An analysis of pterosaurian biogeography: implications for the evolutionary history and fossil record quality of the first flying vertebrates

Figures & data

Figure 1 The pterosaur relationships and stratigraphical/geographical ranges used in the ‘all taxa’ Treefitter data-set. This tree is based on the cladogram presented by Andres et al. (2014) and shows the more basal portion in detail (Eupterodactyloidea has been condensed to a single branch – see Figure 2). The thick branches represent known stratigraphical ranges (based on data in The Paleobiology Database); thin branches represent estimated ghost ranges and connectors used to demarcate phylogenetic relationships. Time-sliced data-sets were derived from this tree by appropriate inclusion/exclusion of taxa. Most stratigraphical stage and taxon abbreviations are listed in the legend of Table 1. Additional abbreviations: CA, Central Asia; CO, Coniacian; EA, East Asia; Eop, Eopterosauria; EU, Europe; Euc, Euctenochasmatia; KI, Kimmeridgian; NA, North America; OX, Oxfordian; RH, Rhaetian; SA (after taxon name), South America; SA (time scale), Santonian; TU, Turonian.

Figure 2 The pterosaurian relationships and stratigraphical/geographical ranges used in the ‘all taxa’ Treefitter data-set. This tree is based on the cladogram presented by Andres et al. (2014) and shows the relationships among Eupterodactyloidea (see Figure 1 for the more basal part of the cladogram). The thick branches represent known stratigraphical ranges (based on data in The Paleobiology Database); thin branches represent estimated ghost ranges and connectors used to demarcate phylogenetic relationships. Time-sliced data-sets were derived from this tree by appropriate inclusion/exclusion of taxa. All abbreviations are listed in the legend of Table 1 and/or Figure 1.

Figure 3 Palaeogeographical map for the Late Triassic (210 Ma) showing the locations of 29 collections of pterosaurian specimens. The map was generated using software available at Fossilworks (Alroy 2013), with collections data downloaded from The Paleobiology Database.

Figure 4 Palaeogeographical map for the Early and Middle Jurassic (170 Ma) showing the locations of 88 collections of pterosaurian specimens. The map was generated using software available at Fossilworks (Alroy 2013), with collections data downloaded from The Paleobiology Database.

Figure 5 Palaeogeographical map for the Late Jurassic (150 Ma) showing the locations of 77 collections of pterosaurian specimens. The map was generated using software available at Fossilworks (Alroy 2013), with collections data downloaded from The Paleobiology Database.

Figure 6 Palaeogeographical map for the Early Cretaceous (130 Ma) showing the locations of 176 collections of pterosaurian specimens. The map was generated using software available at Fossilworks (Alroy 2013), with collections data downloaded from The Paleobiology Database.

Figure 7 Palaeogeographical map for the Late Cretaceous (80 Ma) showing the locations of 182 collections of pterosaurian specimens. The map was generated using software available at Fossilworks (Alroy 2013), with collections data downloaded from The Paleobiology Database.

Table 1 A summary of the geographical and stratigraphic distributions of pterosaur families and other major clades (based on Barrett et al. 2008 modified by Andres, personal data).

Stage	Africa	Antarctica	Australia	Central Asia	East Asia	Europe	India	North America	South America
Car								Eud
Nor						Aus, Eud, Pre, Pte, Pti
Rha						Pti
Het						Dmp, Pti
Sin			Dmp			Dmp		Pti
Pli	Pti						Pti	Pti
Toa						Cmp, Dmp, Rhm,
Aal					Pti			Pti
Baj				Rhm	Rhm
Bat	Pti			Pti	Anu, Pto, Rhm	Pti, Rhm	Pti	Pti
Cal				Rhm	Anu	Pti, Rhm		Pti	Rhm
Oxf				Anu, Pdi, Rhm, Sor	Rhm, Wuk	Pti		Rhm
Kim	Ctn, Pdi, Pti, Ten					Ard, Gal, Ger, Pti, Ptl, Rhm		Ctn, Pdi, Pti, Rhm
Tth					Pti	Anu, Ctn, Gal, Ger, Pdi, Pti, Ptr, Rhm			Pdi
Ber	Ctn, Pti				Ctn, Dsg, Pti	AZH?, Ctn, Lnc, Orc, Pdi, Pti			Ctn, Dsg
Val					Ctn, Orc, Pdi, Pti	Orc
Hau				Orc	Pti	Orc, Pti, PTN			Pdi
Bar				Orc	Anu, Ctn, Hao Orc, Tpj	Isd, Orc, Pdi, Pti, Tpj			Anh
Apt	AZH, PTN			Orc	Anh, Chy, Ctn, Dsg, Isd, Orc, PTN, Tpj,	Pti, PTN		Azh, PTN, Pdi, Pti	Anh, AZH, Ctn, Orc, Pti, Tpj
Alb	Orc		Orc, Pti,	Pdi	PTN	Anh, Azh, Lnc, Orc, Pti		AZH, Azh	Anh, Ctn, Orc, PTN, Pti, Tpj
Cen	Anh, Azh, Orc, Pdi, Tpj			Azh, Orc, Pdi, PTN	Azh, Pdi	Anh, Lnc, Orc, Pti		Pti, Ptn	AZH
Tur				Azh		Pdi		PTN	Pdi
Con						AZH,		Nyc, Ptn
San				Azh, Pti	Azh, Pti	Azh, Tpj		Nyc, Pti, Ptn
Cam	Azh, Pti		Pdi	Azh, PTN	Azh, Pdi	Azh, Pdi, Pti		Azh, Nyc, Pdi, Pti, Ptn	Azh
Maa	Azh	Pdi		Azh		Azh, Pti	Pti	Azh, Pti	Nyc

Notes: Clade names used here are based on those in the reference cladogram in Figures 1 and 2, and therefore often differ in precise definition or content from those used by Barrett et al. (2008). When there is uncertainty in the age of the earliest known member of a taxon, it is shown in the oldest possible stage. Stratigraphic stage abbreviations: Aal, Aalenian; Alb, Albian; Apt, Aptian; Baj, Bajocian; Bar, Barremian; Bat, Bathonian; Ber, Berriasian; Cam, Campanian; Car, Carnian; Can, Cenomanian; Con, Coniacian; Hau, Hauterivian; Het, Hettangian; Kim, Kimmeridgian; Maa, Maastrichtian; Nor, Norian; Ox, Oxfordian; Pli, Pliensbachian; Rha, Rhaetian; San, Santonian; Sin, Sinemurian; Toa, Toarcian; Tth, Tithonian; Tur, Turonian; Val, Valanginian. Taxonomic abbreviations: Anh, Anhangueridae; Anu, Anurognathidae; Ard, Ardeadactyluslongicollum; Aus, Austriadactylus; Azh, Azhdarchidae; AZH, Azhdarchoidea; Chy, Chaoyangopteridae; Cmp, Campylognathoides; Ctn, Ctenochasmatidae; Dmp, Dimorphodon and potentially closely related material; Dsg, Dsungaripteridae; Eud, Eudimorphodon; Gal, Gallodactylidae; Ger, Germanodactylidae; Hao, Haopterus; Isd, Istiodactylidae; Lnc, Lonchodectidae; Nyc, Nyctosauridae; Orc, ‘Ornithocheiridae’ sensulato; Par, Parapsicephalus; Pdi, indeterminate pterodactyloid remains; Pre, Preondactylus; Pte, Peteinosaurus; Pti, indeterminate pterosaur remains (includes trackways); Ptn, Pteranodontidae; PTN, Pteranodontoidea; Pto, Pterorhynchus; Ptr, Pterodactylidae; Rhm, Rhamphorhynchidae; Sor, Sordes; Ten; Tendaguripteridae; Tpj, Tapejaridae sensu lato. Wuk, Wukongopteridae.

Table 2 Summary of results of Treefitter analyses based on SC and MC costs.

Time slice	No. of taxa	No. of areas	SCs	No. of SC optimal trees	SC p-value	MC cost	No. of MC optimal trees	MC p-value
Total data	106	5	52	2	0^*	− 18	8	1.0
Late Triassic–Late Jurassic	42	4	17	1	0.0012^*	− 7	4	1.0
Middle and Late Jurassic	19	4	10	2	0.00056, 0.00057^*	− 6	4	0.98
Late Jurassic	20	3	3	2	0.24	− 3	3	1.0
Late Jurassic–Early Cretaceous	76	4	32	5	0^*	− 12	2	1.0
Early Cretaceous	50	3	18	2	0.0022, 0.0029^*	− 7	1	1.0
Cretaceous	58	4	26	3	0–0.0002^*	− 11	1	1.0

^*Statistically significant p-values.

Table 3 Biogeographical event frequencies.

Time slice	Vicariance	Sympatry	Extinction	Dispersal
Total data	7–10	72–75 (gtr p = 0)	6–12	20–23 (ltr p < 0.029 in one of the four area cladograms)
Late Triassic–Late Jurassic	2	31 (gtr p < 0.0026)	1	8
Middle and Late Jurassic	4–5	18 (gtr p < 0.026–0.028)	2–4	3–4
Late Jurassic	2	13	1	1
Late Jurassic–Early Cretaceous	2–8 (ltr p < 0.0078)	53–55 (gtr p = 0)	0–14 (ltr p < 0.012)	9–16 (ltr p < 0.048)
Early Cretaceous	1–2 (ltr p < 0.036)	37 (gtr p < 0.0018–0.00019)	0–2 (ltr p < 0.031)	8–9
Cretaceous	4–6	38–41 (gtr p = 0–0.0002)	4–14	6–11 (ltr p < 0.03 in one of the three area cladograms)

Notes: All calculations were carried out using SCs and 10,000 pterm randomisations. Only statistically significant p-values are listed. p-Values marked with ‘gtr’ indicate an event type that occurs more often than expected from random data, and those marked with ‘ltr’ indicate event types that occur less often than expected from random data.

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