New species of Kromtitis Müller, 1984 (Decapoda: Brachyura: Dynomenidae) from the Eocene of Iberian Peninsula
Article number: 22.2.47
Copyright Paleontological Society, August 2019
Submission: 13 February 2019. Acceptance: 12 July 2019
A new species of a fossil brachyuran decapod crustacean, Kromtitis lluisprietoi sp. nov. (Dynomenidae, Paradynomeninae), from the Lutetian (middle Eocene) outcrops of Alicante province (Spain), is described. Kromtitis is a genus of dynomenid crabs, known only from the fossil record, characterized by a dorsal carapace with areolate morphology, with rounded tubercles ornate with granules, and irregular lateral teeth. In contrast to its congeners, Kromtitis lluisprietoi sp. nov. has less vaulted carapace with areolate regions bearing raised tubercles ornamented with granules. The genus Kromtitis is well-known from several species reported from the Eocene strata of northern Italy and Hungary. Kromtitis lluisprietoi sp. nov. represents the first record of the genus from the Iberian Peninsula.
Àlex Ossó. Llorenç de Villalonga, 17B, 1er-1ª 43007-Tarragona, Catalonia.
Keywords: Podotremata; Dromioidea; Paradynomeninae; new species; Lutetian; Alicante
Ossó, Àlex. 2019. New species of Kromtitis Müller, 1984 (Decapoda: Brachyura: Dynomenidae) from the Eocene of Iberian Peninsula. Palaeontologia Electronica 22.2.47A 1-9. https://doi.org/10.26879/967
Copyright: August 2019 Paleontological Society.
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The discovery of Kromtitis Müller, 1984, in Lutetian strata of La Vila Joiosa (Alicante) increases the number of decapod species previously reported from the Eocene of Alicante province (Via, 1991; Ossó, 2011). The presence of Kromtitis in the outcrops of Alicante province adds new evidence of the affinity between the decapod fauna of the eastern part of the Iberian Peninsula and the coeval faunas of the Central Pyrenees of Aragon and Catalonia (north and north-eastern parts of the Iberian Peninsula, respectively), and the faunas of northern Italian Peninsula and Hungary. This further supports a relative faunal homogeneity among the westernmost Tethys decapod faunas during the Eocene (e.g., Domínguez and Ossó, 2016; Khodaverdi et al., 2016). Kromtitis, with its oldest representative reported from the Paleocene of Denmark (Collins, 2010), is relatively common in Eocene and Oligocene reefal outcrops of northern Italy (Beschin et al., 2002, 2007, 2018) and Hungary (Müller and Collins, 1991). From the Miocene strata, Kromtitis has been reported from reefal settings of Austria, Hungary and Poland (Bachmayer and Tollmann, 1953; Müller, 1984, 1996), as well as from Jamaica (Portell and Collins, 2004).
The studied specimen was recovered in the municipality of La Vila Joiosa (Marina Baixa, Alicante province, Valencian Community, Spain), in the glades of a place called l’Arginent, in the northern flank of the Alt de l’Olivanet, midway between La Vila Joiosa and Finestrat (Figure 1.1, Figure 2.1). Geologically, the area belongs to the Internal Prebetic System (sensu García-Hernández, 1978) of the Betic Ranges, eastern Iberian Peninsula. L’Arginent outcrop is well known since the nineteenth century as one of the localities within the Alicante province with rich record of Eocene echinoids (e.g., Cotteau, 1890; Vaello López, 2015). However, no specific geological or stratigraphic works have been published on this particular outcrop. The 1981 IGME Chart (847-Villajoyosa) indicates the outcrop’s area as undifferentiated Paleogene series of clayey marls, biocalcarenites and levels of fossiliferous limestones, ranging from Danian to Aquitanian (TA-Ba11-12). Nevertheless, the abundant associate fauna of large foraminifera and echinoids found at l’Arginent, exemplified by Assilina exponens (Sowerby, 1840); Nummulites millecaput Boubée, 1832; Ditremaster nux Desor, 1853; Linthia macphersoni Cotteau, 1889; Arachniopleurus reticulates Duncan and Sladen, 1882; Salenia garciae Cotteau, 1890; or even decapods as Dromilites pastoris Vía, 1959, Lophoranina straeleni Vía, 1959, and Calappilia cf. scopuli Quayle and Collins, 1981, is identical to that of other well-calibrated localities of the Alicante province, such as Orxeta, Busot, Villafranqueza or Agost, which indicates a Lutetian (middle Eocene) age for the l’Arginent outcrop (Jiménez de Cisneros, 1917; Sillero, 1992; Molina et al., 2000; Sillero, 2002; Ossó-Morales, 2011, 2013; Vaello López, 2015) (Figures 2.2-2.5).
Infraorder BRACHYURA Latreille, 1802
Section PODOTREMATA Guinot, 1977
Subsection DYNOMENIFORMIA Guinot, Tavares and Castro, 2013
Superfamily DROMIOIDEA De Haan, 1833
Family DYNOMENIDAE Ortmann, 1892
Subfamily PARADYNOMENINAE Guinot, 2008
Genus KROMTITIS Müller, 1984
Type species. Dromilites koberi Bachmayer and Tollmann, 1953, by monotypy.
Included species. K. bicuspidatus Beschin et al., 2009; K. daniensis Collins, 2010; K. koberi (Bachmayer and Tollmann, 1953); K. koberiformis Beschin et al., 2007; K. levigatus Beschin et al., 2007; K. lluisprietoi sp. nov.; K. pentagonalis Müller and Collins, 1991; K. spinulata Portell and Collins, 2004; K. subovatus Beschin et al., 2007; K. tergospinosus Beschin et al., 2018; K. tetratuberculatus Beschin et al., 2002.
Kromtitis lluisprietoi sp. nov.
Diagnosis. Carapace small, roundish subquadrate, as long as wide, markedly convex, surface areolate, sparsely granulate. Regions swollen, with rounded tubercles peaked with coarse granules and well-defined by grooves. Front with produced and triangular rostrum. Orbits ovate, obliquely arranged, visible dorsally. Anterolateral margin with five irregular teeth (excluding the outer orbital tooth). Posterolateral margin with two teeth, the second being larger. Frontal groove deep. Epigastric lobes swollen with two tubercles. Mesogastric lobe subpentagonally elongate, acute anteriorly with one tubercle, larger posteriorly, with two contiguous tubercles. Metagastric region undifferentiated from mesogastric region. Protogastric lobes swollen, with three tubercles. Urogastric region wide, V-shaped, with two contiguous tubercles. Epibranchial region with two swollen lobes, each with one tubercle. Mesobranchial region with two swollen lobes bearing large tubercle. Metabranchial region incomplete, bearing one tubercle. Cardiac region diamond shaped, swollen, with three tubercles. Hepatic region slightly swollen, with one tubercle. Sub-hepatic region with small tooth. Cervical, post-cervical, branchial and branchiocardiac grooves well-defined.
Kromtitis lluisprietoi sp. nov. is easily distinguishable from all the previously known Kromtitis species in having less globose dorsal carapace and by its particular dorsal aspect, with more areolate regions bearing raised tubercles ornamented with granules. Kromtitis bicuspidatus from the upper Eocene of NE of Italy, differs from the new species in having a pair of semi-fused inner epibranchial swellings, whereas in K. lluisprietoi sp. nov. there is only one swelling; moreover, the former presents more swollen region lobes and less individualized anterolateral teeth than the new species (Beschin et al., 2009, p. 7-10, pl. 1, figs. 3-4). Kromtitis daniensis from the middle Danian of Denmark differs from the new species in having a pair of inner mesobranchial tubercles and fused metabranchial tubercles, whereas in K. lluisprietoi sp. nov., there is only one tubercle on each lobe, and its surface is more granulate (see Collins, 2010, p. 15, figs. 1.1, 2). Kromitis koberi (the type species) from the middle Miocene of Central Europe and K. koberiformis from the lower Eocene of NE Italy differ from the new species by their coarser and tighter swellings and double or semi-fused inner epi- and mesobranchial tubercles (Müller, 1984, p. 63-64, pl. 31, figs. 1-4; Beschin et al., 2007, p. 26-27, pl. 3, figs. 2-4; Schweitzer et al., 2012, p. 33, figs. 21.4a-21.4b; Collins, 2014, p. 36, pl. 2, figs. 7-8). Kromtitis levigatus from the lower Eocene of NE Italy differs from K. lluisprietoi sp. nov. by its smooth dorsal carapace (Beschin et al., 2007, p. 26-27, pl. 3, fig. 5). Kromtitis spinulata from the lower Miocene of Jamaica differs from the new species by its subglobose carapace and more spiny lateral margin than in K. lluisprietoi sp. nov. (Portell and Collins, 2004, p. 11-113, fig. 1.1). Kromtitis subovatus from the lower Eocene of NE Italy differs from K. lluisprietoi sp. nov. in having a subglobose and subpentagonal carapace and more acute lateral teeth (Beschin et al., 2007, p. 26-27, pl. 3, figs. 6-8). Kromtitis tetratuberculatus from the middle Eocene of NE Italy differs clearly from the new species by its coarser and swollen dorsal surface with a very dense granulation (Beschin et al., 2002, p. 12-13, text-fig. 7, pl. 2, figs. 2, 3a-3b). Kromtitis tergospinosus from the upper Eocene of NE Italy differs from K. lluisprietoi sp. nov. by its more flattened dorsal carapace, and regions and lateral margins more spiny than in the new species (Beschin et al., 2018, pl. 58, figs. 63-65). Kromtitis pentagonalis Müller and Collins, 1991 from the upper Eocene of Hungary was transferred to Paradynomene by Beschin et al. (2016a, p. 71), which indicates the strong similarities in respect with dorsal carapace morphology among both genera. However, the species was re-evaluated as a representative of Kromtitis by Beschin et al. (2018, p. 162). It differs from K. lluisprietoi sp. nov. by its more produced frontal margin and rostrum, general outline and more swollen regions (Beschin et al., 2018, p. 160-162, pl. 58, fig. 61). Kromtitis pseudolothi Beschin et al., 2016b, morphologically closely related to Dromilites lothi Förster and Mundlos, 1982 (ibid, p. 26), is not considered herein as Kromtitis, by the same reasons that D. lothi was excluded from Kromtitis (see discussion in Beschin et al., 2016a, p. 71, and references therein).
Etymology. The species is dedicated to the memory of late Lluís Prieto, an enthusiastic collector from Barcelona who discovered the holotype.
Holotype. GCP-CI-4528, near-complete carapace (without rostrum and posterior margin) with well-preserved cuticle, stored at Museo Paleontológico de Elche (Alicante, Spain), under acronym MUPE.
Measurements (in mm). Length=14.5; width=18.0; front-orbital width=9.5; height=7.0.
Description. Carapace small, roundish subquadrate, apparently as long as wide, markedly convex, surface areolate, sparsely granulate. Regions distinct, swollen, with well individualized rounded tubercles peaked with coarse granules, and well-defined by grooves. Front broken, probably with produced and downturned triangular rostrum. Orbits ovate, obliquely arranged, visible dorsally; supraorbital margin not well preserved, granulate; infraorbital margin with a prominent granulate tooth or projection. Anterolateral margin begins below level of postorbital tooth, armed with five irregular teeth (excluding outer orbital tooth), the third being larger, and the fifth broken; among them, irregular acute small projections ornate the margin. Posterolateral margin with two teeth, the first behind the branchial groove, the second being larger, near the posterior corner. Posterior margin not preserved. Frontal groove deep. Epigastric lobes separated by the frontal groove, swollen, elongate, with two tubercles aligned longitudinally, the anterior larger. Mesogastric lobe subpentagonally elongate, anterior portion acute, bearing one tubercle, posterior portion larger, with two contiguous tubercles separated by short and smooth axial groove. Metagastric region undifferentiated from mesogastric region. Protogastric lobes swollen, with three tubercles, two aligned longitudinally on anterior and medial portion of the lobes, the second being larger; the third tubercle placed obliquely below between the second tubercle and the gastrohepatic groove. Urogastric region wide, swollen, V-shaped, separated from mesogastric and metagastric regions by the cervical groove; with two small contiguous tubercles separated by a weak axial depression; small anterior portion of lobe divided transversely, below two gastric pits; small lateral portions of lobe obliquely divided by a groove forming small secondary lobes at each side of the principal lobe. Epibranchial region laterally elongate, bounded anteriorly by the cervical groove and posteriorly by the post-cervical groove; with two separate swollen lobes, inner and outer, each bearing one tubercle, the inner one being larger. Mesobranchial region laterally elongate, bounded anteriorly by the post-cervical groove and posteriorly by the branchial groove; with two separate swollen lobes, each bearing large tubercle, larger than the epibranchial ones, the outer one being very large and prominent. Metabranchial region incomplete, bearing one tubercle. Cardiac region diamond shaped, separated from urogastric region by post-cervical groove and bounded laterally by deep branchiocardiac grooves; swollen, bearing three tightly positioned tubercles, two of them aligned transversely and separated by axial short groove; the third positioned posteriorly. Intestinal region not completely preserved. Hepatic region slightly swollen, with one tubercle surrounded by scattered granules. Sub-hepatic region bearing a tooth below the first anterolateral tooth and the infraorbital projection. Cervical, post-cervical, branchial and branchiocardiac grooves well-defined; cervical and branchial ones visible ventrally.
Remarks. In view of the dorsal and frontal features of the studied specimen, it can be assigned with confidence to the Dynomenidae, and more accurately within the Paradynomeninae. Indeed, regarding its carapace outline, densely ornamented with marked swellings crowned by tubercles, its antero- and posterolateral margins armed with irregular salient teeth, and its produced ventral anterior area, it fits the diagnosis of Paradynomeninae (Guinot, 2008, p. 11-13). The assignment on the genus level is somewhat more complex because it lacks the posterior part of the carapace, obscuring the carapace outline and details of the intestinal region and the posterior margin. However, the roundish subquadrate carapace outline with anterolateral irregular teeth, the presence of posterolateral teeth, the swollen region lobes with rounded tubercles tipped with granules and its granulated surface with well-defined grooves correspond with the diagnosis of the genus Kromtitis (Müller, 1984, p. 64; Schweitzer et al., 2012, p. 33). Simultaneously, the new species also corroborates the diagnosis of Paradynomene, as far as the areolate dorsal carapace surface, densely covered with tubercles and granules, the anterolateral irregular salient teeth and the inflated sub-hepatic area, are concerned (McLay and Ng, 2004). Nevertheless, Paradynomene exhibits usually a more elongate and subquadrangular carapace outline, and anterolateral margins are more parallel to each other (Guinot, 2008, p. 11) than they are in the studied specimen and other species of Kromtitis, in which they are fairly convex. Additionally, the new species exhibits a wide transverse V-shaped urogastric region, which is wider than the cardiac region, a character typical for the genus Kromtitis (Müller 1984, pl. 31, figs. 1-4; Portell and Collins, 2004, fig. 1.1; Beschin et al., 2016a, pl. 8, figs. 4-6). Accordingly, based on the roundish subquadrate carapace and the particular shape of the urogastric region, the studied material is assigned herein to Kromtitis.
The systematic placement of Kromtitis changed through time. Originally it was considered a representative of Dromiidae De Haan, 1833 (Müller, 1984; De Grave et al., 2009). Guinot (2008) re-evaluated Dynomenidae Ortmann, 1892, and presented a key for extant subfamilies, chiefly based on sternal and abdominal characters, which are not preserved in the studied specimen. Therein, she proposed a new subfamily Paradynomeninae for the genus Paradynomene Sakai, 1963, as well as the fossil forms Kieronopsis Davidson, 1966, and Kromtitis, based on the strong similarities in dorsal carapace features in both fossil genera and the extant species of Paradynomene (Guinot, 2008, p. 21). Moreover, the specimen of La Vila Joiosa possesses an areolate dorsal ornamentation and irregularly toothed lateral margins, that strongly recall those of some extant species of Paradynomene (McLay and Ng, 2004), which supports the current subfamilial placement of Kromtitis within Paradynomeninae (Guinot, 2008).
Occurrence. Kromtitis lluisprietoi sp. nov. is known only from the Lutetian (middle Eocene) strata of its type locality, i.e., La Vila Joiosa (Marina Baixa, Alicante province, Valencian Community, Spain).
Guinot (2008, p. 21), when including Kromtitis within Paradynomeninae, pointed out the scarce dorsal divergence between the fossil forms and the extant species of this subfamily. In fact, she considered that the modern Paradynomeninae clearly are “barely modified relicts” (Guinot, 2008), which is also evidenced with the discovery of K. lluisprietoi sp. nov., which shares characters and features with extinct genus Kromtitis and extant species of Paradynomene (McLay and Ng, 2004). In this respect, no significant morphological differences can be found among Eocene and Miocene species of Kromtitis, or even the only Paleocene representative, K. daniensis, although poorly preserved (cast). However, as mentioned above, dorsal regions with pronounced tubercles and the anterolateral outline with prominent irregular teeth of Kromtitis lluisprietoi sp. nov. differ slightly from the typical dorsal pattern of its congeners.
Extant Paradynomeninae are restricted to the Indo-West Pacific, and usually inhabit moderately deep to deep waters (McLay and Ng, 2004, p. 1). In contrast, most fossil representatives of Paradynomeninae lived in shallower reefal environments (e.g., Beschin et al., 2016a).
The genus Kromtitis is reported for the first time from the Iberian Peninsula. The presence of Kromtitis lluisprietoi sp. nov. in the Lutetian strata of the southeastern of the Peninsula, in addition to increasing the number of known decapod species from the respective area, further supports a suggested faunal homogeneity of the western Tethys during the Eocene. Dorsal morphology of Kromtitis lluisprietoi sp. nov. differentiates the species from its congeners; however, it simultaneously shows affinity to extant representatives of the subfamily Paradynomeninae.
My sincere gratitude to M. Sánchez (Barcelona) who generously donated the holotype. I am grateful to A. De Angeli (Vicenza, Italy) for his help with enriching feedback and comparative material, and L. Hernández (Alicante, Spain) for information about the outcrop and his assistance in the field. Thanks to A. Aberásturi (MUPE, Elche, Spain) for the facilities in the donation process, to P. Artal (MGSB, Barcelona, Catalonia) and to an anonymous reviewer, whose constructive reviews improved this article.
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