB (1960) Preliminary descriptions of one new genus twelve species and three new subspecies of Scyl-larid lobsters (Crustacea Decapoda Macrura) ndash Pro-ceedings of the Biological Society of Washington 73 147-154

HoLtHuis LB (1985) A revision of the family Scyllaridae (Crustacea Decapoda Macrura) I Subfamily Ibacinae ndash Zoologische Verhandelingen 218 3-130

HoLtHuis LB (1991) FAO species catalogue Marine lob-sters of the world ndash FAO Fisheries Synopsis 13 1-276

HoLtHuis LB (2002) The Indo-Pacific scyllarine lobsters (Crustacea Decapoda Scyllaridae) ndash Zoosystema 24 499-683

Hu C-H amp tao H-J (1996) Crustacean fossils of Taiwan ndash 228 pp Taipei (Ta-Jen)

JafaRian Ma saMani Pb amp gHobadiPouR M (1999) In-vestigation and introduction of some Eocene fish fossils in Zagros Basin ndash University of Isfahan Journal (Sci-ence) 13 181-196 [in Farsi]

Jones tR (1854) Notes on the Entomostraca of the Wool-wich and Reading Series ndash Quarterly Journal of the Geological Society London 10 1-100

Koumlnig CDE (1825) Icones fossilium sectiles ndash London (GB Sowerby)

LatReiLLe PA (1802-1803) Histoire naturelle geacuteneacuterale et particuliegravere des Crustaceacutes et des Insectes 3 ndash 467 pp Paris (Dufart)

LatReiLLe PA (1825) Familles naturelles du Regravegne Ani-mal exposeacutees succinctement et dans un ordre analy-tique avec lrsquoindication de leurs genres ndash 570 pp Paris (Bailliegravere)

LeacH WE (1817-1818) A general notice of the animals taken by Mr John Cranch during the expedition to ex-plore the source of the River Zaire ndash In tucKey JK (ed) Narrative of an expedition to explore the River Zaire usually called the Congo in South Africa in 1816 under the direction of Captain JK Tuckey RN to which is added the journal of Professor Smith some general observation on the country and its inhabitants and an appendix containg the natural history of that part of the Kingdom of Congo through which the Zaire-flows London (Murray)

MaRsson t (1878) Die Foraminiferen der weissen Schreib-kreide der Insel Ruumlgen ndash Mitteilungen des Naturwis-senschaftlichen Vereins fuumlr Neu-Vorpommern und Ruuml-gen in Greifswald 10 115-196

MiRZaee-MaHMoodabadi R afgHaH M amp saeedi s (2010) High resolution sequence stratigraphy and depo-sitional environment of Pabdeh Formation in Dashte ndash Arjan area (Shiraz Fars Zagros Iran) ndash World Acade-my of Science Engineering and Technology 4 818-822

MoHseni H amp aL-asaM is (2004) Tempestite deposits on a storm-influenced carbonate ramp an example from the Pabdeh Formation (Paleogene) Zagros Basin SW Iran ndash Journal of Petroleum Geology 27 (2) 163-178

MoHseni H beHbaHani R KHodabaKH s amp atasHMaRd Z (2011) Depositional environments and trace fossil as-semblages in the Pabdeh Formation (Paleogene) Zagros Basin Iran ndash Neues Jahrbuch fuumlr Geologie und Palaumlon-tologie Abhandlungen 262 (2) 59-77

MuRRis RJ (1980) Middle East stratigraphic evolution and oil habitat ndash Bulletins of the American Association of Petroleum Geologists 4 597-618

oRtMann A (1898) Gliederfuumlssler Arthropoda ndash In Bronnrsquos Klassen und Ordnungen der Tierreichs 5 (2) Crustacea Malacostraca (47-52) [Systematik] 1057-1168

PaRR WJ (1947) An Australian record of the foraminiferal genus Hantkenina ndash Proceedings of the Royal Society of Victoria 58 45-47

PeacuteReZ faRfante I amp KensLey B (1997) Penaeoid and Sergestoid shrimps and prawns of the world Keys and diagnoses for the families and genera ndash Meacutemoires du Museacuteum national drsquoHistoire naturelle 175 11-223

PeaRson Pn oLsson RK HeMLeben c HubeR bt amp beRggRen Wa (2006) Atlas of Eocene Planktonic Foraminifera ndash Cushman Foundation of Foraminiferal Research Special Publications 41 513 pp Lawrence (Allen Press)

PRieM F (1908) Poissons fossiles de Perse ndash Annales drsquoHistoire Naturelle Paleacuteontologie 1 1-25

quayLe WJ (1987) English Eocene Crustacea (Lobsters and Stomatopod) ndash Palaeontology 30 (3) 581-612


Rafinesque-scHMaLtZ CS (1815) Analyse de la nature ou tableau de lrsquounivers et des corps organiseacutes ndash 224 pp Palermo (Barravecchia)

RatHbun MJ (1935) Fossil Crustacea of the Atlantic and Gulf Coastal Plain ndash Geological Society of America special papers 2 1-160

RogeR J (1946) Les inverteacutebreacutes des couches agrave poissons du Creacutetaceacute supeacuterieur du Liban ndash Meacutemoires de la Societeacute geacuteologique de France 23 1-92

scHoLtZ G amp RicHteR S (1995) Phylogenetic systematics of the reptantian Decapoda (Crustacea malacostraca) ndash Zoological Journal of the Linnean Society 113 289-328

scHWeitZeR CE feLdMann RM gaRassino A Ka-RasaWa H amp scHWeigeRt G (2010) Systematic list of fossil decapod crustacean species ndash Crustaceana Mono-graphs 10 1-222

squiRes RL (2001) Additions to the Eocene Megafossil Fauna of the Llajas Formation Simi Valley southern California ndash Contributions in Science 489 1-40

tiRMiZi NM (1960) Crustacea Penaeidae Part II Series Benthesicymae ndash Scientific Reports of the John Murray Expedition 10 (7) 319-383

Wood-Mason J (1891) Phylum Appendiculata Branch Arthropoda Class Crustacea ndash In Wood-Mason J amp aLcocK A (eds) Natural history notes from HM In-dian marine survey steamer ldquoInvestigatorrdquo Commander RF Hoskyn RN commanding Series II No 1 On the results of deep-sea dredging during the season 1890-91 ndash Annals and Magazine of Natural History 8 (6) 269-286

Woods H (1925-1931) A monograph of the fossil macrur-ous Crustacea of England ndash Palaeontographical Society of London Monographs 127 pp

ZaHedi M (1993) Geological quadrangle map of Iran No E8 sheet Shahr-e-kurd ndash Tehran (Geological Survey of Iran)

Manuscript received May 26th 2014Revised version accepted by the Stuttgart editor August 11th 2014

Addresses of the authors

aLessandRo gaRassino Sezione di Paleontologia degli Invertebrati Museo di Storia Naturale Corso Venezia 55 20121 Milano Italye-mail alegarassinogmailcomaLi baHRaMi MeHdi yaZdi Department of Geology Uni-versity of Isfahan POB 81746-73441 Isfahan IR Irane-mails Bahrami_geoyahoocom mehyazdigmailcomfRancisco J vega Instituto de Geologiacutea Universidad Na-cional Autoacutenoma de Meacutexico Ciudad Universitaria Coyo-acaacuten 04510 Meacutexico DF Mexicoe-mail vegverunammx


includes shale pelagic lime and calciturbidite facies The existence of planktonic bioclasts related to deep sea such as Globigerina drsquooRbigny 1826 Globrotalia

cusHMan 1927 and abundant micrite indicates the de-position of this group in a deep sea environment The existence of glauconite facies (as authigenic and inter-

Fig 1 A ndash General location map of the studied area B ndash Geological map of the Shahr-e-kurd area the rectangle at the upper left of the map shows the studied locality (modified after ZaHedi 1993)






















Fig 4 A-C D-I







































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Fig 5 A-I




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Fig 5 A-I




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Fig 5 A-I




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Fig 5 A-I




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report on decapod crustaceans from the eocene of zagros basin, iran

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