18 FROM: Ubick, D., P. Paquin, P.E. Cushing, andV. Roth (eds). 2005. Spiders of North America: an identification manual. American Arachnological Society. 377 pages. Chapter 2 PHYLOGENY AND CLASSIFICATION OF SPIDERS Jonathan A. Coddington ARACHNIDA eyes, jumping spiders also share many other anatomical, Spiders are one of the eleven orders of the class Arach- behavioral, ecological, and physiological features. Most nida, which also includes groups such as harvestmen (Opil- important for the field arachnologist they all jump, a useful iones), ticks and mites (Acari), scorpions (Scorpiones), false bit of knowledge if you are trying to catch one. Taxonomic scorpions (Pseudoscorpiones), windscorpions (Solifugae), prediction works in reverse as well: that spider bouncing and vinegaroons (Uropygi). All arachnid orders occur in about erratically in the bushes is almost surely a salticid. North America. Arachnida today comprises approximately Another reason that scientists choose to base classifica- 640 families, 9000 genera, and 93,000 described species, but tion on phylogeny is that evolutionary history (like all his- the current estimate is that untold hundreds of thousands tory) is unique: strictly speaking, it only happened once. of new mites, substantially fewer spiders, and several thou- That means there is only one true reconstruction of evolu- sand species in the remaining orders, are still undescribed tionary history and one true phylogeny: the existing clas- (Adis & Harvey 2000, reviewed in Coddington & Colwell sification is either correct, or it is not. In practice it can be 2001, Coddington et ol. 2004). Acari (ticks and mites) are complicated to reconstruct the true phylogeny of spiders by far the most diverse, Araneae (spiders) second, and the and to know whether any given reconstruction (or classifi- remaining taxa orders of magnitude less diverse. Discount- cation) is "true." Indeed, scientists generally regard "truth" ing secondarily freshwater and marine mites, and a few in this absolute sense as beyond their reach. Instead they semi-aquatic or intertidal forms, all extant arachnid taxa strive to make their hypotheses as simple as possible, and as are terrestrial. Arachnida evidently originated in a marine explanatory as possible. Simpler and more general hypoth- habitat (Dunlop & Selden 1998, Dunlop & Webster 1999), eses win. They win through comparison of predictions invaded land independently of other terrestrial arthropod made by the hypothesis to factual observation. Scientific groups such as myriapods, crustaceans, and hexapods hypotheses (e.g., explanations, classifications, taxonomies, (Labandeira 1999), and solved the problems of terres- phylogenies) are constantly tested by discovery of new traits trialization (desiccation, respiration, nitrogenous waste and new species. To the extent that the hypothesis is good, removal without loss of excess water, and reproduction) it accommodates and comfortably explains new data. If the in different ways. Although the phylogeny of Arachnida is new data do not fit the theoretical expectations, sooner or still controversial (Coddington etcd. 2004), specialists agree later a new hypothesis or a revised version of the old one that the closest relative of Araneae is a group of orders col- takes its place. In biological classification, and phylogeny lectively known as Pedipalpi: Amblypygi, Schizomida, and reconstruction in particular, scientists have developed a Uropygi (Shultz 1990). number of technical terms to describe the various ways that classifications or phylogenies do, or do not, correspond to PHYLOGENETIC THEORY AND METHOD fact (Fig. 2.1). Any group in a classification is said to be a Systematics is the study and classification of the differ- taxon (plural taxa) or clade, and in theory corresponds to ent kinds of organisms and the relationships among them. one common ancestral species and all of its descendants. Good classifications are predictive: knowing one feature Such clades are said to be monophyletic ("mono" = single, predicts many others. If one knows that an animal has and "phylum" = race). spinnerets on the end of the abdomen, it will also have In the preceding examples, spiders (Araneae), labido- fangs and poison glands (lost in a few spiders), eight legs, gnath spiders (now called Araneomorphae), sticky-silk two body regions, male palpi modified for sperm transfer, spinners (Araneoidea), jumping spiders (Salticidae) and and it will spin silk: in short, it is a spider. All spiders share wolf spiders (Lycosidae) are all thought to be monophyletic these features because they inherited them from a common groups, clades, and taxa. Each of these groups is distin- ancestor, but today's spiders have evolved to differ among guished by one or more uniquely evolved features or inno- themselves. For example, the earliest spiders had fangs that vations. Such characters are said to be "derived," because worked in parallel (orthognath, like tarantulas and their they are transformations of a more primitive trait. Orthog- allies), but later in spider evolution one lineage developed nath chelicerae is the original, primitive (plesiomorphic) fangs that worked in opposition (labidognath, like the condition for spiders, and labidognath chelicerae is the majority of spiders in North America). Much later within later, derived (apomorphic) condition. The only acceptable the labidognath lineage, some evolved the ability to coat silk evidence for monophyletic groups are shared, derived char- lines with a viscid, semi-liquid glue, useful for entrapping acters, or synapomorphies ("syn" = shared, "apomorphy" = and subduing prey. This nested pattern of branching lin- derived morphology) such as the evolution of viscid silk in eages (phylogeny), results from evolutionary descent with Araneoidea (Fig. 2.1). modification (Fig. 2.1). The vast majority of similarities Sometimes systematists (scientists who infer phylogeny and differences among species are due to phylogeny. Jump- and use the results to classify organisms) make mistakes ing spiders (Salticidae) all have huge anterior median eyes and group taxa based on primitive characters or plesio- because they are relatively closely related, and wolf spider morphies. Such groups, containing a common ancestor (Lycosidae) eyes exhibit their characteristic eye pattern for and some but not all of its descendants, are then termed the same reason. Phylogeny explains more biological pat- paraphyletic. In Figure 2.1, the grouping "Orthognatha" is tern than any other scientific theory (e.g., ecology, physiol- paraphyletic because it is based on a primitive character, ogy, ethology, etc.), and therefore classifications based on orthognath or paraxial chelicerae, and because it includes phylogeny will be maximally predictive. Besides huge front the common ancestor of all spiders but excludes some Spiders of North America • 19 descendants, i.e. the Labidognatha. Even worse, sometimes to the analysis (or even an a priori assumption) one taxon groups don't even include any common ancestor at all and in the analysis is specified to join at the root of the tree, are then termed polyphyletic ("Big Spiders," Lycosidae + and powerful computer algorithms are used to find the Mesothelae in Fig. 2.1 would be polyphyletic). Polyphyletic most plausible tree (or branching diagram, also termed a groups are usually based on convergent features and para- cladogram) that unites all taxa and best explains the data. phyletic groups on primitive features. Systematists adopt the initial null hypothesis that all simi- Classifications (and phylogenies) need not be strictly larities are due to phylogeny. The fit between the tree and binary or dichotomous: in Figure 2.1 the three-way fork the data decreases to the extent that one must suppose the uniting Araneoidea, Lycosidae, and Salticidae intentionally "same" trait arose two or more times independently (con- doesn't indicate which is most closely related to which. If vergent evolution) or was lost secondarily. An example of nodes are dichotomous, the two daughter lineages are often the former might be "big." Not all "big" spiders are each called sister taxa, or, informally, sisters. other's closest relatives (but some are). An example of the In practice systematists infer phylogeny by compiling latter is the absence of true abdominal segmentation in all large tables or matrices of taxa and their traits or fea- spiders. Spiders are arthropods and arthropods typically tures. Traits may be anything presumed to be genetically have segmented abdomens; spider relatives also have seg- determined and heritable, such as morphology, physiol- mented abdomens. Rather than suppose that all arthro- ogy, behavior, or, increasingly, DNA sequences. The ideal pods with segmented abdomens gained the condition approach would encapsulate all comparative knowledge independently, and thus that spiders reflect the ancestral about the group in question. Based on evidence external unsegmented arthropod, it becomes very much simpler (Monophyletic) Entelegynae "A £> ^ :# P # ^ & & ^ ^ <3 Viscid silk (Polyphyletic) O "Big Spiders" o Female fertilization ducts (Paraphyletic) "Orthognatha" Synapomorphy O Orthognath ^Z Cheliceral ^V Labidognath plesiomorphic OriGfltStiOn apomorphic Clades Spinnerets I Time Araneae Fig. 2.1 Taxa are in regular, characters in italic font. Only synapomorphies (shared, derived characters) are valid evidence of monophyletic groups (clades). Paraphyletic groups are usually based on plesiomorphies, polyphyletic groups