Supplementary table 1. Intraspecific variations in testes investment in species with alternative reproductive tactics between which body size are different. Data were taken from texts or estimated from figures drawn in the cited papers. See also Taborsky (1994,1999, 2008) for fish examples.

Testes investment / Body size††
Class / Order / Family / Species / Bourgeois Parasitic / Measurements † / Bourgeois Parasitic / References
Insecta / Coleoptera / Nitidulidae / Librodor japonicus / TA / (2.0) / Okada et al. 2008
Scarabaeidae a / Onthophagus binodis / < / TA / (1.5) / Simmons et al. 1999; Tomkins and Simmons 2002
Onthophagus taurus / = * / TA / (1.5) / Simmons et al. 1999; Tomkins and Simmons 2002
Orthoptera / Anostostomatidae / Hemideina crassidens / = * / TA / (1.2) b / Kelly 2008
Dermaptera / Forficulidae / Forficula auricularia / TA / (1.8) / Tomkins and Simmons 2002
Tymenaptera / Apidae / Amegilla dawsoni, / TA / (2.0) / Simmons et al. 2000; Tomkins and Simmons 2002
Malacostraca / Decapoda / Palaemonidae / Macrobrachium rosenbergii / < / GSI / (6.0) / Sagi & Ra’anan 1988; Sagi et al. 1988
Cephalopoda / Tethida / Loliginidae / Loligo bleekeri / < / GSI / (1.7) b / Wada 2005
Osteichthyes / Batrachoidiformes / Batrachoididae / Porichthys notatus / GSI / (8.8) / Bass and Marchaterre 1989
Cyprinodontiformes / Poecilidae / Xiphophorus nigrensis / = / TA / (1.4) b / Smith and Ryan 2010
Poecilia latipinna / GSI / N/A / Schlupp et al. 2006
Poecilia mexicana / GSI / N/A / Schlupp et al. 2006
Poecilia parae / GSI & TA / (1.3) b / Hurtago-Gonzales and Uy 2009
Perciformes / Bleniedae / Axoclinus carminalis / = / GSI / (1.4) b / Neat 2001
Axoclinus nigricaudus / GSI / (1.6) b / Neat 2001
Parablennius parvicornis / GSI / (2.4) / Miranda et al. 2003
Salaria pavo / GSI / (1.4) b / Ruchon et al., 1995
Scartella cristata / c / GSI / (5.5) / Neat et al. 2003
Tripterygion delaisi / = / GSI / (1.3) b / Jonge and Videler 1989; Jonge et al. 1989
Tripterygion malenurus / d / GSI / Geertjes & Videler 2002
Tripterygion tripteronotus / GSI / (1.3) b / Jonge and Videler 1989; Jonge et al. 1989
Centrarchidae / Lepomis gibbosus / GSI / N/A / Gross 1982; Rios-Cardenas and Webster 2008
Lepomis macrochirus / GSI / (15.8) / Gross 1982
Lepomis megalotis / GSI / (2.0) b / Jennings and Philipp 1992
Lepomis punctatus / GSI / (1.9) b / DeWoody et al. 2000
Cichlidae / Julidochromis ornatus / TA / (1.3) b / Awata et al. 2006
Lamprologus calliptreus / GSI / (40.1) / Schütz et al. 2006
Lamprologus lemairii / TA / (8.6) / Ota, Awata, Morita and Kohda in preparation
Neolamprologus brevis / TA / (4.6) / Ota, Aibara, Awata, Morita and Kohda unpubl. data
Neolamprologus pulcher / > ** / GSI / (1.9) b / Balshine et al. 2001; Fitzpatrick et al. 2006
Telmatochromis temporalis / GSI / (15.1) / Kato et al. 2005; Kato unpubl
Telmatochromis vittatus / *** / GSI, TA / (8.3) / Ota and Kohda 2006; Ota et al. 2010; Ota unpubl
Gobidae / Apollonia melanostoma / GSI / (3.6) / Marentette et al. 2009
Gobius niger / GSI / (5.5) b / Rasotto & Mazzoldi 2002; Scaggiante et al. 2006
Lythrypnus dalli / = * / GSI / (1.3) b / Drilling and Grober 2005
Pomatoschistus minutus / GSI, TA / (2.1) / Kvarnemo et al. 2010
Zosterisessor ophiocephalus / GSI / (8.9) b / Scaggiante et al. 1999, 2006; Mazzoldi et al. 2000
Labridae / Halichoeres bivittatus / GSI / (1.5-2.0) b / Warner and Robertson 1978
Halichoeres maculipinna / GSI / (1.5) b / Warner and Robertson 1978
Halichoeres pictus / GSI / (1.5) b / Warner and Robertson 1978
Halichoeres poeyi / GSI / (1.5-2.0) b / Warner and Robertson 1978
Halichoeres trimaculatus / GSI / (1.7) / Kim et al. 2002
Symphodus melops / GSI / (2.0) / Uglem et al. 2000, 2001
Symphodus ocellatus / GSI / (1.5) b / Warner and Lejeune 1985
Symphodus roissali / GSI / (1.3) b / Warner and Lejeune 1985
Thalassoma bifasciatum / GSI / (1.3-1.5) b / Warner and Robertson 1978
Thalassoma duperrey / GSI / (1.3) b / Warner 1982; Hourigan et al. 1991
Salmonidae / Oncorhynchus kisutch / GSI / (1.5) b / Gross 1985; Leach 1997
Oncorhynchus masou / GSI / (57) / Koseki and Maekawa 2000, 2002
Oncorhynchus mykiss / = / GSI / (3.2) / Liley et al. 2002
Oncorhynchus nerka / GSI / N/A / Foote et al. 1997
Salmo salar / GSI, TA / (153) / Vladic and Jarvi 2001; Tomkins & Simmons 2002
Scaridae / Chlorurus sordidus / GSI / (1.3) b / Gust 2004
Scarus croicensis / GSI / (1.4) b / Robertson and Warner 1978
Scarus ferrugineus / GSI / (1.8) b / Abdel-Aziz et al. 2006
Scarus vetula / GSI / (1.4) b / Robertson and Warner 1978
Amphibia / Anura / Myobatrachidae / Crinia georgiana / = * / TA / (2.0) / Byrne 2004
Aves / Charadriiformes / Scolopacidae / Philomachus pugnax / GSI / (1.1) b / Jukema and Piesma 2006

† Testes investment was compared using gonadosomatic index (GSI) or testes allometry analysis (TA, Tomkins and Simmons 2002).

†† Numbers in parenthesis represent mean difference in body mass between tactics (see caption b).

* Bourgeois males face high sperm competition because parasitic males are abundance.

** Helper males are reproductively suppressed by bourgeois males.

*** A part of bourgeois malesface high sperm competition risk.

a See also Simmons et al. (2007) for other examples in this family.

b The size difference was calculated by body length because body mass data were not available.

c Parasitic males have,not only relatively, but also absolutely larger testes than bourgeois males.

d Larger parasitic males have GSI similar to bourgeois males, and parasitic spawning is rare.

References

Abdel-Aziz SH, Bawazeer FA, Otaibi MA (2006) Histology, ultrastructure and gonadal steroids of two male phenotypes of the protogynous fish, Scarus ferrugineus (Scaridae). JKAU:Mar. Sci. 17: 33–58.

Awata, S., Heg, D., Munehara, H. & Kohda, Awata S, D Heg, H Munehara, Kohda M (2006) Testis size depends on social status and the presence of male helpers in the cooperatively breeding cichlid Julidochromis ornatus. Behav Ecol 17:372–379. doi:10.1093/beheco/arj043

Balshine S, Leach B, Neat F, Reid H, Taborsky M, Werner N (2001) Correlates of group size in a cooperatively breeding cichlid fish (Neolamprologus pulcher). Behav Ecol Sociobiol 50:134-140. doi:10.1007/s002650100343

Bass AH, Marchaterre MA (1989) Sound-generating (sonic) motor system in a teleost fish (Porichthys notatus): sexual polymorphism in the ultrastructure of myofibrils. J Comp Neurol 286:141–153. doi:10.1002/cne.902860202

Byrne PG (2004) Male sperm expenditure under sperm competition risk and intensity in quacking frogs. Behav. Ecol. 15:857-863. doi:10.1093/beheco/arh098

DeWoody JA, Fletcher DE, Mackiewicz M, Wilkins SD, Avise JC (2000) The genetic mating system of spotted sunfish (Lepomis punctatus): mate numbers and the influence of male reproductive parasites. Mol Ecol 9:2119-2128. doi:10.1046/j.1365-294X.2000.01123.x

Drilling CC, Grober MS (2005) An initial description of alternative male reproductive phenotypes in the bluebanded goby, Lythrypnus dalli (Teleostei, Gobiidae). Env Biol Fish 72:361-372. doi:10.1007/s10641-004-2590-5

Fitzpatrick JL, Desjardins JK, Stiver KA, Montgomerie R, Balshine S (2006) Male reproductive suppression in the cooperatively breeding fish Neolamprologus pulcher. Behav Ecol 17:25-33.doi:10.1093/beheco/ari090

Foote CJ, Brown GS, Wood CC (1997) Spawning success of males using alternative mating tactics in sockeye salmon, Oncorhynchus nerka. Can J Fish Aquat Sci 54:1785-1795. doi:10.1139/cjfas-54-8-1785

Geertjes GJ, Videler JJ (2002)A Quantitative Assessment of the Reproductive System of the Mediterranean Cave-Dwelling Triplefin Blenny Tripterygion melanurus. Mar. Ecol. 23: 327-340.doi:10.1046/j.1439-0485.2002.02789.x

Gross MR (1982) Sneakers, satellites and parentals: Polymorphic mating strategies in North American sunfishes. Z Tierpsychol 60:1-26. doi:10.1111/j.1439-0310.1982.tb01073.x

Gross MR (1985) Disruptive selection for alternative life histories in salmon. Nature 313:47-48. doi:10.1038/313047a0

Gust N (2004) Variation in the population of protogynous coral reef fishes over tens of kilometres. Can J Fish Aquat Sci 61:205-218.doi:10.1139/F03-160

Hourigan TF, Nakamura M, YNagahama Y, Yamauchi K, Gordon EG (1991) Histology, ultrastructure, and in vitro steroidogenesis of the testes of two male phenotypes of the protogynous fish, Thalassoma duperrey (Labridae). Gen Comp Endocr 83:193-217.doi:10.1016/0016-6480(91)90023-Y

Hurtado-Gonzales JL, Uy JAC (2009) Alternative mating strategies may favour the persistence of a genetically based colour polymorphism in a pentamorphic fish. Anim. Behav. 77:1187–1194.doi:10.1016/j.anbehav.2008.12.032

Jennings MJ, Philipp DP (1992) Reproductive investment and somatic growth rates in longear sunfish. Env Biol Fish 35:257-271. doi:10.1007/BF00001892

Jonge J, Videler JJ (1989) Differences between the reproductive biologies of Tripterygion tripteronotus and T. delaisi (Pisces, Perciformes, Tripterygiidae) the adaptive significance of an alternative mating strategy and a red instead of a yellow nuptial colour. Mar Biol 100:431–437. doi:10.1007/BF00394818

Jonge J, Ruiter AJH, Hurk R (1989) Testis-testicular gland complex of two Tripterygion species (Blennioidei, Teleostei): differences between territorial and non-territorial males. J Fish Biol 35:497-508.doi:10.1111/j.1095-8649.1989.tb03001.x

Jukema J, Piersma T (2006) Permanent female mimics in a lekking shorebird. Biol Lett 2:161-164. doi:10.1098/rsbl.2005.0416

Kato R, Munehara H, Kohda M (2005) Alternative male mating tactics of the substrate brooding cichlid Telmatochromis temporalis in Lake Tanganyika. Zool Sci 22:555-561.doi:10.2108/zsj.22.555

Kelly CD (2008) Sperm investment in relation to weapon size in a male trimorphic insect? Behav. Ecol. 19:1018-1024.

doi:10.1093/beheco/arn058

Kim SJ, Ogasawara K, Park JG, Takemura A, Nakamura M (2002) Sequence and expression of androgen receptor and estrogen receptor gene in the sex types of protogynous wrasse, Halichoeres trimaculatus. Gen Comp Endocr 127:165-173.

doi:10.1016/S0016-6480(02)00020-5

Koseki Y, Maekawa K (2000) Sexual selection on mature male parr of masu salmon (Oncorhynchus masou) : does sneaking behavior favor small body size and less-developed sexual characters? Behav Ecol Sociobiol 48:211-217. doi:10.1007/s002650000231

Koseki Y, Maekawa K (2002) Differential energy allocation of alternative male tactics in masu salmon (Oncorhynchus masou). Can J Fish Aquat Sci. 59:1717-1723. doi:10.1139/f02-143

Kvarnemo C, Svensson O, Manson W (2010) Investment in testes, sperm-duct glands and lipid reserves differs between male morphs but not between early and late breeding season in Pomatoschistus minutus. J Fish Biol 76:1609–1625.

doi:10.1111/j.1095-8649.2010.02587.x

Leach B (1997) Competition and the evolution of sperm characteristics in Pacific salmon (Oncorhynchus spp). MSc, Queen’s University

Liley NR, Tamkee P, Tsai R, Hoysak DJ (2002) Fertilization dynamics in rainbow trout (Oncorhynchus mykiss): effect of male age, social experience, and sperm concentration and motility on in vitro fertilization. Can J Fish Aq Sci 59:144-152. doi:10.1139/f01-202

Marentette JR, Fitzpatrick JL, Berger RG, Balshine S (2009) Multiple male reproductive morphs in the invasive round goby (Apollonia melanostoma). J Great Lakes Res 35:302-308.doi:10.1016/j.jglr.2009.01.009

Mazzoldi C, Scaggiante M, Ambrosin E, Rasotto MB (2000) Mating system and alternative male mating tactics in the grass goby Zosterisessor ophiocephalus (Teleostei: Gobiidae). Mar Biol 137:1041-1048. doi:10.1007/s002270000417

Miranda JA, Oliveira RF, Carneiro LA, Santos RS, Grober MS (2003) Neurochemical correlates of male polymorphism and alternative reproductive tactics in the Azorean rock-pool blenny, Parablennius parvicornis. Gen comp endocr 132:183-189.doi:10.1016/S0016-6480(03)00063-7

NeatFC (2001) Male Parasitic Spawning in Two Species of Triplefin Blenny (Tripterigiidae): Contrasts in Demography, Behaviour and Gonadal Characteristics Env Biol Fish 61:57-64.doi:10.1023/A:1011074716758

Neat FC, Locatello L, Rasotto MB (2003) Reproductive morphology in relation to alternative male reproductive tactics in Scartella cristata. J Fish Biol 62:1381-1391. doi:10.1046/j.1095-8649.2003.00122.x

Okada K, Miyatake T, Nomura Y, Kuroda K (2008) Fighting, dispersing, and sneaking: body ]size dependent mating tactics by male Librodor japonicus beetles. Ecol. Entomol. 33:269-275. doi:10.1111/j.1365-2311.2007.00965.x

Ota K, Kohda M (2006) Description of alternative male reproductive tactics in a shell-brooding cichlid, Telmatochromis vittatus, in Lake Tanganyika. J Ethol 24:9-15. doi:10.1007/s10164-005-0154-6

Ota K, Heg D, Hori M, Kohda M (2010) Sperm phenotypic plasticity in a cichlid: a territorial male’s counterstrategy to spawning takeover. Behav Ecol 21:1293-1300.doi:10.1093/beheco/arq146

Rasotto M B, Mazzoldi C (2002) Male traits associated with alternative reproductive tactics in Gobius niger. J Fish Biol 6:173-184. doi:10.1111/j.1095-8649.2002.tb01744.x

Rios-Cardenas O, Webster MS (2008) A molecular genetic examination of the mating system of pumpkinseed sunfish reveals high pay-offs for specialized sneakers. Mol Ecol 17: 2310-2320. doi:10.1111/j.1365-294X.2008.03746.x

Robertson DR, Warner RR (1978) Sexual patterns in the labroid fishes of the western Caribbean, II: the parrotfishes (Scaridae). Smithsonian Cont. Zool. 255:1–26.

Ruchon F, Laugier T, Quignard JP (1995) Alternative male reproductive strategies in the peacock blenny. J Fish Biol 47:826–840. doi:10.1111/j.1095-8649.1995.tb06005.x

Sagi A, Milner Y, Cohen D (1988) Spermatogenesis and sperm storage in the testes of the behaviorally distinctive male morphotypes of Macrobrachium rosenbergii (Decapoda, Palaemonidae). Biol. Bull. 174:330-336.

Sagi A, Ra'anan Z (1988) Morphotypic differentiation of males of the fresh-water prawn Macrobrachium rosenbergii: changes in the midgut glands and the reproductive system. J Crustac Biol 8:43-47.

Scaggiante M, Mazzoldi C, Petersen CW, Rasotto MB (1999) Sperm competition and mode of fertilization in the grass goby Zosterisessor ophiocephalus (Teleostei: Gobiidae). J Exp Zool part A 283: 81-90.

doi:10.1002/(SICI)1097-010X(19990101)283:1<81::AID-JEZ9>3.0.CO;2-9

Scaggiante M, Grober M S, Lorenzi V, Rasotto M B (2006) Variability of GnRH secretion in two goby species with socially controlled alternative male mating tactics. Horm behav 50:107-117.doi:10.1016/j.yhbeh.2006.01.011

Schlupp I, Poschadel JR, Tobler M, Plath M (2006) Male size polymorphism and testis weight in two species of mollies (Poecilia latipinna, P. mexicana, Poeciliidae, Teleostei). Z. Fischk. 8:9-16.

Schütz D, Parker GA, Taborsky M, Sato T (2006) An optimality approach to male and female body sizes in an extremely size-dimorphic cichlid fish. Evol Ecol Res 8:1-16.

Simmons LW, Tomkins JL, Hunt J (1999) Sperm competition gamesplayed by dimorphic male beetles. Proc. R. Soc. Lond. B 266:145–150. doi:10.1098/rspb.1999.0614

Simmons LW, Tomkins JL, Alcock JA (2000) Can minor males of Dawson’s burrowing bee, Amegilla dawsoni (Hymenoptera: Anthophorini) compensate for reduced access to virgin females through sperm competition? Behav. Ecol. 11:319–325.doi:10.1093/beheco/11.3.319

Smith CC, Ryan MJ (2010) Evolution of sperm quality but not quantity in the internally fertilized fish Xiphophorus nigrensis. J. Evol. Biol. 23:1759-1771. doi:10.1111/j.1420-9101.2010.02041.x

Taborsky M (1994) Sneakers, satellites, and helpers: parasitic and cooperative behavior in fish reproduction. Adv St Behav 23:1-100.doi:10.1016/S0065-3454(08)60351-4

Taborsky M (1999) Conflict or Cooperation: What determines Optimal Solutions to Competition in Fish Reproduction? In: Almada VC, Oliveira RF, Gonçalves EJ (eds). Behaviour and Conservation of Littoral Fishes. ISPA, Lisboa, pp 301-350.

Taborsky M (2008) Alternative reproductive tactics in fish. In: Oliveira R, Taborsky M, Brockmann HJ (eds). Alternative Reproductive Tactics: An Integrative Approach. Cambridge University Press, Cambridge, pp 251-299.

Tomkins JL, Simmons LW (2002) Measuring relative investment: a case study of testes investment in species with alternative male reproductive tactics. Anim Behav 63:1009–1016.doi:10.1006/anbe.2001.1994

Uglem I, Rosenqvist G, Wasslavik H S (2000) Phenotypic variation between dimorphic males in corkwing wrasse. J Fish Biol 57:1-14.doi:10.1111/j.1095-8649.2000.tb00771.x

Uglem I, Galloway TF, Rosenqvist G, Folstad I (2001) Male dimorphism, sperm traits and immunology in the corkwing wrasse (Symphodusmelops L.). Behav Ecol Sociobiol.50:511-518. doi:10.1007/s002650100392

Vladić TV, Järvi T (2001) Sperm quality in the alternative reproductive tactics of Atlantic salmon: the importance of the loaded raffle mechanism. Proc R Soc Lond B 268:2375-2381. doi:10.1098/rspb.2001.1768

Wada T (2005) Reproductive behavioral ecology in each three species of the loliginid squids and cuttlefishes (Mollusca: Cephalopoda). Ph.D. thesis, Nagasaki University (in Japanese with English summary)

Warner RR (1982) Mating Systems, Sex Change and Sexual Demography in the Rainbow Wrasse, Thalassoma lucasanum. Copeia 1982:653-661.

Warner RR, Robertson DR (1978) Sexual patterns in the labroid fishes of the western Caribbean, I: the Wrasses (Labridae). Smithsonian Cont. Zool. 254:1–27.

Warner RR, Lejeune P (1985) Sex change limited by paternal care: a test using four Mediterranean labrid fishes, genus Symphodus. Mar Biol 87:89-99. doi:10.1007/BF00397010