Review of the genera Berlesia Canestrini, 1884, and Katydiseius Fain & Lukoschus 1983, the subfamily Katydiseiinae Fain & Lukoschus, 1983, and their family group relationships (Acari: Mesostigmata: Gamasina), with description of three new species parasitic on gryllacridid crickets (Orthoptera)
Author
Lindquist, Evert E.
Author
Oconnor, Barry M.
0000-0003-4588-0788
Department of Ecology and Evolutionary Biology, Museum of Zoology, University of Michigan, Ann Arbor, Michigan, 48108, USA. bmoc @ umich. edu, https: // orcid. org / 0000 - 0003 - 4588 - 0788
Author
Shaw, Matthew D.
0000-0002-5037-3740
Terrestrial Invertebrates, South Australian Museum, North Tce, Adelaide 5000, South Australia, Australia matthew. shaw @ samuseum. sa. gov. au, https: // orcid. org / 0000 - 0002 - 5037 - 3740
Author
Sidorchuk, Ekaterina A.
Arthropoda Laboratory, Paleontological Institute, Russian Academy of Sciences, Profsoyuznaya ulitsa 123, 117997 Moscow, Russia. International Complex Research Laboratory for the Study of Climate Change, Land Usage, and Biodiversity, Tyumen State University, 10 Semakova Str., 625003, Tyumen, Russia.
text
Zootaxa
2020
2020-09-28
4857
1
5
70
journal article
9059
10.11646/zootaxa.4857.1.4
c4a1490d-15c3-446f-8676-e6c80096bcb6
1175-5326
4396474
F0AF75AD-BAE2-4B7D-9CCB-3D9477F350BD
Review of
Iphiopsididae
as a definable family-group
The genus
Berlesia
was first placed in the controversial family
Iphiopsidinae
(now
Iphiopsididae
) by
Vitzthum (1943)
where it has remained. This placement is unsatisfactory for several reasons.
Firstly the
Iphiopsididae
itself needs examination.
Kramer (1886)
mentioned the new family as
Iphiopsidae
(sic), just once, after stating that the available gamasid families were not sufficient to encompass known species, and thereby granting family status to
Iphiopsis mirabilis
(
Berlese, 1882
)
.
Kramer (1886)
did not specifically include any other genera in his
Iphiopsidae
, and his proposal of a family for
Iphiopsis
was overlooked until noted by
Vitzthum (1943)
.
Berlese (1892
, p.16) provided a diagram of relationships between genera (Genera Mesostigmatum Secundum Ordinem Naturalem Disposita) which clearly showed that
Berlesia
(along with the ant-associated
Neoberlesia
Berlese, 1892
) was considered to be distinctly separated from
Iphiopsis, Iphis
and genera considered similar at that time. The associated key (p. 37) did not diagnose
Berlesia
,
as the key was explictly restricted to European taxa. However the diagnostic information in that key, separating
Iphiopsis
based on loss of peritreme, re-appears later where it is then used in a much broader geographic context (
Vitzthum 1943
), and thereafter for many more genera (
Evans 1955
).
Oudemans (1904)
was struck by the similarity of female
Berlesia
chelicerae to those of
Varroa
Oudemans, 1904
, and their modifications for parasitism (although which cheliceral digit regresses was noted as dissimilar); however, he rejected a close relationship between these genera because of differences in ventral shields.
Vitzthum (1943)
modified
Kramer’s (1886)
monobasic
Iphiopsididae
, adding
Jacobsonia
Berlese, 1910b
and
Berlesia
to form a subfamily
Iphiopsidinae
within
Laelaptidae
(sic). The only significant character given in sup- port of this subfamilial grouping was the almost absent peritreme (along with a weak attribute of the idiosoma being colourless or brownish). However, this criterion was sometimes clearly subordinated to others, since the genus
Dinogamasus
Kramer, 1898
, which lack peritremes, was placed in the newly-created Hypoaspidinae, while
Neolaelaps
Hirst, 1926
, with similarly vestigial peritremes, was placed in the Laelaptinae.
Keegan (1950)
added
Dynatochela
,
a beetle-associate, to the
Iphiopsidinae
. Its reduced peritreme was the sole basis for subfamily placement, despite other notable discordant features of
Dynatochela
(including well-developed claws on all tarsi).
Clearly, the concept for any supraspecific grouping must include the
type
species.
Evans (1955)
invoked this point at the genus level, noting that the most important characters of
Iphiopsis
were the reduction of the peritreme and the absence of claws on leg I (the species original description by
Berlese, 1882
, indeed states that the first pair of legs are destitute of claws, and his accompanying figure 1, plate 64, clearly shows that leg I lacks claws albeit provided with a well-developed ambulacrum). Potential implications at the subfamily level were avoided by
Evans (1955)
noting that claws of one or more pairs of legs are
usually
reduced in
Iphiopsidinae
(our italics). This qualification allowed Evans to retain
Berlesia
and
Dynatochela
,
and include
Dinogamasus
in Iphiopsidinae. However
, by considering primarily myriapod hosts, Evans’ discussion did not account for the growing heterogeneity in this taxon. Also, iphiopsidines were stated to be “paraphages of arthropods”, thus overlooking statements by previous authors (
Oudemans 1904
,
Vitzthum 1943
) that
Berlesia
is parasitic.
Evans (1955)
makes a strong assertion that
Iphiopsidinae
“with little doubt arose from” the
Hypoaspis
-group—if so, Hypoaspidinae is rendered paraphyletic, and this creates another question as to the validity of
Iphiopsidinae
.
The
Iphiopsidinae
was again defined according to a reduced peritreme for the placement of
Dyscinetonyssus hystricosus
Moss & Funk, 1965
; however, this was qualified with acknowledgement that some authorities considered the characters used to define
Iphiopsidinae
as convergent (
Moss & Funk 1965
).
In the first published cladistic morphological analysis of the
Laelapidae,
Casanueva (1993)
recovered a phylogenetic estimate where all sampled myriapod associates clade together. Thus, the earlier concept of the
Iphiopsidinae
was recovered in part, but became substantially modified by including
Julolaelaps
Berlese, 1916
and
Scissuralaelaps
Womersley, 1945
. The former iphiopsidine
Dyscinetonyssus
was recovered in a separate clade that included all beetle and cockroach associates, while all hymenopteran associates including
Neoberlesia
and
Dinogamasus
were recovered in a greatly expanded Melittiphidinae.
Berlesia
was not analysed. These are interesting results, which might be intuitively appealing if one believes dermanyssines show a largely inflexible pattern of cophylogeny with their hosts at most levels ranging from species to family. However, the combination of character and taxon sampling does not permit full confidence in the results. For instance, while the loss of hypostomatic seta
hp3
has relevance to some myriapod associates, it was used to define a clade with
Julolaelaps
Berlese, 1916
subtending the myriapod-associated
Iphiopsidinae
. The sampling entailed a strong assumption that the exclusion of
Julolaelaps
which retain
hp3
(e.g.,
J. moseri
Hunter & Rosario, 1986
) would not substantially alter the results, including grouping with the similar
Hypoaspis
as previously hypothesized (
Evans 1955
). This result and similar ones appear to be reinforced by sampling taxa in a way that creates superficially distinct characters. For instance, the defining synapomorphy reconstructed for most of the Hypoaspidini is loss of podonotal
z3
which is a poor, mostly unrepresentative attribute to characterize many taxa intended for this clade including
Hypoaspis
sensu stricto
. Podonotal
z3
is an ontogenetically weak deutonymphal seta whose suppression is subject to much homoplasy (
Lindquist & Evans 1965
), so it would be surprising if its loss could define any sizeable clade. Also the five putative synapomorphies that define a clade equivalent to
Iphiopsidinae
(i.e. myriapod associates other than
Julolaelaps
and
Scissuralaelaps
) plus an additional four other putative synapomorphies that largely define a yet more inclusive clade, are all regressive leg characters that are also found in hypotrichous melittiphines, notably
Myrmozercon brevipes
Berlese, 1902
(with this genus actually not included in the analysis). Note that the leg chaetotaxy of
Berlesia
species is relatively holotrichous compared to most of the myriapod-associated
Iphiopsididae
sampled by
Casanueva (1993)
and bears only two of the nine reductive characters reconstructed by Casaneuva (1993;
Fig 8
, Stem 3) as putative synapomorphies for the relevant group containing the
type
species (Group VII Iphiopsini). However, an alleged sister group to the main iphiopsidid clade proposed by
Casanueva (1993)
is relevant to consider with respect to
Berlesia
as it contains potential relatives but also reveals possible pitfalls in uncovering these with morphological analysis. In this clade the mygalomorph-associated genus
Ljunghia
Oudemans, 1932
and the crab-associated genus
Cyclothorax
von Frauenfeld, 1868
are recovered together. Casaneuva (1993) noted that the synapomorphies linking
Ljunghia
to iphiopsidids were weak with low confidence in this expansion proposed for the
Iphiopsididae
.
Ljunghia
has similarities with
Berlesia
which may or may not indicate relationship. Deserving attention amongst these are: the usually pos- teriorly-positioned paranal setae, the shared absence of
gv3
on the anal shield, and the stout movable digit. For the latter, however, it is important to note that the movable digit of
Berlesia
appears rotated laterally, with its teeth flared away from the fixed digit, whose base is swollen (somewhat as in
Ixodida
). This contrasts with
Ljunghia
where the teeth on the movable digit still oppose what remains of the fixed digit, and the fixed digit base is not swollen. Thus
Ljunghia
movable digits lack some special features found in
Berlesia
that would make it more difficult to argue for similarity due to relationship under a structural criterion (
Riedl 1978
, p 34).
The above highlights the rampant homoplasy due to convergence and parallelism that plagues all potential candidates for
Iphiopsididae
. When both
Moraza & Kazemi (2012)
and
Seeman & Alberti (2015)
tried to apply characters proposed by Casaneuva (1993) to define/diagnose
Iphiopsididae
, they found that these characters had widespread occurrences elsewhere in the
Laelapidae
, such that both sets of authors rejected the family-ranking of
Iphiopsididae
.
Nemati
et al.
(2015)
also found the inclusion of
Julolaelaps
to be problematic.
Characters such as hypotrichy, reduced peritremes and reduced claws found in the “
Iphiopsididae
”, are known to be correlated with a wide variety of intimate symbioses in various
Dermanyssoidea
and are regressive in nature (as realized 65 years ago by
Evans 1955
).
Iphiopsididae
, at whatever rank, will remain an unstable concept, unless future molecular analyses, backed by unexpected morphological attributes indicate otherwise. Currently, it can only be considered satisfactory for taxa that are highly similar morphologically to the
type
genus
Iphiopsis
. Better knowledge of the
type
species would be useful here (
Halliday & Juvara-Bals 2016
), insofar as clarifying the core taxa related to
Iphiopsis
. Other piecemeal renovations to
Iphiopsididae
will not help, unless they include new diagnostic characters that reliably link included iphiopsidid taxa. However, new suprageneric groupings of taxa that are clearly not related to
Iphiopsis
, when these are supported by convincing synapomorphies, will advance overall systematic perspectives partly because they will reduce the number of taxa to be considered for placement in
Iphiopsididae
.