Oviparous (egglaying) Sharks

After Compagno 1984b, Compagno 1988, Compagno 1990a, Dulvy 1998.

Shark eggcases are very popular when on exhbit in the Kelp Lab of the Monterey Bay Aquarium. I thought that eggcases occured in 4 shark families (but it is 6 with one species each for 2 families) but did not realize that there were over a hundred sharks (~25% of all sharks) which lay eggs. Here I'm presenting a summary of all oviparous sharks.
According to Compagno 1990, 43% of all cartilaginous fishes are oviparous. All sharks in the order Heterodontiformes are oviparous, 60-70 % in the order Orectolobioformes , and ~40-50% in the order Carcharhiniformes . [All rays in the order Rajiformes (=skates) are oviparous. In discussing the advantages of viviparity, Compagno 1990 points out that it is paradoxical that the skates, which are the most speciose order of cartilagious fishes, retained (extended) oviparity.] The gestation time is usually between 6-12 months, but can be much longer for deep sea catsharks which lay their eggs in very cold water, e.g. 27 months for the brown catshark (Van Dykhuizen 1992). Gestation times much less than 5 months are an indication that eggcases are retained in the uteri before being laid. For some sharks, with little information available, it may be difficult to determine if the reproductive mode is oviparous or viviparous (yolksac viviparity (YSV), previously known a ovoviviparity).
Oviparity is only one of 6-8 modes of reproduction in elasmobranchs.

To learn more about this interesting subject I recommend:
Wourms, J. P. 1981. Viviparity: The maternal-fetal relationship in fishes. American Zoologist 21: 473-515.
Compagno, L. J. V. 1990. Alternative lif-history styles of cartilaginous fishes in time an space. Environ. Biol. Fishes 28: 33-75.
Otake, T. 1990. Classification of reproductive modes in sharks with comments on female reproductive tissues and structures. NOAA Tech. Rep. NMFS 90: 111-130. complete reference,
Wourms, J. P. 1993. Maximization of evolutionary trends for placental viviparity in the spadenose shark, Scoliodon laticaudus. Env. Biol. Fish. 38(1-3): 269-294.
Dulvy, N. K. & Reynolds J. D. 1997. Evolutionary transitions among egg-laying, live-bearing and maternal imputs in sharks and rays. Proc. R. Soc. Lond. B 264: 1309-1315.
Dulvy, N.K. 1998. Evolution and ecology of sharks and rays. Unpublished Ph.D. thesis, University of East Anglia, England. Abstract and Appendix I
Hamlett, W. C. 1997. Reproductive modes of elsmobranchs. Shark News 9:1-3 (June 1997).

See also homepage by Peter Bor on egglaying skates and sharks.


Dim,,dimensions of eggcase (length x width/diameter) ; Gest, gestation time (months); TLaH, total length at hatching (cm); XX, sharks and/or eggcases on display at Monterey Bay Aquarium; ?in Dim column indicates that oviparity is not certain, eggcases could be retained in uteri until the young hatch.
No. Scientific name Common name Dim (cm) Gest
(mo)		 TLaH (cm) Max. size (cm)
Order Heterodontiformes (Bullhead sharks), Familiy Heterodontidae (Bullhead or Horn sharks)
These sharks are oviparous, producing eggs in unique, large, spiral-flanged egg cases. These are laid in specfic "nesting" sites in at least two species. Eggs may take over 5 months to hatch, young hatch at a large size (>14 cm).
001 Heterodontus francisci (Girard, 1854) Horn shark XX ~11 x 4.5;
12.5 (Cox, 1963) 		7-9 15-16 122
002 H. galeatus (Günther, 1870) Crested bullhead shark 11 (4.5")
Whitley 1940		 >5 ~17 152
003 H. japanonicus (Maclay & Macleay, 1884) Japanese bullhead shark   ~12 ~18 120
004 H. mexicanus Taylor & Castro-Aguiree 1972 Mexicn hornshark       70
005 H. portusjacksoni (Meyer, 1793) Port Jackson shark 13-17 x 5-7 9-12 23-24 165
006 H. quoyi (Freminville, 1840) Galapagos bullhead shark     17? 59
007 H. ramalheira (Smith, 1949) Whitespotted bullhead shark       83
008 H. zebra (Gray, 1831) Zebra bullhead shark       122
Order Orectolobiformes (Carpet sharks), Familiy Parascylliidae (Collared carpetsharks)
At least some of the species are ovoparous, depositing eggs in elongated, flattened egg cases on the bottom. One genus (Parascyllium)  is confined to Australian waters, while the second (Cirrhoscyllium) occurs in the China Sea north to Japan and Taiwan Island.
009 Cirrhoscyllium expolitum Smith & Radcliffe, 1913 Barbelthroat carpetshark ?     33.5+
010 C. formosanum Teng, 1959 Taiwan saddled carpetshark ?     39
011 C. japonicum Kamohara, 1943 Saddle carpetshark ?     49
012 Parascyllium collare Ramsay & Ogilby, 1888 Collared carpetshark       87
013 P. ferrugineum McCulloch, 1911 Rusty carpetshark     ~17 75
014 P. multimaculatum Scott, 1911 Tasmanian carpetshark ?     75
015 P. variolatum (Duméril, 1853) Necklace carpetshark       91
Order Orectolobiformes (Carpet sharks), Familiy Brachaeluridae (Blind sharks)
  Brachaelurus waddi (Bloch & Schneider, 1801) Blind shark YSV   15-18 122
016 Heteroscyllium colcloughi (Ogilby, 1908) Bluegray carpetshark ?     ~60
Order Orectolobiformes (Carpet sharks), Familiy Hemiscylliidae (Bamboo sharks)
At least some of these species are oviparous, depositing eggs on the bottom in oval eggcases. One genus (Chiloscyllium)  is wide-ranging, the second (Hemiscyllium)  is confined to Australia and New Guinea.
No. Scientific name Common name Dim (cm) Gest
(mo)		 TLaH (cm) Max. size (cm)
017 Chiloscyllium arabicum Gubanov, 1980 Arabian carpetshark 6.0-7.0 x 3.0-3.5
Peter Bor 		    70
018 C. caerulopunctatum Pellegrin, 1914 Bluespotted bambooshark ?     67
019 C. griseum Müller & Henle, 1838 Grey bambooshark 7.0-7.8 x 3.2-3.6
Ray 1914		     74
020 C. indicum (Gmelin, 1789) Slender bambooshark       ~65
021 C. plagiosum Bennett, 1830 Whitespotted bambooshark 8-10 cm long
x 5.0-5.5 (P. Bor)		 2-2.5   95
022 C. punctatum Müller & Henle, 1838 Brownbanded bambooshark 8.6-9.6 x 4.4-5.0
P. Bor		     104
023 Hemiscyllium freycineti (Quoy & Gaimard, 1824) Indonesian speckled carpetshark ?     46
024 H. hallstromi Whitley, 1967 Papuan epaulette shark ?     75
025 H. ocellatum (Bonnaterre, 1788) Epaulette shark 8.2-11.0 x 3.5-4.6 (W&C 1991) 4.1-4.7   107
026 H. strahani Whitley, 1967 Hooded carpetshark ?     75
027 H. trispeculare& Richardson, 1843 Speckled carpetshark ?     64
Order Orectolobiformes (Carpet sharks), Familiy Stegostomatidae (Zebra sharks)
028 Stegostoma fasciatum (Hermann, 1783) Zebra shark 17 x 8 x 5     354
Order Orectolobiformes (Carpet sharks), Familiy Ginglymostamatidae (Nurse sharks)
028a Ginglymostoma brevicaudatum   Guenther, 1866 Shorttail nurse shark oviparous     75
028b Ginglymostoma cirratum   (Bonnaterre, 1788) Nurse shark ovoviviparous*     2.80-3.04 m
*Reported to have laid eggs in the Curacao Sea Aquarium (Dutch Antilles). After laying 25 empty eggs the animal produced normal live young (21 born between 6 Nov 98 and 18 Jan 99). Gerard Visser, 1999. Nieuwe verpleegsterhaaien voor het Oceanium. Blijdorpblad 1999 no. 6:10. Specimen at Rotterdam Zoo was observed to lay one egg-case (9.5 x 4.9 cm), perhaps aborted. Location of capture is being tracked (Peter Bor).
028c Nebrius ferrugineus   (Lesson, 1830) Tawny nurse shark 8.9 x5.0 (Whitley 1938, 1940) *      3.14-3.20 m
* Reported as ovoviviparous or oviparous (Compagno 1984, p. 203).
Order Orectolobiformes (Carpet sharks), Familiy Rhiniodontidae (Whale sharks)
  Rhincodon typus Smith, 1828 Whale shark 30 x 14 x 9 (YSV)   58-64
(~1 kg) 		13.7-18? m
Until recently, the reproductive mode of the whale shark was uncertain, oviparous or ovoviparous.
Acc. to Compagno 1984 : "In 1953 a large eggcase, 30 cm long, 14 cm wide and 9 cm thick containing a nearly full-term 36 cm embryo whale shark was collected from the Gulf of Mecico, and the assumption was made that the species is oviparous. (Baughman 1955, Garrick 1964, Bass et al. 1975). However, the rarity of 'free-living' whale shark eggs, the extreme thinness and lack of tendrils on the only known case, the considerable yolk and partially developed gill sieve in the only known embryo, and the presence of umbilcal scars on larger free living specimens 55 cm long suggest the whale shark is ovoviviparous."
A recently harpooned pregnant female contained about 300 embryos in the uteri. About 15 of the pups were alive and put into containers with sea water. Only 2 could be kept alive , one in Japan, the other in Taiwan. C. T. Chen reported at the 1996 AES meetin in New Orleans that the pup in Taiwan subsequently died (Abstract presented by W.-B. Chang at the 4th International Aquarium Congress Tokyo, June 23-27, 1996. The success of keeping a baby whale shark from its fetal stage in Taiwan. L.-S. Fang, M.-Y. Leu and W.-B. Chang.
Order Carcharhinifomres (Ground Sharks), Familiy Scyliorhididae (Catsharks)
Many species show the primitive single oviparity, in which only one fertilized egg enters each oviduct and is deposited on the substrate at a time. The large eggs, encapsulated in tough egg cases with corner tendrils to achor them, have most of their embryonic development outside the mother shark and may take nearly a year to produce a hatchling shark.
029 Apristurus atlanticus (Koefoed, 1932) Atlantic ghost catshark       25+
New A. aphyodes Nakaya & Stehman 1998   ~4.3-5.2 x 1.6 x 1.0
(Iglésias et al. 2002) 		     
030 A. brunneus  (Gibert, 1892) Brown catshark XX 5 x 2.5 27 7.6-8.5 68
031 A. canutus Springer & Heemstra, 1979) Hoary catshark       45.5
032 A. herklotsi (Fowler, 1934) Longfin catshark     . 31+
033 A. indicus (Brauer, 1906) Smallbelly catshark       34+
034 A. investigatoris (Misra, 1962) Broadnose catshark       26+
035 A. japonicus Nakaya, 1975 Japnanese catshark       71
036 A.kampae Taylor, 1972 Longnose catshark       52+
037 A. laurussonii (Saemundson, 1922) Iceland catshark ~6-7 x 2.4-2.7 x 1.0
(Iglésias et al. 2002) 		    68
038 A. longicephalus Nakaya, 1975 Longhead catshark       37.5+
039 A. macrorhynchus (Tanaka, 1909) Flathead catshark 5.4 x 2.3
(Nakaya 1975)		     66
No. Scientific name Common name Dim (cm) Gest
(mo)		 TLaH (cm) Max. size (cm)
040 A. maderensix Cadenat & Maul, 1966 Madeira catshark       68.1+
041 A. manis (Springer, 1979) Ghost catshark       85.2
042 A. microps (Gilchrist, 1922) Smalleye catshark       54++
043 A. nasutus de Buen, 1959 Largenose catshark       59
044 A. parvipinnis Springer & Heemstra, 1979 Smallfin catshark       52.1
045 A. platyrhynchus (Tanaka, 1909) Spatulasnout catshark 7.9 x 1.9
(Nakaya 1975)		     80
046 A. profundorum (Goode & Bean, 1886) Deepwater catshark       51++
047 A. riveri Bigelow & Schroeder, 1944 Broadgill catshark 5.0 x 1.5
Springer 1965		     46
048 A. saldanha (Barnard, 1925) Saldanha catshark       81
049 A. sibogae (Weber, 1913) Pale catshark       21++
050 A. sinensis Chus & Hu, 1981 South China catshark       41.7+
051 A. spongiceps (Gilbert, 1895) Spongehead catshark       50+
052 A. stenseni (Springer, 1979 Panama ghost catshark       23+
053 A. verweyi (Fowler, 1934) Borneo catshark       29.7+
054 Asymbolus analis (Ogilby, 1885) Austarlain spotted catshark       61
055 A. vincenti (Zeitz, 1908) Gulf catshark 5.2 x 2.1 x 1.05
5.7 g (Hale 1935)		     61
056a Atelomycterus fasciatus Compagno & Stevens, 1993 New catshark 6.7 x 2.0 (C&S 1993)     45.1+
056b Atelomycterus macleayi Whitley, 1939 Australian marbled catshark 7.0 x 2.5   ~10 60
057 A. marmoratus (Bennett, 1830) Coral catshark 7.1-7.9 x 2.3
P. Bor		     70
058 Aulohalaelurus labiosus Waite, 1905 Blackspotted catshark       67
No. Scientific name Common name Dim (cm) Gest
(mo)		 TLaH (cm) Max. size (cm)
059 Cephaloscyllium fasciatum Chan, 1966 Reticulated swellshark       42+
060 C. isabellum (Bonnaterre, 1788) Draughtsboard shark       100+
061 C. laticeps (Duméril, 1853) Australian swellshark Springer 1979     97+
062 C. nascione Whitley, 1932 Whitefinned swellshark       ~100
063 C. silasi (Talwar, 1974) Indian swellshark       36
064 C. sufflans (Regan, 1921) Balloon swellshark       106
065 C. ventriosum (Garman, 1880) Swellshark XX 9.0-12.5x2.8-5.5
Cox 1964		 7.5-10 13-15 (13-?g) 100+
066 Cephalurus cephalus (Gilbert, 1892) Lollipop catshark YSV?     28
067 Galeus arae (Nichols, 1927) Roughtail catshark * 1.5-2.0 ED 14 43
*"Mode of development uncertain; possibly oviparous in the Caribbean island subspecies G. a. antillensis, but possibly ovovivparous in the continental  G. a. arae." (Compagno 1984). See Iglésias et al. 2002 for review.
068 G. atlanticus (Vaillant, 1888) Atlantic sawtail catshark ?      
069 G. boardmani (Whitley, 1928) Australian sawtail catshark ?     61
070 G. eastmani (Jordan & Snyder, 1904) Gecko catshark 6.0 x 1.6
(Nakaya 1975)		     50
071 G. melastomus Rafinesque, 1810 Blackmouth catshark 6.0-6.4 x 2.0-3.0; Review P. Bor; Iglésias et al. 2002     90
072 G. murinus (Collett, 1904) Mouse catshark ~5.5 x 1.6 x 1.0
(Iglésias et al. 2002)		     63+
073 G. nipponensi Nakaya, 1975 Broadfin sawtail catshark 8.9 x 2.0
(Nakaya 1975)		     65.6
074 G. piperatus Springer & Wagner, 1966 Peppered catshark       30
075 G. polli Cadenat, 1959 African sawtail catshark YSV   ~12 42
076 G. sauteri (Jordan & Richardson, 1909) Blacktip sawtail catshark 3.5 x 1.5
(Nakaya 1975)		   12 45
077 G. schultzi Springer, 1979 Dwarf sawtail catshark     12 29.7
No. Scientific name Common name Dim (cm) Gest
(mo)		 TLaH (cm) Max. size (cm)
078 Halaelurus alcocki ? Garman, 1913 Arabian catshark       <30?
079 H. boesemani Springer & D'Aubrey, 1972 Speckled catshark ?   >7 48
080 H. buerger (Müller & Henle, 1838) Blackspotted catshark 5.1 x 2.1
(Nakaya 1975)
 short   49
081 H. canescens (Günther, 1878) Dusky catshark 6.6 x 3.2
(Springer 1979)		     70
082 H. dawsoni Springer, 1971 New Zealand catshark ?     34.9
083 H. hispidus (Alcock, 1891) Bristly catshark ?     29
084 H. immaculatus Chu & Meng, 1982 Spotless catshark ?     76
085 H. lineatus Bass, D'Aubrey & Kistnasamy, 1975 Lined catshark   1 (20C)   56
  H. lutarius Springer & D'Aubrey, 1972 Mud catshark YSV   10 39
086 H. natalenis (Regan, 1904) Tiger catshark 4 x 1.5 short   47
087 H. quagga (Alcock, 1899) Quagga catshark     ~8 35
088 Haploblepharus edwardsii (Voigt, 1832) Puffadder shyshark 3.5-5.0 x 1.5-3.0     60
089 H. fuscus (Smith, 1950) Brown shyshark       56
090 H. pictus (Müller &Henle, 1838) Dark shyshark 6 x 3   ~11 73
091 Holohalaelurus punctatus (Gilchrist, 1914) African spotted catshark       34
092 H. regani (Gilchrist, 1922) Izak catshark 3.5 x 1.5     61
093 Parmaturus campechiensis Springer, 1979 Campeche catshark       16++
094 P. macmillani Hardy, 1985 New Zealand filetail        
095 P. melanobranchius Chan, 1966 Blackgill catshark       85
096 P. pilosus& Garman, 1906 Salamander catshark       64
097 P. xaniurus  (Gilbert, 1892) Filetail catshark XX 3-4 x 7-11 >24? 7.5-8.5 55
098 Pentanchus profundicolus Smith & Radcliffe, 1912 Onefin catshark ?     50.8+
No. Scientific name Common name Dim (cm) Gest
(mo)		 TLaH (cm) Max. size (cm)
099 Poroderma africanum (Gmelin, 1789) Striped or pyjama catshark 9.5 x 4.5
S & G 1997		 5.5 14-15;
11 (AD)
7.86 g		 101
100 P. pantherinum (Smith, 1838) Leopard catshark 8.2-8.5 x 2.9
P. Bor		     84
101 Schroederichthys bivius (Smith, 1838) Narrowmouthed catshark 2.1 x 6.6
P. Bor		     70
102 S. chilensis (Guichenot, 1848) Redspotted catshark       62
103 S. maculatus Springer, 1966 Narrowtail catshark 1.4 x 4.4
Springer 1965		     34
104 S. tenuis Springer, 1966 Slender catshark 1.7 x 3.9
Gomes 1995		     70
105 Scyliorhinus besnardi Springer & Sadowsky, 1970 Polkdadot catshark 2.1-2.3 x 5.6-6.3
P. Bor		     47
106 S. boa Goode & Ben, 1896 Boa catshark       54
107 S. canicula (Linnaeus, 1758) Small-spotted catshark (Med. Sea) 4 x 2 8-9 10.0-11.5
(3.0-3.5 g) 		 
  S. canicula (Linnaeus, 1758) Small-spotted catshark (North Sea) 5-7 x 2-3 8-9 9-10 100
108 S. capensis (Smith, 1838) Yellowspotted catshark 8 x 3   <30 122
109 S. cervigoni Maurin & Bonnet, 1970 West African catshark 7-8 x 3     76
110 S. garmani (Fowler, 1934) Brownspotted catshark       36+?
111 S. haeckeli (Ribeiro, 1907) Freckled catshark 6-7 x 2-3
Gomes 1995 		  10-13 35+
112 S. hesperius (Ribeiro, 1907) Whitesaddled catshark       47+
113 S. meadi Springer, 1966 Blotched catshark 4.0 x 1.4
Springer 1965		     49++
114 S. retifer (Garman, 1881) Chain catshark 5.3 x 2.0
Nichols 1931		     47
115 S. stellaris (Linnaeus, 1758) Nursehound or large-spotted catshark 10-13 x 3.5-4.0
Bor (1998) 		9? 16 162
116 S. torazame (Tanaka, 1908) Cloudy catshark 5.5 x 1.9
(Nakaya 1975)		   8+ 48
117 S. torrei Howell-Rivero, 1936 Dwarf catshark       32
Order Carcharhinifomres (Ground Sharks), Familiy Proscylliidae (Finback catsharks)
Most species are ovoviviparous, except for the oviparous Proscyllium haberei
118 Proscyllium haberei Hilgendorf, 1904 Graceful catshark       65

Bor, P. H. F. 1998. Eikapsels van Haaien en Roggen. Wetenschappelijke Mededeling KNNV, nr. 223, 48 p
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Cox, K.W. 1963. Egg-cases of some elasmobranchs and a cyclostome from californian waters. Californian Fish and Game 49: 271-289.
Gomes, U .L .1995. Egg capsules of Schroederichtys tenuis and Scyliorhinus haeckelii (Chondrichthyes, Scyliorhinidae). Copeia 1995: 232-236.
Funicelli, N. A, 1972. Eggcases of cartilaginous fishes of the western north atlantic. MsC thesis Long Island University: 57 pp.
Hale, H. M. 1935. The egg-case of a cat shark, scyliorhinus vincenti (Zietz). Records S. Australian Museum 5: 367-368.
Iglésias, S. P., Du Buit M.-H., and Nakaya K. 2002. Egg capsules of deep-sea catsharks from eastern North Atlantic, with first description of the capsule of Galeus murinus and Apristurus aphyodes (Chondrichthyes: Scyliorhinidae). Cymbium 26, 59-63.
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Nakaya, K. 1975. Taxonomy, comparitive anatomy and phylogeny of Japanese catsharks, Scyliorhiniidae. Mem. Fac. Fish. Hokkaido Univ. 23, 1-94.
Nichols, J. T. 1931. Egg capsule of Scyliorhinus retifer. Copeia 1931: 38-39.
Raj, B. S. 1914. Note on the breeding of Chiloscyllium griseum, Mull. and Henle. Rec. Ind. Mus. Calcutta 10: 318-319.
Springer 1965. A review of western atlantic sharks, Scyliorhinidae with descriptions of a new genus and five new species. Fish. Bull. 65: 581-642.
Springer, S. 1979. A revision of the catsharks, Familie Scyliorhinus. NOAA technical report NMFS circular 422, 152pp.
West, J. G. & Carter S. 1991. Observations on the development and growth of the epaulette shark Hemiscyllium ocellatum (Bonnaterre) in captivity. Journal of Aquariculture and Aquatic Sciences 5 (4): 111-117.
Whitley, G. P. 1940. The fishes of Australia, Part 1, the Sharks, rays, devil-fish, and other primitive fishes of Australia and New Zealand. Royal zoological society of New South Wales, Sydney, Australia, 280pp.

Created January 1998; revised December 2009. Back to previous page
Please send comments or corrections to henry@elasmollet.org