Tracking the trends of karyotype differentiation in the phylogenetic context of Gint, a scorpion genus endemic to the Horn of Africa (Scorpiones: Buthidae)
Author
Just, Pavel
Author
Šťáhlavský, František
Author
Kovařík, František
Author
Štundlová, Jana
text
Zoological Journal of the Linnean Society
2022
2022-10-01
196
2
885
901
https://academic.oup.com/zoolinnean/article/196/2/885/6632614
journal article
164206
10.1093/zoolinnean/zlac049
e68e772f-ce7f-4254-a5c4-57779e7f4d28
0024-4082
7184492
GINT
GUBANENSIS
Karyotype of
G. gubanensis
comprises 2
n
= 45 chromosomes, which decrease in length from 2.97% to 1.02% of DSL (Supporting Information,
Fig. S2G
; Table S2). Overall, 21 bivalents and a single trivalent, composed of chromosomes 7, 33 and 45, are detected in post-pachytene nuclei (
Fig. 3I
; Supporting Information,
Fig. S2G
). One pair of 18S rDNA loci is found on a terminal part of the heteromorphic chromosome pair (chromosomes 26 and 44). A single large 18S rDNA cluster is located on the overhanging part of chromosome 26 (the larger chromosome of the heteromorphic pair) (
Fig. 3J
; Supporting Information,
Fig. S2G
).
MTDNA DIVERSITY AND PHYLOGENY
The sequences for both 16S and
COI
were obtained from 22
Gint
individuals. In
G. calviceps
, only the 16S gene fragment was successfully sequenced. The final alignment with a total length of 963 bp (16S: 358 bp;
COI
: 605 bp) consisted of 765 conserved sites, 194 variable sites and 172 parsimony-informative sites.
Based on
COI
data,
Gint
exhibits genetic variation at both inter- and intraspecific levels, with the pairwise genetic distances ranging from 3.28% to 11.52% between species, and from 0.00% to 2.64% within species (Supporting Information, Tables S3, S4). The haplotype network reconstruction showed a presence of 15
COI
sequence haplotypes in
Gint
(Supporting Information, (
Fig. S3
; Table S3). Within the species studied,
G. banfasae
shows the highest within-location haplotype diversity (four haplotypes per locality), but with minor pairwise genetic differences (
p
-distance: ≤0.66%). In this species, all individuals of the same cytotype have distinct haplotypes. The haplotypes of cytotype I individuals (S1531 and S1534) are separated from one another by four mutational steps. Cytotype II (S1532) shares its haplotype with the cytotype III individual (S1530), which is one mutational step distant from the haplotype of the cytotype III specimen (S1533) (Supporting Information,
Fig. S3
).
Gint dabakalo
, comprising three haplotypes, shows the highest intraspecific genetic variation with a
p
-distance of 0.17–2.64%. In this species, the genetic distance between individuals of cytotype I and cytotype II from the same locality is higher than between each of them and a specimen of cytotype III from a distinct locality (Supporting Information,
Fig. S3
; Table S1).
Gint maidensis
, a species with a stable karyotype 2
n
= 34, is represented by three haplotypes with a
p
-distance of 0.33–0.50%.
Gintgaitako
presentstwohaplotypes, where one haplogroup, formed by
five specimens
, is distant from the other by one mutational step (
p
-distance: 0.17). In
G. amoudensis
, individuals of cytotype II, cytotype III and cytotype IV share the same haplotype, whereas the specimen of cytotype I differs from this haplogroup by eight point mutations (
p
-distance: 1.49%).
Both BI and ML mtDNA analyses provide similar tree topologies and nodal supports for clades. Phylogenetic reconstruction depicts
Gint
as a monophyletic group (
Fig. 5
). Most of the
Gint
species
form well-supported clades [bootstrap support (bs) ≥ 83, posterior probability (PP) ≥ 0.98]. The monophyly of the
G. banfasae
specimens remains unresolved due to low support values.
Gint maidensis
forms an early-branching clade, which is sister to the clade comprising all of the remaining
Gint
species.
This clade is further subdivided into two groups: (1) a clade in which
G. gubanensis
is a sister-species to
G. amoudensis
,
and (2) a clade consisting of
G. gaitako
,
G. calviceps
,
G. dabakalo
and
G. banfasae
. In the latter,
G. gaitako
forms a sister-lineage to the clade comprising
G. calviceps
,
G. dabakalo
and
G. banfasae
. The mutual relationships between the species in this group remain unresolved due to low support values.
THE RECONSTRUCTION OF ANCESTRAL CHROMOSOME NUMBER AND POSITION OF 18S RDNA GENE CLUSTERS
Maximum parsimony analysis inferred 2
n
= 32–34 as the ancestral chromosome number for the genus
Gint
(
Fig. 6A
). Two independent increases of 2
n
were detected within
Gint
: (1) the clade containing all cytotypes of
G. amoudensis
and
G. gubanensis
,
and (2) cytotype III within the
G. dabakalo
clade. When compared with the ancestral 2
n
, a tendency of decreasing of 2
n
from 30 to 18 chromosomes can be observed in
G. gaitako
,
G. dabakalo
and
G. banfasae
. Concerning the position of 18S rRNA gene clusters, maximum parsimony inferred their terminal position as the ancestral state for the genus
Gint
(
Fig. 6B
). Two independent changes of the ancestral terminal position to a subterminal position were identified in
G. amoudensis
and in the clade containing
G. dabakalo
and
G. banfasae
.