Contributions to Zoology, 69 (4) 251-258 (2000) SPB Academic Publishing bv, The Hague fossil record: the test Testing cladograms by ghost range Alexandr+P. Rasnitsyn Paleontological Institute, Russian Academy ofSciences, Profsoyuznaya Street 123, 117647 Moscow, Russia words: fossil Key phylogeny, cladogram, ghost range, record, Vespida, Hymenoptera Abstract The fossil record is a source of the evolutionary information that is not strictly dependent on our A method ofthe calculation is ghost range proposed to assess evolutionary hypotheses, and so in principle it may the between and the fossil congruence a cladogram record and be used as an external standard in the comparison this basis. The method is tested to compare cladograms on on a of Of the fossil record is im- cladograms. course, set of cladograms developed recently to reveal the phylogeny perfect, but so are all our sources of evidence. This ofthe hymenopterous insects (Order Vespida), and the results is true for cladistics as well, for the character are discussed. set is never exhaustive, and transformation series com- position and polarity are always hypothetical. That Contents is why both sources of the evolutionary informa- the fossil tion, record and the cladograms, possess of Introduction 251 only degrees ‘perfection’, but even so are wor- Material 251 thy of comparison in testing phylogenies. Methods 252 There are methods proposed to assess quality Results and discussion 254 of the fossil how it record based on agrees with Conclusion 257 the cladogram of taxa Acknowledgements 258 respective (reviewed by References 258 Benton and Hitchin 1997). Unfortunately, these metrics do not fit the reverse task, that is, to test for their different cladograms of a particular taxon Introduction extent of congruence with the fossil record. To- ward this end, 1 am proposing here a similar but research In It is an ordinary condition in cladistic that slightly different approach. its present form the method is rather but whenever found cladograms developed from different data for the rough, ap- it be for same set of taxa often differ significantly from each propriate, may easily improved more pre- other. This is crucial since the cladogram compari- cise calculation. the son is aimed at selecting most correct one. There are a number of indices that have been developed to estimate statistical features that reflect devia- Material tion of the cladogram from the most parsimonious one (Quicke, 1996: 3.2.5). It is not so evident, Fossil record of the hymenopterous insects (Order that the does = is selected however, evolutionary process strictly Vespida Hymenoptera) to test the observe the parsimony principle. That is why1 it method for the following reasons. Incomplete like seems reasonable to seek a method of assessment any fossil history, the hymenopteran record is rich 54 less dependent on particular evolutionary concepts. enough to represent as many as 51 out of ex- Downloaded from Brill.com10/01/2021 06:30:10AM via free access 252 A.P. Rasnitsyn - Testing cladograms by fossil record tant families (e.g.. Fig. 1; superfamily Chalcidoidea ing more than a connection oftwo parts of a hori- is used here as a single taxon because there is no zontal line. available it of cladogram for yet). As many as 38 these families are recorded starting from the Me- sozoic, a time wherein many high-level phyloge- Methods netic events occurred, and 20 more families are known be extinct since the When to (mostly Mesozoic). we convert a cladogram into a phylogenetic A been wealth of cladograms have proposed re- tree, we fit the intemode length to the paleontologi- the cently for family level phylogeny of the order, cally confirmed minimum duration of a particular allows select of them have that and this variety us to enough subclade. In some cases we to infer a The in 1-6. to are clade earlier than the fossil record would test. examples displayed Figs. appears 1 in Fig. is based on Rasnitsyn (1988), with several suggest (vertical thin lines Figs. 1-6). These minor additions and modifications in explained inferred cryptic intervals of existence were termed the 2 shows the results of the caption. Fig. recent ‘ghost ranges’ (Norell, 1992). When more than one re-consideration of the Ras- parsimony concept by cladogram is developed for a particular set of taxa, nitsyn (1988) performed by Ronquist, Rasnitsyn, they usually differ in their configuration as well Eriksson and 3-5 in the extension of Roy, Lindgren (1999). Figs. as ghost ranges displayed by are re-drawn from the cladograms in Vilhelmsen the respective trees. It is apparent that the smaller (1997: fig. 2), Whitfield (1992: fig. 5), and Brothers these extensions, the better the cladogram fits the and 6 Carpenter (1993: fig. 11), respectively. Fig. fossil record, i. e., the more parsimonious it is in molecular represents the apocritan phylogeny that respect due to a lower degree of inference for Dillon and Bartow- performed by Dowton, Austin, the duration of taxa. This is a reason to try and use with the families, the for and evalu- sky (1997: fig. 2), symphytan ghost range concept comparison Orussidae and Stephanidae added after the work ation of competing cladograms. of Dowton and Austin (1994). All dendrograms Thus we might suggest calculating the ghost are unified in their with the known duration in million either their dura- style, ranges years, as total of indicated thick taxa being by lines, relationships tion as expressed on the cladogram, or their aver- horizontal thin and the by lines, implied ghost ranges age duration per node per terminal branch. How- vertical thin lines; the vertical thin (see below) by ever, the absolute duration of the geochronologi- line horizontal noth- cal units connecting two ones means is still an extensively debatable issue, full Fig. I. Relation and duration ofthe hymenopteran families (superfamily for Chalcidoidea) (modified from Rasnitsyn, 1988). Thick - horizontal thin vertical thin lines - double thin lines - lines show known longevity, lines relationship, ghost ranges, long ghost - than before fossil Periods: T J - K - ranges (more two geochronological epochs Ipntering record). Triassic, Jurassic, Cretaceous, P - - Paleogene, N Neogene,R - contemporary(Holocene). Epochs are shown by subscript indices: , - Lower, - Middle, - Upper (in - - - - P P N - Known for the Cenozoic, : Paleocene, P, Eocene, Oligocene, ( Miocene,N, Pliocene). ranges are modified some taxa comparing Rasnitsyn (1988) basing on following sources, Argidae and Sapygidae are found in the Upper Eocene Baltic amber (old data Brischke 1886, in the Lower Cretaceous of China et ah, by ignored by Rasnitsyn 1988), Megalodontesidae (Ren 1995; my Jibaissodes Guo Ji of Evaniidae in the Burmese amber of identificationof Ren, et as a genus Megalodontesidae), disputably Upper Cretaceous amber (Bashibuyuk et al., 2000), Monomachidae in the Lower Cretaceous of Koonwarra, Australia (Jell and Duncan 1986: 66 Scelionidae and in the Cretaceous of fig. F; my identification), Chalcidoidea(new family) lowermost Mongolia (my identification), Platygastridae in the Upper Cretaceous amber of New Jersey (identified by L. Masner, personal communication by D.A. Grimaldi), Mymarommatidae in the Lower Cretaceous (Aptian) amber ofAlava, Spain (my identification), Diapriidae and Bethylonymidae in the Lower Cretaceous of England (Rasnitsyn et ah, 1998), Sclerogibbidae in the Miocene Dominican amber (my identification), Embolemidae in the Lower Cretaceous ofTransbaicalia and Mongolia (Rasnitsyn 1995b), Scolebythidae in the Upper Eocene Baltic amber (Brothers and Janzen, 1999), Plumariidae in the Upper Cretaceous New Jersey amber (my identification), Rhopalosomatidae and Tiphiidae in the Lower Cretaceous ofBrazil (Darling and Sharkey 1990), Sierolomorphidae in the Upper Cretaceous amber of New Jersey and Falsiformicidae in the Lower Cretaceous Lebanese amber (my identification), Formicidae in the Lower Cretaceous of East Siberia (Dlussky 1999). Superfamilies are outlined by boundary lines, except that sometimes members of one and the same and and the of and other superfamily (Karatavitidae Ephialtitidae, Serphitidae rest Platygastroidea, Diapriidae Proctotrupoidea) appear families separated in the cladogram to simplify it visually. These “orphan” are marked by a broken arrow. Downloaded from Brill.com10/01/2021 06:30:10AM via free access Contributions to Zoology, 69 (4) - 2000 253 Downloaded from Brill.com10/01/2021 06:30:10AM via free access 254 A.P. Rasnitsyn - Testing cladograms by fossil record and for To avoid Mai- of gaps traps a non-expert user. Stephanidae, Ichneumonoidea, Trigonalidae, these traps, the calculation should be preceded by metshidae, Ceraphronidae, and Apidae) above those in Vilhelmsen’s special efforts to evaluate the absolute time ranges seen Fig. 1. cladogram (Fig. 3) of the involved taxa. However, this sophistication considers only the lower hymenopterans (13 ter- additional of approach does not seem necessary now, when minal taxa) and yields two long ghost and Whitfield’s the matter of issue is a testing of the method itself ranges (Cimbicidae Cephidae). rather than fine tuning a particular phylogeny. cladogram deals mostly with superfamilies; his tree Indeed, if the method is found workable in its sim- (Fig. 4) also includes 13 terminal taxa and six long in 1. plest form, it will be the more so even after rea- ghost ranges, all additional to those seen Fig. sonable complication. The cladogram from Brothers and Carpenter (Fig. is the of aculeate It a tree The simple metrics applied here sum 5) concerns only wasps. produces This with 19 terminal taxa and four the ‘long ghost ranges’ per cladogram. means long ghost ranges, counting the cases when a terminal taxon or a all being the same as in Fig. 1 (the long ghost ranges subclade is inferred to originate prior to its first of Pompilidae and Sapygidae are longer here than time in while in the case of the fossil appearance for more than a particular Fig. 1, Bradynobaenidae unit. For the present, to satisfy the definition of the opposite may be true).