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Tshudy et al.—New Oligocene lobster from Hungary


be referred to Hoploparia” (Tshudy and Sorhannus, 2003, p. 701; supporting details in their paper). 3. Hoploparia is paraphyletic. Cladistic results showed


other nephropid genera scattered throughout the Hoploparia results. To these points, we here add: 4. The range of variation in Hoploparia is inconsistent with


that in Recent genera (i.e., species of Hoploparia are morpho- logically more different from each other than are some other nephropid genera). For example, the modern Thymopides Burukovsky and Averin, 1977, and Thymops Holthuis, 1974 are, in external morphology, more closely similar to each other (seen best in a quick, comparative glance at each, but for details, see Tshudy, 1993) than many species of Hoploparia are to each other. The same is true for Homarus and Homarinus Kornfield, Williams, and Steneck, 1995 (likewise, and details in Tshudy, 1993), and especially Thaumastocheles Wood-Mason, 1874 and Thaumastochelopsis Bruce, 1988. Tshudy and Sorhannus (2003) did not propose a solution to


the problem (i.e., they did not present a revision of Hoploparia in that particular paper, but hoped to stimulate discussion and new characters and cladograms that would lead to a con- sensus and, subsequently, revision). Feldmann et al. (2007, p. 702–703) rebuffed the idea that Hoploparia was a waste- basket genus, noting that they were “struck by how similar the species currently referred to Hoploparia are to one another.” But they did not comment directly on the points (above) made by Tshudy and Sorhannus (2003). Thus, in our opinion, the definition and species composition of Hoploparia remain problematic. It would seem that a consensus from species-level


cladistic analyses would be a reasonable way of dividing the current contents of Hoploparia into smaller groups. We do not believe that a single published cladistic analysis or cluster analysis is a firm enough basis for disrupting the taxonomic stability of fossil lobsters. We cite, as an example of the apparent fallibility of computer analyses, a recently published (Schweitzer and Feldmann, 2014) cluster analysis of extant lobster genera. Among the curious results is that Thaumas- tocheles and Thaumastochelopsis are well separated, and each is paired with a lobster that looks much less like it than does its former family-mate (former family Thaumastocheli- dae). Indeed, Ahyong et al. (2007, p. 206–207) opined that the “differences between Thaumastocheles and Thaumas- tochelopsis are minor, and whether or not both genera should be recognized requires further study.” With results this strange, it seems unreasonable to make sweeping, formal changes to taxonomy on the basis of a single, computerized analysis.


Tshudy (1993) noted that, despite the range of variation


among species of Hoploparia, there were no obvious correla- tions between character states that would make subdividing the genus easy. In other words, he did not (and we do not) see certain spines associated with certain thoracic carinae, etc. Instead, each species of Hoploparia seems like a random sample from a grab bag of character states. Feldmann et al. (2007, p. 703) independently found the same. Comparison of the new species to Miocene (23.03–5.33 Myr) Hoploparia species.—Hoploparia gazdzicki Feldmann


175


and Crame, 1998, from the lower Miocene of Antarctica, is a good Hoploparia that differs from the new species in having a granulose cephalothorax, a much more distinct ventral extension of the branchiocardiac groove, and a pleon with the tergum-pleuron surface separated by a distinct ridge. Comparison of the new species to Eocene


(56–33.9 Myr) Hoploparia species.—Hoploparia alpinus (Van Straelen, 1936) from the lower Upper Eocene (‘Auversien’)of Switzerland is fairly complete and well preserved, and “clearly a Hoploparia” (Tshudy, 1993, p. 73), but we cannot verify species identification without having seen the actual specimen or at least a whitened photograph. Tshudy (1993) was unable to locate the holotype and sole specimen known. Van Straelen (1936) cited ‘Le Musée d’Histoire naturelle de la Ville de Neuchâtel’ as the repository. From the description and photocopied photographs in Van Straelen (1936, figs. 1, 2), Hoploparia alpinus has pleonal pleura 3–5 narrower near the terminations than those of the new species. Hoploparia alpinus also appears to have relatively smaller chelipeds than those on the new species. Chelipeds on the new species are approximately 1.5x longer, or more, than the cephalothorax. Hoploparia corneti Van Straelen, 1921, from the lower


Eocene (Ypresian) of Belgium, is assigned to Hoploparia, based on granulation of the skeleton and the ventral extension of the branchiocardiac groove (Tshudy, 1993, p. 109). Van Straelen (1921, p. 138) wrote that H. corneti most closely resembles H. gammaroides. Tshudy (1993) was unable to locate material during his dissertation work; H. corneti differs from the new species in having granulation and a distinct branchiocardiac groove.


Hoploparia gammaroides M’Coy, 1849, from the lower


Eocene of England and Belgium, is a good Hoploparia that differs from the new species in having a granulose cepha- lothorax and pleonal pleura that are more sickle-shaped/hook- like (see especially pleuron 3 [e.g., NHMUK 59127 and 59118] and with posterior margins more indented). Hoploparia groenlandica Ravn, 1903 has been


recorded from the lower Eocene of Greenland.We have exam- ined a latex cast; cephalothorax, pleon, and claws all support identification as Hoploparia. It differs from the new species in having a distinct ventral extension of the branchiocardiac groove, a carapace surface covered by slightly squamiform granulations, and perhaps a more distinct sculpture (grooves) on the pleon.


Hoploparia johnsoni Rathbun, 1935, from the Middle


Eocene of Alabama, U.S.A., is a good Hoploparia with a granular cephalothorax. It differs from the newspecies in having a granular cephalothorax, a better-defined ventral extension of the branchiocardiac groove, a hepatic spine, some spines on the postantennal region, and a better-defined sculpture (grooves) of the pleon.


Hoploparia wardi Quayle, 1987, from the lower Eocene of


southern England, also is a good Hoploparia with a granular cephalothorax; Quayle removed it from H. gammaroides. It can be differentiated from the new species in having a granular cephalothorax, a hepatic spine, spines on the antennal region, and a more distinct ventral extension of the branchiocardiac groove.


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