Revision of Chinese mainland Hybovalgus Kolbe, 1904, with description of a new species, and Excisivalgus Endrödi, 1952 reduced to synonymy with Hybovalgus (Coleoptera: Scarabaeidae)
Author
Ricchiardi, Enrico
F09BCF76-35DE-4821-91C4-EEDC5114810A&2DE7F73C-A515-418C-A2AC-5394D5AB44C1
Corso A. Tassoni 79 / 4, 10143 Torino, Italy. Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese Academy of Sciences, Box 92, Beichen West road, Chaoyang District, Beijing, 100101, China; University of the Chinese Academy of Sciences, Yuquan road, Shijingshan, Beijing, 100039, China.
alericor@fastwebnet.it&545407454@qq.com
text
European Journal of Taxonomy
2017
2017-08-01
340
340
1
32
journal article
22065
10.5852/ejt.2017.340
0be43b36-2678-45a5-acfc-bec202ecb14a
2118-9773
3832552
2C6DC937-E543-4B8D-BF24-CED437F97676
Hybovalgus
Kolbe, 1904
Hybovalgus
Kolbe, 1904: 55
.
Excisivalgus
Endrödi, 1952: 62
,
syn. nov.
Type
species
Hybovalgus bioculatus
Kolbe, 1904
by monotypy.
Diagnosis
In the years after 1904 some entomologists tried to better define
Hybovalgus
(
Arrow 1944
;
Paulian 1961
;
Krikken 1978
;
Miyake 1985
), but could not find even one morphological synapomorphy.
Krikken (1984)
and
Miyake (1985)
wrote that the shape of the parameters was peculiar to
Hybovalgus
.
Miyake (1985)
distinguished
Hybovalgus
from
Dasyvalgus
as follows: “Parameres more or less prolonged, not confluent each other, with a excision on each inner edge,
Hybovalgus
. Parameres vestigial, sometimes confluent each other,
Dasyvalgus
”. The paramere shape of each genus is different, and easily distinguishable, although the meaning given to the word “vestigial” by Miyake is not clear.
After studying many specimens (including all the species of
Hybovalgus
in
Taiwan)
, we conclude that the males of
Hybovalgus
can be distinguished from those of
Dasyvalgus
by a series of non-exclusive characters and by the shape of the parameres (
Fig. 3
A–F). The special shape of
Hybovalgus
parameres is distinctive compared with the variable parameres of males of
Dasyvalgus
species. This paramere shape, that we define as “
Hybovalgus
shape”, within the Chinese
Valgina
is shared only with
Dasyvalgoides denticulatus
Endrödi, 1952
, in a monotypic genus. The relationship between these two genera must still be investigated, but the latter species is readily distinguishable from any
Hybovalgus
by the protibia with six teeth; and from
H. sexdentatus
by its flattened body with inconspicuous scales tufts.
The series of non-exclusive characters that separate
Hybovalgus
from the other Oriental
Valgina
genera are:
- Protibial external teeth 1–3 long and sharp in males, where tooth 2 is sometime a bit shorter than 1 and 3; the same for females but often they are worn by digging (see remarks on page 7 about differences between the two sexes) and in some cases tooth 2 is slightly shorter than 1 or 3 (
Fig. 1
).
- Both sexes have 5–6 protibial teeth.
- Suctorial brush long in males, short in females (
Ricchiardi & Li 2015
). See remarks below about differences between the two sexes.
- Anterior margin of clypeus noticeably sinuate in both sexes, a bit less in females of
H. thibetanus
.
- Pronotal lateral margins more or less crenate in both sexes.
- Pronotal lateral margin depression always present in both sexes.
- Pronotal posterior margin of most females (but not all) more or less excavated.
- Parameres of
Hybovalgus
shape (
Fig. 3
).
Fig. 1.
Five-toothed protibia.
A
. LT ♂ of
Hybovalgus fraternus
Moser, 1915
(MNHUB).
B
. LT ♂ of
Hybovalgus fulvosquamosus
Moser, 1904
(MNHUB).
- Protibia with five teeth on the external margin, in one case six (
H. sexdentatus
).
- Pygidium of males always narrowed from the middle to the apex; apex rounded in caudal view, not prominent in lateral view. Pygidium of females often (but not always) modified at apex (see Remarks under species accounts below).
- Pygidium of males with a ventral small, flat or slightly hollowed surface with posterior margin rounded or pointed at center. In females this area is bigger and deeply hollowed.
- Pygidium of females without pygidial stylus.
- Propygidial scale tufts of females darker than those of males.
Some species have protibial teeth 1–2 and/or 3–4 closer to one another than to the others. This character is not consistent among all the specimens of
Hybovalgus
as it varies within the same species.
Fig. 2.
Mating pair of
Hybovalgus tonkinensis
Moser, 1904
, from Fujian (IZAS).
A
. ♂.
B– C
. ♀.
Another character that we decided not to use for the separation of species is the presence or absence of cretaceous maculae on the propygidium or pygidium of males. In some species those maculae are really difficult to see because they are covered with thick scales or by the built-up body grease.
Identification of females
Generally, it is easy to separate females from males because the pronotal posterior margin is more or less deeply excavated (
H. bioculatus
,
H. fraternus
,
H. thibetanus
,
H. tonkinensis
) and/or the apex of the pygidium is modified into a pointed projection (
H. fraternus
,
H. sexdentatus
,
H. thibetanus
,
H. tonkinensis
), a rounded projection (
H. bioculatus
), or the apex is simply rounded (
H. calvus
sp. nov.
).
Often, females are larger than males. All the females we studied bear black propygidial scale tufts, even when their male counterparts have testaceous propygidial scale tufts.
Females of two species,
H. fulvosquamosus
and
H. jordansi
, are unknown.
Variability of paramere shape
After analysis of the males of a many
Valgina
belonging to hundreds of species, we concluded that the study of the shape of the parameres is a reliable way to define species. There are, however, a couple of notable exceptions.
Jameson & Swoboda (2005)
, revising the New World
Valgina
, wrote that: “However, one species of New World valgine,
Valgus seticollis
Beauvois, 1807
, has extremely variable male genitalia although there is an overall similarity in pattern” (
Jameson & Swoboda 2005
: figs 31–35); “Other characters that we examined are consistent within this species, including form of the tarsomeres, pygidial spine, pronotum, sternites, and spiracles. Lacking other morphological characters to separate the genitalic forms, we conservatively group all the forms as one species. A thorough study of the genitalic forms of
V. seticollis
would be an excellent topic for a molecular analysis and could reveal interesting evolutionary patterns”.
Kolbe (1897)
described a new
Valgina
genus,
Homovalgus
, to include
Valgus seticollis
.
Jameson & Swoboda (2005)
clarified the nomenclature of the species and placed it under
Valgus
. It is our opinion that because the shape of the parameres is similar to that of
Hybovalgus
/
Dasyvalgoides
, and the lack of a pygidial stylus in females, this species must be separated from
Valgus
and that its valid name is
Homovalgus seticollis
. The relationships of this species to the Oriental
Valgina
could add appeal to further study.
In the Oriental Region, another species,
Hybovalgus fraternus
, is a similar case to that of
Homovalgus seticollis
.
Hybovalgus fraternus
also exhibits limited variation in morphology and great variation in paramere shape. This is why we decided to conservatively avoid splitting this species merely on the basis of variation in paramere shape.
Description of aedeagus
Figure 3
shows several examples of the aedeagus of Chinese
Hybovalgus
species along with some nomenclature. The outline of the aedeagus is distinctive. The phallobase is always V-shaped ventroapically (vs) and with a sclerotized upper anterior part (spb) notched, flat or pointed, sometime variable within the same species. The parameres are prominent, with the apex extended and with different shapes, with or without a sub-apical external or internal tooth; basal lobe more or less developed and in some cases variable in shape within a species. In lateral view the parameres shows a more or less deep, basomedial notch (bmn).
Fig. 3.
Different shapes of aedeagus.
A
.
Hybovalgus fraternus
Moser, 1904
.
B
.
H. thibetanus
(
Nonfried, 1891
)
.
C
.
H. tonkinensis
Moser, 1904
. Proposed nomenclature of
Hybovalgus
aedeagus: aap = anterior apex of phallobase; ap = apex of parameres; bl = basal lobe; bmn = baso-medial notch; etp = sub-apical external tooth of parameres; iml = internal medial lobe; itp = sub-apical internal tooth of parameres; ma = membrane of aedeagus; pb = phallobase; spb = sclerotized upper anterior part of phallobase; vs = V-shaped ventro-apically membranous region of aedeagus. Drawings: grey parts: parameres; white parts: phallobase.
Distribution
Hybovalgus
species are mostly distributed in south-central continental
China
, with most species extending to northern
Laos
, northern
Vietnam
, Hainan Island (5–6 species) and
Taiwan
. In
Taiwan
there are six species:
Hybovalgus kamiyai
(Sawada, 1941)
;
H. major
Sawada, 1939
;
H. matsudai
Miyake, 1985
;
H. obsoletosquamosus
Miyake, 1985
;
H. takasagoensis
(Sawada, 1941)
and
H. thoracicus
Moser 1911
. These six Taiwanese species are considered endemic, but their relationships with continental
Hybovalgus
must still be analyzed.
Figure 4
shows the approximate distribution of
Hybovalgus
species. In the west, the genus inhabits Yunnan and Sichuan (there are no records so far for
Myanmar
). In the east some species reach
Shandong
, Jangsu and
Zhejiang
. In the north there are extensions into the southestern part of Palaearctic
China
(
sensu
Chen
et al.
2008
). We found no records of
Hybovalgus
from
Henan
and
Anhui
provinces or
Chongqing
Municipality, but this may be due to a lack of collecting.
Summary of Chinese species distribution
Hybovalgus bioculatus
Kolbe, 1904
:
Laos
,
Vietnam
,
China
(
Hainan
,
Guangxi
).
Hybovalgus calvus
sp. nov.
:
Vietnam
,
China
(
Hainan
).
Hybovalgus fraternus
Moser, 1915
:
Vietnam
,
Laos
,
China
(
Fujian
,
Zhejiang
,
Sichuan
,
Guangxi
, Jangxi,
Hunan
,
Guizhou
).
Hybovalgus fulvosquamosus
Moser, 1904
:
Laos
,
Vietnam
,
China
(
Hainan
,
Hunan
,
Guangdong
).
Hybovalgus jordansi
(
Endrödi, 1952
)
:
China
(
Fujian
).
Fig. 4.
Dotted area: approximate distribution of
Hybovalgus
. Original map from http://www.d-maps. com/pays.php?num_pay=77&lang=it.
Hybovalgus sexdentatus
Arrow, 1944
:
Vietnam
,
Laos
,
China
(
Guangxi
,
Yunnan
,
Sichuan
,
Hainan
,
Shandong
?).
Hybovalgus thibetanus
(
Nonfried,1891
)
:
China
(
Yunnan
,
Sichuan
, Jangsu,
Shaanxi
,
Jiangxi
,
Hubei
,
Gansu
,
Zhejiang
).
Hybovalgus tonkinensis
Moser, 1904
:
Vietnam
,
China
(
Fujian
,
Zhejiang
,
Jiangxi
,
Guangxi
,
Hunan
,
Guizhou
,
Hainan
,
Gansu
,
Sichuan
).