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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,

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 (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-

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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 ). 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 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). , , , 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, and 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), (Ren 1995; my

Jibaissodes Guo Ji of in the 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 () amber ofAlava, Spain (my identification), and Bethylonymidae in the Lower Cretaceous of England (Rasnitsyn et ah, 1998), 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), and in the Lower Cretaceous ofBrazil (Darling and Sharkey 1990), 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 , Diapriidae ) appear

families separated in the cladogram to simplify it visually. These “orphan” are marked by a broken arrow.

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and for To avoid Mai- of gaps traps a non-expert user. Stephanidae, , , these traps, the calculation should be preceded by metshidae, , and ) 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 ( ). 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 . 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, unit. For the present, to satisfy the definition of the opposite may be true). The tree based on the

the time interval should molecular from Dowton with the “long” ghost range, composite cladograms considers 24 terminal and be equal to, or longer than, two geochronological co-authors (Fig. 6) taxa

in of which epochs (two steps of the time grid as displayed produces eight long ghost ranges, seven

shown there do not those in Figs. 1-6). Thus, all ghost ranges are repeat Fig. 1.

The above results indicate that the by thin vertical lines, while long ghost ranges are cladograms represented by double thin lines. The dashed double by Rasnitsyn (1988) and Brothers and Carpenter

record lines that in 2-6 indicate the are more with the fossil appears figs long ghost (1993) congruent

1. than the others. This follows from the observation ranges absent in Fig.

that these two cladograms (each within its own

of scope) infer the minimum number ghost ranges

Results and discussion when corresponding sections of the considered

cladograms are compared. The results deserve some

additional however. Figs. 1-6 show considerable differences between discussion,

The is not the cladograms in regards to the occurrence ofthe high score gained by my cladogram

of long ghost ranges. Fig. 1 depicts the tree with 76 peculiar. My cladogram acknowledges presence

that ancestral char- terminal taxa and eight long ghost ranges taxa (those lacking autapomorphic

acter The other follow the strict concern three families lacking fossil record (Pergi- states). cladograms

that dae, and Bradynobaenidae), five cladistic concept ancestors can never be iden- others (, clade Electrotomidae + tified as such. Therefore, only sister group rela-

Argidae + , Sclerogibbidae, Pompilidae, tionships are available for these cladograms. Unlike sister and Sapygidae) that enter the fossil record in the mother-and-daughter taxa, the groups always

So inclusions in the Baltic have when we take a for Late Eocene (e. g., as amber) equal age. daughter

later. These individual be found in a sister, we infer it to have an earlier and or cases may origin all other trees tested, whenever they consider these therefore a longer ghost range. I have discussed at taxa. Furthermore, I shall only discuss the “long” length elsewhere the reasons why it is necessary ghost ranges that appear in addition to these eight. to acknowledge phylogenetic ancestors (Rasnitsyn,

al. 1996). In fact, in 2), The cladogram from Ronquist et (Fig. 2) pro- Ronquist’s cladogram (Fig. four of duces a tree with 75 terminal taxa and eight addi- long ghost ranges (cases Cimbicidae, Mega-

and tional long ghost ranges (Cimbicidae, Argidae, lodontesidae, Ceraphronidae, Apidae) appear

formatted after Fig. 2. Relation ofthe hymenopteran taxa as calculated by Ronquist et al. (1999: figs. 2, 5 and 9, combined), Fig. I.

Dashed double line additional those in - long ghost ranges to seen Fig. 1.

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Fig. 3. Relation ofthe symphytan taxa as calculated by Vilhelmsen (1997: fig. 2), formatted after Figs. 1, 2

Fig. 4. Relation of the parasitic taxa as presented by Whitfield (1992: fig. 5), formatted after Figs. 1, 2.

Fig. 5. Relation of the aculeate taxa as calculated by Brothers and Carpenter (1993: fig. 11), formatted after Figs. 1,2,

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for Fig. 6. Molecular phylogeny of hymenopteran taxa as calculated by Dowton and Austin (1994; fig. 3), symphytan families,

Orussidae and Stephanidae, and by Dowton et al. (1997: fig. 2) for the apocritans, formatted after Figs. 1, 2.

solely because they are considered to be sister and Conclusion

of Praesirici- not daughter groups ,

dae, , and , respectively. The The above observations indicate that the total

different same is true for three cases (Cimbicidae, Megalo- amount of ghost ranges displayed by

dontesidae and Cephidae) in Vilhelmsen’s cla- phylogenetic trees for the same set of taxa does

is dogram (Fig. 3); in my cladogram Cephidae the allow us to compare and evaluate the cladograms

daughter taxon of Scpulcidae. Of six long ghost forming the background of these trees. Moreover,

ranges in Whitfield’s tree (Fig. 4), five are absent the results ofapplication of the method are under-

in because and Cera- least in my cladogram Trigonalidae standable and possible to explain, at part.

fits better phronoidea (s.str.) are suggested there to descend That the cladogram from Rasnitsyn (1988)

directly from , Platygastroidea + Chal- to the fossil record than the majority of others,

- of cidoidea and from the ancestral depends at least partially on the appreciation additional the proctotrupoid family Mesoserphidae, and Ichneu- mother and daughter groups in to

monoidea from these latter the + Ephialtitidae. In Dowton sister groups, being only legitimate

there in of the and Austin’s cladogram (Fig. 6), are seven ones cladistics. At the same time, testing

1. Brothers and indi- additional long ghost ranges comparing to Fig. cladogram by Carpenter (1993)

Four of them result from the same causes: in my cates that the high score received by Rasnitsyn’s

cladogram, Tenthredinoideaoriginate directly from (1988) cladogram from the present test is not a

Xyclidae, Cephidae from Sepuicidae, Trigonalidae straightforward result of the method used there,

from Megalyridae, and Evaniidae + i.e., the making of a cladogram by the method of

from Praeaulacidae. Flennig (“paper and pencil” approach) instead of

and The reasons why other long ghost ranges ap- the automatic parsimony calculation, an ap-

and in addition those and descendant pear in Figs. 2, 3, 4 6 to seen preciation of the ancestor taxa.

in the not evident. are Several to further of the Fig. 1, are so Possibly they ways development ghost

relevant the of rather method be The most evident to phylogeny Hymenoptera range may proposed.

than and so to methodology of phylogenetics, they one is the calculation of the length rather than the

will be considered elsewhere. this would simple count of ghost ranges: make the

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Molecular of the the Procto- method more precise. However, there is yet another phylogeny apocritan wasps: and Ent. 22: 245-255. method trupomorpha Evaniomorpha. Syst. possibility. As proposed, the ghost range Jell PA, Duncan PM. 1986. Invertebrates, mainly insects, is designed to test competing cladograms in regards from the freshwater, Lower Cretaceous, Koonwarra Fos- their with the fossil record. to congruence Instead, sil Bed (Korumburra Group), South Gippsland, Victoria. it might be found useful to use the geochronologi- Mem. Ass. Australas. Palaeontologs 3: 111-205. cal data directly in the parsimony calculation, to Norrel MA. 1992. Taxic origin and temporal diversity: the

effect of In: Novacek MJ, Wheeler QD, eds. include the information from fossil record into the phylogeny. Extinction and phylogeny. New York: Columbia Univ. cladistic procedure. However, even it its simplest Press: 89-118. form, the ghost range method might be a useful Quicke D. 1996. Principles and techniques of contemporary test for cladograms. London etc.: Blackie Academic and Professional. .

Ren D, Lu L, Guo Z, Ji Sh. 1995. Faunae and stratigraphy

Jurassic-Cretaceous and the of in Beijing adjacent areas.

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