Contributions to Zoology, 69 (4) (2000)Alexandr P. Rasnitsyn: Testing cladograms by fossil record: the ghost range test

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Material

Fossil record of the hymenopterous insects (Order Vespida = Hymenoptera) is selected to test the method for the following reasons. Incomplete like any fossil history, the hymenopteran record is rich enough to represent as many as 51 out of 54 extant families (e.g., Fig. 1; superfamily Chalcidoidea is used here as a single taxon because there is no cladogram available for it yet). As many as 38 of these families are recorded starting from the Mesozoic, a time wherein many high-level phylogenetic events occurred, and 20 more families are known to be extinct (mostly since the Mesozoic). A wealth of cladograms have been proposed recently for the family level phylogeny of the order, and this variety allows us to select enough of them to test. The examples are displayed in Figs. 1-6.

FIG2

Fig. 1. Relation and duration of the hymenopteran families (superfamily for Chalcidoidea) (modified from Rasnitsyn, 1988). Thick lines show known longevity, horizontal thin lines – relationship, vertical thin lines – ghost ranges, double thin lines – long ghost ranges (more than two geochronological epochs before entering fossil record). Periods: T – Triassic, J – Jurassic, K – Cretaceous, P – Paleogene, N – Neogene, R – contemporary (Holocene). Epochs are shown by subscript indices: 1 – Lower, 2 – Middle, 3 – Upper (in the Cenozoic, P1 – Paleocene, P2 – Eocene, P3 – Oligocene, N1 – Miocene, N2 – Pliocene). Known ranges are modified for some taxa comparing Rasnitsyn (1988) basing on following sources. Argidae and Sapygidae are found in the Upper Eocene Baltic amber (old data by Brischke 1886, ignored by Rasnitsyn 1988), Megalodontesidae in the Lower Cretaceous of China (Ren et al., 1995; my identification of Jibaissodes Ren, Guo et Ji as a genus of Megalodontesidae), Evaniidae in the Burmese amber of disputably Upper Cretaceous amber (Bashibuyuk et al., 2000), Monomachidae in the Lower Cretaceous of Koonwarra, Australia (Jell and Duncan 1986: fig. 66 F; my identification), Scelionidae and Chalcidoidea (new family) in the lowermost Cretaceous of 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 of Alava, Spain (my identification), Diapriidae and Bethylonymidae in the Lower Cretaceous of England (Rasnitsyn et al., 1998), Sclerogibbidae in the Miocene Dominican amber (my identification), Embolemidae in the Lower Cretaceous of Transbaicalia and Mongolia (Rasnitsyn 1996b), 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 of Brazil (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 superfamily (Karatavitidae and Ephialtitidae, Serphitidae and the rest of Platygastroidea, Diapriidae and other Proctotrupoidea) appear separated in the cladogram to simplify it visually. These “orphan” families are marked by a broken arrow.

Fig. 1 is based on Rasnitsyn (1988), with several minor additions and modifications explained in the caption. Fig. 2 shows the results of the recent parsimony re-consideration of the concept by Rasnitsyn (1988) performed by Ronquist, Rasnitsyn, Roy, Eriksson and Lindgren (1999). Figs. 3-5 are re-drawn from the cladograms in Vilhelmsen (1997: fig. 2), Whitfield (1992: fig. 5), and Brothers and Carpenter (1993: fig. 11), respectively. Fig. 6 represents the apocritan molecular phylogeny performed by Dowton, Austin, Dillon and Bartowsky (1997: fig. 2), with the symphytan families, Orussidae and Stephanidae added after the work of Dowton and Austin (1994). All dendrograms are unified in their style, with the known duration of taxa being indicated by thick lines, relationships by horizontal thin lines, and the implied ghost ranges (see below) by vertical thin lines; the vertical thin line connecting two horizontal ones means nothing more than a connection of two parts of a horizontal line.

FIG2

Fig. 2. Relation of the hymenopteran taxa as calculated by Ronquist et al. (1999: figs. 2, 5 and 9, combined), formatted after Fig. 1. Dashed double line – long ghost ranges additional to those seen in Fig 1