Estimation of apparent survival probability of the harvestman Paranemastoma sillii sillii (Herman, 1871) from two caves

in Animal Biology
Restricted Access
Get Access to Full Text
Rent on DeepDyve

Have an Access Token?

Enter your access token to activate and access content online.

Please login and go to your personal user account to enter your access token.


Have Institutional Access?

Access content through your institution. Any other coaching guidance?


Reliable estimates of population parameters are lacking for most cave-dwelling species. This lack of knowledge may hinder the appropriate management of caves and populations of cave-dwelling species. Using monthly capture-recapture data and Cormack-Jolly-Seber models, we (i) estimated the apparent survival of individuals in two cave populations of the harvestman Paranemastoma sillii sillii (Herman, 1871) from the Mehedinti Mountains in south-western Romania; (ii) investigated temporal variation in apparent survival; (iii) tested if surface weather conditions affect apparent survival of cave-dwelling harvestmen through their influence upon cave environmental conditions and (iv) tested for sex differences in apparent survival. Our results show that the apparent monthly survival estimates were high for both studied cave populations and there was a significant sex effect on survival. Males had lower survival than females, and the survival difference between caves was larger in males than in females. Temporal (i.e., monthly) variation in apparent survival was low and the weather conditions at the surface had little influence on apparent survival as the environment inside the caves is well buffered against weather fluctuations outside the caves. Our results indicate that caves stabilize survival of facultative cave-dwelling species and may serve as microrefugia for epigean species. We suggest that caves should be considered for conservation because they may serve as a refuge for some epigean species during harsh weather conditions.



AvramŞ.DumitrescuD. (1969) Contributii la cunoasterea raspindirii geografice si a ecologiei opilionidelor cavernicole, endogee si epigee, din Romania. Trav. Inst. Spéol. E. Racovitza, 8, 99-145.

AvramŞ. (1973) Contribution à la connaissance du développement embryonnaire et postembryonnaire chez Nemastoma cf. sillii Herman (Opiliones, Nemastomatidae). In: Livre Cent. Trav. Inst. Spéol. E. Racovitza, pp.  269-303.

BadinoG. (2010) Underground meteorology – “What’s the weather underground?”. Acta Carsol., 39, 427-448.

BalogováM.JelićD.KyselováM.UhrinM. (2017) Subterranean systems provide a suitable overwintering habitat for Salamandra. I.J.S., 46(3).

BurnhamK.P.AndersonD.R.WhiteG.C.BrownieC.PollockK.H. (1987) Design and Analysis Methods for Fish Survival Experiments Based on Release-Recapture. American Fisheries Society Monograph 5. Bethesda, Maryland.

BurnhamK.P.AndersonD.R. (1998) Model Selection and Inference. Springer, New York.

BurtonJ.L.LanceR.F.WesterveltJ.D.LebergP.J. (2012) An individual-based model for metapopulations on patchy landscapes-genetics and demography (IMPL-GD). In: VesterveltJ.D.CohenG.L. (Eds) Ecologist-Developed Spatially-Explicit Dynamic Landscape Models. Modeling Dynamic Systems, pp.  197-209. Springer Science + Business Media LLC, New York.

BuzattoB.A.RequenaG.S.MartinsE.G.MachadoG. (2007) Effects of maternal care on the lifetime reproductive success of females in a Neotropical harvestman. J. Anim. Ecol., 76, 937-945.

BuzattoB.A.TomkinsJ.L.SimmonsL.W.MachadoG. (2014) Correlated evolution of sexual dimorphism and male dimorphism in a clade of Neotropical harvestmen. Evolution, 68, 1671-1686.

CignaA.A. (2002) Modern trend in cave monitoring. Acta Carsologica, 3, 35-54.

ConstantinS.LauritzenS.E.LundbergJ. (2006) New data on the chronology of the Termination II and paleoclimate during MIS 5, based on the study of a stalagmite from Closani Cave (SW Romania). In: OnacB.P.TamasT.ConstantinS.PersoiuA. (Eds) Archives of Climate Change in Karst. Special Publication 10, pp.  98-100. Karst Waters Institute, USA.

CostacheI. (2011) Flora şi Vegetaţia Bazinului Hidrografic Inferior al Râului Motru, I, Flora. Universitaria, Craiova.

CroneE.E. (2001) Is survivorship a better fitness surrogate than fecundity? Evolution, 55, 2611-2614.

CulverD.C. (1982) Cave Life: Evolution and Ecology. Harvard University Press, Massachussets and London.

CulverD.C.PipanT. (2009) The Biology of Caves and Other Subterranean Habitats. Oxford University Press, New York.

EmlenS.T.OringL.W. (1977) Ecology, sexual selection and the evolution of mating systems. Science, 197, 215-222.

FicetolaG.F.PennatiR.ManentiR. (2012) Do cave salamanders occur randomly in cavities? An analysis with Hydromantes strinatii. Amphib-Reptil., 33, 251-259.

FrederiksenM.LebretonJ.D.PradelR.ChoquetR.GimenezO. (2014) Identifying links between vital rates and environment: a toolbox for the applied ecologist. J. Appl. Ecol., 51, 71-81.

GouldW.R.NicholsJ.D. (1998) Estimation of temporal variability of survival in animal populations. Ecology, 79, 2531-2538.

HoyleS.D.PopleA.R.ToopG.J. (2001) Mark-recapture may reveal more about ecology than about population trends: demography of a threatened ghost bat (Macroderma gigas) population. Austral. Ecol., 26, 80-92.

InchaustiP.HalleyJ. (2003) On the relation between temporal variability and persistence time in animal populations. J. Anim. Ecol., 72, 899-908.

JohnsonH.E.MillsL.S.StephensonT.R.WehausenJ.D. (2010) Population-specific vital rate contributions influence management of an endangered ungulate. Ecol. Appl., 20, 1753-1765.

JoyceS.J.JamiesonI.G.BarkerR. (2004) Survival of adult mountain stone weta Hemideina maori (Orthoptera: Anostostomatidae) along an altitude gradient as determined by mark-recapture. New Zeal. J. Ecol., 28, 55-61.

LaakeJ.RexstadE. (2008) RMark – an alternative to building linear models in MARK. In: CoochE.WhiteG. (Eds) Program MARK: a Gentle Introduction. 9th Edition. Available at (Accessed 1 October 2015).

LambertD.M.HoenigJ.M.LipiciusR.N. (2006) Tag return estimation of annual and semiannual survival rates of adult female blue crabs. Trans. Am. Fish. Soc., 135, 1592-1603.

LebretonJ.D.BurnhamK.P.ClobertJ.AndersonD.R. (1992) Modeling survival and testing biological hypothesis using marked animals: a unified approach with case studies. Ecol. Monogr., 62, 67-118.

LimaS.L.DillL.M. (1990) Behavioral decisions made under the risk of predation: a review and prospectus. Can. J. Zool., 68, 619-640.

LunghiE.ManentiR.FicetolaG.F. (2014) Do cave features affect underground habitat exploitation by non-troglobite species? Acta. Oecol., 55, 29-35.

MacNamaraJ.M.HoustonA.I. (1994) The effect of a change in foraging options on intake rate and predation rate. Am. Nat., 144, 978-1000.

ManentiR.FicetolaG.F.BianchiB.De BernardiF. (2009) Habitat features and distribution of Salamandra salamandra in underground springs. Acta. Herpetol., 4, 143-151.

ManentiR.FicetolaG.F.MarieniA.De BernardiF. (2011) Caves as breeding sites for Salamandra salamandra: habitat selection, larval development and conservation issues. North-West. J. Zool., 7, 304-309.

ManentiR.SiesaM.E.FicetolaG.F. (2013) Odonata occurrence in caves: active or accidentals? A new case study. J. Cave. Karst. Stud., 75, 205-209.

MestreL.A.M.Pinto-da-RochaR. (2004) Population dynamics of an isolated population of the Harvestmen Ilhaia cuspidata (Opiliones, Gonyleptidae), in Araucaria Forest (Curitiba, Paraná, Brazil). J. Arachnol., 32, 208-220.

PabstW. (1953) Zur Biologie der mitteleuropäischen Troguliden. Zool. Jb. Abt. Syst. Ökol. U. Geog. Tiere., 82, 1-156.

PaintingC.J.ProbertA.F.TownsendD.J.HolwellG.I. (2015) Multiple exaggerated weapon morphs: a novel form of male polymorphism in harvestmen. Sci. Rep., 5, 16368.

PipanT.CulverD.C. (2012) Convergence and divergence in the subterranean realm: a reassessment. Biol. J. Linnean Soc., 107, 1-14.

ProtasM.JefferyW.R. (2012) Evolution and development in cave animals: from fish to crustaceans. Wiley Interdiscip. Rev. Dev. Biol., 1, 823-845.

R Development Core Team (2015) R: a Language and Environment for Statistical Computing. R Foundation for Statistical Computing, Vienna. Available at (Accessed 1 October 2015).

RaceyP.A.EntwistleA.C. (2000) Life history and reproductive strategies of bats. In: CrichtonE.G.KrutzschP. (Eds) The Reproductive Biology of Bats, pp.  363-414. Academic Press, London.

ReddellJ.R. (2012) Spiders and related groups. In: CulverC.WhiteW.B. (Eds) Encyclopedia of Caves, 786-797. Elsevier.

RequenaG.S.BuzattoB.A.MartinsE.G.MachadoG. (2012) Paternal care decreases foraging activity and body condition, but does not impose survival costs to caring males in a Neotropical arachnid. PLoS ONE, 7, e46701.

SandelB. (2014) Towards a taxonomy of spatial scale-dependence. Ecography, 38, 358-369.

SchmidtB.R.SchaubM.AnholtB.R. (2002) Why you should use capture-recapture methods when estimating survival and breeding probabilities: on bias, temporary emigration, overdispersion and common toads. Amphib-Reptil., 23, 375-388.

SchmidtB.R.SchaubM.SteinfartzS. (2007) Apparent survival of the salamander Salamandra salamandra is low because of high migratory activity. Front. Zool., 4, 19.

SendorT.SimonM. (2003) Population dynamics of the pipistrelle bat: effects of sex, age and winter weather on seasonal survival. J. Anim. Ecol., 72, 308-320.

SimõesM.H.Souza-SilvaM.FerreiraR.L. (2015) Cave physical attributes influencing the structure of terrestrial invertebrate communities in Neotropics. Subterr. Biol., 16, 103-121.

TalleyT.S. (2007) Which spatial heterogeneity framework? Consequences for conclusions about patchy population. Ecology, 88, 1476-1489.

TaylorS.J.KrejcaJ.K.NiemillerM.L.DreslikM.J.PhillipsC.A. (2015) Life history and demographic differences between cave and surface populations of the western slimy salamander, Plethodon albagula (Caudata: Plethodontidae), in central Texas. Herpetol. Conserv. Biol., 10, 740-752.

VillellaR.F.SmithD.R.LemarieD.P. (2004) Estimating survival and recruitment in a freshwater mussel population using mark-recapture techniques. Am. Midl. Nat., 151, 114-133.

WhiteG.C.BurnhamK.P. (1999) Program MARK: survival estimation for populations of marked animals. Bird Study., 46(sup001), 120-138.

WillemartR.H.GnaspiniP. (2004) Spatial distribution, mobility, gregariousness, and defensive behavior in Brazilian cave harvestman Goniosoma albiscriptum (Arachnida, Opiliones, Gonyleptidae). Anim. Biol., 54, 221-235.

WillemartR.H.OssesF.CheliniM.C.Macías-OrdóñezR.MachadoG. (2009) Sexually dimorphic legs in a Neotropical harvestman (Arachnida, Opiliones): ornament or weapon? Behav. Processes, 80, 51-59.

ZajitschekF.BrassilC.BondurianskyR.BrooksR.C. (2009) Sex effects on life span and senescence in the wild when dates of birth and death are unknown. Ecology, 90, 1698-1707.


  • Estimated monthly apparent survival probabilities (± SE) of harvestmen in Closani and Lazului caves. Circles are females and triangles are males. Estimates are based on the most parsimonious mark-recapture model (Model 11 in table 2).

    View in gallery


Content Metrics

Content Metrics

All Time Past Year Past 30 Days
Abstract Views 23 23 12
Full Text Views 12 12 7
PDF Downloads 0 0 0
EPUB Downloads 4 4 3