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M. Korn et al. • Sister species within Triops cancriformis


            Table 5 Splits of divergence of selected clades, subclades or haplotype groups of Triops cancriformis, estimated from applying the slowest and
            the fastest (Schubart et al. 2000) inferred molecular clock for Crustacea to pairwise 16S rDNA genetic distances.

                                Myr BP at 0.65% per Myr                Myr BP at 0.9% per Myr
                                p-distance         ML-distance         p-distance          ML-distance
                                mean   min   max   mean    min   max   mean   min    max   mean   min   max
            T.c.c. to T.c.m. + T.c.s.  4.5  3.2  7.2  5.1  3.5   8.9   3.2    2.3    5.2   3.7    2.6   6.4
            within T.c.m. + T.c.s.:
             ‘S. Spain’ to ‘Gitanilla’  3.8  3.6  4.0  4.1  3.9  4.5   2.7    2.6    2.9   3.0    2.8   3.2
             ‘S. Spain’ to ‘Portugal’  2.5  2.5  2.6  2.7  2.7   2.8   1.8    1.8    1.9   1.9    1.9   2.0
             ‘S. Spain’ to T.c.s.  2.2  2.2  2.2   2.2     2.2   2.3   1.6    1.6    1.6   1.6    1.6   1.7
             ‘S. Spain’ to T.c.m. Morocco  1.9  1.4  2.6  2.0  1.5  2.7  1.4  1.0    1.9   1.4    1.1   1.9
             T.c.s. to T.c.m. Morocco  2.4  1.4  3.2  2.5  1.5   3.5   1.7    1.0    2.3   1.8    1.1   2.5
             ‘Gitanilla’ to ‘Portugal’  4.1  3.9  4.3  4.7  4.5  4.9   3.0    2.9    3.1   3.4    3.2   3.5
            within T.c.c.:
             ‘C. Eur.’ to ‘Russia’  0.7  0.4  1.1  0.7     0.4   1.1   0.5    0.3    0.8   0.5    0.3   0.8
             ‘C. Eur.’ to ‘Austria’  0.4  0.0  1.1  0.4    0.0   1.1   0.3    0.0    0.8   0.3    0.0   0.8
             ‘C. Eur.’ to ‘Sicily’  0.4  0.4  1.1  0.4     0.4   1.1   0.3    0.3    0.8   0.3    0.3   0.8
             ‘C. Eur.’ to ‘Hungary’  0.4  0.4  0.7  0.4    0.4   0.7   0.3    0.3    0.5   0.3    0.3   0.5


            Table 4. Within Triops the lowest value of divergence between  Tunisia (#101, 102) and Italy (#34; Fig. 4A). For purposes of
            recognized species was observed between  T.  longicaudatus  morphological and taxonomic comparison, however, the
            (New World and Japan) and Triops australiensis (Australia):  former are assigned to a different haplotype group (‘Northern
            6.1% p-distance (12S; Table 4c). For comparison, the p-distance  Spain’) than the latter (‘Central European’). Nevertheless,
            between the two lineages found within T. cancriformis (Fig. 4),  they are not a separate lineage and the group consisting of
            i.e. T. c. cancriformis and T. c. mauritanicus + T. c. simplex, is  ‘T. c. simplex’ from Girona (northern Spain) and T. c. cancriformis
            4.1% (12S; Table 4c). Within Lepidurus the lowest values of  is referred to below simply as the T. c. cancriformis clade.
            divergence (p-distance) are observed between L. a. apus and  Genetic data also demonstrate that the Japanese specimen
            Lepidurus arcticus: 2.8% (16S; Table 4b) and 4.6% (12S;  belongs to the T. c. cancriformis lineage (Fig. 4, labelled T.c.
            Table 4d). These values are in the range of the 2.9% (16S;  AB084514). Certain samples from Sicily (but not all from this
            Table 4a) and 4.1% (12S; Table 4c) observed among the two  island), Ustica and Tunisia (but not all from this country)
            lineages within T. cancriformis.                 form a separate subclade, which is named the ‘Sicilian’ haplo-
             The estimates of divergence times of selected clades within  type group. Similarly, the samples from different localities
            T. cancriformis are presented in Table 5. Splits within the  in Hungary cluster together, illustrating the existence of the
            T. c. cancriformis lineage are clearly more recent than those  haplotype group ‘Hungary’.
            within the lineage of  T. c.  mauritanicus  plus  T. c.  simplex.
            Although the two haplotype groups ‘Portugal’ and ‘Gitanilla’  The Triops cancriformis simplex group
            form a monophyletic clade in the reconstructions with suffi-  Triops cancriformis simplex consist of two genetically very
            cient resolution, the split among them is nearly as high as  distinct lineages, one forming part of the T. c. cancriformis
            between the two main lineages within T. cancriformis.  lineage (see above), the other grouping within T. c. mauritanicus.
                                                             Thus, genetic data indicate that a separate T. c. simplex lineage
            The Triops cancriformis cancriformis lineage     as indicated by morphological similarities (see below) and
            The Russian samples branch off first within this lineage,  most recent classification as a separate subspecies (with
            illustrating the ‘Russian’ haplotype group. Next to branch  populations occurring in northern Africa as well as in Iberia)
            off is the ‘Austrian’ haplotype group, including the sample  does not exist.
            from the United Arab Emirates (and one sample from Italy,
            interpreted as an artefactual cluster formation, details in  The Triops cancriformis mauritanicus lineage
            Discussion section). The sequences of Triops c. simplex from  Triops cancriformis mauritanicus is paraphyletic in both the MP
            Girona (northern Spain) genetically belong to the same  and ML reconstructions as it includes those North African
            haplotype group as those of T. c. cancriformis samples from  samples assigned to T. c. simplex (Fig. 4). The T. c. mauritanicus
            Malta, Serbia, Germany as well as certain samples from  samples form three separate monophyletic clusters (Fig. 4A):


            © 2006 The Authors. Journal compilation © 2006 The Norwegian Academy of Science and Letters • Zoologica Scripta, 35, 4, July 2006, pp301–322  309
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