Boletín de la Sociedad Geológica Mexicana Volumen 76, núm. 1, A200324, 2024 http://dx.doi.org/10.18268/BSGM2024v76n1a200324

New report of decapod crustaceans from the Miocene of Iran

Nuevo reporte de crustáceos decápodos del Mioceno de Irán

Alessandro Garassino¹, Giovanni Pasini², Majid Mirzaie Ataabadi³, Mehdi Hadi⁴, Meraj Parsazad³, Torrey Nyborg¹, Francisco J. Vega^5,*

¹ Department of Earth and Biological Sciences, Loma Linda University, Loma Linda, CA 92350, USA.

² Via Alessandro Volta 16, 22070 Appiano Gentile (Como), Italy.

³ Department of Geology, Faculty of Science, University of Zanjan, Zanjan, Iran.

⁴ Independent Researcher.

⁵ Instituto de Geología, Universidad Nacional Autónoma de México, Ciudad Universitaria, 04510 Coyoacán, CDMX, Mexico.

* Corresponding author: (F.J. Vega) This email address is being protected from spambots. You need JavaScript enabled to view it.

How to cite this article:

Garassino, A., Pasini, G., Mirzaie Ataabadi, M., Hadi, M., Parsazad, M., Nyborg, T., Vega, F.J., 2024, New report of decapod crustaceans from the Miocene of Iran: Boletín de la Sociedad Geológica Mexicana, 76 (1), A200324. http://dx.doi.org/10.18268/BSGM2024v76n1a200324

Manuscript received: March 1, 2024; Corrected manuscript received: March 20, 2024; Manuscript accepted: March 25, 2024.

ABSTRACT

We report some brachyuran crabs from the late Miocene of Dar Pahn unit, Makran Province, Iran. The studied specimens have been assigned to Ixoides MacGilchrist, 1905, with I. miocenicus n. sp. (Leucosiidae Samouelle, 1819) and Persianus arcuatus n. gen., n. sp. (Portunidae Rafinesque, 1815). Ixoides miocenicus n. sp. represents the first fossil representative of the genus, enlarging its stratigraphic range to the Miocene. Persianus arcuatus n. gen., n. sp. represents the first report of a portunid crab from the Miocene of Iran. We also report an indeterminate majoid crab, representing the first report for the Majoidea from the Miocene of Iran. Finally, one portunid crab is compared with Persianus n. gen. based on some similar morphological characters.

Keywords: Crustacea, Brachyura, Leucosioidea, Majoidea, Portunoidea, Miocene, taxonomy.

RESUMEN

Reportamos algunos cangrejos braquiuros del Miocene tardío de la unidad Dar Pahn, Makran, Irán. Los especímenes estudiados son asignados a Ixoides MacGilchrist, 1905, con I. miocenicus n. sp. (Leucosiidae Samouelle, 1819) y Persianus arcuatus n. gen., n. sp. (Portunidae Rafinesque, 1815). Ixoides miocenicus n. sp. representa el primer fósil del género, extendiendo su alcance estratigráfico al Mioceno. Persianus arcuatus n. gen., n. sp. representa un reporte adicional de cangrejos portunoides del Mioceno de Iran. También se reporta un cangrejo majoideo indeterminado, el cual es el primer reporte de Majoidea del Mioceno de Irán. Por último, un cangrejo portúnido es comparado con Persianus n. gen. con base en características morfológicas similares.

Palabras clave: Crustacea, Brachyura, Leucosioidea, Majoidea, Portunoidea, Miocene, taxonomía.

1. Introduction

The fossil record of decapod crustaceans from Iran starts with Jurassic and Cretaceous taxa. Eryma bedelta (Quenstedt, 1857) was reported from the Middle Jurassic (Aalenian) of northern Iran (Förster and Seyed-Emami, 1982). Joeranina xizangensis Wang, 1981 and Huhatanka iranica Yazdi et al., 2010 were reported from the early Cretaceous (Albian) of Central Iran (Wang, 1981; Yazdi et al., 2009, 2010). Finally, Paraclytia valashtensis McCobb and Hairapetian, 2009 was reported from the Late Cretaceous of northern Iran (McCobb and Hairapetian, 2009).

Decapod crustaceans are also present in the Cenozoic of Iran. Toraby and Yazdi (2002) reported a portunid crab from the Miocene in an abstract, but a full description was never provided. Vega et al. (2010) reported Leucosia persica, Philyra hormozganensis, and Harpactocarcinus miocenicus from the Miocene of southern Iran. Hyžný et al. (2013) reported Glypturus persicus from the middle-late Miocene of the Mishan Formation (southwestern Iran). Vega et al. (2010) also recorded Arcania assigning it to the genus level due to its poor preservation. Finally, Key Jr. et al. (2017) and Khosravi et al. (2022) reported several brachyurans (mostly leucosiid crabs), including Leucosia persica, Myra sp., and Galene dashtbani from the Miocene Mishan Formation in the Zagros Moutains (southwestern Iran).

The purpose of this paper is the description of some new decapod crustaceans from the late Miocene of the Dar Pahn unit in the Makran Geological province, Southeastern Iran.

The studied specimens have been assigned to Ixoides miocenicus n. sp., the first representative of this genus in the fossil record and to Persianus arcuatus n. gen., n. sp., the first report of a portunid crab from the Miocene of Iran. In addition, an indeterminate majoid crab is reported.

2. Geological setting

The study area is part of the Makran geological zone in southeast Iran (Figure 1). The Makran zone is a large accretionary wedge, formed by the convergence between the Eurasian and the Arabian plates, resulting in a sediment thickness of about 7000 m (Falcon, 1947; Farhoudi and Karig, 1977; Fruehn et al., 1997; Kopp et al., 2000). This zone is separated from the Zagros zone to the west, by the Minab‐Zendan Transform Fault and to the east by the Chaman Transform Fault System in Pakistan (Stöcklin 1968; Bird et al., 1975). The tectonic and structural styles of the Makran accretionary complex developed during the Cenozoic at the convergent plate boundary resulting from the subduction of the oceanic part of the Arabian plate beneath the Lut and Afghan blocks of Eurasia (Byrne et al., 1992; Kopp et al., 2000; Haghipour, 2014). This zone contains sedimentary and igneous associations divided into four main sub-divisions (North, Inner, Outer, and Coastal Makran), which reflect different stages in the evolution of the Makran accretionary wedge (Dolati, 2010).

Figure 1. Location map of the Rudig section, east of Chahbahar, southeastern Iran.

The studied stratigraphic section is located about 6 km south of Rudig village and 50 km east of Chabahar Sistan and Baluchistan province (Figure 1). This 100 m thick fossiliferous section is composed of different lithological units including sandstone, silty marl, marly limestone, and siltstone (Figure 2), the “Dar Pahn” unit (McCall, 1985) within the Coastal Makran area (Figure 3). Previous studies attributed the Dar Pahn unit deposits to the Middle-Upper Miocene to Pliocene (McCall, 1985, 1997; McCall and Kidd, 1992; McCall, et al., 1994; Dolati, 2010). The lower boundary of Dar Pahn unit is cut-off by the Chah Khan Thrust in the northern areas (ca. 70- 80 km north of Rudig), while this marl-dominated member is covered unconformably by the continental deposits of the Pliocene–Pleistocene Nahang Unit (Dolati, 2010).

Figure 2. Stratigraphic column of the Rudig section (Dar Pahn unit, Late Miocene, Tortonian), in Makran zone of southeast Iran coast. Calcareous nannofosils zones, and the position of crustacean fauna are indicated (modified after Hadi et al., 2024).

The crustacean-bearing fossil layer is approximately at the 90 m level of the Rudig section at the one-meter interval of marly limestone and silty marl of Tortonian age. Fossils were discovered during excavation of a fossil whale from the surrounding sediment layers. Decapod crustaceans were obtained in-situ while cracking and opening the marly limestone layers. The measured beds contain late Miocene calcareous nannofossils. They are diverse and hence provide a high-resolution biostratigraphy (Hadi et al., 2024). Based on these fossils, the measured interval and its crustacean fauna is confined to the Tortonian stage. The calcareous nannofossils are correlative with the NN9 to NN11 biozones (Figure 2).

Figure 3. Geological map of southeastern Iran with the Cenozoic lithostratigraphic units (modified after Dolati, 2010).

3. Material

Four three-dimensionally preserved specimens, lacking appendages and ventral surfaces. The specimens are stored at the Paleontological Laboratory (PL) of Zanjan University (ZNU), Zanjan, Iran (hence ZNUPL). They are coded and numbered as Chabahar Crab (CCXX).

For the higher-level classification, we follow the recent arrangement proposed by Karasawa et al. (2019) and Schweitzer et al. (2020, 2021). For the terminology of leucosiid crab, we follow Galil (2001), whereas for the portunid crab we follow Schweitzer et al. (2021).

4. Systematic palaeontology

Superfamily Leucosioidea Samouelle, 1819

Family Leucosiidae Samouelle, 1819

Subfamily Ebaliinae Stimpson, 1871

Genus Ixoides MacGilchrist, 1905

Type species: Ixoides cornutus MacGilchrist, 1905 by monotypy.

Included fossil species: Ixoides miocenicus n. sp. (this study).

Ixoides miocenicus Garassino, Pasini, Mirzaie Ataabadi and Nyborg n. sp.

Figure 4

Figure 4. Ixoides miocenicus n. sp., Holotype ZNUPL-CC2 (A), same specimen outside matrix (B), line drawing (C). Scale bar equals 10 mm.

Etymology: The trivial name alludes to the Miocene.

Diagnosis: Carapace rhomboidal wider than long, globose; front with a shallow median depression; rounded orbital margin with two triangular lobes separated by deep diagonal grooves; stout epibranchial spine; hepatic region with a rounded bulge; wide bulge-shaped intestinal spine.

Type material: Holotype ZNUPL-CC2.

Type locality: Rudig village 50 km east of Chabahar.

Type age: Miocene (Tortonian).

Material and measurements: One three-dimensionally specimen, preserving a nearly complete carapace with broken right corner, in dorsal view. [ZNUPL-CC2 – lcxp: 34 mm (as preserved); wcxp: 44 mm (excluding epibranchial spine)].

Description: Carapace rhomboidal wider than long, globose; carapace dorsal surface finely granulated, granules somewhat larger on gastric and cardiac regions; frontal margin poorly preserved; front weakly produced anteriorly with a shallow axial depression flanked by a pair of short ridge; rounded orbital margins with two triangular ridges separated by deep diagonal grooves flanked by one ridge; sinuous, oblique anterolateral margins, diverging posteriorly, creating a strong inflexion at level of hepatic bulge; elongate epibranchial spines, lacking distal portion; sinuous, oblique posterolateral margins, narrowing posteriorly, creating an inflexion at level of mesobrachial regions; regions poorly distinct; hepatic regions with a rounded bulge; rounded, swollen intestinal region, laterally defined by weak grooves; wide bulge-shaped intestinal spines rounded distally.

Discussion: Based on MacGilchrist (1905: 255), the studied specimen shows the main distinctive proxy-characters of Ixoides MacGilchrist, 1905 in having a rhomboidal carapace, front moderately prominent, stout epibranchial spine, and a bulge-shaped intestinal spine.

Galil (2001: 173) synonymised Ixoides under Arcania Leach, 1817 and transferred its type species there. Several authors concurred with this systematic arrangement (Schweitzer et al., 2010; Karasawa, 2014; Moazzam and Kazmi, 2016; Karasawa et al., 2019, among others). However, Ng et al. (2017: 48) did not concur with this systematic arrangement, highlighting that the external morphology of Ixoides is quite different, and therefore the genus should be recognized as distinct. This suggestion is supported by WoRMS that considers Arcania cornuta as a superseded combination (AphiaID - urn:lsid:marinespecies.org:taxname:441088), keeping valid the original combination Ixoides cornutus (AphiaID - urn:lsid:marinespecies.org:taxname:453515). In conclusion, based on WoRMS Ixoides must be considered as a valid genus within the Leucosiidae, distinct from Arcania (AphiaID - urn:lsid:marinespecies.org:taxname:450157). The original systematic arrangement was also supported by Galil and Ng (2023: 74).

Ixoides is known to date with the sole extant species I. cornutus, which has a widespread distribution in Fiji Island, New Caledonia, Papua New Guinea, Vanuatu, Japan, China, Philippines, Vietnam, Persian Gulf, Madagascar, Mozambique Channel, west coast India and Pakistan (Galil, 2001: 176; Moazzam and Kazmi, 2016: 441; Galil and Ng, 2023: 76). We justify the description of Ixoides miocenicus n. sp. not only from a stratigraphic point of view, but also in having sinuous, oblique posterolateral margins, narrowing posteriorly, creating an inflexion at level of mesobrachial regions (vs oblique posterolateral margin in I. cornutus) and narrow, bulge-shaped intestinal spine rounded distally (vs intestinal spine a short stump flanked by posterior spines papillate in I. cornutus) (Galil, 2001: 174, Figure 1B).

In conclusion, Ixoides miocenicus n. sp. represents the first fossil representative of the genus, enlarging its stratigraphic range back to the Miocene. Finally, the new species is the second report of the Leucosiidae from the Miocene of Iran after Leucosia persica Vega, Gholamalian and Bahrami, 2010. Vega et al. (2010: 491, Figure 6d) reported also a poorly preserved specimen assigning it to Arcania sp. We raise some doubts about the assignment to this genus, lacking the main proxy-characters of Arcania, such as the prominent bilobate front and the margins of carapace spinose.

Superfamily Majoidea Samouelle, 1819

Family, subfamily and genus indeterminate

Figure 5

Figure 5. Majoidea indeterminate, ZNUPL-CC3, dorsal view (A), frontal view showing the incomplete fronto-orbital margin (B). Scale bar equals 10 mm.

Locality: Rudig village 50 km east of Chabahar.

Material and measurements: One three-dimensionally preserved specimen, lacking front and orbits [ZNUPL-CC3 – lcxp: 42 mm (as preserved); 45 wcxp: mm].

Description: Subpentagonal carapace, widest at level of branchial regions; fronto-orbital margin poorly preserved; straight anterolateral margins with two small pointed spines; epibranchial spines stronger than anterolateral spines directed outward; smooth, curved posterolateral margins narrowing posteriorly; posterior margin poorly preserved; inflated regions; epigastric regions with one axial tubercle; protogastric regions with one lateral tubercle; mesogastric region with rhomboidal-shaped tubercles, two axial and two lateral; meta- and urogatric regions with one axial tubercle respectively; pentagonal-shaped cardiac region with one tubercle posteriorly; depressed intestinal region; epibranchial regions with three aligned tubercles forming an arched ridge; smooth meso- and metabranchial regions; deep, sinuous cervical and branchiocardiac grooves, delimiting gastric, urogastric, and cardiac regions respectively; smooth dorsal surface.

Discussion: Based on Schweitzer et al. (2020) the studied specimen can be assigned to the Majoidea Samoulle, 1819 for two characters, as follows: the carapace widest in branchial regions and anterolateral margins with spines. Unfortunately, the lack of the fronto-orbital margins does not allow us to assign the studied specimen to a family and a subfamily. Indeed, the lack of a bifid or singular rostrum, the presence or absence of the intercalated spine, and the presence or absent of the supraorbital eave preclude the possibility to assign the studied specimen to the family level within the Majoidea. However, we can presume that it could represent a new genus within this superfamily in having strong tubercles in gastric, cardiac, and epibranchial regions and three anterolateral spines (including the epibranchial spines) not shared with any fossil genera known to date.

In conclusion, though the studied specimen clearly shows some peculiar characters, enough to justify the description of a new genus within the Majoidea, we prefer to leave it in open nomenclature, waiting for better preserved specimen useful to clarify its systematic assignment.

Superfamily Portunoidea Rafinesque, 1815

Family Portunidae Rafinesque, 1815

Subfamily Portuninae Rafinesque, 1815

Genus Persianus Garassino, Pasini, Mirzaie Ataabadi and Nyborg nov.

Type species: Persianus arcuatus n. gen., n. sp. by monotypy.

Etymology: From Persia, ancient name of Iran. Gender: masculine.

Diagnosis: Subhexagonal carapace, much wider than long; flat, bilobate front with axial notch; supraorbital margin with two fissures; short, convex anterolateral margins with three spines (excluding outer-orbital spines); very elongate epibranchial spine gently curved upward; oblique posterolateral margins longer than the anterolateral with P5 concave recess; granulated, short arcuate epigastric ridges; granulated, transverse protogastric ridge; mesobranchial region depressed; wide epibranchial regions with one arched transverse ridge.

Persianus arcuatus Garassino, Pasini, Mirzaie Ataabadi and Nyborg n. sp.

Figure 6A-B

Figure 6. Persianus arcuatus n. gen., n. sp., ZNUPL-CC1, holotype (A), line drawing (B), Persianus cfr. P. arcuatus, ZNUPL-CC6 (C). Scale bar equals 10 mm.

Etymology: The trivial name alludes to the elongate curved epibranchial spine.

Diagnosis: As for the genus.

Type material: Holotype, ZNUPL-CC1.

Type locality: Rudig village 50 km east of Chabahar.

Type age: Miocene (Tortonian).

Material and measurements: One carapace in dorsal view, lacking the posterior margin partially, chelipeds, ambulatory legs, and ventral surfaces [ZNUPL-CC1 – lcxp: 16 mm (as preserved); wcxp: 35 mm].

Description: Subhexagonal smooth carapace, much wider than long; flat, bilobate front with axial notch; wide curved orbits nearly 50% wider than the frontal width; orbits delimited by inner- and extra-orbital spines; granulated supraorbital margins with two deep fissures, first medially and second one at basis of the outer-orbital spines; short, convex, anterolateral margins with three spines (excluding outer-orbital spines), almost equal in size and slightly directed forward; very elongate epibranchial spines gently curved upward; oblique posterolateral margins longer than anterolateral with distinct P5 concave recess; posterior margin not preserved; short, granulated epigastric ridges interrupted medially; elongate, granulated, continuous protogastric ridge; mesobranchial region depressed; flat hepatic regions; urogastric, cardiac, intestinal regions poorly preserved; wide epibranchial regions with one arcuate transverse ridge respectively; flat meso- and metabranchial regions; smooth dorsal surface.

Discussion: Based on Schweitzer et al. (2021), the studied specimen can be assigned to the Portunoidea for some characters, such as the subhexagonal carapace, the lobate front, the spiny anterolateral margins, and the arcuate epibranchial ridges.

Based on Schweitzer et al. (2021), the Portunoidea includes seven families, as follows: Carcineretidae Beurlen, 1930; Carcinidae MacLeay, 1838; Geryonidae Colosi, 1924; Lithophylacidae Van Straelen, 1936; Longusorbiidae Karasawa, Schweitzer and Feldmann, 2008; Portunidae Rafinesque, 1815; and Psammocarcinidae Beurlen, 1930. Some characters of the studied specimen, such the shape of the carapace, the lobate front, the shape of the orbit, the number of anterolateral spines, and the presence of ridges in some regions of the carapace, allow us a comparison with the above-mentioned families for its systematic assignment at family and subfamily levels.

The subhexagonal carapace and the front without spines rule out the belonging of the studied specimen to the Carcineretidae. The lobate front of the studied specimen excludes its belonging to the Carcinidae. The oval or subhexagonal carapace, the front with three spines, and the anterolateral margins with alternate pointed spines rule out the belonging of the studied specimen to the Geryonidae. The orbits extremely wide, occupying entire maximum width of carapace, which converge posteriorly, exclude the belonging of the studied specimen to the Lithophylacidae. The rostrum axially sulcate and strongly downturned distally and the spineless anterolateral margins rule out the belonging of the studied specimen to the Longusorbiidae. The front with three spines (excluding the inner-orbital spines) and the anterolateral margins with five spines (including the outer-orbital spines) exclude the belonging of the studied specimen to the Psammocarcinidae.

Finally, the subhexagonal carapace, the anterolateral margins with 3-9 spines (including the outer-orbital spines), and the well-developed epibranchial ridges allow us to assign the studied specimen to the Portunidae.

Based on Schweitzer et al. (2021), the Portunidae includes six subfamilies, as follows: Carupinae Paul’son, 1875; Necronectinae Glaessner, 1928; Lupocyclinae Paul’son, 1875; Podophthalminae Dana, 1851; Portuninae Rafinesque, 1815; and Thalamitinae Paul’son, 1875. The ovate carapace and the absence of ridges on the carapace regions rule out the belonging of the studied specimen to the Carupinae and Necronectinae. Though the Lupocyclinae shares with the studied specimen the gastric and epibranchial ridges, the anterolateral margins lacking the elongate epibranchial spine exclude the belonging of the studied specimen to this subfamily. The narrow front and the orbits extremely wide rule out the belonging of the studied specimen to the Podophthalminae. The less wide carapace and the lack of a distinct elongate epibranchial spine rule out the belonging of the studied specimen to the Thalamiitinae.

Based on the diagnosis of the Portuninae provided by Schweitzer et al. (2021), the studied specimen is assigned to this subfamily for the number of the anterolateral spines and arcuate epibranchial ridge. The Portuninae includes seven genera, as follows: Portunus Weber, 1795; Acanthoportunus Schweitzer and Feldmann, 2002; Arenaeus Dana, 1851; Callinectes Stimpson, 1862; Colneptunus Lőrenthey in Lőrenthey and Beurlen, 1929; Pseudoachelous Portell and Collins, 2004; and Rathbunites Schweitzer, Dworschak and Martin, 2011.

The anterolateral margins with five spines (Rathbunites), with eight spines (Acanthoportunus, Colneptunus, Pseudoachelous) and with nine spines (Arenaeus, Callinectes, Portunus) exclude the belonging of the studied specimen to these genera.

In conclusion, based on the above observations, we justify the description of the new genus Persianus within the Portuninae in having the anterolateral margins with only three flat spines slightly directed forward and the elongate epibranchial spine curved forward.

Portunid crabs from the Miocene of Iran were reported by Toraby and Yazdi (2002) in an abstract, and by Heidari et al. (2012). The record of Persianus n. gen. represents formally the first report of a portunid crab from the Miocene of Iran.

Persianus cf. P. arcuatus n. sp.

Figure 6C

Locality: Rudig village 50 km east of Chabahar.

Material and measurements: One incomplete carapace in dorsal view, lacking chelipeds, ambulatory legs, and ventral surfaces [ZNUPL-CC6 – lcxp: 29 mm (as preserved); wcxp: 32 mm].

Description: Subhexagonal carapace wider than long; frontal margin poorly preserved; straight wide orbits (as preserved); short, convex anterolateral margins with serrate four anterolateral spines (excluding outer-orbital spines); epibranchial spine gently curved upward lacking distal portion; oblique posterolateral margin longer than anterolateral margins with distinct P5 concave recess; straight posterior margin; dorsal regions poorly preserved, convex transversally; elongate, granulated, continuous protogastric ridge; wide epibranchial regions with one arched ridge respectively.

Discussion

The studied specimen has a poorly preserved front and dorsal regions. In addition, the incomplete epibranchial spines do not allow a complete assignment of the studied specimen. However, the specimen does have some characters that resemble Persianus arcuatus n. sp., such as: the shape of carapace and the anterolateral spines arrangement; the presence of a continuous protogastric ridge; and an elongate epibranchial spine. The proportionally wider carapace size (as preserved) could be interpreted as an intraspecific character, whereas the presence of four serrate anterolateral spines (vs three smooth spines in P. arcuatus n. sp.) suggest perhaps another systematic assignment, therefore, we choose to leave the specimen in open nomenclature, waiting for more complete preserved specimens useful for a closer comparison.

Contributions of authors

Alessandro Garassino and Giovanni Pasini - identification and description of specimens. Majid Mirzaie Ataabadi, Mehdi Hadi and Meraj Parsazad - fieldwork and sampling, conceptualization, investigation. Torrey Nyborg and Francisco J. Vega writing original draft.

Financing

This work did not receive any financial support.

Acknowledgements

We wish to thank two anonymous reviewers for careful review and criticism of the manuscript. MMA appreciate support from Iran’s Department of Environment (Sistan and Baluchistan province and Chabahar offices) and University of Zanjan.

Conflicts of interest

The authors state that there are no conflicts of interest.

Handling editor

Antoni Camprubí.

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