**Author details**

Paulo Henriques<sup>1</sup> \*, João Delgado2,3 and Ricardo Sousa<sup>4</sup>

\*Address all correspondence to: paulomvh@gmail.com

1 University of Madeira, Campus da Penteada, Funchal, Madeira, Portugal

2 Service Directorate of Research (DSI)—Regional Directorate of Fisheries, Estrada da Pontinha, Funchal, Madeira, Portugal

3 Interdisciplinary Centre of Marine and Environmental Research (CIIMAR/CIMAR), Rua dos Bragas, Porto, Portugal

4 Oceanic Observatory of Madeira (OOM)—Edifício Madeira Tecnopolo, Funchal, Madeira, Portugal

### **References**


[16] Jenkins SR, Coleman RA, Burrows MT, Hartnoll RG, Hawkins SJ. Regional scale differences in determinism of limpet grazing effects. Mar Ecol Prog Ser. 2005;**287**:77-86. DOI: 10.3354/meps287077

**References**

1992. 249 p

84 Organismal and Molecular Malacology

1971.11447402

2010.00410.x

2013;**40**:686

1973;**182**(3):1305-1392

Soc Lond. 1934;**21**(3):217-222

Museum of Natural History; 1973. 132 p

Biol. 1975;**31**:63-67. DOI: 10.1007/BF00390648

tropoda: Lottiidae). Biodivers J. 2016;**7**(2):287-293

DOI: 10.1098/rstb.1998.0316

[1] Bowman RS, Lewis JR. Geographical variation in the breeding cycles and recruitment of

[2] Stearns SC. The evolution of life histories (1st ed.). Oxford: Oxford University Press;

[3] Begon M, Harper JL, Townsend CR. Ecology: individuals, populations and communities

[4] Ridgway SA, Reid DG, Taylor JD, Branch GM, Hodgson AN. A cladistic phylogeny of the family Patellidae (Mollusca: Gastropoda). Phil Trans R Soc B. 1998;**353**:1645-1671.

[5] Koufopanou V, Reid DG, Ridgway SA, Thomas RH. A molecular phylogeny of the Patellid limpets (Gastropoda: Patellidae) and its implications for the origins of their antitropical distribution. Mol Phylogenet Evol. 1999;**11**(1):138-156. DOI: 10.1006/mpev.1998.0557 [6] WoRMS Editorial Board. World register of marine species [Internet]. 2017. Available

[7] Branch GM. The ecology of *Patella* Linnaeus from the Cape Peninsula, South Africa. I. Zonation, movements, and feeding. Zool Afr. 1971;**6**:1-38. DOI: 10.1080/00445096.

[8] Christiaens J. Révision du genre *Patella* (Mollusca, Gastropoda). B Mus Natl Hist Nat.

[9] Powell AWR. The patellid limpets of the world (Patellidae) (1st ed.). Delaware: Delaware

[10] Espinosa F, Nakano T, Guerra-García JM, García-Gómez JC. Population genetic structure of the endangered limpet *Cymbula nigra* in a temperate Northern hemisphere region: influence of palaeoclimatic events? Mar Ecol. 2010;**32**:1-5. DOI: 10.1111/j.1439-0485.

[11] Moore HB. The relation of shell growth to environment in *Patella vulgata*. Proc Malacol

[12] Davies PS. Effect of environment on metabolic activity and morphology of Mediterranean

[13] Bannister JV. Shell parameters in relation to zonation in Mediterranean limpets. Mar

[14] Boukhicha J, Kalthoum O, Hassine B, Tlig-Zouari S. Morphological evidence for adaptive diversification of sympatric Mediterranean *Patella* limpets. Rapp Comm Int Mer Medit.

[15] Scuderi D, Eernisse DJ. A new alien limpet for the Mediterranean: *Lottia* sp. (Patellogas-

*Patella* spp. Hydrobiologia. 1986;**142**:41-56. DOI: 10.1007/BF00026746

(2nd ed.). Oxford: Blackwell Scientific Publications; 1996. 945 p

from: http://www.marinespecies.org [Accessed: 24-01-2017]

and British species of *Patella*. Pubbl Stn Zool. 1969;**37**:641-656


[46] Branch GM. The biology of limpets: physical factors, energy flow and ecological interactions. Oceanogr Mar Biol Annu Rev. 1981;**19**:235-380

[30] Bree PJH. Homing-gedrag van *Patella vulgata* L. Verslagen van de gewone vergadering der Afdeeling natuurkunde K. Nederlandse Akademie van wetenschappen. 1959;**68**:106-108

[31] Cook A, Bamford OS, Freeman JDB, Teidman DJ. A study on the homing habit of the

[32] Evans MR, Williams GA. Time partitioning of foraging in the limpet *Patella vulgata*.

[33] Espinosa F, Guerra-García JM, Fa D, García-Gómez JC. Aspects of reproduction and their implications for the conservationof the endangered limpet, *Patella ferruginea*. Invertebr

[34] Sebastián CR, Steffani CN, Branch GM. Homing and movement patterns of a South African limpet *Scutellastra argenvillei* in an area invaded by an alien *mussel Mytilus galloprovincialis*.

[35] Branch GM, Cherry MI. Activity rhythms of the pulmonated limpet *Siphonaria capensis* Q. and G. as an adaptation to osmotic stress, predation and wave action. J Exp Mar Biol

[36] Iwasaki K. Intra- and interspecific variation in activity patterns of intertidal limpets.

[37] Mackay DA, Underwood AJ. Experimental studies on homing in the intertidal patellid

[38] Branch GM. The response of South African patellid limpets to invertebrate predators.

[39] Garrity SD, Levings SC. Homing to scars as a defence against predators in the pulmonate limpet *Siphonaria gigas* (Gastropoda). Mar Biol. 1983;**72**:319-324. DOI: 10.1007/BF00396838

[40] Iwasaki K. Analyses of limpet defence and predator offense in the field. Mar Biol.

[41] Branch GM. Activity rhythms in *Siphonaria thersites*. In: Chelazzi G, Vannini M, editors. Behavioural adaptation to intertidal life NATO ASI Series (Vol. 151. 1st ed.). New York:

[42] Gray DR, Hodgson AN. Foraging and homing behaviour in the high-shore, crevicedwelling limpet *Helcion pectunculus* (Prosobranchia: Patellidae). Mar Biol. 1998;**132**:

[43] Stimson J. Territorial behaviour of the owl limpet *Lottia gigantea*. Ecology. 1970;**51**:113-118.

[44] Branch GM. Mechanisms reducing intraspecific competition in *Patella* spp.: migration,

[45] Funke W. Heimfindevermögen und Ortstreue bei *Patella* L. (Gastropoda: Prosobranchia).

differentiation and territorial behaviour. J Anim Ecol. 1975;**44**:575-600

limpet. Anim Behav. 1969;**17**:330-339. DOI: 10.1016/0003-3472(69)90019-0

Reprod Dev. 2006;**49**:85-92. DOI: 10.1080/07924259.2006.9652197

Mar Ecol Prog Ser. 2002;**243**:111-122. DOI: 10.3354/meps243111

limpet *Cellana tramoserica* (Sowerby). Oecol. 1977;**30**:215-237

Zool Afr. 1978;**13**:221-232. DOI: 10.1080/00445096.1978.11447624

J Anim Ecol. 1991;**60**(2):563-575. DOI: 10.2307/5298

Ecol. 1985;**87**:153-168

86 Organismal and Molecular Malacology

Venus Jpn J Malacol. 1994;**53**:85-104

1993;**116**:277-289. DOI: 10.1007/BF00350018

Plenum Press; 1988. pp. 27-44

DOI: 10.2307/1933604

283-294. DOI: 10.1007/s002270050394

Oecol. 1968;**2**:19-142. DOI: 10.1007/BF00394506


[76] Boaventura D, Fonseca LC, Hawkins SJ. Size matters: competition within populations of the limpet *Patella depressa*. J Anim Ecol. 2003;**72**:435-446. DOI: 10.1046/j. 1365-2656.2003.00713.x

[60] Brown KM, Quinn JF. The effect of wave action on growth in three species of intertidal

[61] Lewis JR, Bowman RS. Local habitat-induced variations in the population dynamics of

[62] Branch GM. The ecology of *Patella Linnaeus* from the Cape peninsula, South Africa. 3.

[63] Black R. Population regulation in the intertidal limpet *Patelloida alticostata* (Angas, 1865).

[64] Gray DR. Studies of the biology and ecology of the high shore South African limpet, *Helcion pectunculus* (Mollusca: Patellogastropoda) [thesis]. South Africa: Rhodes University;

[65] Etim L, Sankare Y. Growth and mortality, recruitment and yield of the fresh-water shrimp, *Macrobrachium vollenhovenii*, Herklots, 1851 (Crustacea, Palaemonidae) in the Fahe reservoir, Côte d'Ivoire, West Africa. Fish Res. 1988;**38**:211-223. DOI: 10.1016/

[66] Pauly D, Munro JL. Once more on the comparison of growth in fish and invertebrates.

[67] Sparre P, Ursin E, Venema SC. Introduction to tropical fish stock assessment, part 1

[68] Orton JH, Southward AJ, Dodd JM. Studies on the biology of limpets II. The breeding of *Patella vulgata* L. in Britain. J Mar Biol Assoc UK. 1956;**35**:149-176. DOI: 10.1017/

[69] Hyman LH. The invertebrates VI – mollusca I: aplacophora, polyplacophora, monoplacophora, gastropoda (1st ed.). New York: McGraw-Hill Book Company; 1967. 792 p [70] Fretter V, Graham A. The prosobranch mollucs of Britain and Denmark. I. Pleuroto-

[71] Orton JH. Observations on *Patella vulgata*. Part III. Habitat and habits. J Mar Biol Assoc

[72] Orton JH, Southward AJ. Studies on the biology of limpets IV. The breeding of *Patella depressa* pennant on the north Cornish coast. J Mar Biol Assoc UK. 1961;**41**(3):653-662.

[74] Henninger TO, Hodgson AN. The reproductive cycle of *Helcion pruinosus* (Patellogastropoda) on two South African boulder shores. J Molluscan Stud. 2001;**67**:

[75] Hawkins SJ, Côrte-Real HBSM, Pannacciulli FG, Weber LC, Bishop JDD. Thoughts on the ecology and evolution of the intertidal biota of the Azores and other Atlantic Islands.

mariacea, Fissurellacea and Patellacea. J Molluscan Stud Supp. 1976;**1**:1-37

manual. Rome: FAO Fisheries Technical Paper; 1989. 337 p

UK. 1929;**16**:277-288. DOI: 10.1017/S0025315400029805

[73] Orton JH. Biology of *Patella* in Great Britain. Nature. 1946;**158**:173-174

Hydrobiologia. 2000;**440**:3-17. DOI: 10.1023/A:1004118220083

gastropods. Oecol. 1988;**75**(3):420-425

*Patella vulgata* L. J Exp Mar Biol Ecol. 1975;**17**:165-203

Growth rates. Trans R Soc S Afr. 1974;**41**:161-193

Oecol. 1977;**30**:9-22. DOI: 10.1007/BF00344888

1996. 304 p

88 Organismal and Molecular Malacology

S0165-7836(98)00161-1

S0025315400009036

385-394

ICLARM Fishbyte. 1984;**2**:21

DOI: 10.1017/S0025315400016210


[105] Lasiak TA. The susceptibility and/or resilience of rocky littoral molluscs to stock depletion by the indigenous coastal people of Transkei, southern Africa. Biol Cons. 1991;**56**:245-264. DOI: 10.1016/0006-3207(91)90060-M

[92] Warner RR, Fitch DL, Standish JD. Social control of sex change in the shelf limpet, *Crepidula norrisiarum*: size-specific responses to local group composition. J Exp Mar Biol

[93] Munday PL, Buston PM, Warner RR. Diversity and flexibility of sex-change strategies

[94] Marra S, de Lucia GA, Camedda A, Esinosa F, Coppa S. New records of the distribution and conservation status of the endangered limpet *Patella ferruginea* in Sardinia (Italy, W Mediterranean). Aquat Conserv Mar Freshw Ecosys. 2015;**26**(3):607-612. DOI: 10.1002/

[95] Durán LR, Castilla JC. Variation and persistence of the middle rocky intertidal community of central Chile, with and without human harvesting. Mar Biol. 1989;**103**:555-562.

[96] Lasiak TA. Multivariate comparisons of rocky infratidal macrofaunal assemblages from replicate exploited and non-exploited localities on the Transkei coast of South Africa.

[97] Sagarin RD, Ambrose RF, Becker BJ, Engle JM, Kido J, Lee SF, Miner CM, Murray SN, Raimondi PT, Richards DV, Roe C. Ecological impacts on the limpet *Lottia gigantea* populations: human pressure over a broad scale on islands and mainland intertidal zones.

[98] Fenberg PB, Roy K. Ecological and evolutionary consequences of size-selective harvesting: how much do we know? Mol Ecol. 2008;**17**:209-220. DOI: 10.1111/j.1365-

[99] Eekhout S, Raubenheimer CM, Branch GM, Bosman AL, Bergh MO. A holistic approach to the exploitation of intertidal stocks: limpets as a case study. Afr J Mar Sci.

[100] Griffiths CL, Branch GM. The exploitation of coastal invertebrates and seaweeds in South Africa: historical trends, ecological impacts and implications for management.

[101] Pombo OA, Escofet A. Effect of exploitation on the limpet *Lottia gigantea*: a field study in Baja California (Mexico) and California (U.S.A.). Pac Sci. 1996;**50**:393-403. DOI: 10125/2914

[102] Turrero P, Munoz Colmenero AM, Prado A, García-Vázquez E. Long-term impacts of human harvesting on shellfish: North Iberian top shells and limpets from the upper palaeolithic to the present. J Mar Syst. 2014;**139**:51-57. DOI: 10.1016/j.jmarsys.2014.05.011

[103] McCoy M. Hawaiian limpet harvesting in historical perspective: a review of modern and archaeological data on *Cellana* spp. From the Kalaupapa Peninsula, Moloka'i Island. Pac Sci. 2008;**62**(1):28-38. DOI: 10.2984/1534-6188(2008)62[21:HLHIHP]2.0.CO;2

[104] Keough MJ, Quinn GP, King A. Correlations between human collecting and intertidal

mollusc populations on rocky shores. Conserv Biol. 1993;**7**:378-390

Trans R Soc S Afr. 1997;**52**:121-148. DOI: 10.1080/00359199709520619

in animals. Trends Ecol Evol. 2006;**21**:89-95. DOI: 10.1016/j.tree.2005.10.020

Ecol. 1996;**204**:155-167. DOI: 10.1016/0022-0981(96)02582-8

Mar Ecol Prog Ser. 1998;**167**:15-23. DOI: 10.3354/meps167015

Mar Biol. 2007;**150**:399-413. DOI: 10.1007/s00227-006-0341-1

aqc.2615

90 Organismal and Molecular Malacology

DOI: 10.1007/BF00399588

294X.2007.03522.x

1992;**12**:1017-1029


[135] Pelc RA, Warmer RR, Gaines SD, Paris CB. Detecting larval export from marine reserves. Proc Natl Acad Sci USA. 2010;**107**:18266-18271. DOI: 10.1073/pnas.0907368107

[119] Alonzo SH, Mangel M. The effects of size selective fisheries on the stock dynamics of

[120] Baskett ML, Levin SA, Gaines SD, Dushoff J. Marine reserve design and the evolution of size at maturation in harvested fish. Ecol Appl. 2005;**15**:882-901. DOI: 10.1890/04-0723

[121] Heppell SS, Heppell SA, Coleman FC, Koenig CC. Models to compare management options for a protogynous fish. Ecol Appl. 2006;**16**:238-249. DOI: 10.1890/04-1113 [122] Hamilton SL, Caselle JE, Standish JD, Schroeder DM, Love MS, Rosales-Casian JA, Sosa-Nishizaki O. Size-selective harvesting alters life histories of a sex-changing fish.

[123] Martins GM, Jenkins SR, Hawkins SJ, Neto AI, Medeiros AR, Thompson RC. Illegal harvesting affects the success of fishing closure areas. J Mar Biol Assoc UK. 2011;**91**:

[124] Costello C, Gaines SD, Lynham J. Can catch shares prevent fisheries collapse? Science.

[125] Baxter JM. Establishing management schemes on marine special areas of conservation in Scotland. Aquat Conserv Mar Freshw Ecosys. 2001;**11**:261-265. DOI: 10.1002/aqc.465

[126] Thompson RC, Crowe TP, Hawkins SJ. Rocky intertidal communities: past environmental changes, present status and predictions for the next 25 years. Environ Conserv.

[127] Ballantine B. Marine reserves for New Zealand. Leigh Laboratory bulletin (Vol. 25).

[128] Zann LP. Our sea, our future. Major findings of the state of the marine environment report for Australia (1st ed.). QLD: Great Barrier Reef Marine Park Authority; 1995

[129] Halpern BS. The impact of marine reserves: do reserves work and does reserve size matter? Ecol Appl. 2003;**13**:117-137. DOI: 10.1890/1051-0761(2003)013[0117:TIOMRD]2

[130] Roberts CM, Hawkins JP. Extinction risk in the sea. Trends Ecol Evol. 1999;**14**:241-246

[131] Lubchenco J, Palumbi SR, Gaines SD, Andelman S. Plugging a hole in the ocean: the energy science of marine reserves. Ecol Appl. 2003;**13**:S3–S7. DOI: 10.1890/1051-0761

[132] Gell FR, Roberts CM. Benefits beyond boundaries: the fishery effects of marine reserves.

[133] Halpern BS, Warner RR. Matching marine reserve design to reserve objectives. Proc R

[134] Pelc RA, Baskett ML, Tanci T, Gaines SD, Warner RR. Quantifying larval export from South African marine reserves. Mar Ecol Prog Ser. 2009;**394**:65-78. DOI: 10.3354/meps08326

Trends Ecol Evol. 2003;**18**:448-455. DOI: 10.1016/S0169-5347(03)00189-7

Soc Lond Ser B Biol Sci. 2003;**270**:1871-1878. DOI: 10.1098/rspb.2003.2405

and sperm limitation in sex-changing fish. Fish Bull. 2004;**102**:1-13

Ecol Appl. 2007;**17**:2268-2280. DOI: 10.1890/06-1930.1

929-937. DOI: 10.1017/S0025315410001189

92 Organismal and Molecular Malacology

2008;**321**:1678-1681. DOI: 10.1126/science.1159478

2002;**29**:168-191. DOI: 10.1017/S0376892902000115

Auckland: University of Auckland; 1991. pp. 1-196

(2003)013[0003:PAHITO]2.0.CO;2

.0.CO;2


[162] Boersma DP, Parrish JK. Limiting abuse: marine protected areas, a limiting solution. Ecol Econ. 1999;**31**:287-304. DOI: 10.1016/S0921-8009(99)00085-3

[148] Bastida-Zavala JR, García-Madrigal MS, Rosas-Alquicira EF, López-Pérez RA, Benítez-Villalobos F, Meraz-Hernando JF, Torres-Huerta M, Montoya-Márquez A, Barrientos-Luján NA. Marine and coastal biodiversity of Oaxaca. Mexico Check List. 2013;**9**:329-390.

[149] Halpern BS, Warner RR. Marine reserves have rapid and lasting effects. Ecol Lett.

[150] Hockey PAR, Bosman AL. Man as an intertidal predator in Transkei: disturbance, community convergence and management of a natural food resource. Oikos. 1986;**46**:3-14.

[151] Roberts C. Marine fishery reserves for the Caribbean. Caribbean Parks Protect Area

[152] Russ G, Alcala A. Natural fishing experiments in marine reserves 1983-1993: community and trophic responses. Coral Reefs. 1998;**17**:383-397. DOI: 10.1007/s003380050144

[153] Polacheck T. Year around closed areas as a management tool. Nat Resour Model.

[154] Carr MH, Reed DC. Conceptual issues relevant to marine harvest refuges: examples from temperature reef fishes. Can J Fish Aquat Sci. 1993;**50**:2019-2028. DOI: 10.1139/

[155] Rowley RJ. Marine reserves in fisheries management. Aquat Conserv Mar Freshw

[156] Palumbi SR. Marine reserves and ocean neighborhoods: the spatial scale of marine populations and their management. Annu Rev Environ Resour. 2004;**29**:31-68. DOI:

[157] Hockey PAR, Branch GM. Conserving marine biodiversity on the African coast: implications of a terrestrial perspective. Aquat Conserv Mar Freshw Ecosys. 1994;**4**:345-362.

[158] Caley MJ, Carr MH, Hixon MA, Hughes TP, Jones GP, Menge BA. Recruitment and the local dynamics of open marine populations. Annu Rev Ecol Syst. 1996;**27**:477-500. DOI:

[159] Coppa S, Lucia GA, Massaro G, Magni P. Density and distribution of *Patella ferruginea* in a marine protected area (western Sardinia, Italy): constraint analysis for population

[160] Carr MH. Marine protected areas: challenges and opportunities for understanding and

[161] Claudet J, Guidetti P, Mouillot D, Shears NT, Micheli F. Ecological effects of marine protected areas: conservation, restoration and functioning. In: Claudel J, editor. Marine protected areas: a multidisciplinary approach (1st ed.). Cambridge, UK: Cambridge

conservation. Mediterr Mar Sci. 2012;**13**(1):108-117. DOI: 10.12681/mms.27

conserving coastal marine ecosystems. Environ Conserv. 2000;**27**:106-109

University Press; 2011. DOI: 10.1017/CBO9781139049382.005

2002;**5**:361-366. DOI: 10.1046/j.1461-0248.2002.00326.x

Ecosys. 1994;**4**:233-254. DOI: 10.1002/aqc.3270040305

10.1146/annurev.energy.29.062403.102254

DOI: 10.1002/aqc.3270040406

10.1146/annurev.ecolsys.27.1.477

DOI: 10.15560/9.2.329

94 Organismal and Molecular Malacology

DOI: 10.2307/3565373

Bull. 1995;**5**(2):8-11

1990;**4**:327-353

f93-226


**Cellular and Molecular Malacology**
