Metacryphaeus
is a calmoniid trilobite genus from the Devonian Malvinokaffric Realm,
exclusive to the Gondwanan regions. It includes eleven species, which
are for the first time included here in a single phylogenetic analysis.
The resulting hypotheses establish relations among the Metacryphaeus species with few ambiguities, also suggesting the inclusion of both Plesiomalvinella pujravii and P. boulei
within the genus, as originally considered. The results of
palaeobiogeographic analyses employing the
Dispersal-Extinction-Cladogenesis (DEC) model reinforce the hypothesis
that Bolivia and Peru form the ancestral home of Metacryphaeus.
The radiation of the genus to other Gondwanan areas took place during
transgressive eustatic episodes during the Lochkovian–Pragian. The
Lochkovian dispersal occurred from Bolivia and Peru to Brazil (Paraná
and Parnaíba basins) and the Falklands, and Pragian dispersal occurred
towards South Africa. Dispersal events from Bolivia and Peru to the
Parnaíba Basin (Brazil) were identified during the Lochkovian–Pragian,
suggesting the presence of marine connections between those areas
earlier than previously thought.
Introduction
The Malvinokaffric Realm includes a plethora of trilobites, including the Calmoniidae, which is composed of several genera (e.g., Calmonia Clarke, 1913, Typhloniscus Salter, 1856, Plesioconvexa Lieberman, 1993, Punillaspis Baldis & Longobucco, 1977, Eldredgeia Lieberman, 1993, Clarkeaspis Lieberman, 1993, Malvinocooperella Lieberman, 1993, Wolfartaspis Cooper, 1982, Metacryphaeus Reed, 1907) reported from the Devonian rocks of Brazil, Argentina, Bolivia, Peru, Falkland Islands, and South Africa1,2,3,4,5,6,7,8,9,10. The present work focuses on the genus Metacryphaeus, which only includes Gondwanan species, namely: M. tuberculatus (Kozłowski, 1923), M. kegeli Carvalho et al., 1997, M. meloi Carvalho et al., 1997, M. parana (Kozłowski, 1923), M. giganteus (Ulrich, 1892), M. convexus (Ulrich, 1892), M. curvigena Lieberman, 1993, M. branisai Lieberman, 1993, M. caffer (Salter, 1856), M. australis (Clarke, 1913), and M. allardyceae (Clarke, 1913).
During the 1990s, Lieberman5 presented the first phylogenetic analysis of the group including Metacryphaeus, represented by M. parana, M. convexus, M. curvigena, M. branisai, M. giganteus, and M. tuberculatus, among other calmoniids (Fig. 1a). Later, Carvalho et al.6 (Fig. 1b) conducted a phylogenetic study of that genus, represented by M. parana, M. australis, M. caffer, M. allardyceae, M. tuberculatus, and M. meloi. More recently, Abe & Lieberman9 presented a palaeobiogeographical area-taxon cladogram including all Metacryphaeus species, based on the tree provided by Lieberman5,
with the manual insertion of additional species (i.e., without carrying
out a new phylogenetic analysis). Those phylogenetic studies did not
include all the species of Metacryphaeus. Accordingly, this study
provides a new phylogenetic analysis including all species, in order to
perform a new palaeobiogeographic analysis for the distribution of the
genus.
Figure 1
Previous phylogenetic models including Metacryphaeus: (a) Lieberman5; and (b) Carvalho et al.6.
The genus Metacryphaeus
occurs in many Gondwanan geological units of Devonian age, including
those in Brazil, Bolivia, Falkland Islands, Peru, and South Africa,
spanning the Pragian to the Givetian–Frasnian3,4,5,6,11,12 (Figs 2 and 3).
It has been suggested that the genus originated and diversified in
small basins of the Malvinokaffric Realm in Bolivia and Peru9. The records in this area are from the Pragian to the Givetian, including M. giganteus, M. tuberculatus, M. parana, M. convexus, M. curvigena, and M. branisai.
Figure 2
Palaeobiogeographic distribution of the genus Metacryphaeus during the Early and Middle Devonian. Areas were divided into (A–E) for the palaeobiogeographic analysis. Note: Plesiomalvinella boulei and P. pujravii have been reassigned here to Metacryphaeus.
Chronostratigraphic distribution of Metacryphaeus. Note: Plesiomalvinella boulei and P. pujravii have been reassigned here to Metacryphaeus.
Abbreviations: Loch., Lochkovian; Prag., Pragian; Emsi., Emsian; Eife.,
Efelian; Give., Givetian; Fras., Frasnian; Fame., Famennian.
The parsimony analysis resulted in two MPTs of 132 steps (consistency index = 0.41 and retention index = 0,52; Fig. 4). The only topological difference between these two trees is the placement of Metacryphaeus branisai. The strict consensus is presented in Fig. 5, along with bootstrap probabilities and Bremer decay indices for each node.
Figure 4
(a,b) Two most parsimonious trees (132 long and consistency index of 0.41) calculated in the present phylogenetic analysis.
Plesiomalvinella boulei and P. pujravii were found deeply nested within a clade of Metacryphaeus species. Accordingly, those two species are here referred to that genus, as previously proposed by Wolfart13. Metacryphaeus (including M. boulei and M. pujravii)
is here supported by two synapomorphies: frontal lobe projecting beyond
the cephalic anterior border in dorsal view (character 4) and uniformly
divergent axial furrows from SO to the cephalic margin (character 19).
In contrast to Lieberman5, Clarkeaspis gouldi (Lieberman, 1993) and C. padillaensis (Lieberman, 1993) were grouped into a clade supported by four synapomorphies (Figs 4 and 5):
cephalic anterior border (cranidial) extended and pointed (characters 2
and 3); pentagonal glabella (character 6); 60 to 70% ratio between the
basal glabellar width and the glabellar length (character 9). Clarkeaspis is here placed closer to Metacryphaeus (as its sister group) than in Lieberman5. The Metacryphaeus + Clarkeaspis clade is supported by a single synapomorphy (character 9, 0 → 1) and shows low bootstrap support (Fig. 5). The placement of Malvinocooperella pregiganteus (Lieberman, 1993) and Wolfartaspis cornutus (Wolfart 1968) as successive outgroups of the Metacryphaeus + Clarkeaspis (Figs 4 and 5), also differs from the arrangement seen in Lieberman5.
The present analysis recovered the clades formed by Metacryphaeus giganteus + M. parana (Figs 4 and 5) and M. boulei + M. pujravii (Figs 4 and 5), previously recognized by Lieberman5. Synapomorphies of the M. giganteus + M. parana clade are: 60 to 70% ratio between the basal glabellar width and the glabellar length (character 9), convergently acquired in Clarkeaspis;
slender genal spine (character 36); dorsoventral height of the pygidium
gradually decreasing posteriorly (character 39); 0.65 to 0.80 ratio
between the maximum pygidial axial width and the maximum pygidial axial
length (character 42). The M. boulei + M. pujravii clade
is supported by six synapomorphies which are related to the presence of
two symmetrical rows of sagittal spines on the posterior part of the
glabella (character 15), the presence of one or two spines on L1 and L2
(characters 17 and 18), 0.15 and 0.25 ratio between the distance of
posterior margin of the eyes to the axial furrow and the maximum
glabellar width (character 25), the presence of four or five spines on
the thoracic axial rings (character 37), and the prosopon covered by
spines (character 48).
Our study also recovered new hypotheses for the relationships of Metacryphaeus, including a clade formed by M. allardyceae, M. caffer, M. australis, M. meloi, M. kegeli, and M. tuberculatus.
This is supported by four synapomorphies related to the shape and
extension of the (cranidial) cephalic anterior border (characters 2 and
3), the ratio between the sagittal length of L1 and the glabellar
sagittal length (character 14), and the incision of the occipital furrow
medially (character 29). The clade including M. caffer, M. australis, M. meloi, M. kegeli, and M. tuberculatus is supported by four synapomorphies (Fig. 4a):
glabella posteriorly elevated and declined anteriorly to S3 (character
8); 65 to 75° α angle (character 22); rounded pygidial terminus
(character 45); no spine on the pygidial terminus (character 46). Also,
the clade formed by M. tuberculatus, M. meloi, and M. kegeli
is supported by four synapomorphies related to L2 and L3 that do not
merge distally (character 13), 55 to 64° β angles (character 23), the
connection of S2 and the axial furrow (character 24), and the lack of
connection between the anterior margin of the eyes and the axial furrow
(character 26) (Fig. 5). Two synapomorphies support the M. caffer plus M. australis
clade: characters 9 (reverted to the plesiomorphic condition) and 41,
which are respectively related to a ratio greater than 80% between the
basal glabellar width and the glabellar length, and to 0.25 to 0.35
ratios between the maximum pygidial axial width and the maximum pygidial
width.
The clade that includes all Metacryphaeus except for M. convexus, M. curvigena, and M. branisai (Fig. 4a)
is supported by three synapomorphies related to a 0.15 to 0.25 ratio
between the distance from the posterior margin of the eyes to the axial
furrow and the maximum glabellar width (character 25), occipital furrow
weakly incised medially (character 29), and 130 to 160° γ angle
(character 34). Three synapomorphies support the group formed by M. giganteus, M. parana, M. allardyceae, M. australis, M. caffer, M. meloi, M. kegeli, and M. tuberculatus (Fig. 4a):
0.25 to 0.34 ratio between sagittal length of L1 glabellar lobe and
glabellar sagittal length (character 14), 0.3 to 0.4 ratio between the
maximum exsagittal eyes length and the glabellar sagittal length
(character 27), 0.60 to 0.80 ratio between maximal sagittal pygidial
length and maximal transverse pygidial width (character 40). The
position of Metacryphaeus branisai is variable in the two MPTs (Fig. 4), probably because its pygidium is unknown, implying the non-codification for characters 38 to 47.
In the phylogeny modelled by Lieberman5, Metacryphaeus convexus and M. curvigena are not considered sister taxa to all other Metacryphaeus. Instead, M. curvigena is considered the sister taxon to M. branisai and M. convexus the sister taxon to both (Fig. 1a). In our analysis, the clade formed by M. convexus and M. curvigena
is supported by five synapomorphies: inclination of 10–20° of S3 in
relation to SO (character 12); L2 and L3 not merged distally (character
13); cephalic axial furrows deep and broad (characters 20 and 21);
evident connection between S2 and the axial furrow. Likewise, the
affinities of M. meloi and M. kegeli are supported by four
synapomorphies. This is interesting because these species are endemic
to the Parnaíba Basin (Brazil), as is their sister-taxon M. tuberculatus, the only other species of the genus known to that basin.
Palaeobiogeography
Likelihood Ratio Test supports DEC M2 (w and j set as free parameters) as the best-fit model to our data (Table 1).
The palaeobiogeographic reconstructions differ only slightly for the
two MPTs, so we focus the discussion on the first MPT. The summary of
biogeographic stochastic mapping (BSM) counts (Table 2)
shows a predominance of dispersals among range change events (33.6% of
total events) and, among those, founder events (19.6%) are slightly more
frequent than anagenetic dispersals (14.1%). Vicariance was very
uncommon according to our model, accounting only for 3.9% of the events
(Table 2).
Most dispersals occurred from Bolivia and Peru (A) to other areas, more
frequently to the Paraná (B) and Parnaíba (E) basins (Table 3).
Table 1 Pairwise comparison of the results of the ancestral area reconstructions of nested DEC models on tree 1.
Table
2 Summary of BSM (Biogeographic Stochastic Mapping) counts based on DEC
M2 model showing the mean, standard deviations (SD), and percentage of
different types of biogeographic events.
Table
3 Counts (and standard deviations in parentheses) of dispersal events
averaged across 100 biogeographics stochastic mappings based on the
biogeographic history of Metacrypheus according to DEC M2 model.
All three models estimate a 100% probability for Bolivia and Peru (A) as the ancestral area for the Metacryphaeus clade, as well as for most of its internal clades (Fig. 6; Supplementary Supple 3). The earliest Metacryphaeus records in this area are from the early Pragian4,5, but three range changes were estimated to have occurred earlier, during the late Lochkovian (Fig. 6): 1- the ancestor of M. parana and M. giganteus
expanded its occurrence to encompass the Paraná Basin (B), with the
former species maintaining this broader distribution and the latter
restricted to B (subset sympatry) - in an alternative scenario, the
ancestor of this clade is present only in Bolivia and Peru (A), with M. parana expanding its range to also the Paraná Basin (B); 2- M. allardyceae dispersed to the Falklands area (D); 3- the ancestor of M. australis and M. caffer dispersed to the Paraná Basin (B). During the early Pragian, M. caffer
dispersed from the Paraná Basin to South Africa (C). It is interesting
to note that those dispersal and expansion events likely occurred before
the transgressive events on western Gondwana14,15,16,17 dated between the late Pragian and the early Emsian (Fig. 6). Those areas (A, B, C, D) were eventually connected by transgressive-regressive cycles (Fig. 6), which promoted the faunal similarity observed among the Malvinokaffric fauna of the Early Devonian15,18.
Figure 6
Ancestral
area reconstructions based on DEC M2 model on the tree 1 (top),
sea-level changes curves from Lochkovian to Frasnian (middle) based on
Haq & Schutter55, and Lower Devonian palaeomap of Southern Gondwana (bottom) modified from Torsvik & Cocks56. Arrows on the palaeomap indicate inferred Lochkovian (full arrow) and Pragian (dashed arrow) dispersal routes for Metacryphaeus
taxa. Additional abbreviations: DML, Dronning Maud Land, Antarctica;
EWM, Ellsworth-Whitmore Mountains, Antarctica; MT, Mexican terranes; P,
Precordillera Terrane, Argentina; Pat., Patagonia.
The last common ancestor of Metacryphaeus meloi, M. kegeli, and M. tuberculatus,
and the node including only the latter two taxa were reconstructed with
two almost equal probable ranges, either restricted to Bolivia and Peru
(A) or a joint distribution (Fig. 6)
also including the Parnaíba and Paraná basins (ABE). These different
ancestral range reconstructions imply distinct processes of range
changes, respectively: 1 - successive dispersals from Bolivia and Peru
to the other areas (for an ancestral with distribution restricted to A),
2 - distribution expansions inferred as founder events (for an
ancestral widely distributed in ABE). Nevertheless, in all cases M. meloi and M. kegeli became restricted to the Parnaíba Basin (E), whereas M. tuberculatus
maintained (or reached) a widespread distribution (ABE), even though
its earliest records, dated as late Eifelian and early Givetian, do not
include the Parnaíba Basin4,5,6,11,12. Alternatively, but with lower statistical support, the ancestral range reconstruction hypothesized for the clades M. meloi + (M. tuberculatus + M. kegeli) and M. tuberculatus + M. kegeli
could be AB, encompassing only their older records. This would imply
expansion events towards the Parnaíba Basin (E) after the arrival of
ancestors in the Paraná Basin (B).
The arrival of Metacryphaeus in the Parnaíba Basin may have occurred via two alternative routes (Fig. 6).
A northern route (surrounding the northern margin of the South American
continent) would impose no continental (landmass) barriers, but there
would be climatic barriers related to the warmer waters the animals
would need to overcome, as the Malvinokaffric Realm marks cooler areas.
Also, faunas of this age on the northern margin of South-America belong
to other realms, which lack Metacryphaeus. On the other hand, a route through the Amazon Basin (Fig. 6) would have presented no climatic or faunal barriers (cf.15,18,19).
Even a continental barrier might not have been in place, as there were
transgression events possibly connecting that basin to Bolivia and Peru.
The lack of fossils of this age in the Amazon Basin, which could
confirm such a dispersal route, is related to the depositional gap
present in the upper Lochkovian and lower Emsian of the basin (cf.20,21,22,23,24,25). This absence of Lochkovian–lower Emsian rocks is also observed in the Parnaíba Basin20,21,24, which hinders palaeobiogeographical inferences related to the presence/absence of Metacryphaeus in the Lower Devonian of this basin.
Other trilobite genera also have a broad Gondwanan distribution during the Devonian, e.g. the calmoniid Eldredgeia, with occurrences in the Bolivia, Brazil (Amazon and Parnaíba basins), and South Africa, and the homalonotid Burmeisteria, with records in the Brazil (Amazon, Parnaíba, and Paraná basins), Falkland Islands, South Africa, and Ghana1,15,19,26. Furthermore, the distribution of the brachiopods Tropidoleptus carinatus (Conrad, 1839) and Australocoelia palmata (Mooris & Sharpe, 1846), and the crinoids Exaesiodiscus Moore & Jeffords, 1968, Laudonomphalus Moore & Jeffords, 1968, Monstrocrinus Schmidt, 1941, and Marettocrinus Le Menn15,27,28,29,30,31,32,33,34,
also reinforce that connections between the Bolivian-Peruvian region
and the Amazon, Parnaíba, and Paraná basins were recurrent by the Middle
Devonian (e.g.15,26). However, the dispersal and range expansion events highlighted in our biogeographic analyses (except that related to M. caffer dispersal from the Paraná Basin to South Africa) occurred during the late Lochkovian (Fig. 6). As such, our data suggest an earlier connection between all those Gondwanan regions, allowing Metacryphaeus trilobites to expand into the Paraná and Parnaíba basins via southeastern and northern/northeastern routes, respectively (Fig. 6). Another interesting fact is the diversification of Metacryphaeus in South America occurring earlier than its dispersal to South Africa (where it is represented by M. caffer).
This was temporally the latest dispersal of the genus, taking place
during the Pragian, and a separate event from the dispersal of M. allardyceae in the same direction (to the Falkland Islands), which occurred earlier.
Methods
Phylogenetic analysis
The phylogenetic analysis conducted here was based on the phylogeny of Lieberman5, with extra characters and species added to the data matrix. The added species were Metacryphaeus australis, M. caffer, M. kegeli, M. meloi, and M. allardyceae,
as to encompass all valid species of the genus. Other ingroup taxa were
defined according to the phylogenetic hypothesis of Lieberman5 consisting of Plesiomalvinella boulei, P. pujravii, Wolfartaspis cornutus, Malvinocooperella pregiganteus, Clarkeaspis gouldi, and C. padillaensis. Also, according to Lieberman5, Kozlowskiaspis (K.) superna Braniša & Vaněk, 1973 was used to root of the phylogenetic trees.
Among the 48 characters employed here (see Appendix 1), 33 were taken or modified from Lieberman5
and 15 are new (characters 7, 9, 12, 13, 14, 22, 23, 24, 25, 26, 27,
34, 40, 41, and 42), although based on characters used in phylogenetic
analyses of other trilobite groups (e.g.35,36,37,38,39,40,41,42). The morphological elements of the exoskeleton are shown in Fig. 7 and all morphological relations/angles used in the 15 newly proposed characters were measured as indicated in Fig. 8.
Figure 7
Schematic drawing showing the major exoskeleton elements of the dorsal surface of Metacryphaeus.
Measurements
used: mgwwfl = maximum glabelar width without consider the frontal
lobe; mfll = maximum frontal lobe length; mcl = maximum cephalic length;
dpmeaf = distance of posterior margin of the eyes to the axial furrow;
mele = maximum exsagittal length of the eyes; bgtw = basal glabellar
transverse width; gsl = glabellar sagittal length; gslwfl = glabellar
sagittal length without consider the frontal lobe; L1sl = sagittal
length of L1 glabellar lobe; mtpaw = maximum transverse pygidial axis
width; mtpw = transverse maximum pygidial width; mspl = sagittal maximum
pygidial length; mpal = maximum pygidial axis length; α = angle between
the axial furrow and the furrow of cephalic posterior border; β = angle
between the cephalic posterior border furrow and a line traced from the
posterior margin of the axial furrow to the anterior margin of the
eyes; γ = angle between a straight line traced adjacent to the lateral
genae (from the contact with the cephalic posterior furrow) and a line
traced from the anterior part of the genae (from the contact of the
axial furrow) in direction to the medial-posterior part of the genae;
Ω = S3 inclination in relation to SO.
Among the characters taken from Lieberman5,
some scores were changed for some taxa based on our own
interpretations. This is the case for characters 5 (changed from 0 to 1
in Malvinocooperella pregiganteus, Metacryphaeus giganteus, and Me. branisai), 18 (changed from 0 to 1 in Me. branisai), and 19 (changed from 0 to 1 in Me. giganteus). Other characters from Lieberman5, e.g.
characters 9, 10, 12, 13, 19, 23, 24, and 34, were not used here
because they either have too much variation between individuals of the
same species or can be easily affected by taphonomic deformation. Some
characters from Lieberman5 were split into two or more characters, as in case of characters 2 and 3 (=character 1 of Lieberman5), 20 and 21 (=character 18 of Lieberman5), 30 and 31 (=character 25 of Lieberman5), 35 and 36 (=character 29 of Lieberman5), and 45, 46, and 47 (=character 36 of Lieberman5), following a contingential approach43.
Characters
1 to 36 are related to the cephalon, character 37 to the thorax, 38 to
47 to the pygidium, and 48 to the prosopon (Appendix 1 and 2).
All characters are related to the dorsal surface of the exoskeleton and
were treated as ordered. The data matrix was analyzed in search of the
Most Parsimonious Trees (MPTs) using the software TNT version 1.144. A heuristic search was conducted with 1,000 replicates, random addition of taxa (random seed 0), Tree Bisection and Reconnection
(TBR) as branch swapping algorithm, and “hold” of 10 trees per replica.
The recovered MPTs were summarized in a strict consensus tree. Bremer45 decay indices and bootstrap proportions46
were calculated using scripts incorporated in TNT. The data matrix was
compiled in NEXUS format using the software Mesquite version 3.03 (702)
and the tree images were generated with the software FigTree version
1.4.2.
Palaeobiogeographical analysis
We conducted palaeobiogeographic analyses to explore the distribution dynamic and biogeographical events that affected Metacryphaeus
distribution through time in five areas pre-defined based on the known
occurrences genus: Bolivia and Peru (A); Paraná Basin, Brazil (B); South
Africa (C); Falkland Islands (D); and Parnaíba Basin, Brazil (E).
Bolivia and Peru were treated as a single area due to their geographical
proximity, strong palaeontological association, and co-occurrence of
endemic species2,47.
Only fossil taxa with accurate occurrence data and taxonomic
identification were included. For this reason, taxa with doubtful
assignation (cf., aff.) were not considered in our analyses (e.g.48).
Ancestral area reconstructions were conducted using R (R Development Core Team 2013) package BioGeoBEARS49, which allows comparing the likelihood of our data given distinct models, choosing that with better fit50. We tested three nested models based on the LAGRANGE Dispersal-Extinction-Cladogenesis (DEC) model51,52: M0 contains the default parameters of the DEC models49; M1 has the addition of the free parameter w; and M2 has the addition of the free parameters w and j. The free parameter w is a multiplier of the dispersal matrices and when set to 1 (e.g. in M0) the probabilities of dispersal events are based solely on the dispersal matrices and equal across all events53. The founder-event parameter j
(included only in M2 and set to 0 in M0 and M1) allows range changes to
areas distinct to that of the ancestor during a cladogenetic event49. We employed the Likelihood Ratio Test (LRT) to select the best model.
We
used time-calibrated versions of the two MPTs, dividing them into two
time slices, Silurian to Lower Devonian (430–395 Ma) and Middle to Upper
Devonian (395–382 Ma). Based on that, we conducted a time-stratified
analyses using time-specific dispersal multiplier and area matrices (see
Supplementary supple 2 and supple 3).
This allowed changing the distances and probabilities between the areas
along these periods, simulating the continental transformations.
We also conducted a biogeographic stochastic mapping (BSM) on BioGeoBEARS54
to estimate the number and type of biogeographical events. We conducted
the BSM only for the first MPT, as the ancestral area reconstruction of
both MPTs differ only slightly, and employed the parameters of the
best-fit model of the ancestral area reconstruction53.
The mean and standard deviation of event counts of 100 BSMs were used
to estimate the frequencies of range change between the considered areas
and of each kind of biogeographic event.
Conclusions
This work provides new phylogenetic hypotheses for the relationships of all species within the genus Metacryphaeus, including the identification of the clades composed of (1) M. caffer and M. australis, (2) M. tuberculatus, M. meloi, and M. kegeli, (3) M. tuberculatus and M. kegeli, (4) M. curvigena and M. convexus, the latter two as sites clades. The position of M. branisai varied in the two recovered MPTs, probably due to the unknown pygidium for this specie. As Plesiomalvinella pujravii and P. boulei were positioned within the Metacryphaeus clade, these species were reinserted in that genus, as originally suggested by Wolfart13. Finally, the genus Clarkeaspis represents the immediate outgroup to Metacryphaeus.
The results of the palaeobiogeographic analyses with DEC models reinforce the interpretations of Lieberman5 and Abe & Lieberman9 that Metacryphaeus
originated in the Lower Devonian of Bolivia and Peru, where they are
represented by a higher taxonomic diversity. The radiation of Metacryphaeus
to other Gondwanan regions probably occurred during the transgressive
events in the Lochkovian–Pragian. In the Lochkovian, dispersals would
have occurred to the Paraná Basin, in Brazil (M. parana, M. australis, M. tuberculatus), as well as to the Falklands area (M. allardyceae) and the Parnaíba Basin (M. meloi, M. kegeli, M. tuberculatus). Pragian dispersal events were reconstructed only towards South Africa (M. caffer).
The ancestral area reconstructions for Metacryphaeus show dispersal events occurring earlier than expected, i.e.
during the Early Devonian, even though the faunal similarities of
Bolivia and Peru with the Parnaíba and Amazon basins are more prominent
in the Middle Devonian, with the sharing of brachiopod (Tropidoleptus and Australocoelia), crinoid (Exaesiodiscus, Laudonomphalus, Monstrocrinus, and Marettocrinus), and other trilobite (Eldredgeia and Burmeisteria)
taxa. The results presented here indicate that these areas were also
somehow connected during the beginning of the Devonian, as to allow the
dispersal of Metacryphaeus.
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We
would like to thank Fundação de Amparo à Pesquisa do Estado de São
Paulo [FAPESP 2013/09683-3 (F.A.C.), 2014/03825-3 (M.C.L.), 2016/18275-4
(R.P.G.), 2015/10788-0 (S.S.N.), 2014/25379-5 (G.S.F.), 2016/03934-2
(G.S.F.)] for their financial support. S. S. Nihei thanks the financial
support from CNPq (403165/2016-4, 303615/2015-2). We would like to thank
Nicholas J. Matzke for the help with BioGeoBEARS.
Author information
Affiliations
Departamento
de Ciências Biológicas, Faculdade de Ciências de Bauru, Universidade
Estadual Paulista Júlio de Mesquita Filho – UNESP, Av. Eng. Luiz Edmundo
Carrijo Coube, 14-01, CEP 17033-360, Bauru/SP, Brazil
Fábio Augusto Carbonaro
& Renato Pirani Ghilardi
Laboratório
de Paleontologia, Faculdade de Filosofia, Ciências e Letras de Ribeirão
Preto, Universidade de São Paulo – USP, Av. Bandeirantes, 3900, CEP
14040-901, Ribeirão Preto/SP, Brazil
Max Cardoso Langer
& Gabriel de Souza Ferreira
Departamento
de Zoologia, Instituto de Biociências, Universidade de São Paulo – USP,
Rua do Matão, Travessa 14, 101, Butantã, CEP 05508-090, São Paulo/SP,
Brazil
Silvio Shigueo Nihei
Fachbereich Geowissenschaften der Eberhard Karls Universität Tübingen, Hölderlinstraße 12, 72074, Tübingen, Germany
Gabriel de Souza Ferreira
Contributions
F.A.C.
wrote the main manuscript text, prepared figures, conducted the
phylogeny and paleobiogeography analyses. G.S.F. conducted the
BioGeoBEARS analysis, wrote part of the Methods and Results (related to
Palaeobiogeography), helped in the Figure 6 and in the
palaeobiogeographical interpretation. S.S.N. helped in
palaeobiogeography (methods and interpretation). M.C.L. and R.P.G.
helped in taxonomy, phylogeny, and palaeobiogeography interpretations.
All authors reviewed the manuscript.
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International License, which permits use, sharing, adaptation,
distribution and reproduction in any medium or format, as long as you
give appropriate credit to the original author(s) and the source,
provide a link to the Creative Commons license, and indicate if changes
were made. The images or other third party material in this article are
included in the article’s Creative Commons license, unless indicated
otherwise in a credit line to the material. If material is not included
in the article’s Creative Commons license and your intended use is not
permitted by statutory regulation or exceeds the permitted use, you will
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