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Novttates PUBLISHED by the AMERICAN MUSEUM of NATURAL HISTORY CENTRAL PARK WEST at 79TH STREET NEW YORK, N.Y AMERICAN MUSEUM Novttates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET NEW YORK, N.Y. 10024 U.S.A. NUMBER 2627 JULY 7, 1977 NORMAN I. PLATNICK The Hypochiloid Spiders: A Cladistic Analysis, With Notes on the Atypoidea (Arachnida, Araneae) AMERICAN MUSEUM Novitates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 2627, pp. 1-23, figs. 1-31 July 7, 1977 The Hypochiloid Spiders: A Cladistic Analysis, With Notes on the Atypoidea (Arachnida, Araneae) NORMAN I. PLATNICK1 "The value of any classification rests on the soundness of the princi- ples underlying it." -A. Petrunkevitch, 1933, p. 303 "But I shall certainly admit a system as empirical or scientific only if it is capable of being tested by experience. These considerations suggest that not the verifiability but the falsifiability of a system is to be taken as a criterion of demarcation. In other words: . it must be possible for an empirical scientific system to be refuted by experience." -K. R. Popper, 1968, p. 40 ABSTRACT A hypothesis of interrelationships of the prim- ordinated classifications derived from this clado- itive araneomorph spiders based on shared de- gram are presented for purposes of comparison rived characters is presented. The genera Hy- and evaluation. The family Ectatostictidae pochilus and Ectatosticta are regarded as sister Lehtinen is newly synonymized with the Hypo- groups and as constituting the sister group of all chilidae. Serrula morphology suggests that the remaining araneomorphs. Three other genera superfamily Atypoidea is not monophyletic and (Hickmania, Gradungula, and Thaida) are sequen- that the Mecicobothriidae are more closely re- tially considered plesiomorphic sister groups of lated to the Dipluridae than to the Antrodiaeti- the remaining araneomorphs. Sequenced and sub- dae or Atypidae. INTRODUCTION The present paper represents an attempt to (1965, 1966), and (2) with the establishment of examine the higher classification of araneomorph a classification of these spiders that reflects the spiders from the viewpoint of phylogenetic sys- hypotheses of relationship thus generated. In tematics. As such, it is primarily concerned (1) addition, some data collected on mygalomorph with the placement of the "hypochiloids," the spiders for purposes of out-group comparison, most primitive of the true spiders, in monophy- and relevant to a phylogenetic analysis of that letic groups on the basis of shared derived charac- group, are also provided. ters with the methods developed by Hennig One opinion occasionally expressed is that the 'Associate Curator, Department of Entomology, the American Museum of Natural History. Copyright © The American Museum of Natural History 1977 ISSN 0003-0082 / Price $1.65 2 AMERICAN MUSEUM NOVITATES NO. 2627 higher classification of many or most groups rep- 31 represents a hypothesis that liphistiids (Meso- resents a consensus view based on consideration thelae) are more closely related to opisthothe- of many characters by many workers, and that lines than either of those groups are to amblypy- no substantial improvements can be made in gids. Brief inspection will show that phylogenies those classifications unless newly available or pre- are merely internested series of such three-taxon viously ignored characters are brought to bear on statements that proceed from greater to lesser them. If previous classifications of the hypo- levels of universality in terms of the number of chiloid spiders are representative, and there taxa they include. seems to be no reason to suspect that they are As stressed by Hennig, the only evidence we not, this argument is indefensible. Although can have that taxa A and B are each other's clos- some new observations are added below, this in- est relatives is that they share a uniquely derived formation only corroborates hypotheses for (synapomorphic) character state that they do not which sufficient indicatory data have been avail- also share with C. For any three-taxon statement able in the literature at least since the work of there are three possible explicit (dichotomous) Marples (1968). That a satisfactory hypothesis of hypotheses of relationship (A and B are closest hypochiloid interrelationships has not yet been relatives, A and C are closest relatives, or B and C presented seems to indicate that arachnologists are closest relatives) and one possible general (tri- have lacked a satisfactory method of analyzing chotomous) hypothesis of relationship (A, B, and their data, not that they have lacked a sufficient C are closest relatives, as compared with D). If all data base on which to reach a consensus. Cladis- we have available is a synapomorphic character tic analysis provides that sorely needed metho- state shared by A, B, and C, we must choose the dology. general hypothesis. If we also have available a Taking to heart the statement Petrunkevitch synapomorphic character state shared by only made when he began his famous study on the two of the three taxa, we can choose one of the internal anatomy of spiders, and which is quoted three explicit hypotheses, not because we have above, a brief discussion of the principles under- demonstrated it to be true, but because we have lying this study is in order. The point of depar- falsified the two alternate explicit hypotheses ture is the view expressed by Popper (1968) that (such falsifications are apparent and not abso- the line of demarcation between science and lute, of course; absolute falsification seems as non-science is falsifiability, that only a statement theoretically unobtainable as absolute truth). that can potentially be shown to be wrong by Given a set of purportedly synapomorphic char- some possible observation of the real world quali- acter states that do not have perfectly congruent fies as a scientific hypothesis. Thus if classifica- distributions among the taxa considered, par- tions are to be scientific they must be falsifiable simony dictates that we must always choose the hypotheses. Since the hypothesis involved in hypothesis that appears to have been falsified the grouping a set of taxa together is one of relation- least number of times. Thus if A and B share six ship (namely that those taxa so grouped are more purportedly synapomorphic character states, and closely related to each other than to taxa ex- A and C share two purportedly synapomorphic cluded from the group), the process of construct- character states, we must choose the hypothesis ing a scientific classification (as opposed to a that A and B are closest relatives, the implication classification designed only as a technique for being that the two incongruent character states information storage and retrieval) necessarily are not actually synapomorphic. They may have involves an attempt at reconstructing the inter- been derived more than once by parallelism, they relationships of the organisms concerned. If life may be primitive rather than derived, or they on earth had a common origin, all organisms are may indeed not be homologous in all the taxa related, and hypotheses of relationship must be considered. The crucial point is that such a deci- comparative. The most explicit possible compara- sion, once made, is always open to falsification tive hypothesis is a three-taxon statement: taxon by simply finding a larger number of purportedly A is more closely related to B than either of them synapomorphic character states in favor of one are to C. For example, the circled area of figure of the alternate hypotheses (Gaffney, MS). 1 977 PLATNICK: HYPOCHILOID SPIDERS 3 Polarity judgments, statements that one given there is no possible combination of shared de- character state, such as paraxial chelicerae, is rived characters that can falsify a hypothesized primitive (plesiomorphic) at a given level of uni- ancestor-descendant relationship (autapomorphic versality and that an alternate character state, character states found in a purported ancestor such as diaxial chelicerae, is derived at that level but not in a purported descendant could be the of universality, are thus themselves subsidiary result of character reversal within the lineage). hypotheses that can be falsified by incongruence Thus a cladogram uniting A and B may mean with an otherwise corroborated hypothesis of that they had a common ancestor (X) that under- relationship. Primitive character states shared by went a speciation event, or it may mean that A is two out of three taxa cannot be used to choose the ancestor of B, or that B is the ancestor of A. between alternate three-taxon statements be- A cladogram containing fossil taxa may include cause they do not falsify any of the alternate branching points that represent no speciation hypotheses, but all shared primitive character events at all, but only ancestor-descendant rela- states do represent synapomorphies at higher lev- tionships. Similarly, in a cladogram containing els of universality. Shared character states de- only Recent taxa a common ancestor may be rived separately in different lineages (i.e., paral- identical with one of its descendants (if, for ex- lelisms) cannot be used to choose between alter- ample, one parasitic species gives rise sympatri- nate three-taxon statements and do not represent cally to another by a host change without itself synapomorphies at higher levels of universality; undergoing any genetic change) or not (if one they can, however, be detected by incongruence species gives rise allopatrically to two or
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