A stable phylogenomic classification of Travunioidea (Arachnida, Opiliones, Laniatores) based on sequence capture of ultraconserved elements
Author
Derkarabetian, Shahan
Author
Starrett, James
Author
Tsurusaki, Nobuo
Author
Ubick, Darrell
Author
Castillo, Stephanie
Author
Hedin, Marshal
text
ZooKeys
2018
760
1
36
http://dx.doi.org/10.3897/zookeys.760.24937
journal article
http://dx.doi.org/10.3897/zookeys.760.24937
1313-2970-760-1
0B57270DC24C4D4DA04F15CA442E0A07
Family
PARANONYCHIDAE Briggs, 1971
Type genus.
Paranonychus
Briggs, 1971
Type species.
Paranonychus brunneus
(Banks, 1893).
Diagnosis.
The
Paranonychidae
can be diagnosed by their relatively complex glans (except
Paranonychus
) (Figure 7 and Suppl. material 2: Figure 4), and by their intestinal complex (Suppl. material 2: Figure 2). For all taxa that have been examined, the paranonychids possess a small D1 that is circular to subtriangular, and a simple and shorter OD3. The paranonychids are restricted to western North America and East Asia. In southern Japan the paranonychids are sympatric with
Yuria
and can be diagnosed by several characteristics:
Yuria
possesses a free ninth tergite, and the penis has a dorsal plate with fused stylus; the paranonychids do not have a free ninth tergite and the penis glans lacks a dorsal plate. In western North America, the paranonychids are sympatric and syntopic in surface habitats with the
Cladonychiidae
(
Briggsus
,
Isolachus
) and
Cryptomastridae
(
Cryptomaster
). The paranonychids can be differentiated from these families by the structure of D1: paranonychids possess a small circular to subtriangular unbranched D1, while
Cladonychiidae
and
Cryptomastridae
possess an elongate, triangular, and branched D1.
Included genera and species.
Paranonychus
(Figure 1K). This trans-Beringian genus includes three known species:
P. brunneus
(Banks, 1893) distributed in the Coast and Cascade Ranges of Oregon and Washington with records extending north to Alaska;
P. concolor
Briggs, 1971, recorded from a single location in the southern Cascade Range of Oregon; and
P. fuscus
found throughout northern Honshu in Japan.
Metanonychus
Briggs, 1971. This genus and all species were described by Briggs (1971) and are restricted to the moist forests of the Pacific Northwest of North America.
Metanonychus
includes three species:
M. nigricans
Briggs, 1971 with two subspecies,
M. n. nigricans
and
M. n. oregonus
, found in Oregon;
M. setulus
Briggs, 1971 with five subspecies,
M. s. setulus
,
M. s. cascadus
,
M. s. mazamus
,
M. s. navarrus
, and
M. s. obrieni
, found in Oregon, Washington, and northern California; and
M. idahoensis
Briggs, 1971 found in northern Idaho.
Sclerobunus
Banks, 1893 (Figure 1L). Recently revised by
Derkarabetian and Hedin (2014)
,
Sclerobunus
is distributed throughout western North America and currently includes 12 species divided into three species groups. The nondimorphicus group includes
S. nondimorphicus
Briggs, 1971 from Oregon, Washington, and British Columbia, and
S. idahoensis
Briggs, 1971 from northern Idaho. The cave-obligate cavicolens group includes:
Sclerobunus cavicolens
(Banks, 1905) restricted to Lewis and Clark Caverns, Montana;
Sclerobunus ungulatus
(Briggs, 1971) from caves in Great Basin National Park, Nevada;
Sclerobunus madhousensis
(Briggs, 1971) from caves near Provo, Utah. The robustus group includes the widespread
S. robustus
(Packard, 1877),
S. glorietus
Briggs, 1971, and
S. skywalkeri
Derkarabetian & Hedin, 2014, all distributed throughout the high elevation forests of the southwestern United States, and
S. jemez
Derkarabetian & Hedin, 2014,
S. klomax
Derkarabetian & Hedin, 2014,
S. speoventus
Derkarabetian & Hedin, 2014, and
S. steinmanni
Derkarabetian & Hedin, 2014, which are all troglomorphic species restricted to cave and talus habitats along the eastern edge of the southern Rocky Mountains in New Mexico and Colorado.
Kaolinonychus
Suzuki, 1975. This monotypic genus endemic to South Korea is recorded mostly from caves.
Kaolinonychus coreanus
(Suzuki, 1966) includes two subspecies
K. c. coreanus
and
K. c. longipes
.
Metanippononychus
Suzuki, 1975. (Figure 1M). Endemic to Japan,
Metanippononychus
is restricted to southern Honshu, Shikoku, and Kyushu and includes four species:
M. daisenensis
Suzuki, 1975;
M. iriei
Suzuki, 1975, with two subspecies
M. i. iriei
and
M. i. yakuensis
;
M. iyanus
Suzuki, 1975;
M. tomishimai
Suzuki, 1975, with two subspecies
M. t. tomishimai
and
M. t. awanus
.
Nippononychus
Suzuki, 1975. A monotypic genus endemic to Japan,
Nippononychus japonicus
(Miyosi, 1957) is restricted to southern Honshu and Shikoku.
Zuma
(Figure 1N).
Zuma
includes two species restricted to forests of central and northern California:
Zuma acuta
Goodnight & Goodnight, 1942 restricted to the coastal forests south of San Francisco;
Zuma tioga
Briggs, 1971 found in the central and northern Sierra Nevada range.
Izunonychus
Suzuki, 1975. A monotypic genus endemic to Japan,
Izunonychus ohruii
Suzuki, 1975 is restricted to the Izu peninsula and Hakone area in central Honshu.
Kainonychus
Suzuki, 1975 (Figure 1O). A monotypic genus endemic to Japan,
Kainonychus akamai
(Suzuki, 1972) includes two subspecies,
K. a. akamai
distributed throughout northern Honshu and
K. a. esoensis
restricted to Hokkaido.
Remarks.
In this study all genera in the
Paranonychidae
have been sampled and the generic relationships are consistent and highly supported across all analyses (Figs 4, 5). Although the study of
Derkarabetian et al. (2010)
only included North American taxa, the relationships of paranonychids recovered here are the same, notably
Paranonychus
as the earliest diverging genus, and a sister relationship between
Sclerobunus
and
Metanonychus
. The familial name
Sclerobunidae
has been used previously (
Giribet et al. 2010
) for the "northern triaenonychids". However,
Paranonychidae
and the subfamily
Paranonychinae
Briggs, 1971 have priority over the names
Sclerobunidae
and
Sclerobuninae
Dumitrescu 1976
.
The Japanese genera
Metanippononychus
and
Nippononychus
show levels of UCE divergence consistent with congeners (Figs 4, 5). Intermediate morphological forms between
Nippononychus japonicus
and
Metanippononychus daisenensis
can be found where the two species come into contact (Tsurusaki pers. obs.). These genera are differentiated only by tarsal claw structure:
Metanippononychus
possessing a ventral tooth on the median prong of the hind claws. The original drawings of male genitalia show that
M. daisenensis
and
N. japonicus
differ in the width of the stylus (
Suzuki 1975b
). However, the penis of
N. japonicus
is highly similar to that of the geographically proximate
M. tomishimai tomishimai
.
Kury et al. (2014)
includes
Paranonychus fuscus
(formerly
Mutsunonychus fuscus
) as a synonym of
Paranonychus brunneus
(Banks, 1893) based on
Shear's
(1986)
statement "
Paranonychus brunneus
(=
Mutsunonychus fuscus
Suzuki;
Paranonychidae
)". Later in
Shear and Derkarabetian (2008)
, the genus
Mutsunonychus
was formally synonymized under
Paranonychus
, and although a potential species level synonymy was noted, it was not formally established. The levels of UCE divergence between
P. brunneus
and
P. fuscus
are consistent with species level divergences compared to other pairs of congeneric taxa included (Figs 4 and 5), and as such,
P. fuscus
is again treated as a distinct species here.