Phylogeny of extant nephilid orb‐weaving spiders (Araneae, Nephilidae): testing morphological and ethological homologies

The Pantropical spider clade Nephilidae is famous for its extreme sexual size dimorphism, for constructing the largest orb‐webs known, and for unusual sexual behaviors, which include emasculation and extreme polygamy. We synthesize the available data for the generaNephila,Nephilengys,HerenniaandClitaetrato produce the first species level phylogeny of the family. We score 231 characters (197 morphological, 34 behavioral) for 61 taxa: 32 of the 37 known nephilid species plus twoPhonognathaand oneDeliochusspecies, 10 tetragnathid outgroups, nine araneids, and one genus each of Nesticidae, Theridiidae, Theridiosomatidae, Linyphiidae, Pimoidae, Uloboridae and Deinopidae. Four most parsimonious trees resulted, among which successive weighting preferred one ingroup topology. Neither an analysis of an alternative data set based on different morphological interpretations, nor separate analyses of morphology and behavior are superior to the total evidence analysis, which we therefore propose as the working hypothesis of nephilid relationships, and the basis for classification. Ingroup generic relationships are (Clitaetra(Herennia(Nephila,Nephilengys))). DeliochusandPhonognathagroup with Araneidae rather than Nephilidae. Nephilidae is sister to all other araneoids (contramost recent literature). Ethological data, although difficult to obtain and thus frequently missing for rare taxa, are phylogenetically informative. We explore the evolution of selected morphological and behavioral characters, discuss and redefine the homology of palpal sclerites, disprovesemientelegynyin spiders, trace the newly interpreted evolution of the orb web, and show that nephilid genital morphologies coevolve with sexual behaviors and extreme sexual size dimorphism. Phylogenetic interpretations of behavior suggest new insights into spider biology and avenues for future research.