Download from the Early Cretaceous in China

Zootaxa 3273: 63–68 (2012)
www.mapress.com / zootaxa/
Copyright © 2012 · Magnolia Press
ISSN 1175-5326 (print edition)
Article
ZOOTAXA
ISSN 1175-5334 (online edition)
A new fossil Saldidae (Hemiptera: Heteroptera: Leptopodomorpha)
from the Early Cretaceous in China
WEITING ZHANG1, JINGJING SONG1, YUNZHI YAO1,2,3 & DONG REN1
1
Key Lab of Insect Evolution and Environmental Changes, Capital Normal University, Beijing 100048, China;
State Key Laboratory of Palaeobiology and Stratigraphy (Nanjing Institute of Geology and Palaeontology, CAS), Nanjing 210008,
China
3
Corresponding author. E-mail: [email protected]
2
Abstract
Venustsalda locella gen. et sp. nov. is described and illustrated from the Lower Cretaceous Yixian Formation at Huangbanjigou Village, Liaoning Province, China. The new genus is established based on its unusual six cells on the membrane,
with the second cell smallest.
Key words: Heteroptera, Saldidae, fossil, Early Cretaceous, Yixian Formation
Introduction
Saldidae, commonly called shore bugs, is the largest family in the Leptopodomorpha: about 335 species are
recognized. They are distributed worldwide, especially in the Northern Hemisphere (Schuh et al.1987; Schuh &
Polhemus 2009). Saldids are usually found on rocky shores of streams and lakes (Brooks & Kelton 1967; Poinar &
Buckley 2009). All saldids are predatory and most of them feed on organisms, found in the damp surface layers of
the substrate (Brooks & Kelton 1967; Polhemus & Chapman 1979).
Schuh and Polhemus (1980) present a hierarchic classification in Leptopodomorpha and considered the
Aepophilidae to be the sister group of Saldidae. Some subfamilies such as Aepophilinae, Leptosaldinae,
Saldoniinae, were formerly erected in Saldidae (Cobben 1959, 1971; Popov 1973), but they are finally removed
(Leston 1956; Schuh & Polhemus 1980; Popov 1985). The widely accepted classification of the family Saldidae is
that it contains two subfamilies: Chiloxanthinae and Saldinae (Schuh & Slater 1995).
Records of fossil Saldidae are scarce and only seven species have been reported (Carpenter 1992). Only one
incontrovertible Mesozoic species (Brevrimatus pulchalifer Zhang, Yao & Ren, 2011) from the Early Cretaceous
Yixian Formation of Duolun County, Inner Mongolia, China, has been reported (Zhang et al. 2011). Most of the
fossil species are found from the Cenozoic, such as the Eocene species, Salda exigua Germar & Berendt, 1856
from Baltic amber; the Upper Oligocene species, Oligosaldina rottensis Statz & Wagner, 1950, O. rhenana Statz &
Wagner, 1950, and O. aquatilis Statz & Wagner, 1950 from Germany; the Miocene species, Propentacora
froeschneri (Lewis, 1969) (= Oreokora froeschneri) from USA, and Salda littoralis found in recent Late Glacial
clay (Jessen 1923). However, two genera and four species assigned to this group previously: Leptosalda chiapensis
Cobben, 1971 from Tertiary (late Oligocene or early Miocene age) amber of Chiapas, Mexico amber; Saldonia
rasnitsyni Popov, 1973, S. sibirica Popov, 1985, and S. maculata Popov, 1985 from the Lower or Middle Jurassic in
Transbaikalia of Russian, were transferred to Leptopodidae and Archegocimicidae respectively (Cobben 1971,
1987; Popov 1973, 1985; Polhemus 1977; Schuh & Polhemus 1980).
In the present paper, a new genus and species of Saldidae is described from the Lower Cretaceous Yixian
Formation (about 125 Ma; Swisher et al. 1999; Zhou et al. 2003; Xing et al. 2005) in Huangbanjigou, Chaomidian
Village, Beipiao City, Liaoning Province, China. The Yixian Formation is considered part of the Jehol Biota,
yielding beautifully preserved insects, reptiles, birds (Lei et al. 2005; Wang & Ren 2006; Ren et al. 2010; Yao et al.
2011). The stable ecosystem, abundant vegetation, and various species suggest that the region as a whole had a
Accepted by C. Schaefer: 14 Mar. 2012; published: 19 Apr. 2012
63
warm, humid climate (Liu et al. 2009). Because this species cannot be assigned to any known genera of Saldidae,
we erected a new genus to accommodate it.
Material and methods
This fossil material is housed in the Key Laboratory of Insect Evolution & Environmental Changes, Capital
Normal University, Beijing, China. The specimen was examined with a Leica MZ12.5 dissecting microscope.
Photo was taken by a Nikon Digital Camera DXM1200C. Outline was drawed with Adobe Photoshop CS2. Basic
terminology and the classification system follow Polhemus (1977) and Schuh and Polhemus (2009).
Body length was measured along the midline from the anterior margin of the head to the apex of the abdomen.
Body width was measured at the maximal width of the body. The lengths of pronotum and scutellum were
measured at the midline. Wing length was measured from the basal to the apex of anterior margin. Wing width was
measured at the maximal width of the wing. All metrics are provided in millimeters.
Systematic paleontology
Order Hemiptera Linnaeus, 1758
Suborder Heteroptera Latreille, 1810
Infraorder Leptopodomorpha Popov, 1971
Family Saldidae Amyot & Serville, 1843
Subfamily Chiloxanthinae Cobben, 1959
Genus Venustsalda gen. nov.
Type species. Venustsalda locella gen. et sp. nov.
Diagnosis. Compound eyes large, covering most of head in lateral view. Rostrum reaching to the base of hind
coxae. Posterior pronotal margin indented distinctly. Forewings covering abdomen, macropterous; medial fracture
and costal fracture long; hypocostal ridge and associated secondary hypocostal ridge present on hemelytra; corial
vein reaching to outermost cell; veins on membrane forming six closed cells. In male, parandrium tapered.
Etymology. The generic name is a combination of the Latin “venust-” (meaning “charming”) and “Salda” (the
type genus of this family). Gender feminine.
Distribution. China.
Remark. It is indubitable to classify this species into the family Saldidae based on the combination of characters: compound eyes large, rostrum long, posterior margin of pronotum concave, hemelytra with costal fracture and
medial fracture, membrane with few cells. The new genus has six cells on its membrane, which is different from
the normal cell number in Saldidae. In most saldids, the innermost cell (first cell) is longest, whereas in this new
genus the normal innermost cell seems to be divided into two cells, and then finally form six cells on membrane.
Therefore, the six-cells system is speculated to be derived from the five-cells system. According to its long medial
fracture and costal fracture, we intend to put this genus in Chiloxanthinae.
Both Venustsalda gen. nov. and Brevrimatus Zhang, Yao & Ren, 2011 come from the same formation; but the
former easily differs from the latter in body size less than 6.5 mm (vs. reaches to 8.0 mm), medial fracture fused
together with costal fracture (vs. medial fracture short, not connect with costal fracture), membrane with six closed
cells (vs. membrane with five closed cells), and forewings surpassing the end of abdomen (vs. forewings at most
reach to the end of abdomen).
Venustsalda gen. nov. is similar to Oligosaldina Statz & Wagner, 1950 in body size, but can be distinguished
from the latter: forewing with costal fracture (vs. without costal fracture), body dorsal surface smooth, without long
hair (vs. body covered by long hair), membrane with six cells (vs. five cells).
64 · Zootaxa 3273 © 2012 Magnolia Press
ZHANG ET AL.
FIGURE 1. Line drawing of Venustsalda locella gen. et sp. nov.
A NEW FOSSIL SALDIDAE FROM THE EARLY CRETACEOUS
Zootaxa 3273 © 2012 Magnolia Press ·
65
PLATE 1. Photograph of Venustsalda locella gen. et sp. nov., Holotype, CNU-HET-LB2010335.
66 · Zootaxa 3273 © 2012 Magnolia Press
ZHANG ET AL.
Propentacora froeschneri (Lewis), initially assigned to the genus Oreokora (Lewis 1969) in the subfamily Saldinae, was transfered to the genus Propentacora in Chiloxanthinae (Polhemus 1977, 1985). Venustsalda gen. nov.
compared with Propentacora (Lewis): length of hemelytra over 4 mm (vs. less 3mm), membrane with six cells (vs.
five cells), the innermost cell (first cell) shorter than the third cell (vs. the innermost cell longest, cells reduced
gradually from the inner to the outer as the normal arrangement), and corial vein reaching to the outermost cell (vs.
corial vein continue between cells three and four).
Salda is an extant genus with two fossil species. Most members of Salda have four cells and the fossil species
S. exigua Germar & Berendt, 1856 has only three cells. Venustsalda gen. nov. has six cells, more than that in Salda.
Venustsalda locella gen. et sp. nov.
(Fig.1, Plate 1)
Description. Body small, oval, 2.0 times as long as wide. Head short and broad, 1.7 times as wide as long, length
of head almost subequal to length of pronotum on midline. Antennae long, slender, four-segmented, ratio of antennal segments I:II:III:IV=1:2.8:1.9:1.8. Pronotum subtrapezoid, 3.1 times as wide as long, moderately narrowed
towards the front, anterior margin narrower than head, lateral margins nearly straight, posterior margin concave,
posterior angles feebly rounded. Scutellum longer than pronotum on midline, triangle, and width about 1.4 times of
length. Fore femora stout, subequal to length of corresponding tibiae, tibiae 1.9 times as long as tarsi; mid tibiae 2.0
times as long as tarsi; hind legs longest, tibiae about 1.7 times as long as femora, and 2.3 times as long as tarsi, tibiae with a row of stout setae on lateral margin, tarsi three-segmented, tarsomere I shortest, tarsomere II 1.3 times as
long as tarsomere III; all tarsi with two claws. Hemelytra 0.8 times as long as body, covering apex of abdomen, clavus about half length of fore wing, claval commissure shorter than scutellum length at median line, corium with
embolium, embolium similar in color to corium; outer edge of corium with a white spot at base of hemelytra; hypocostal ridge and associated secondary hypocostal ridge present; corial vein reaching to outermost cell; membrane
with six cells, second cell smallest, less than half length of third cell, outermost cell reduced, about half length of
adjacent cell.
Measurements. Body length 4.81, width 2.39. Head length 0.66, width 1.10mm. Antenna length I–IV: 0.38,
1.07, 0.75, 0.72. Rostrum length 2.92. Scutellum length 0.85, width 1.23. Length of fore leg: femur 1.19, tibia 1.23,
tarsus 0.63; length of middle leg: tibia 1.51, tarsus 0.75; length of hind leg: femur 1.63, tibia 2.75, tarsomere I–III:
0.08, 0.62, 0.46. Length of hemelytra 4.09, width 1.29.
Type material. Holotype, ♂, CNU-HET-LB2010335.
Type locality and horizon. Huangbanjigou Village, Beipiao City, Liaoning Province, China, Yixian Formation. Early Cretaceous.
Etymology. The species name is derived from Latin word “locella,” which indicated the species with small
cells. Gender feminine.
Acknowledgements
This research was funded by grants from the National Basic Research Program of China (973 Program) (No.
2012CB821900); National Natural Science Foundation of China (No. 30800095, 31071964, 40872022); Nature
Science Foundation of Beijing (No. 5082002); the PHR Project of Beijing Municipal Commission of Education
(No. 20090509, 201107120), the General Program of Science and Technology Development Project of Beijing
Municipal Education Commission of China (No. KM201210028016), Fok Ying-Tong Education Foundation for
Young Teachers in the Higher Education Institutions of China (No.131021), and the State Key Laboratory of Palaeobiology and Stratigraphy (Nanjing Institute of Geology and Palaeontology, CAS) (No.123114).
References
Brooks, A.R. & Kelton, L.A. (1967) Aquatic and semiaquatic Heteroptera of Alberta, Saskatchewan, and Manitoba
(Hemiptera). Memoirs of the Entomological Society of Canada, 51, 1–92.
A NEW FOSSIL SALDIDAE FROM THE EARLY CRETACEOUS
Zootaxa 3273 © 2012 Magnolia Press ·
67
Cobben, R.H. (1959) Notes on the classification of Saldidae with the description of a new species from Spain. Zoologische
Mededelingen, 36 (22), 303–316.
Cobben, R.H. (1971) A fossil shore bug from the Tertiary amber of Chiapas Mexico (Heteroptera, Saldidae). University of California Publications in Entomology, 63, 49–56.
Cobben, R.H. (1987) New African Leptopodomorpha (Heteroptera: Saldidae, Omaniidae, Leptopodidae), with an annotated
checklist of Saldidae from Africa. I. New species of the genus Saldula (Saldidae). Revue de Zoologie Africaine, 100,
399–421.
Carpenter, F.M. (1992) Treatise on invertebrate paleontology, Part R, Arthropoda 4, Volume 3: Superclass Hexapoda. Geological Society of America and University of Kansas Press, Lawrence, 655 pp.
Jessen, K. (1923) A submerged peat-bog in the harbour of Rungsted and changes of sea-level in the sound in the late-glacial
period. Dansk Geologisk Forening, 6 (14), 1–18.
Lei, G.Z. (2005) The origin and discovery of Jehol Entomofauna. Journal of Microbiology, 25(1), 110–112.
Leston, D. (1956) Systematics of the Marine-Bug. Nature, 178, 427–428
Lewis, S.E. (1969) Fossil Insects of the Latah Formation (Miocene) of Eastern Washington and Northern Idaho. Northwest Science, 41 (3), 99–115.
Liu, P.J., Huang, J.D. & Ren, D. (2009) Palaeocology and structure of entomofauna in the Yixian Formation of northern Hebei
and western Liaoning. Journal of Environmental Entomology, 31(3), 254–274 [in Chinese with English abstract].
Poinar, G. & Buckley, R. (2009) Palaeoleptus burmanicus n. gen., n. sp., an Early Cretaceous shore bug (Hemiptera: Palaeoleptidae n. fam.) in Burmese amber. Cretaceous Research, 30, 1000–1004.
Polhemus, J.T. (1977) The biology and systematics of the Saldidae of Mexico and middle America. PhD thesis, University of
Colorado, Boulder, 606 pp.
Polhemus, J.T. & Chapman H.C. (1979) Family Saldidae/Shore Bugs. In: Menke, A.S. (Ed.) The Semiaquatic and Aquatic
Hemiptera of California (Heteroptera: Hemiptera). Bulletin of the California Insect Survey, Vol. 21. University of California Press, Berkeley, pp. 16–33.
Polhemus, J.T. 1985. Shore bugs (Heteroptera, Hemiptera; Saldidae): a world overview and taxonomy of Middle American
forms. Different Drummer, Englewood, 252 pp.
Popov, Yu.A. (1973) The first discovery of the hemipteran family Saldidae (Heteroptera) in the Mesozoic of Siberia. Doklady
Akademii Nauk SSSR, 209, 703–705 [in Russian].
Popov, Yu.A. (1985) Jurassic bugs and Coleorrhyncha of southern Siberia and western Mongolia. In: Rasnitsyn, A.P. (Ed.),
Jurassic insects of Siberia and Mongolia, Trudy Paleontologicheskogo Instituta Akademii Nauk SSSR, 211, 28–47 [in Russian].
Ren, D., Shih, C.K., Gao, T.P., Yao, Y.Z. & Zhao, Y.Y. (2010) Silent Stories--Insect Fossil Treasures from Dinosaur Era of the
Northeastern China. Science Press, Beijing, 322 pp.
Schuh, R.T. & Polhemus, J.T. (1980) Analysis of taxonomic congruence among morphological, ecological, and biogeographic
data sets for the Leptopodomorpha (Hemiptera). Systematic Zoology, 29 (1), 1–26.
Schuh, R.T. & Polhemus, J.T. (2009) Revision and analysis of Pseudosaldula Cobben (Insecta: Hemiptera: Saldidae): a group
with a classic Andean distribution. Bulletin of the American Museum of Natural History, 323, 1–102.
Schuh, R.T. & Slater, J.A. (1995) True Bugs of the World (Hemiptera: Heteroptera). Classification and Natural History. Cornell University Press, Ithaca, New York, 336 pp.
Schuh, R.T., Galil, B. & Polhemus, J.T. (1987) Catalog and bibliography of Leptopodomorpha (Heteroptera). American
Museum of Natural History, 185, 243–406.
Statz, G. & Wagner, E. (1950) Geocorisae (Landwanzen) aus den Oberoligocäner Ablagerungen von Rott. Palaeontographica
(abt. A), 98, 97–136.
Swisher, C.C., Wang, Y.Q., Wang, X.L., Xu, X. & Wang, Y. (1999) Cretaceous age for the feathered dinosaurs of Liaoning,
China, Nature, 400, 58–61.
Wang Y. & Ren, D. (2007) Two new genera of fossil palaeontinids from the Middle Jurassic in Daohugou, Inner Mongolia,
China (Hemiptera, Palaeontinidae). Zootaxa, 1390, 41–49
Xing, D.H., Sun, C.L., Sun, Y.W., Zhang, L.D., Peng, Y.D. & Chen, S.W. (2005) New knowledge on Yixian Formation. Acta
geoscientica sinica, 26 (1), 25–30 [in Chinese with English abstract].
Yao, Y.Z., Zhang, W.T., Ren, D. & Shih, C.K. (2011) New fossil Ochteridae (Hemiptera: Heteroptera: Ochteroidea) from the
Upper Mesozoic of northeastern China, with phylogeny of the family. Systematic Entomology, 36, 589–600.
Zhang, W.T., Yao, Y.Z. & Ren, D. (2011) New shore bug (Hemiptera, Heteroptera, Saldidae) from the Early Cretaceous of
China with phylogenetic analyses. Zookeys, 130, 185–198.
Zhou, Z.H., Barrett, P.M. & Hilton, J. (2003) An exceptionally preserved lower Cretaceous ecosystem. Nature, 421, 807–814.
68 · Zootaxa 3273 © 2012 Magnolia Press
ZHANG ET AL.