Premaxillary crest variation within the Wukongopteridae (Reptilia, Pterosauria) and comments on cranial structures in pterosaurs

Cranial crests show considerable variation within the Pterosauria, a group of flying reptiles that developed powered flight. This includes the Wukongopteridae, a clade of non-pterodactyloids, where the presence or absence of such head structures, allied with variation in the pelvic canal, have been regarded as evidence for sexual dimorphism. Here we discuss the cranial crest variation within wukongopterids and briefly report on a new specimen (IVPP V 17957). We also show that there is no significant variation in the anatomy of the pelvis of crested and crestless specimens. We further revisit the discussion regarding the function of cranial structures in pterosaurs and argue that they cannot be dismissed a priori as a valuable tool for species recognition.

Key words:
Wukongopteridae; Pterosauria; cranial crest; sexual dimorphism; China

Authorship

XIN CHENG

Key Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, P.O. Box 643, 100044, Beijing, ChinaChinese Academy of SciencesChinaBeijing, ChinaKey Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, P.O. Box 643, 100044, Beijing, China

Laboratory of Systematics and Taphonomy of Fossil Vertebrates, Department of Geology and Paleontology, Museu Nacional/Universidade Federal do Rio de Janeiro, Quinta da Boa Vista, s/n, São Cristóvão, 20940-040 Rio de Janeiro, RJ, BrazilUniversidade Federal do Rio de JaneiroBrazilRio de Janeiro, RJ, BrazilLaboratory of Systematics and Taphonomy of Fossil Vertebrates, Department of Geology and Paleontology, Museu Nacional/Universidade Federal do Rio de Janeiro, Quinta da Boa Vista, s/n, São Cristóvão, 20940-040 Rio de Janeiro, RJ, Brazil

SHUNXING JIANG

XIAOLIN WANG

University of Chinese Academy of Sciences, 100049, Beijing, ChinaUniversity of Chinese Academy of SciencesChinaBeijing, ChinaUniversity of Chinese Academy of Sciences, 100049, Beijing, China

ALEXANDER W.A. KELLNER

Correspondence to: Xiaolin Wang E-mail: wangxiaolin@ivpp.ac.cn

SCIMAGO INSTITUTIONS RANKINGS

University of Chinese Academy of Sciences, 100049, Beijing, ChinaUniversity of Chinese Academy of SciencesChinaBeijing, ChinaUniversity of Chinese Academy of Sciences, 100049, Beijing, China

Figures | Tables

Figures (2)
Tables (3)

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Figure 1
The variation of cranial crest within the Wukongopteridae. Kunpengopterus sinensis (IVPP V 16047), (a) complete skull and (b) close-up of the area in the white frame of a, showing the flat crestless skull roof; holotype of Darwinopterus linglongtaensis (IVPP V 16049), (c) complete skull and (d) close-up of the area in the white frame of c; new specimen (IVPP V 17957), (e) complete skull (inverted), and (f) close-up of the area in the white frame of e; IVPP V 17959 (g) complete skull (inverted), and (h) close-up of the area in the white frame of g, with arrow 1 pointing at the low premaxillary crest and arrow 2 pointing at the crestless middle portion of the premaxilla; Darwinopterus robustodens (holotype, HGM 41HIII-0309A) (i) complete skull based on Lü et al. 2011bLÜ JC, XU L, CHANG HL and ZHANG XL. 2011b. A new darwinopterid pterosaur from the Middle Jurassic of western Liaoning, northeastern China and its ecological implications. Acta Geol Sin-Engl 85: 507-514.), and (j) close-up of the area in the white frame of i. Scale bar: 50mm in a, c, e, g, i; 10mm in b, d, f, h, j.

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Figure 2
Reconstruction of the various cranial crests of the Wukongopteridae. (a) Kunpengopterus sinensis (based on Wang et al. 2010WANG XL, KELLNER AWA, JIANG SX, CHENG X, MENG X and RODRIGUES T. 2010. New long-tailed pterosaurs (Wukongopteridae) from western Liaoning, China. An Acad Bras Cienc 82: 1045-1062.); (b) Darwinopterus linglongtaensis (based on Wang et al. 2010); (c) new specimen (IVPP V 17957); (d) ZMNH M8802 (female, modified Lü et al. 2011aLÜ JC, UNWIN DM, DEEMING DC, JIN XS, LIU YQ and JI Q. 2011a. An egg-adult association, gender, and reproduction in pterosaurs. Science 331: 321-324.); (e) IVPP V 17959 (based on Cheng et al. 2016CHENG X, JIANG SX, WANG XL and KELLNER AWA. 2016. New information on the Wukongopteridae (Pterosauria) revealed by a new specimen from the Jurassic of China. PeerJ 4: e2177.); (f) Darwinopterus robustodens (based on Lü et al. 2011b, (g) Darwinopterus modularis (holotype, based on Lü et al. 2010). Reconstruction of the crests in grey. Scale bar: 50mm.

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TABLE I
Measurements of the length of skull, humerus and ulna of the Wukongopteridae, from the smallest to the largest individual based on wingspan (in mm).

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TABLE II
Measurements (in mm) and ratio of the pelvis and humerus of Darwinopterus modulari s, Darwinopterus linglongtaensis, and one female wukongopterid.

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TABLE III
Measurements of IVPP V 17957 and other published wukongopterid specimens (in mm).

Figure 1 The variation of cranial crest within the Wukongopteridae. Kunpengopterus sinensis (IVPP V 16047), (a) complete skull and (b) close-up of the area in the white frame of a, showing the flat crestless skull roof; holotype of Darwinopterus linglongtaensis (IVPP V 16049), (c) complete skull and (d) close-up of the area in the white frame of c; new specimen (IVPP V 17957), (e) complete skull (inverted), and (f) close-up of the area in the white frame of e; IVPP V 17959 (g) complete skull (inverted), and (h) close-up of the area in the white frame of g, with arrow 1 pointing at the low premaxillary crest and arrow 2 pointing at the crestless middle portion of the premaxilla; Darwinopterus robustodens (holotype, HGM 41HIII-0309A) (i) complete skull based on Lü et al. 2011bLÜ JC, XU L, CHANG HL and ZHANG XL. 2011b. A new darwinopterid pterosaur from the Middle Jurassic of western Liaoning, northeastern China and its ecological implications. Acta Geol Sin-Engl 85: 507-514.), and (j) close-up of the area in the white frame of i. Scale bar: 50mm in a, c, e, g, i; 10mm in b, d, f, h, j.

Figure 2 Reconstruction of the various cranial crests of the Wukongopteridae. (a) Kunpengopterus sinensis (based on Wang et al. 2010WANG XL, KELLNER AWA, JIANG SX, CHENG X, MENG X and RODRIGUES T. 2010. New long-tailed pterosaurs (Wukongopteridae) from western Liaoning, China. An Acad Bras Cienc 82: 1045-1062.); (b) Darwinopterus linglongtaensis (based on Wang et al. 2010); (c) new specimen (IVPP V 17957); (d) ZMNH M8802 (female, modified Lü et al. 2011aLÜ JC, UNWIN DM, DEEMING DC, JIN XS, LIU YQ and JI Q. 2011a. An egg-adult association, gender, and reproduction in pterosaurs. Science 331: 321-324.); (e) IVPP V 17959 (based on Cheng et al. 2016CHENG X, JIANG SX, WANG XL and KELLNER AWA. 2016. New information on the Wukongopteridae (Pterosauria) revealed by a new specimen from the Jurassic of China. PeerJ 4: e2177.); (f) Darwinopterus robustodens (based on Lü et al. 2011b, (g) Darwinopterus modularis (holotype, based on Lü et al. 2010). Reconstruction of the crests in grey. Scale bar: 50mm.

TABLE I Measurements of the length of skull, humerus and ulna of the Wukongopteridae, from the smallest to the largest individual based on wingspan (in mm).

Taxa	sq-pm	hu	ul/ra	maxws	Reference
Darwinopterus linglongtaensis (IVPP V 16049)	119.2	40.4	58.0	708.2	Wang et al. 2010WANG XL, KELLNER AWA, JIANG SX, CHENG X, MENG X and RODRIGUES T. 2010. New long-tailed pterosaurs (Wukongopteridae) from western Liaoning, China. An Acad Bras Cienc 82: 1045-1062. and this paper (Table III)
Wukongopterus lii (IVPP V 15113)	-	~38.7	~62.1	~734.0	Wang et al. 2010WANG XL, KELLNER AWA, JIANG SX, CHENG X, MENG X and RODRIGUES T. 2010. New long-tailed pterosaurs (Wukongopteridae) from western Liaoning, China. An Acad Bras Cienc 82: 1045-1062. and this paper (Table III)
Kunpengopterus sinensis (IVPP V 16047)	106.9	~36.2	59.2	~734.4	Wang et al. 2010WANG XL, KELLNER AWA, JIANG SX, CHENG X, MENG X and RODRIGUES T. 2010. New long-tailed pterosaurs (Wukongopteridae) from western Liaoning, China. An Acad Bras Cienc 82: 1045-1062. and this paper (Table III)
Darwinopterus modularis (referred specimen, YH-2000)	est. 140	44	64	760.8	Lü et al. 2010LÜ JC, UNWIN DM, JIN XS, LIU YQ and JI Q. 2010. Evidence for modular evolution in a long-tailed pterosaur with a pterodactyloid skull. Proc Roy Soc B-Biol Sci 277: 383-389. and this paper (Table III)
IVPP V 17957	est. 130.0	44.9	62.8	778.8	This paper (Table III)
IVPP V 17959	est. 145.0	est. 41.0	67.3	est. 782.2	Cheng et al. 2016CHENG X, JIANG SX, WANG XL and KELLNER AWA. 2016. New information on the Wukongopteridae (Pterosauria) revealed by a new specimen from the Jurassic of China. PeerJ 4: e2177. and this paper (Table III)
ZMNH M8802/IVPP V 18043	150	52	70	est. 879.8	Lü et al. 2011aLÜ JC, UNWIN DM, DEEMING DC, JIN XS, LIU YQ and JI Q. 2011a. An egg-adult association, gender, and reproduction in pterosaurs. Science 331: 321-324.
Darwinopterus robustodens (HGM 41HIII-0309A)	173.2	52.8	85.4	996.6	This paper (Table III)
Darwinopterus modularis (holotype, ZMNH M8782)	185	59	87	est. 1074.9	Lü et al. 2010LÜ JC, UNWIN DM, JIN XS, LIU YQ and JI Q. 2010. Evidence for modular evolution in a long-tailed pterosaur with a pterodactyloid skull. Proc Roy Soc B-Biol Sci 277: 383-389.

-, not preserved; est, estimated value. Abbreviations: hu, humerus; maxws, maximized wingspan; sq-pm, squamosal to the tip of premaxilla; ul, ulna. The maximized wingspan (maxws=cor+hu+ul/ra+car+mcIV+ph1d4+ph2d4+ph3d4+ph4d4, Kellner et al. 2013KELLNER AWA, CAMPOS DA, SAYÃO JM, SARAIVA AAF, RODRIGUES T, OLIVEIRA G, CRUZ LA, COSTA FR, SILVA HP and FERREIRA JS. 2013. The largest flying reptile from Gondwana: a new specimen of Tropeognathus cf. T. mesembrinus Wellnhofer, 1987 (Pterodactyloidea, Anhangueridae) and other large pterosaurs from the Romualdo Formation, Lower Cretaceous, Brazil. An Acad Bras Cienc 85: 113-135.; abbreviations and measurements see Table III) of Kunpengopterus sinensis, Darwinopterus linglongtaensis, Wukongopterus lii, YH-2000, and Darwinopterus robustodens is based on complete forelimb elements plus the coracoid, while that of IVPP V 17959, ZMNH M8802/IVPP V 18043, and the holotype of Darwinopterus modularis is estimated by the ratio between the preserved elements and maximized wingspan compared to complete specimens.

TABLE II Measurements (in mm) and ratio of the pelvis and humerus of Darwinopterus modulari s, Darwinopterus linglongtaensis, and one female wukongopterid.

	hu	sl	pl	pd	pw	pl/pd	pw/pd	pl/pw	sl/pd
supposed male Darwinopterus modularis (holotype, ZMNH M8782)	59.0	est. 22.0	37.5	20.0	19.5	1.86	0.98	1.92	1.10	Lü et al. 2011aLÜ JC, UNWIN DM, DEEMING DC, JIN XS, LIU YQ and JI Q. 2011a. An egg-adult association, gender, and reproduction in pterosaurs. Science 331: 321-324. and this paper
supposed female “Darwinopterus modularis” (ZMNH M8802 or IVPP V 18403)	52.0	est. 19.0	36.0	19.5	20.0	1.85	1.03	1.80	0.97	Lü et al. 2011aLÜ JC, UNWIN DM, DEEMING DC, JIN XS, LIU YQ and JI Q. 2011a. An egg-adult association, gender, and reproduction in pterosaurs. Science 331: 321-324. and this paper
Darwinopterus linglongtaensis (holotype, IVPP V 16049)	40.4	16.3	30.2	20.6	~18.0	1.47	0.87	1.68	0.79	This paper
ZMNH M8782 / ZMNH M8802 or IVPP V 18403	1.13	1.16	1.04	1.03	0.98					Lü et al. 2011aLÜ JC, UNWIN DM, DEEMING DC, JIN XS, LIU YQ and JI Q. 2011a. An egg-adult association, gender, and reproduction in pterosaurs. Science 331: 321-324. and this paper
ZMNH M8782 / IVPP V 16049	1.46	1.35	1.24	0.97	1.08					This paper
ZMNH M8802 or IVPP V 18403 / IVPP V 16049	1.29	1.17	1.19	0.95	1.11					This paper

~, estimate based on the third sacral vertebra; est, estimated value based on pictures. Abbreviations: hu, humerus; pd, pelvis depth from dorsal margin of the ilium to the ventral margin of the puboischiadic plate at level of the acetabulum; pl, pelvis length from anterior tip of preacetabular process of ilium to posterior tip of postacetabular process of ilium; pw, pelvis width between the acetabula; sl, length of sacral vertebrae.

TABLE III Measurements of IVPP V 17957 and other published wukongopterid specimens (in mm).

Bone/Taxa	W. lii IVPP V 15113	D. modularis ZMNH M8782	D. modularis YH-2000	D. linglongtaensis IVPP V 16049	D. robustodens HGM 41HIII-0309A	K. sinensis IVPP V 16047	ZMNH M8802 /IVPP V 18403	IVPP V 17959	IVPP V 17957
sq-pm	-	185	est 140	~119.2	173.2	106.9	150	est 145	est. 130
ros	32.6			~30.3	61.7	~40.5	46	-	40.1
naof	-	82	-	~57.9	77.7	~37.9	*72	est 80	-
lower jaw	103.2	157	115	102.4	151.3	89.5	est 125	est 130	106.8
man.sys	19.5			~23.6	-	-	34	est 30	25.5
sca	34.1(r) 32.9(l)	>25	27	31.1(r)	37.3(r)	~28.8(r)	34	-	31.8(r) 32.3(l)
cor	~25.3(r)	34	26	24.9(r) 26.3(l)	34.6(r)	~23.8(l)		-	28.4(r)
hu	~38.7(r)	59	44	40.4(r) 39.9(l)	52.8(r) 53.3(l)	~36.2(r)	52	est 41(r)	44.9(r) 43.6(l)
ul/ra	~62.1(r)	87	64	58.0(r) 61.2(l)	85.4(r) 82.3(l)	~59.2(l)	70	67.3(r)	62.8(r) 62.4(l)
car	5.7(r)		5.4	4.6(r) 4.5(l)	7.4(r) 7.2(l)	4.8(r)	-	4.3(l)	5.2(l)
mcIV	22.9(l)	36	26	22.5(r) 23.6(l)	33.1(r) 30.7(l)	~23.0(l)		26.5(r) 27.6(l)	26.6(l)
ph1d4	~45.7(r)	75	48	46.2(r) 44.4(l)	63.9(r) 61.1(l)	54.2(r)		~55.3(r)	54.1(r) 53.3(l)
ph2d4	56.8(r)	-	55	50.7(r) 52.1(l)	75.0(r) 74.3(l)	58.0(r)		~59.8(r)	59.5(r) 58.6(l)
ph3d4	58.3(r) 59.3(l)	-	59	53.2(r) 53.7(l)	78.1(r) 75.7(l)	59.2(r)		-	58.5(r) 57.7(l)
ph4d4	51.5(l)	-	53	53.6(r) 53.7(l)	68.0(r)	~48.8(l)		-	49.4(r)
fe	33.3(r) 35.6(l)	-	40	39.4(r) 40.1(l)	43.5(r)	~39.3(r) ~41.1(l)	41	-	42.2(r) 42.7(l)
ti	52.8(r)	-	54	49.8(r) 49.6(l)	61.4(l)	54.5(l)	61	-	-
mt3	~17.2(r) 16.8(l)	23	17	17.0(r)	21.5(r) 20.6(l)	21.8(r) 20.7(l)	20	-	-
mt4	~14.5(r) 15.0(l)			14.2(r) 14.2(l)	18.5(r) 17.6(l)	17.9(r)	16	15.1	-
pph1d5	13.9(r) 13.7(l)			12.8(r) 12.1(l)	18.1(r) 18.0(l)	11.2(r)	18.5	-	-
ptd	>7.3(r)	-	26	28.9(l)	34.0(l)	~25.8(l)	25	24.9(l)	est 31(r)
	Wang et al. 2010WANG XL, KELLNER AWA, JIANG SX, CHENG X, MENG X and RODRIGUES T. 2010. New long-tailed pterosaurs (Wukongopteridae) from western Liaoning, China. An Acad Bras Cienc 82: 1045-1062. and this paper	Lü et al. 2010LÜ JC, UNWIN DM, JIN XS, LIU YQ and JI Q. 2010. Evidence for modular evolution in a long-tailed pterosaur with a pterodactyloid skull. Proc Roy Soc B-Biol Sci 277: 383-389.	Lü et al. 2010LÜ JC, UNWIN DM, JIN XS, LIU YQ and JI Q. 2010. Evidence for modular evolution in a long-tailed pterosaur with a pterodactyloid skull. Proc Roy Soc B-Biol Sci 277: 383-389. and this paper	Wang et al. 2010WANG XL, KELLNER AWA, JIANG SX, CHENG X, MENG X and RODRIGUES T. 2010. New long-tailed pterosaurs (Wukongopteridae) from western Liaoning, China. An Acad Bras Cienc 82: 1045-1062. and this paper	This paper	Wang et al. 2010WANG XL, KELLNER AWA, JIANG SX, CHENG X, MENG X and RODRIGUES T. 2010. New long-tailed pterosaurs (Wukongopteridae) from western Liaoning, China. An Acad Bras Cienc 82: 1045-1062. and this paper	Lü et al. 2011aLÜ JC, UNWIN DM, DEEMING DC, JIN XS, LIU YQ and JI Q. 2011a. An egg-adult association, gender, and reproduction in pterosaurs. Science 331: 321-324. and this paper	Cheng et al. 2016CHENG X, JIANG SX, WANG XL and KELLNER AWA. 2016. New information on the Wukongopteridae (Pterosauria) revealed by a new specimen from the Jurassic of China. PeerJ 4: e2177. and this paper	This paper

-, not preserved; *, corrected in this paper based on pictures; est, estimated value. Abbreviations: car, carpus; cor, coracoid; fe, femur; hu, humerus; l, left; man.sys, mandibular symphysis; mcIV, metacarple IV; mt3-4, metatarsle 3-4; naof, nasoantorbital fenestra; ph1-4d4, first to fourth phalanx of manual digit IV; pph1d5, first phalanx of pedal digit V; ptd, pteroid; r, right; ra, radius; ros, rostrum; sca, scapula; sq-pm, tip of premaxilla to posterior end of squamosal; ti, tibia; ul, ulna.

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Premaxillary crest variation within the Wukongopteridae (Reptilia, Pterosauria) and comments on cranial structures in pterosaurs

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[1] Correspondence to: Xiaolin Wang E-mail: wangxiaolin@ivpp.ac.cn