© Author(s) 2020. CC Atribution 4.0 License
On the systematic placement of Pyreneplax Ossó,  
Domínguez & Artal, 2014 (Decapoda, Brachyura, Vultocinidae) 
Taksonomska uvrstitev rodu Pyreneplax Ossó, Domínguez & Artal, 2014  
(Decapoda, Brachyura, Vultocinidae) 
Àlex OSSÓ1 & José Luis DOMÍNGUEZ2
1Llorenç de Villalonga, 17B, 1-1 43007 Tarragona, Catalonia, and Museu de Ciències Naturals de Barcelona, Pale-
ontologia, Barcelona, Catalonia; e-mail: aosso@comt.cat
2Padre Manjón, 12 50010 Zaragoza, Spain; e-mail: jl.domin@hotmail.com
Prejeto / Received 14. 10. 2019; Sprejeto / Accepted 2. 4. 2020; Objavljeno na spletu / Published online 22. 4. 2020
Key words: Goneplacoidea, Martinocarcinidae, systematics, Eocene, Priabonian, Europe
Ključne besede: Goneplacoidea, Martinocarcinidae, sistematika, eocen, priabonij, Evropa
Abstract
Examination of the thoracic sternum and pleonal elements of a new male specimen of Pyreneplax basaensis 
Ossó, Domínguez & Artal, 2014 from the upper Eocene of northern Spain confirms its assignment to the family 
Vultocinidae Ng & Manuel-Santos, 2007 and reveals the presence of an old lineage (Pyreneplax and possible 
allies) that appeared during the Eocene, persisting to the present day.
Izvleček
V prispevku opisujemo nove fosilne ostanke primerka samca vrste Pyreneplax basaensis Ossó, Domínguez & 
Artal, 2014 iz zgornjega eocena na severu Španije. Ohranjene podrobnosti delov oprsja in repa potrjujejo njegovo 
uvrstitev v družino Vultocinidae Ng & Manuel-Santos, 2007 in razkrivajo prisotnost stare linije (Pyreneplax in 
sorodni taksoni), ki se je pojavila v eocenu in je prisotna še danes.
GEOLOGIJA 63/1, 125-131, Ljubljana 2020
https://doi.org/10.5474/geologija.2020.013
Introduction
A new specimen of Pyreneplax basaensis 
with complete sternal and pleonal features and 
remains of ambulatory legs reveals additional 
features that were not seen in the type series. It 
has allowed observation of additional similar-
ities to the extant species Vultocinus anfractus 
Ng & Manuel-Santos, 2007 and a more detailed 
comparison between both taxa (see Domínguez 
& Ossó, 2019).
The genus Pyreneplax Ossó, Domínguez & 
Artal, 2014 was erected to accommodate P. ba-
saensis from the Priabonian of the south Pyrene-
an basins of Spain. Subsequently, other, closely 
similar species have been recorded from the Eo-
cene of the Atlantic Coast of North America and 
from northern Italy. On the basis of dorsal cara-
pace morphology, fossil crab species such as Lo-
bonotus saundersi (Blow & Manning, 1997) from 
South Carolina (USA) and L. granosus (Beschin, 
Busulini, De Angeli & Tessier, 2002) and L. som-
marugai Beschin, Busulini & Tessier, 2009 from 
northern Italy, have lately been transferred to the 
genus Pyreneplax (see Ossó, Domínguez & Artal, 
2014; De Angeli, 2014). 
Dorsal and ventral (thoracic sternum, pleon) 
features of Pyreneplax basaensis confirm its 
family relationship with Vultocinus anfractus, in 
spite of the time span that separates both spe-
126 Àlex OSSÓ & José Luis DOMÍNGUEZ
cies, placing the origins of the family Vultocini-
dae in the late Eocene. Furthermore, new DNA 
molecular studies of Vultocinus anfractus have 
revealed that, “it comes out as a long branch in-
side the Heterotremata, far away from the rest 
of the Goneplacoidea” (Ng & Tsang, pers. comm., 
June/2019).
The studied material is housed at Museo de 
Ciencias Naturales de la Universidad de Zarago-
za (Spain), under acronym MPZ.
Systematic palaeontology
Order Decapoda Latreille, 1802
Infraorder Brachyura Latreille, 1802
Section Eubrachyura de Saint-Laurent, 1980
Subsection Heterotremata Guinot, 1977
Superfamily Goneplacoidea MacLeay, 1838
Family Vultocinidae Ng & Manuel-Santos, 2007
Genus Pyreneplax Ossó, Domínguez & Artal, 
2014 , emend.
Type species: Pyreneplax basaensis Ossó, 
Domínguez & Artal, 2014.
Other species included: Pyreneplax granosa 
(Beschin, Busulini, De Angeli & Tessier, 2002), 
Pyreneplax saundersi (Blow & Manning, 1997) 
and Pyreneplax sommarugai (Beschin, Busulini 
& Tessier, 2009).
Diagnosis (emended): Small- to medium-sized 
carapace, suboctagonal, from wider than long to 
slightly wider than long, slightly convex in an-
terior third, widest at level of third anterolater-
al tooth. Dorsal regions well defined, elevated, 
ornamented with granules; delimited by large 
and smooth grooves. Frontal margin bilobed, 
slightly advanced, edge granulated. Orbits oval, 
oblique, separated from frontal margin by deep 
fold; supraorbital margin with three teeth sepa-
rated by two notches, inner orbital tooth subtri-
angular, prominent. Anterolateral margins with 
four rounded and granulated teeth (outer orbital 
spine excluded); first one smallest, at lower lev-
el. Posterolateral margins slightly convex, orna-
mented with granules. Posterior margin slightly 
convex, medially depressed, rimmed. Cervical 
and hepato-gastric grooves well marked, broad, 
smooth. Gastric process well marked; epigastric 
lobes swollen; protogastric lobes, swollen, oval, 
U-shaped, anterior portion medially depressed; 
mesogastric lobe broad posteriorly; anterior por-
tion slender, low, long, reaching basal portion 
of epigastric lobes; metagastric lobe indistinct; 
urogastric region swollen, well delimited from 
meso-metagastric lobe by narrow groove with 
gastric pits. Cardiac region swollen, broadly 
T-shaped. Intestinal region transversely elon-
gate, inflated, narrow, subparallel along poste-
rior margin, medially divided by small smooth 
depression. Hepatic region inflated. Branchial 
regions well defined by swollen lobes, separated 
by broad, shallow, smooth grooves; epibranchi-
al lobe subdivided into two: supra-epibranchial 
lobe transversely elongate, from horizontal to 
oblique, directed to fourth anterolateral tooth; 
sub-epibranchial lobe from rounded to triangu-
lar; both delimited by shallow smooth groove; 
mesobranchial lobe inflated. Male thoracic ster-
num flattened, covered by coarse granules; ster-
nopleonal cavity narrow, deep; sternite 3 with a 
shallow longitudinal median groove connecting 
with sternopleonal cavity, reaching end of ster-
nite 4; sternites 1 and 2 fused, subtriangular; 
sternite 3 subtriangular; sternite 4 subtrapezoi-
dal, with prominent lateral edges, with marked 
grooves paralleling edges; sternites 5, 6 and 7 
subtrapezoidal, elongate; sternite 7 shorter than 
sternite 6; suture 1/2 absent; suture 2/3 complete; 
suture 3/4 distinct, defined by groove, suture vis-
ible only laterally; sutures 4/5, 5/6 medially in-
terrupted; episternites not laterally extended, 
episternite 7 strongly produced, spur shaped, 
reaching coxa of P5. Male pleon narrow, with free 
somites, axially vaulted; somites 1, 2 not folded 
ventrally, visible dorsally; somites 1 to 5 subrec-
tangular, transversely narrow, somite 6 almost 
as long as broad; somite 3 largest, reaching coxa 
of P5; somites 4, 5, 6, with slightly concave upper 
and lateral margins covered by uniformly dis-
tributed granules; telson subtriangular, rounded 
tip. Ischium of third maxilliped subrectangular 
with median sulcus, inner margin convex, outer 
margin concave, covered by scarce granules; ex-
ognath slender; merus subquadrate. Ambulatory 
legs keeled, spiny.
Remarks: At the time, the dorso-ventral sim-
ilarities highlighted by Ossó et al. (2014, pp. 36-
38) were considered sufficient to place the Late 
Eocene Pyreneplax within the extant family 
Vultocinidae (Fig. 1); the additional sternal and 
pleonal features observed in the new specimen 
(Fig. 2) confirm this course of action (Domínguez 
& Ossó, 2019, pp. 70-72). Indeed, in the new spec-
imen, a male, the sternopleonal cavity is deep 
and relatively narrow, almost reaching the end of 
sternite 4, as in Vultocinus (Ng & Manuel-San-
tos, 2007, p. 43, figs. 12A, 9A). The position of the 
press-button in Vultocinus, considered impor-
tant by Ng & Manuel-Santos (2007, p. 42), cannot 
be observed in the new specimen due to preserva-
tion; however, in view of the position of the pleon, 
127On the systematic placement of Pyreneplax Ossó, Domínguez & Artal, 2014 (Decapoda, Brachyura, Vultocinidae)
Fig. 1.: Pyreneplax basaensis Ossó, Domínguez & Artal, 2014 from the Priabonian (Upper Eocene) of the central Pyrenees of 
Huesca (Aragón, Spain). A: holotype MPZ 2013/80, dorsal view; B: ventral view of holotype. C: paratype MPZ 2013/82, dorsal 
view. D: paratype MPZ 2013/83, ventral view. Scale bars equal 10 mm.
slightly shifted down, it is believed that it is on 
the posterior part of sternite 5, close to sternite 6, 
as in Vultocinus (Figs. 2B, C).
It should be noted that the male pleonal somites 
1 and 2 of the new specimen do not appear to be 
folded ventrally and are therefore located in dor-
sal position, as in Vultocinus anfractus (Figs. 2A, 
D; compare Ng & Manuel-Santos, 2007, figs. 1B-
C, 2). Likewise, it presents a pleonal pattern sim-
ilar to that of Vultocinus anfractus, namely free 
pleonal somites, axially vaulted, pleonal somites 
4 to 6 showing slightly concave upper and lateral 
margins and pleonal somite 6 almost as long as 
broad (Figs. 2C, D; Ng & Manuel-Santos, 2007, 
figs. 5B, 9A, 10A, 11A; Ng & Richer de Forges, 
2009, fig. 1B).  
128 Àlex OSSÓ & José Luis DOMÍNGUEZ
Fig. 2.
129On the systematic placement of Pyreneplax Ossó, Domínguez & Artal, 2014 (Decapoda, Brachyura, Vultocinidae)
In addition, the posterior edge of episternite 
7 in Pyreneplax basaensis is, similar to Vultoci-
nus anfractus, “strongly produced posteriorly to 
form a spur which reaches coxa of P5” (Ng & Ma-
nuel-Santos, 2007, p. 44). This character, consid-
ered “unusual” by Ng & Manuel-Santos (2007), 
which together the lateral expansion of pleonal 
somite 3 covers the penis completely, differen-
tiates the Vultocinidae from other goneplacoid 
families (Ng & Manuel-Santos, 2007, pp. 44, 49, 
figs. 13A-B; Castro et al., 2010). It is also seen in 
Pyreneplax (Fig. 2E). 
Discussion: Penis protection is present, in a 
range of expressions, among the Heterotremata 
(Guinot et al., 2013, pp. 84-90). In most gonepla-
coid families, to which the vultocinids were ini-
tially compared, it is usually present as an exten-
sion of sternite 7. For example, Davie et al. (2015) 
noted this condition in the families Acidopsidae 
Števčić, 2005, Chasmocarcinidae Serène, 1964, 
Conleyidae Števčić, 2005, Euryplacidae Stimp-
son, 1871, Goneplacidae MacLeay, 1838, Lito-
cheiridae Števčić, 2005, Progeryonidae Števčić, 
2005 (just touching coxa P5) and Scalopidiidae 
Števčić, 2005. This sternal protection is not pres-
ent in the Mathildellidae Karasawa & Kato, 2003, 
Progeryonidae Števčić, 2005 and Sotoplacidae 
Castro, Guinot & Ng, 2010. However, none of the 
families that have penis protection possess the 
spur-shaped prolongation that is seen in Vulto-
cinus anfractus and Pyreneplax basaensis. This 
unique character, shared by both genera, in addi-
tion to the set of the above-mentioned characters, 
shows their clear family relationship.
In this respect, the Eocene Martinocarcinus 
ickeae Böhm, 1922 (family Martinocarcinidae 
Schweitzer, Feldmann & Bonadio, 2009 within 
the Goneplacoidea) also deserves attention. In-
deed, Schweitzer et al. (2009, p. 4) already point-
ed out the striking similarities in sternal and 
pleonal features of Martinocarcinus and Vultoci-
nus (Schweitzer et al., 2009, pl. 1, fig. 4; pl. 2, figs. 
1-5), but in view of the substantial differences in 
dorsal carapace and chelae, they did not conclude 
that there was a close relationship between them. 
Ossó et al. (2014, p. 38) also noted the sternal and 
pleonal similarities between Martinocarcinus 
and Pyreneplax but argued against a family rela-
tionship, based on differences of dorsal carapace 
morphology. However, in view of the new pieces 
of evidence provided by the new specimen of Pyr-
eneplax, regarding penis protection, a re-exami-
nation of the holotype of Martinocarcinus ickeae 
would appear to be interesting, in particular to 
see whether or not it has the same pattern of pe-
nis protection as in Vultocinus anfractus and P. 
basaensis; this cannot be seen in the illustrations 
of Schweitzer et al. (2009). This character may 
well connect these three taxa phylogenetically. 
Another Eocene taxon, Agostella terrersensis 
Ossó-Morales, 2011 (Goneplacoidea, incertae se-
dis), reveals an extension of sternite 7, as a plate 
that reaches the coxa of P5, with the lateral mar-
gin of pleonal somite 3 completely covering the 
penis (Ossó, 2011, fig. 4.3). This pattern of penis 
protection is similar to that seen in some gone-
placoid families (Ng & Manuel-Santos, 2007, fig. 
10), which supports its original placement within 
this superfamily and consequently rejects inclu-
sion in the Tumidocarcinidae Schweitzer, 2005 
(see Schweitzer et al., 2018, pp. 10-12, fig. 8/1a, b). 
The nomenclature of Pyreneplax saundersi 
(Blow & Manning, 1997) (formerly Eohalimede 
saundersi) must be retained as it was original-
ly spelled by Blow & Manning (1997), instead of 
the correct sandersi (see Blow & Manning, 1998). 
This change of spelling is not allowed under the 
current Code (ICZN, 1999), in accordance with 
articles 32.5.1 and 32.5.1.1 (Ng, pers. comm., No-
vember/2019).
Conclusions
Molecular studies carried out recently, using 
several mitochondrial and nuclear genes, have 
demonstrated that Vultocinus sits in its own deep 
lineage within the Heterotremata and is not re-
lated to any of the known goneplacoids, includ-
ing those without living relatives (Ng & Tsang, 
pers. comm., October/2019). These results are 
consistent with what had already been stated in 
the original paper by Ng & Manuel-Santos (2007, 
p. 40) that, “Vultocinus, new genus, possesses a 
suite of unusual characters that make its precise 
affinities difficult to ascertain”.
The fossil evidence suggests that we are deal-
ing with a case of an extinct family with an ex-
Fig. 2. Pyreneplax basaensis Ossó, Domínguez & Artal, 2014, MPZ 2019/265, from the Priabonian of the central Pyrenees 
of Huesca (Aragón, Spain). A: dorsal view; B: ventral view; C: closeup view of pleon; D: closeup view of posterior margin of 
carapace; E: closeup view of the spur-shaped prolongation of sternite 7 (episternite). Abbreviations: a - pleonal somites; cxP5 
- coxa of fifth pereiopod; es - episternite; P - pereiopods; st - thoracic sternites; t - telson. Scale bar equals 10 mm.
130 Àlex OSSÓ & José Luis DOMÍNGUEZ
tant representative rather than an extant family 
with fossil representatives. The dorsal morpholo-
gy of Pyreneplax is relatively common in numer-
ous Eocene genera, but only preserved sternal 
and pleonal features can establish the relation-
ship among these (Ossó et al., 2014, p. 41; Jagt et 
al., 2015, p. 883). The persistence of this dorsal 
carapace pattern is interpreted either as evolu-
tionary success or an example of convergence. In 
the case of the Vultocinidae and in view of the 
preserved ventral features now observed in Pyr-
eneplax basaensis, this indicates that the unu-
sual penis protection structures are a successful 
adaptation and hence persisted over time. Future 
works and new discoveries will be expected to 
shed light on the suprafamily relationships of 
this family and their possible allies. 
Acknowledgements
We thank Peter K.L. Ng (National University of 
Singapore) and Ling M. Tsang (Chinese University of 
Hong Kong) for providing crucial information and use-
ful comments on molecular studies and the first-na-
med also for advice on rules of nomenclature (P.K.L. 
Ng) and we are grateful to Danièle Guinot (MNHN, 
Paris, France) and Peter K.L. Ng (NUS, Singapore), 
whose accurate reviews have greatly improved the 
present note. 
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