Invertebrate Systematics Invertebrate Systematics Society
Systematics, phylogeny and biogeography

Phylogeny accurately predicts behaviour in Indian Ocean Clitaetra spiders (Araneae : Nephilidae)

Matjaž Kuntner A B E and Ingi Agnarsson A B C D

A Institute of Biology, Scientific Research Centre, Slovenian Academy of Sciences and Arts, Novi trg 2, PO Box 306, SI-1001 Ljubljana, Slovenia.

B Department of Entomology, National Museum of Natural History, Smithsonian Institution, NHB-105, PO Box 37012, Washington, DC 20013-7012, USA.

C Department of Biology and Integrated Bioscience Program, University of Akron, Akron, OH 44325-3908, USA.

D Department of Biology, University of Puerto Rico, PO Box 23360, San Juan, PR 00931-3360, USA.

E Corresponding author. Email:

Invertebrate Systematics 23(3) 193-204
Submitted: 15 January 2009  Accepted: 27 April 2009   Published: 21 July 2009


Phylogenies are underutilised, powerful predictors of traits in unstudied species. We tested phylogenetic predictions of web-related behaviour in Clitaetra Simon, 1889, an Afro-Indian spider genus of the family Nephilidae. Clitaetra is phylogenetically sister to all other nephilids and thus important for understanding ancestral traits. Behavioural information on Clitaetra has been limited to only C. irenae Kuntner, 2006 from South Africa which constructs ladder webs. A resolved species-level phylogeny unambiguously optimised Clitaetra behavioural biology and predicted web traits in five unstudied species and a uniform intrageneric nephilid web biology. We tested these predictions by studying the ecology and web biology of C. perroti Simon, 1894 on Madagascar and C. episinoides Simon, 1889 on Mayotte. We confirm predicted arboricolous web architecture in these species. The expected ontogenetic allometric transition from orbs in juveniles to elongate ladder webs in adults was statistically significant in C. perroti, whereas marginally not significant in C. episinoides. We demonstrate the persistence of the temporary spiral in finished Clitaetra webs. A morphological and behavioural phylogenetic analysis resulted in unchanged topology and persisting unambiguous behavioural synapomorphies. Our results support the homology of Clitaetra hub reinforcement with the nephilid hub-cup. In Clitaetra, behaviour was highly predictable and remained consistent with new observations. Our results confirm that nephilid web biology is evolutionarily conserved within genera.

Additional keywords: homology, Comoros, Madagascar, Mayotte, orb web, phylogenetic prediction, web architecture, web allometry.


Agnarsson I. Kuntner M. 2005 Madagascar: an unexpected hotspot of social Anelosimus spider diversity (Araneae: Theridiidae). Systematic Entomology 30 575 592 doi:10.1111/j.1365-3113.2005.00289.x

ap Rhisiart A. Vollrath F. 1994 Design features of the orb web of the spider, Araneus diadematus. Behavioral Ecology 5 280 287 doi:10.1093/beheco/5.3.280

Blackledge T. A. Gillespie R. G. 2002 Estimation of capture areas of spider orb webs in relation to asymmetry. Journal of Arachnology 30 70 77 doi:10.1636/0161-8202(2002)030[0070:EOCAOS]2.0.CO;2

Blackledge T. A. Hayashi C. Y. 2006 Silken toolkits: biomechanics of silk fibers spun by the orb web spider Argiope argentata (Fabricius 1775). Journal of Experimental Biology 209 2452 2461 doi:10.1242/jeb.02275

Coddington J. A. Agnarsson I. 2006 Subsociality in Helvibis thorelli Keyserling 1884 (Araneae, Theridiidae, Theridiinae) from French Guiana. Journal of Arachnology 34 642 645 doi:10.1636/05-84.1

Coddington J. A. Hormiga G. Scharff N. 1997 Giant female or dwarf male spiders? Nature 385 687 688 doi:10.1038/385687a0

Cranston P. S. 2008 Phylogeny predicts larval biology in a wood-mining chironomid (Diptera: Chironomidae). African Entomology 16 1 6 doi:10.4001/1021-3589-16.1.1

Eberhard W. G. 1976 Physical properties of sticky spirals and their connections: sliding connections in orb webs. Journal of Natural History 10 481 488 doi:10.1080/00222937600770391

Eberhard W. G. 1986 Ontogenetic changes in the web of Epeirotypus sp. (Araneae, Theridiosomatidae). Journal of Arachnology 14 125 128

Eberhard W. G. 2000 Breaking the mold: behavioral variation and evolutionary innovation in Wendildarda spiders (Araneae Theridiosomatidae). Ethology Ecology and Evolution 12 223 235

Eberhard W. G. Agnarsson I. Levi H. W. 2008 a Web forms and the phylogeny of theridiid spiders (Araneae: Theridiidae): chaos from order. Systematics and Biodiversity 6 415 475 doi:10.1017/S1477200008002855

Eberhard W. G. Barrantes G. Madrigal-Brenes R. 2008 b Vestiges of an orb-weaving ancestor? The ‘biogenetic law’ and ontogenetic changes in the webs and building behavior of the black widow spider Latrodectus geometricus (Araneae: Theridiidae). Ethology Ecology and Evolution 20 211 244

Felsenstein J. 1985 Phylogenies and the comparative method. American Naturalist 125 1 15 doi:10.1086/284325

Fromhage L. Schneider J. M. 2005 a Safer sex with feeding females: sexual conflict in a cannibalistic spider. Behavioral Ecology 16 377 382 doi:10.1093/beheco/ari011

Fromhage L. Schneider J. M. 2005 b Virgin doves and mated hawks: contest behaviour in a spider. Animal Behaviour 70 1099 1104 doi:10.1016/j.anbehav.2005.02.020

Fromhage L. Schneider J. M. 2006 Emasculation to plug up females: the significance of pedipalp damage in Nephila fenestrata. Behavioral Ecology 17 353 357 doi:10.1093/beheco/arj037

Harvey P. H. , and Pagel M. D. (1998). ‘The Comparative Method in Evolutionary Biology.’ (Oxford University Press: Oxford.)

Harvey M. S. Austin A. D. Adams M. 2007 The systematics and biology of the spider genus Nephila (Araneae: Nephilidae) in the Australasian region. Invertebrate Systematics 21 407 451 doi:10.1071/IS05016

Higgins L. E. 2002 Female gigantism in a New Guinea population of the spider Nephila maculata. Oikos 99 377 385 doi:10.1034/j.1600-0706.2002.990220.x

Hubert N. Duponchelle F. Nunez J. Garcia-Davila C. Paugy D. Renno J. F. 2007 Phylogeography of the piranha genera Serrasalmus and Pygocentrus: implications for the diversification of the Neotropical ichthyofauna. Molecular Ecology 16 2115 2136 doi:10.1111/j.1365-294X.2007.03267.x

Japyassu H. F. Ades C. 1998 From complete orb to semi-orb webs: developmental transitions in the web of Nephilengys cruentata (Araneae: Tetragnathidae). Behaviour 135 931 956

Kiontke K. Gavin N. P. Raynes Y. Roehrig C. Piano F. Fitch D. H. A. 2004 Caenorhabditis phylogeny predicts convergence of hermaphroditism and extensive intron loss. Proceedings of the National Academy of Sciences, USA 101 9003 9008 doi:10.1073/pnas.0403094101

Kuntner M. 2005 A revision of Herennia (Araneae: Nephilidae: Nephilinae), the Australasian ‘coin spiders’. Invertebrate Systematics 19 391 436 doi:10.1071/IS05024

Kuntner M. 2006 Phylogenetic systematics of the Gondwanan nephilid spider lineage Clitaetrinae (Araneae, Nephilidae). Zoologica Scripta 35 19 62 doi:10.1111/j.1463-6409.2006.00220.x

Kuntner M. 2007 A monograph of Nephilengys, the pantropical ‘hermit spiders’ (Araneae, Nephilidae, Nephilinae). Systematic Entomology 32 95 135 doi:10.1111/j.1365-3113.2006.00348.x

Kuntner M. Coddington J. A. Hormiga G. 2008 a Phylogeny of extant nephilid orb-weaving spiders (Araneae, Nephilidae): testing morphological and ethological homologies. Cladistics 24 147 217 doi:10.1111/j.1096-0031.2007.00176.x

Kuntner M. Haddad C. R. Aljančič G. Blejec A. 2008 b Ecology and web allometry of Clitaetra irenae, an arboricolous African orb-weaving spider (Araneae, Araneoidea, Nephilidae). The Journal of Arachnology 36 583 594 doi:10.1636/T07-54.1

Kuntner M. Coddington J. A. Schneider J. M. 2009 a Intersexual arms race? Genital coevolution in nephilid spiders (Araneae, Nephilidae). Evolution 63 1451 1463 doi:10.1111/j.1558-5646.2009.00634.x

Kuntner M. Kralj-Fišer S. Schneider J. M. Li D. 2009 b Mate plugging via genital mutilation in nephilid spiders: an evolutionary hypothesis. Journal of Zoology 277 257 266 doi:10.1111/j.1469-7998.2008.00533.x

Masters W. M. Moffat A. J. M. 1983 A functional explanation of top–bottom asymmetry in vertical orbwebs. Animal Behaviour 31 1043 1046 doi:10.1016/S0003-3472(83)80010-4

Miyashita T. 2005 Contrasting patch residence strategy in two species of sit-and-wait foragers under the same environment: a constraint by life history? Ethology 111 159 167 doi:10.1111/j.1439-0310.2004.01048.x

Robinson M. H. Lubin Y. D. 1979 Specialists and generalists: the ecology and behavior of some web-building spiders from Papua New Guinea, 1. Herennia ornatissima, Argiope ocyaloides and Arachnura melanura (Araneae: Araneidae). Pacific Insects 21 97 132

Schneider J. M. Elgar M. A. 2001 Sexual cannibalism and sperm competition in the golden orb-web spider Nephila plumipes (Araneoidea): female and male perspectives. Behavioral Ecology 12 547 552 doi:10.1093/beheco/12.5.547

Schneider J. M. Elgar M. A. 2002 Sexual cannibalism in Nephila plumipes as a consequence of female life history strategies. Journal of Evolutionary Biology 15 84 91 doi:10.1046/j.1420-9101.2002.00363.x

Schneider J. M. Elgar M. A. 2005 The combined effects of pre- and post-insemination sexual selection on extreme variation in male body size. Evolutionary Ecology 19 419 433 doi:10.1007/s10682-005-8310-6

Schuh R. T. (2000). ‘Biological Systematics – Principles and Applications.’ (Cornell University Press: Ithaca, NY.)

Tso I. M. 2004 The effect of food and silk reserve manipulation on decoration-building of Argiope aetheroides. Behaviour 141 603 616 doi:10.1163/1568539041166690

Tukey J. W. (1977). ‘Exploratory Data Analysis.’ (Addison-Wesley Publishing: Reading, MA.)

Uhl G. Vollrath F. 2000 Extreme body size variability in the golden silk spider (Nephila edulis) does not extend to genitalia. Journal of Zoology 251 7 14 doi:10.1111/j.1469-7998.2000.tb00587.x

van der Heijden R. T. J. M. Snel B. van Noort V. Huynen M. A. 2007 Orthology prediction at scalable resolution by phylogenetic tree analysis. BMC Bioinformatics 8 83 doi:10.1186/1471-2105-8-83

Vollrath F. Parker G. A. 1992 Sexual dimorphism and distorted sex ratios in spiders. Nature 360 156 159 doi:10.1038/360156a0

Warren D. L. Glor R. E. Turelli M. 2008 Environmental niche equivalency versus conservatism: quantitative approaches to niche evolution. Evolution 62 2868 2883 doi:10.1111/j.1558-5646.2008.00482.x

Ye K. Vriend G. Ijzerman A. P. 2008 Tracing evolutionary pressure. Bioinformatics 24 908 915 doi:10.1093/bioinformatics/btn057

Zschokke S. 1993 The influence of the auxiliary spiral on the capture spiral in Araneus diadematus Clerck (Araneidae). Bulletin of the British Arachnological Society 9 169 173

Zschokke S. 1999 Nomenclature of the orb-web. Journal of Arachnology 27 542 546

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