Glomeris undulata Koch and G. conspersa Koch are conspecific. - Enzyme electrophoretic evidence and taxonomical consequences (Diplopoda: Glomeridae)

19 Pages·1999·5.9 MB·

by  R Hoess

Revue suisse deZoologie 106 (3): 643-661; septembre 1999
Glomeris undulata Koch and G. conspersa Koch are conspecifîc. -
Enzyme electrophoretic evidence and taxonomical consequences
(Diplopoda: Glomeridae)
René HOESS & AdolfSCHOLL
University ofBerne, Institute ofZoology, Division ofPopulation Biology, Baltzerstr.
3, CH-3012 Berne, Switzerland.
Glomeris undulata Koch and G. conspersa Koch are conspecific. -
Enzyme electrophoretic evidence and taxonomical consequences
(Diplopoda: Glomeridae. - Syntopic and allotopic populations of Glomeris
undulata and G. conspersa have been investigated with enzyme electropho-
resis. There was no evidence for separate gene pools of these taxa. However,
G. romana, a species supposed to be closely related to G. undulata and G.
conspersa, prooved to be well differentiated based on allozyme data. We
therefore conclude that the taxa undulata and conspersa are conspecific.
Consequently, G. conspersa C. L. Koch, 1847, is ajunior subjective synonym
of G. undulata C. L. Koch, 1844 (nov. syn.). An analysis ofthe colour pattern
provides arguments forthe discussion why no intermediate forms exist.
Key-words: Diplopoda - Glomeris - revision - local alleles - morphology -
distribution - new synonymy.
INTRODUCTION
Glomeris undulata C.L.Koch, 1844, and G. conspersa C.L.Koch, 1847, are
traditionally treated as two closely related but distinct species (cf. Verhoeff 1911).
They may be separated with reliability only by the form of the dorsal dark spots on
the segments 2 to 12. These spots are parallel-sided in G. undulata and convergent
posteriorly in G. conspersa (seen in walking position). Transitional forms that could
indicate interbreeding of these taxa are very rare and have been named G. undulata
var. pseudoconspersa (but see also Verhoeff (1928b) for a case of assumed
hybridisation). G. undulata and G. conspersa are often found syntopic, and large parts
oftheir ranges overlap. Within the scope of an analysis of the genetic differentiation
of the Central European Glomeris species by means of enzyme electrophoretic
methods, we also examined syntopic and allotopic populations of G. undulata and G.
conspersa. For comparison we included G. romana Verhoeff, 1900, a species which
is supposed to be closely related to G. undulata and G. conspersa (cf. Verhoeff
1911) but with a mainly allopatric distribution.
Manuscriptaccepted 12.01.1999
644 RENÉHOESS &ADOLFSCHOLL
MATERIAL AND METHODS
Population samples of Glomeris undulata, G. conspersa, and G. romana were
analyzed genetically by using routine enzyme electrophoretic methods of our labo-
ratory (Scholl et al. 1978). Vertical starch gel electrophoresis was conducted using
the same buffer systems as in previous studies on Glomeris (Hoess et al. 1997). 18
enzyme loci were analyzed. The enzymes investigated and the loci scored (in
brackets) are: aspartate aminotransferase (Aat-1, Aat-2), glyceraldehyd-3-phosphate
dehydrogenase (Gapdh), glucose-6-phosphate isomerase (Gpi), hexokinase (Hk),
leucine aminopeptidase (Lap), L-lactate dehydrogenase (Ldh-1, Ldh-2), malate
dehydrogenase (Mdh-1, Mdh-2), malic enzyme (Me), mannose-6-phosphate isome-
rase (Mpi), peptidase (Pep), 6-phosphogluconate dehydrogenase (Pgd6), phospho-
glucomutase (Pgm), superoxide dismutase (Sod-1, Sod-2) and sorbitol dehydrogenase
(Sodh).
The zymograms were photographed on Polaroid for reference. We refer to
observed electromorphs as alleles which are identified by their electrophoretic
mobility (in mm) relative to electromorphs ofpreviously studied species (Hoess etal.
1997). Mendelian inheritance of the alleles at a given locus was not tested by cross-
breeding experiments but was concluded by analogy to results of previous studies of
our laboratory (e.g. Zimmermann & Scholl 1993). Allele frequencies and genetic
distances were calculated with the BIOSYS-1 programme package (Swofford &
Selander 1989). Nei-distance D (Nei 1978) was used for the construction of an
UPGMA dendrogram (Sneath & Sokal 1973). Bootstrap analysis was conducted
with the PHYL1P programme package (Felsenstein 1986-95). The bootstrap values
are estimates from 100 replicates (bootstrap values levelled out at 100 replicates) with
UPGMA
as clusteralgorithm and an unrooted consensus tree.
The following population samples (Fig. 1) were examined (number of speci-
mens in brackets): Glomeris undulata: Switzerland: Bösingen (13), Brunnen (9),
Ingenbohler Berg (15), Loucherhorn (6), Malans (6), Meride (7), Merishausen (8),
Niderhorn (6), St-Gingolph (15); Germany: Ehingen (17); G. conspersa: Switzerland:
Biel (10), Bösingen (20), Castagnola (3), Châtollion (14), Densbüren (10), Ingen-
bohler Berg (11), Loucherhorn (8), Malans (12), Meride (8), Merishausen (13),
St-Gingolph (11), Valangin (10); Germany: Ehingen (20); France: Cruseilles (13);
G. romana: Italy: Camaiore (18), Lucca(4).
Several other population samples have been collected (often with low sample
sizes) and have been used in addition to literature data to produce a distribution map
ofthe taxa undulata and conspersa.
Tergites oftwo selected specimens ofG. undulata were drawn using a camera
lucida. The specimens were turned around theiraxes in orderto draw every part ofthe
tergite in vertical view.
RESULTS
Allele frequencies of the 18 loci of all samples are given in Table 1. For
unknown reasons, several specimens in most samples failed to show activity at some
REVISION OFGLOMER1SCONSPERSA 645
Fig. 1
Collecting sites in Switzerlandandadjacentcountries.
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