Dulvy, N. K. (1998). Evolution and Ecology of Sharks and Rays. Unpublished Ph.D. thesis, University of East Anglia, Norwich NR47TJ, UK.

These are the data sumarised in Dulvy, N. K. and J. D. Reynolds. 1997. Evolutionary transitions among egg-laying, live-bearing and maternal inputs in sharks and rays. Proceedings of the Royal Society of London. B 264: 1309-1315.

Abstract
Sharks and rays are thought to have a large number of independent origins of live-bearing. We examined evolutionary transitions to live-bearing and maternal input in this subclass by optimizing reproductive characters onto a composite phylogeny. Egg-laying (40% of all species) is the likely ancestral reproductive mode for this clade, and there is evidence that live-bearing has evolved independently 9-10 times and maternal input 4-5 times. Most transitions (12-15) have been toward live-bearing with provisioning limited to yolk. These have occured from egg-laying ancestors or live-bearing taxa that provide maternal input to embryos. Only 2-3 transitions have occured in the other direction, i. e. away from yolk-only live-bearing. Egg-laying has evolved from live-bearing ancestors in skates, Rajidae (25% of all species) and possibly in the zebra shark, Stegostoma fasciata. Thus, although there has been an overall trend toward the evolution of live-bearing in elasmobranchs, the evolution of additional maternal input has been extremely labile.

Appendix 1. Reproductive modes of sharks, rays and their closest relatives. Taxonomy follows Nelson (1994) and Mould (1995). Species numbers per genera follow Compagno (1984a & b). CS stands for the character state used in these analyses (see Table 1 of paper). Extinct taxa are represented by *. The groupings of the carcharhinid complex are as suggested by Compagno (1988). References to data represent only main data sources. Numbers in square brackets indicate the number of species for which data were available. The number of genera & species in each taxon are shown in round brackets.


Name of taxonMode of reproductionCSReferences
AgnathaEgg-laying, oviparity 1(Cox 1963; Breder and Rosen 1966; Dodd and Dodd 1985)
TeleostomiThe ancestral state is probably egg-laying 1(Breder and Rosen 1966; Wourms 1981; Wourms 1994; Wourms and Lombardi 1992)
Placodermi * (7 orders) Unknown, but there is evidence for internal fertilisation?(Breder and Rosen 1966)
Rhinochimaeridae (3 genera, 6 spp.)Egg-laying, single oviparity? 1(Dean 1906)
Chimeridae (2 genera, 21 spp.) [1]Egg-laying, single oviparity? 1(Dean 1906; Cox 1963)
Callorhynchidae (1 genera, 4 spp.) [1]Egg-laying, single oviparity? 1(Dean 1906; Giacomo 1994)
Delphyodontus dacriformes * (1 sp.)Live-bearing; intra-uterine feeding and probably leicithotrophy5(Lund 1980)
Myliobatoidei (6 families, 22 genera, 158 spp.) [6]Live-bearing; leicithotrophic with additional nutrition supplied by histotroph or 'uterine milk'. Trophonemata present 4(Capape 1993; Daiber and Booth 1960; Martin and Cailliet 1988; Snelson, et al. 1988; Wourms 1977; Wourms 1981; Wourms, et al. 1988)
Rajidae (18 genera, 200+ spp.) [24]Egg-laying, single oviparity1(Cox 1963; Fitz and Daiber 1963; Holden 1971; Ishiyama 1958; Richards, et al. 1963)
Rhinobatoidea (2 families, 9 genera, 51 spp.) [2] Live-bearing; leicithotrophic with additional nutrition supplied by histotroph or 'uterine milk'. Posesses invaginated uterus walls covered with glandular (possible secretive) cells in R. hynnicephalus. Uterine villi present in R. rhinobatos 4(Bigelow and Schroeder 1953; Abdel-Aziz, et al. 1994; Compagno 1984a; Wenbin and Shuyuan 1993)
Torpedinoidei (2 families, 11 genera, 38 spp.) [3]Live-bearing; leicithotrophic with additional nutrition supplied by histotrophe or 'uterine milk'. Uterine villi present 4(Bigelow and Schroeder 1953; Abdel-Aziz 1994; Wourms 1981; Wourms, et al. 1988) Pristiodei
Pristiodei (1 family, 1 genus, 6 spp.) [1] Live-bearing; leicithotrophic with additional nutrition supplied by histotroph or 'uterine milk'. Trophonemata present 4(Compagno 1984a; Cook, et al. 1995)
Pristiophoriformes (2 families, 2 genera, 5spp.) [4]Yolk-only live-bearing, leicithotrophy or ovoviviparity 3(Compagno 1984a)
Squatiniformes (1 genus, 12 spp.) [7]Yolk-only live-bearing, leicithotrophy or ovoviviparity 3(Compagno 1984a; Otake 1990)
Squaliformes (4 families, 23 genera, 74 spp.) [41] Yolk-only live-bearing, leicithotrophy or ovoviviparity3(Breder and Rosen 1966; Wourms 1977; Wourms 1981; Wourms, et al. 1988; Compagno 1984a; Dodd and Dodd 1985; Natanson and Cailliet 1986; Silva 1989; Otake 1990; Vooren 1992)
Hexanchiformes (4 genera, 5 spp.) [5]Yolk-only live-bearing, leicithotrophy or ovoviviparity 3(Compagno 1984a)
Heterodontiformes (1 genus, 8 spp.) [8] Egg-laying, single oviparity 1(Cox 1963; Compagno 1984a)
Orectolobiformes (7 families, 14 genera, 31 spp.) [28]This order exhibits a variety of transistional reproductive forms from oviparity to leicithotrophic live-bearing..
Hemiscyllium (5 spp.) [5]Egg-laying, but not confirmed in all spp. 1(Compagno 1984a)
Pseudoginglymostoma brevicaudatumLive-bearing; multiple oviparity but egg cases retained, hatching long before partuition2(Compagno 1984a)
Ginglymostoma cirratumLive-bearing; multiple oviparity but egg cases retained, hatching long before partuition2 (Compagno 1984a; Wourms 1977; Wourms, et al. 1988)
Nebrius ferrugineusLive-bearing; multiple oviparity but egg cases retained, hatching just before partuition2(Breder and Rosen 1966; Compagno 1984a; Wourms, et al. 1988)
Stegastoma fasciataEgg-laying 1(Breder and Rosen 1966; Compagno 1984a)
Rhincodon typusLive-bearing?; delayed multiple oviparity. Previously thought to be oviparous but the jury is still out2(Baughman 1955; Breder and Rosen 1966; Compagno 1984a; Wourms, et al. 1988; Young, et al. 1996)
Chiloscyllium (6 spp.) [8]Egg-laying, but not confirmed in all spp. 1(Smedley 1931; Compagno 1984a)
Parascyllium (4 spp.) [3]Egg-laying 1(Compagno 1984a)
Cirroscyllium ( spp.) [3]Probably egg-laying 1(Compagno 1984a)
Brachaelurus waddiYolk-only live-bearing; leicithotrophy or ovoviviparity 3(Compagno 1984a)
Sutorectus tentaculatusPresumably yolk-only live-bearing; leicithotrophy or ovoviviparity3[Compagno, 1984a; Dingerkus, 1986]
Eucrossorhinus dasypogonPresumably yolk-only live-bearing; leicithotrophy or ovoviviparity3[Compagno, 1984a; Dingerkus, 1986]
Orectolobus (4 spp.) [4]Yolk-only live-bearing; leicithotrophy or ovoviviparity. 3[Compagno, 1984a; Dingerkus, 1986; Otake 1990]
Lamniformes ( 7 families, 10 genera, 16 spp.) [15]Live-bearing; nutrition to embryo characterised by either oophagy or adelphophagy (intra-uterine cannibalism). ..
Mitsukurina owstoniUnknown but presumably vivparous 5(Compagno 1984a; Wourms, et al. 1988)
Carcharias (=Eugomphodus) (2 spp.) [1]Live-bearing; leicithotrophy then adelphophagy followed by oophagy . Little known about C. tricuspidatus 6(Compagno 1984a; Gilmore 1993; Springer 1948; Wourms, et al. 1988)
Odontaspis (2 spp.) [2]Live-bearing, leicithotrophy followed by oophagy5(Compagno 1984)
Pseudocarcharias kamoharaiLive-bearing, leicithotrophy followed by oophagy5(Compagno 1984; Gilmore 1993)
Megachasma pelagiosLive-bearing, leicithotrophy followed by oophagy5(Castro, et al. 1996; Compagno 1984a)
Alopius (3 spp.) [2]Live-bearing, leicithotrophy followed by oophagy, Gilmore suggests that these spp. may be capable of adelphophagy but there is no direct evidence5(Compagno 1984a; Wourms, et al. 1988; Otake 1990; Gilmore 1993)
Cetorhinus maximusLive-bearing, leicithotrophy followed by oophagy5(Compagno 1984a; Gilmore 1993; Wourms 1977; Wourms, et al. 1988)
Lamna (2 spp.) [2]Live-bearing, leicithotrophy followed by oophagy; adelphophagy unlikely5(Wourms 1977; Wourms, et al. 1988; Compagno 1984a; Otake 1990; Gilmore 1993)
Carcharodon carchariasLive-bearing, leicithotrophy followed by oophagy; adelphophagy unlikely5(Compagno 1984a; Francis 1996; Gilmore 1993; Wourms, et al. 1988)
Isurus (2 spp.) [2]Live-bearing, leicithotrophy followed by oophagy; adelphophagy unlikely5(Compagno 1984a; Gilmore 1993; Wourms, et al. 1988)
Carcharhiniformes (7 families, 47 genera, 208 spp.) [148]...
Apristurus (25 spp.) [10]Single oviparity1(Cox 1963; Breder and Rosen 1966; Compagno 1984b; Compagno 1988; Nakaya 1975; Wourms, et al. 1988)
Cephalurus cephalusLive-bearing; single oviparity but eggs are retained until hatch1(Compagno 1984b; Compagno 1988)
Parmaturus (4 spp.) [1]Single oviparity1(Cox 1963; Compagno 1984b; Compagno 1988)
Galeus (10 spp.) [7] [1]Single oviparity1(Compagno 1984b; Compagno 1988; Nakaya 1975; Wourms 1981)
G. polliYolk-only live-bearing; eggs retained until hatch1(Compagno 1984b; Compagno 1988)
G. araeMultiple oviparity2see refs above
G. melastomusMultiple oviparity2see refs above
Asymbolus (2 spp.) [2]Egg-laying, single oviparity1(Compagno 1984b; Compagno 1988)
Halaelurus 11 spp. in total) [4]..(Compagno 1984b; Compagno 1988; Nakaya 1975; Wourms, et al. 1988)
H. boesemaniMultiple oviparity with egg retention2Compagno 1984b
H. buergeriMultiple oviparity, egg capsules retained until an advanced stage of development is reached whereupon they are laid2Nakaya 1975; Compagno 1984b
Halaelurus subgenus Bythaelurus (6spp.) [1]Single and multiple oviparity1,2See refs above
Haploblepharus (3 spp.) [3]Egg-laying, single oviparity1(Breder and Rosen 1966; Compagno 1984b; Compagno 1988)
Holohalaelurus (2 spp.) [2]Egg-laying, single oviparity1( Compagno 1984b; Compagno 1988)
Schroederichthyinae (4 spp.) [3]Egg-laying, single oviparity1(Compagno 1984b; Compagno 1988; Gomes & de Carvalho 1995)
Scyliorhinus (13 spp.) [11] Egg-laying, single oviparity1(Compagno 1984b; Compagno 1988; Nakaya 1975; Wourms, et al. 1988; Gomes & de Carvalho 1995)
Cephaloscyllium (7 spp.) [5]Egg-laying, single oviparity1(Cox 1963; Nakaya 1975; Compagno 1984b; Compagno 1988; Otake 1990)
Poroderma (3 spp.) [2]Egg-laying, single oviparity1(Compagno 1984b; Compagno 1988)
Atelomycterini (2 genera, 2 spp) [3]Egg-laying, single oviparity1(Breder and Rosen 1966; Compagno 1984; Compagno 1988)
Proscyllium habereriEgg-laying, single oviparity1(Compagno 1984; Compagno 1988; Wourms, et al. 1988)
Ctenacis fehlmanniEgg-laying, possibly retained for some time before laying 1 (2?)(Compagno 1984; Compagno 1988)
Eridacnis (3 spp.) [3]..(Compagno 1984b; Compagno 1988)
E. barbouriYolk-only live-bearing, leicithotrophy or ovoviviparity3See refs above
E. radcliffiYolk-only live-bearing, leicithotrophy or ovoviviparity3See refs above
E. sinuansYolk-only live-bearing, leicithotrophy or ovoviviparity3See refs above
Gollum attenuatusLive-bearing; matrotrophic, oophagy & uterine milk. Feeds on yolk of broken down eggs within egg sac5(Compagno 1984; Compagno 1988; Yano 1993)
Pseudotriakis microdonLive-bearing; matrotrophic, oophagy & uterine milk5 (Compagno 1984b; Compagno 1988; Yano 1992)
Leptocharias smithiiLive-bearing; matrotrophy via globular or spherical placenta7(Compagno 1984b; Compagno 1988; Wourms, et al. 1988)
Triakis (5 spp.) [5]Yolk-only live-bearing, leicithotrophy or ovoviviparity3(Compagno 1984b; Compagno 1988; Otake 1990)
Scylliogaleus queckettiYolk-only live-bearing, leicithotrophy or ovoviviparity3(Compagno 1984b; Compagno 1988)
Mustelus (20 spp.) [20]Yolk-only live-bearing, leicithotrophy or ovoviviparity3(Nakaya 1975; Teshima 1975; Teshima 1981; Wourms 1977; Wourms, et al. 1988; Compagno 1984b; Compagno 1988; Dodd and Dodd 1985; Otake 1990; Vooren 1992)
M. antarticusLive-bearing; matrotrophy, placenta & uterine villi8See refs. above
M. canisLive-bearing; matrotrophy, placental & uterine milk8See refs. above
M. mustelusLive-bearing; matrotrophy, placental nutrition8See refs. above
M. fasciatus?7See refs. above
M. griseusLive-bearing; matrotrophy, highly active secretive intra-uterine epithelium & placenta. Egg membrane incorporated into placenta7(Teshima 1975; Teshima 1981; Otake 1990)
M. henlei?7See refs. above
Golgolia filewoodi Yolk-only live-bearing, leicithotrophy or ovoviviparity3(Compagno 1984b; Compagno 1988; Vooren 1992)
Hemitriakis (2 spp.) [2]Yolk-only live-bearing, leicithotrophy or ovoviviparity3(Compagno 1984b; Compagno 1988)
Iago (2 spp.) [2]Live-bearing; matrotrophic, placenta7 (Compagno 1984b; Compagno 1988)
Furgaleus mackiYolk-only live-bearing, leicithotrophy or ovoviviparity3(Compagno 1984b; Compagno 1988)
Paragaleus (3 spp.) [3]Live-bearing; matrotrophic, placenta & appendiculae8 (Compagno 1984b; Compagno 1988)
Hemigaleus microstomaLive-bearing; matrotrophic, placenta & appendiculae8(Compagno 1984b; Compagno 1988)
Chaenogaleus macrostomaLive-bearing; matrotrophic, placenta7(Compagno 1984b; Compagno 1988)
Hemipristis elongatusLive-bearing; matrotrophic, placenta & appendiculae8(Compagno 1984b; Compagno 1988)
Hypogaleus hyugaensisLive-bearing, placenta7(Compagno 1984b)
Galeorhinus galeusYolk-only live-bearing, leicithotrophy or ovoviviparity3(Compagno 1984b; Vooren 1992)
Galeocerdo cuvierYolk-only live-bearing, placenta possibly lost. This species is thought to be matrotrophic but the mechanism of nutrient transfer is unknown3(Compagno 1984b; Compagno 1988; Wourms, J. P. pers. comm.)
Scoliodon laticaudusLive-bearing; matrotrophic, placenta (trophonematal cup) & appendiculae. Uterine epithelium has high secretory activity8(Compagno 1988; Wourms, et al. 1988; Wourms 1993; Otake 1990)
Eusphyrna blochiiLive-bearing; matrotrophic, placenta & appendiculae8(Compagno 1984b; Stevens and Lyle 1989)
Sphyrnidae (8 spp.) [7]Live-bearing; matrotrophic, placenta & appendiculae8(Schlernitzauer and Gilbert ????; Compagno 1984b; Compagno 1988; Dodd and Dodd 1985; Hamlett 1986; Stevens and Lyle 1989)
Rhiziprionodon (7 spp.) [6]Live-bearing; matrotrophic, placenta & appendiculae8(Compagno 1984b; Compagno 1988; Hamlett 1986; Wourms 1977)
Loxodon macrorhinusLive-bearing; matrotrophic, placenta & appendiculae8(Compagno 1984b; Compagno 1988)
Isogomodon oxyrhynchusLive-bearing; matrotrophic, placenta7(Compagno 1984b; Compagno 1988)
Trianodon obesusLive-bearing; matrotrophic, placenta7(Compagno 1984b; Compagno 1988)
Carcharhinus porosus group (8 spp.) [6]Live-bearing; matrotrophic, placenta7(Compagno 1984b; Compagno 1988)
Carcharhinus acronotus group (6 spp.) [6]Live-bearing; matrotrophic, placenta7(Compagno 1984b; Compagno 1988)
Nasolamia veloxLive-bearing; matrotrophic, placenta7(Compagno 1984b; Compagno 1988)
Carcharhinus obscurus group (6 spp.) [5]Live-bearing; matrotrophic, placenta7(Compagno 1984b; Compagno 1988)
Carcharhinus ambionensis group (2 spp.) [1]Live-bearing; matrotrophic, placenta7(Compagno 1984b; Compagno 1988)
Negaprion (2 spp.) [2]Live-bearing; matrotrophic, placenta7(Compagno 1984b; Compagno 1988)
Lamniopsis temminckiLive-bearing; matrotrophic, placenta7(Compagno 1984b; Compagno 1988)
Glyphis (2 spp.) [1] Live-bearing; matrotrophic, placenta7(Compagno 1984b; Compagno 1988)
Prionace glaucaLive-bearing; matrotrophic, placenta & trophonemata. Intra-uterine epithelium is non-secretory, but may be important for uterine fluid osmoregulation 8(Compagno 1984b; Compagno 1988; Wourms, et al. 1988; Otake 1990)
Carcharhinus brevipinna group (5 spp.) [3]Live-bearing; matrotrophic, placenta7(Compagno 1984b; Compagno 1988)
Carcharhinus perezi group (3 spp.) [2]Live-bearing; matrotrophic, placenta7(Compagno 1984b; Compagno 1988)

References

  • Abdel-Aziz, S. H. 1994. Observations on the biology of the Common Torpedo (Torpedo torpedo, Linnaeus, 1758) and Marbled Electric ray (Torpedo marmorata, Risso 1810) from Egyptian Mediterranean waters. Australian Journal of Marine and Freshwater Research 45: 693-704.

  • Abdel-Aziz, S. H., A. N. Khalil and S. A. Abdel-maguid. 1994. Reproductive cycle of the common Guitarfish, Rhinobatos rhinobatos (Linnaeus 1758), in Alexandria waters, Mediterranean Sea. Australian Journal of Marine and Freshwater Research 44: 507-517.

  • Baughman, J. L. 1955. The oviparity of the Whale shark, Rhineodon typus, with records of this and other fishes in Texas waters. Copeia: 54-56

  • Bigelow, H. B. and W. C. Schroeder 1953. Fishes of the Western North Atlantic. Part 2: Sawfishes, Guitarfishes, Skates and Rays. Yale University, New Haven.

  • Breder, C. M. and D. E. Rosen 1966. Modes of reproduction in fishes. TFH, New Jersey.

  • Capape, C. 1993. New data on the reproductive biology of the thorny stingray, Dasyatis centroura (Pisces : Dasyatidae) from off the Tunisian coasts. Australian Journal of Marine and Freshwater Research 38: 73-80.

  • Compagno, L. J. V. 1984a. Sharks of the world. An annotated & illustrated catalogue of shark species known to date. Part1. Hexanchiformes to Lamniformes. FAO, Rome.

  • Compagno, L. J. V. 1984b. Sharks of the world. An annotated & illustrated catalogue of shark species known to date. Part 2. Carcharhiniformes. FAO, Rome.

  • Compagno, L. J. V. 1988. Sharks of the order Carcharhiniformes. Princeton University Press, New Jersey.

  • Cook, S. F., L. V. J. Compagno and M. Oetinger. 1995. Status of the largetooth sawfish, Pristis perotteti, Muller and Henley, 1841. Shark News 4: 5.

  • Cox, K. W. 1955. Egg-cases of some elasmobranchs and cyclostome from Californian waters. Calif. Fish and Game: 271-289

  • Daiber, F. C. and R. A. Booth. 1960. Notes on the biology of Butterfly rays, Gymnura altavela and Gymnura micrura. Copeia 137-139.

  • Dean, B. 1906. Chimeroid fishes and their development. Publication of the Carnegie Institute, Washington.

  • Dingerkus, G. 1986. Interrelationships of Orectolobiform sharks (Chondrichthyes: Selachii). pp. 227-245 in T. Uyeno, R. Arai, T. Taniuchi and K. Matsuura eds. Indo-Pacific fish biology : Proc. 2nd Int. Conf. on Indo - Pacific fishes. Icthyological Society of Japan, Japan.

  • Dodd, J. M. and M. H. I. Dodd 1985. Evolutionary aspects of reproduction on cyclostomes and cartilaginous fishes. pp. ? in R. E. Foreman, A. Gorbman, J. M. Dodd and R. Olsson eds. Evolutionary biology of primitive fishes. NATO 103

  • Fitz, E. S. and F. C. Daiber. 1963. An introduction to the biology of Raja eglanteria (Bosc) 1802 and Raja erinacea (Mitchill) 1825 as they occur in Delaware bay. Bulletin of the Bingham Oceanographic collection 18: 69-97.

  • Francis, M. P. 1996. Observations of a pregnant white shark from New Zealand. Pp. in A. P. Klimley and D. G. Ainley eds. The biology of the White Shark, Carcharodon carcharias. Academic Press, New York.

  • Giacomo, E. E. D. 1994. Reproductive biology of the cockfish, Callorhynchus callorhynchus (Holocephali: Callorhychidae), in Patagonian waters (Argentina). Fishery Bulletin 92: 531-539.

  • Gilbert, P. W. and Schlernitzauer, D. W. 1966. The placenta and gravid uterus of Carcharhinus falciformis. Copeia: 451-457.

  • Gilmore, R. G. 1993. Reproductive biology of lamnoid sharks. Environmental Biology of Fishes 38: 95-114.

  • Gomes, U. L. and de Carvalho, M. R. 1995. Egg capsules of Schroederichthyes tenuis and Scyliorhinus haekeli (Chondrichthyes: Scyliorhinidae). Copeia 232-235.

  • Holden, M. J. 1971. The rate of egg laying by three species of ray. Journal du Conseil International de l' Exploration de la Mer 33: 335-339.

  • Ishiyama, R. 1958. Observations on the egg-capsules of the skates of the family Rajidae, found in Japan and its adjacent waters. Bulletin of Museum of Comparative Zoology 118: 1-24.

  • Lund, R. 1980. Viviparity and intrauterine feeding in a new Holocephalan fish from the lower carboniferous of Montana. Science 209: 697-699.

  • Martin, L. K. and G. M. Cailliet. 1988. Aspects of reproduction of the Bat ray Myliobatis california, in Central California. Copeia 754-762.
    Mould, B. 1995. The nomenclature and distribution of the recent Elasmobranchii. University of Nottingham (unpublished).

  • Nakaya, N. 1975. Taxonomy, comparative anatomy and phylogeny of Japanese catsharks, Scyliorhinidae. Memoirs of the Faculty of Fisheries, Hokkaido University 23: 1-94.

  • Natanson, L. J. and G. M. Cailliet. 1986. Reproduction and development of the Pacific Angel shark, Squatina californica, off Santa Barbara, California. Copeia 987-994.

  • Otake, T. 1990. Classification of reproductive modes in sharks with comments on female reproductive tissues and structures. Pp. 111-130 in H. L Pratt, S. H. Gruber and T. Taniuchi eds. Elasmobranchs as living resources: Advances in the biology, ecology, systematics, and the status of the fisheries. NOAA Technical Report NMFS 90.

  • Richards, S. W., D. Merriman and L. H. Calhoun. 1963. Studies on the marine resources of southern New England. IX. The Biology of the little skate, Raja erinacea Mitchill. Bulletin of the Bingham Oceanographic collection 18: 5-68.

  • Schlernitzauer, D. A. and Gilbert, P. W. ????. Placentation and associated aspects of gestation in the Bonnethead shark, Sphyrna tiburo. Journal of Morphology 120: 219-232

  • Silva, H. M. d. 1989. Studies made on the biology and stock assesment of elasmobranchs from mainland Portugal and Azores, with reference to not readily available information. ICES ?

  • Smedley, N. 1931. Notes on elasmobranchs eggs. Bulletin of the Raffles Museum 5: 63-66.

  • Snelson, F. F. J., S. E. Williams-Hooper and T. H. Schmid. 1988. Reproduction and ecology of the Atlantic stingray, Dasyatis sabina in Florida coastal Lagoons. Copeia 729-739.

  • Springer, S. 1948. Oviphagous embryos of the sand shark Carcharis taurus. Copeia 153-157.

  • Stevens, J. D. and J. M. Lyle. 1989. Biology of three Hammerhead sharks (Eusphyra blochii, Sphyrna mokarran, S. Lewini) from Nothern Australia. Australian Journal of Marine and Freshwater Research 40: 129-146.

  • Teshima 1975. Studies on sharks: VIII: Placentation in Mustelus griseus. Japanese Journal of Ichthyology 22 (1): 7-12

  • Teshima 1981. Studies on the reproduction of Japanese smooth dogfishes, Mustelus manazo and M. griseus. Journal Shimonoseki Univ. Fish. 29 (2): 113-199.

  • Vooren, C. M. 1992. Reproductive strategies of eight species of viviparous elasmobranchs from southern Brazil. Bulletin de la Socitie zoologique de France 117: 303-312.

  • Wenbin, Z. and Q. Shuyuan. 1993. Reproductive biology of the Guitarfish, Rhinobatos hynnicephalus. Environmental Biology of Fishes 38: 81-93.

  • Wourms, J. P. 1977. Reproduction and development in chondrichthyan fishes. American Zoologist 17: 379-410.

  • Wourms, J. P. 1981. Viviparity: Maternal - fetal relationships in fishes. American Zoologist 21: 473-515.

  • Wourms, J. P. 1993. Maximisation of evolutionary trends for placental viviparity in the spadenose shark, Scoliodon laticaudus. Environmental Biology of Fishes 38: 269-294.

  • Wourms, J. P. 1994. The challenges of piscine viviparity. Israel Journal of Zoology 40: 551-568.

  • Wourms, J. P., B. D. Grove and J. Lombardi 1988. The maternal-embryonic relationship in viviparous fishes. Pp. 1-134 in W. S. Hoar and D. J. Randall eds. Fish Physiology Academic press, London.

  • Wourms, J. P. and J. Lombardi. 1992. Reflections on the evolution of piscine viviparity. American Zoologist 32: 276-293.

  • Yano, K. 1992. Comments on the reproductive mode of the false cat shark Pseudotriakis microdon. Copeia 460-468.

  • Yano, K. 1993. Reproductive biology of the slender smoothhound, Gollum attenuatus, collected from New Zealand waters. Environmental Biology of Fishes. 38: 59-71.

    Created February 1998; revised Februray 1998.

    There are zillions of references missing from this list, apologies to those I have missed. We originally wanted to publish this appendix with the paper but it was too big! Therefore, I would be grateful if you could send comments, corrections and updates to n.k.dulvy@ncl.ac.uk or nick_dulvy@hotmail.com

    Back to previous page