SEARCH

SEARCH BY CITATION

Keywords:

  • Heteroptera;
  • Dipsocoromorpha;
  • Schizopteridae;
  • mid-Cretaceous;
  • French amber;
  • Burmese amber;
  • palaeoecology

Abstract

  1. Top of page
  2. Abstract
  3. Localities and stratigraphy
  4. Material
  5. Systematic palaeontology
  6. Discussion
  7. References

Abstract:  Bugs of two new genera and species are described as Buzinia couillardi and Tanaia burmitica. They are preserved in mid-Cretaceous amber from south-west France and northern Myanmar (Burma), respectively (c. 100 Ma). These are the first formally described fossils of the heteropteran family Schizopteridae. Both belong to the subfamily Hypselosomatinae and are very similar to the extant genus Hypselosoma Reuter, providing evidence for the antiquity and morphological stability of this small bug family and the infraorder Dipsocoromorpha. Given the putative ecology of the fossils, a discussion is provided on the French and Burmese amber forest ecosystems. The geological setting of La Buzinie, a new amber deposit in south-west France that yielded the two specimens of Buzinia couillardi, is outlined.

The Dipsocoromorpha is the smallest and most basal infraorder of Euheteroptera (sensuWheeler et al. 1993; Schuh and Slater 1995) within the Heteroptera (or true, sucking bugs) without the family Enicocephalidae (Text-fig. 1). Though minute insects like these commonly have highly reduced wing venation, the dipsocoromorphs have retained a more extensive venation than most other bugs, reflecting their basal position. Perhaps because of their small size (0·5–4 mm) and cryptic habits, their Mesozoic record has been extremely limited hitherto. Dipsocoromorpha is basal to the Gerromorpha (water striders and relatives) and Nepomorpha (predatory water bugs, water scorpions, backswimmers, and similar), but the two latter infraorders have significant to excellent Mesozoic records. The Nepomorpha range back to the Late Triassic (Carnian) (Grimaldi and Engel 2005), so the Dipsocoromorpha would be expected to be at least that old. Other than an apparent dipsocoromorph from the Triassic (Grimaldi and Engel 2005, fig. 8.60), however, there are almost no other Mesozoic records of the infraorder apart from those reported from Early Cretaceous Lebanese and Burmese ambers (Text-fig. 1). All of these fossils, as well as the very few known from the Cenozoic, are still undescribed. The discovery of two Cretaceous dipsocoromorph genera that are very closely related to the extant schizopterid genus Hypselosoma is consistent with an Early Mesozoic age of the infraorder.

image

Figure TEXT-FIG. 1..  Phylogeny of the heteropteran infraorders (based on Wheeler et al. 1993), showing the fossil record of Dipsocoromorpha and earliest fossil records of Gerromorpha and Nepomorpha. 1, undescribed ?Dipsocoromorpha, Late Triassic, Virginia, USA (Grimaldi and Engel 2005, fig. 8.60). 2–3, undescribed Dipsocoridae and Schizopteridae, Neocomian Lebanese amber (Grimaldi and Engel 2005). 4–5, undescribed Dipsocoridae and Tanaia gen. nov. (Schizopteridae), Albian Burmese amber (Grimaldi et al. 2002; this paper). 6, Buzinia gen. nov. (Schizopteridae), Albian and Cenomanian French amber (this paper). 7, undescribed Schizopteridae, Late Eocene Baltic amber (Popov and Herczek 1993; Shcherbakov and Popov 2002, fig. 200). 8–9, undescribed Schizopteridae, Early Miocene Dominican and Mexican amber (Poinar 1992; Engel 2004). 10, Mesoveliidae, Late Jurassic, Karatau, Kazakhstan (Grimaldi and Engel 2005). 11, undescribed Belostomatidae and Naucoridae, Late Triassic, Virginia, USA (Grimaldi and Engel 2005).

Download figure to PowerPoint

Recent Schizopteridae comprise approximately 120 living species in 35 genera of predominantly tropical distribution. According to Hill (1984), Hypselosoma is the most basal genus of the family, and the most recent generic radiation of the Hypselosomatinae probably occurred in Australia during the Miocene–Pliocene changes in vegetation, as reflected by the almost exclusively Australasian distribution of the extant genera. Indeed, Australia and the Pacific Islands appear to have been a major area of evolution and refuge for the subfamily. The occurrence of isolated genera in Madagascar and North America, together with the French amber fossils described here, merely reflects a relict distribution.

Localities and stratigraphy

  1. Top of page
  2. Abstract
  3. Localities and stratigraphy
  4. Material
  5. Systematic palaeontology
  6. Discussion
  7. References

Two of the fossil bugs described in this paper were found in Late Albian amber from Archingeay-Les Nouillers in south-west France. Detailed descriptions of the geology and taphonomy of this amber deposit were given previously by Néraudeau et al. (2002) and Perrichot (2004, 2005). Two other specimens belonging to the same new species were found in amber from a recently discovered amber deposit named La Buzinie at Champniers, near Angoulême (Text-fig. 2A). This deposit is assumed to be Early Cenomanian in age because it corresponds to black lignitic marls (lithological subunit A2; Text-fig. 2B) located under a beige limestone (subunit B1) that yields the Early Cenomanian foraminiferan Orbitolina concavaLamarck, 1816. The marls are progressively enriched in carbonate, so that the boundary between A2 and B1 is inconspicuous locally; the base of the limestone is grey. Both the black marls and the grey limestone of A2 yield lignitic matter and amber. A second amber-bearing stratum occurs above, at the base of green sands (subunit B2) overlying B1. The section ends with a series of limestones yielding O. concava (subunit B3). The amber containing the specimens described herein comes from the black marls (A2). In the Charentes region, subunits A2, B1, B2 and B3 constitute the whole of the Lower Cenomanian succession (sensuNéraudeau et al. 1997). The lowermost amber deposit from La Buzinie, located at the A2/B1 boundary, is stratigraphically younger than the Upper Albian amber (subunit A1) from Archingeay-Les Nouillers but older than that from the Lower Cenomanian (subunit B2) of Fouras, which was described some years ago (Néraudeau et al. 2003; Text-fig. 2A).

image

Figure TEXT-FIG. 2..  Geological setting of the Cretaceous amber deposits in France. A, simplified geological map of the Charentes region, with location of Cretaceous amber deposits. B, stratigraphic section of La Buzinie at Champniers, with location of the two amber-bearing strata.

Download figure to PowerPoint

Descriptions of the geology of amber deposits in Myanmar have been provided by Zherikhin and Ross (2000), Grimaldi et al. (2002) and Cruickshank and Ko (2003). According to them, the amber is Albian in age, making it contemporaneous with the amber of Archingeay-Les Nouillers.

Material

  1. Top of page
  2. Abstract
  3. Localities and stratigraphy
  4. Material
  5. Systematic palaeontology
  6. Discussion
  7. References

The two specimens from Archingeay-Les Nouillers were found together with 76 synclusions in a single piece of ‘litter amber’ (sensuPerrichot 2004). These bugs were hitherto incorrectly figured and recorded by Néraudeau et al. (2002, figs 4, 6.7) and Perrichot (2004, table 2) as homopteran Membracidae. The two specimens from La Buzinie were found together with 16 other arthropods also in a single piece of ‘litter amber’. They were separated from other synclusions using a lancet as a microsaw, then embedded in Canada balsam between cover glasses for examination.

Two amber pieces provided the representatives from Myanmar. The best preserved one is fossilized with a cockroach nymph in a thin piece of polished amber. It was previously figured by Grimaldi et al. (2002, fig. 25c) and Grimaldi and Engel (2005, fig. 8.61) but not described. The other specimen is incomplete and poorly preserved in a thin piece of amber that was embedded in a clear, synthetic resin block for study and conservation.

The specimens upon which this paper is based are deposited in the Muséum National d'Histoire Naturelle (MNHN), Paris, and the American Museum of Natural History (AMNH), New York. The terminology used in the systematic part follows that of Schuh and Slater (1995). The descriptions refer to the male holotypes unless otherwise noted.

Systematic palaeontology

  1. Top of page
  2. Abstract
  3. Localities and stratigraphy
  4. Material
  5. Systematic palaeontology
  6. Discussion
  7. References

Family SCHIZOPTERIDAE Reuter, 1891 Subfamily HYPSELOSOMATINAE Esaki and Miyamoto, 1959

Type genus. HypselosomaReuter, 1891.

Diagnosis.  Minute bugs with exceedingly large eyes, overlapping anterolateral margin of pronotum; forewing with extensive longitudinal veins; male mesotarsi trimerous but incrassate sensuHill (2004) (= ‘4-segmented’sensuEsaki and Miyamoto 1959; ‘pseudotetramerous’sensuEmsley 1969), but the latter indicated that this last character can be found in at least the non-hypselosomatine genus SilhouettanusEmsley, 1969; male fore- and mid-legs with arolia.

NB: Hill (1980) gave a new diagnosis of the Hypselosomatinae based on the very large eyes and some other characters not visible in our fossils. He added that this definition is not very satisfactory when compared with those for the Schizopterinae and Ogeriinae (an enigmatic subfamily not accepted by Schuh and Slater 1995). However, our fossils are easily distinguished from the latter by the three-segmented labium (Hill 1989, 1990, 2004) and the presence of four pairs of abdominal spiracles (visible in Tanaia gen. nov.).

Included genera. Hypselosoma (five Recent species in Australia and isolated species in New Zealand, New Caledonia, New Guinea, Solomon Islands, Indonesia, the Philippines, Japan, China, Madagascar), GlyptocombusHeideman, 1906 (one species, USA), OmmatidesUhler, 1894 (one species, Lesser Antilles), PateenaHill, 1980 (four species, Australia), RectilaminaHill, 1984 (seven species, Australia), CryptomannusHill, 1984 (two species, Australia), MacromannusHill, 1984 (one species, Australia), OrdireteHill, 1984 (one species, Australia), LativenaHill, 1984 (two species, Australia), DuonotaHill, 1984 (12 species, Australia), Buzinia gen. nov. (one species, French amber), and Tanaia gen. nov. (one species, Burmese amber).

Genus BUZINIA gen. nov.

Type species. Buzinia couillardi sp. nov.

Derivation of name.  After the locality of La Buzinie where the amber containing the allotype and one paratype was found.

Diagnosis.  Closely similar to the extant genus Hypselosoma, especially in the following characters: forewing with veins R, M1, M2, Cu, 1V, 2V present, defining four marginal cells and three narrow, elongate, median cells; male and female tarsal formulae 3/3/3 and 2/2/3, respectively. Distinguished from Hypselosoma by the absence of a trapezoidal cell on the forewing and the presence of three macrosetae on the labrum, instead of five, and from all other Hypselosomatinae by its forewing venation.

Buzinia couillardi sp. nov. Text-figures 3–4
image

Figure TEXT-FIG. 3.. Buzinia couillardi gen. et sp. nov. in French Cretaceous amber. A, C, Albian amber from Archingeay-Les Nouillers; B, D, Cenomanian amber from La Buzinie. A, male holotype, MNHN Arc 115.9. B, female allotype, MNHN Buz 1.5. C, female paratype, MNHN Arc 115.7. D, female paratype, MNHN Buz 1.6. Scale bars represent 0·5 mm.

Download figure to PowerPoint

image

Figure TEXT-FIG. 4.. Buzinia couillardi gen. et sp. nov. in Albian–Cenomanian French amber (male holotype MNHN Arc 115.9). A, dorsal habitus. Scale bar represents 0·5 mm. B, interpretation of the forewing venation.

Download figure to PowerPoint

Derivation of name.  After Mr Alain Couillard who discovered the amber deposit of La Buzinie.

Types.  Holotype male MNHN Arc 115.9; allotype female MNHN Buz 1.5; paratype female MNHN Arc 115.7; other female specimen MNHN Buz 1.6; in mid-Cretaceous (Upper Albian–Lower Cenomanian) amber from Archingeay-Les Nouillers (Arc) and La Buzinie (Buz) at Champniers, Charentes, south-west France.

Diagnosis.  As for genus.

Description

Body length 1·23–1·30 mm (from head to wing apex). Head strongly declivous when viewed from side, 0·1 mm long, 0·5 mm wide; two long trichobothria, or ocular bristles, between eyes; ocelli not visible; antenna four-jointed, with several erect setae; first and second joints the shortest, length of joints 0·06/0·08/0·3/0·3 mm, respectively; eyes large and broad, overlapping anterolateral margin of pronotum; three macrosetae on labrum; labium three-segmented, length 0·18 mm; frons not delimited.

Thorax.  Pronotum transverse, without longitudinal ridges, length 0·15 mm, width 0·53 mm; scutellum short, length 0·22 mm, width 0·29 mm; proepisternal lobe broad, extending below two thirds of eyes.

Wings.  Forewing tegminal, 1·00 mm long; clavus large, 0·40 mm long, 0·1 mm wide; no costal fracture; embolium broad; veins R, M1, M2, Cu, 1V, 2V present, defining four marginal cells and three narrow, elongate, median cells; trapezoidal cell absent, fused with second median cell. Hindwings present, composed of a net of small cells and at least one longitudinal vein, but size and shape undetermined.

Legs.  Tarsal formula 3/3/3, with mesotarsi trimerous but incrassate. Fore coxae extending below mid-length of eyes; long macrosetae on fore femora; fore tarsi without ventral spines; arolia on fore- and mid-legs. Leg measurements (in mm): fore femur 0·28, tibia 0·33, tarsi 0·03/0·05/0·05; mid femur and tibia 0·33, tarsi 0·04/0·05/0·05; hind femur 0·37, tibia 0·54, tarsi 0·05/0·07/0·07.

Abdomen.  Length 0·33 mm, width 0·37 mm, setose; spiracles not visible; male genital structure hardly visible, except the long, curled vesica and two small, rounded structures on each side of abdomen, corresponding to either the apices of parameres or paraproctal appendages.

NB: The females are of similar size but differ by the tarsal formula which is 2/2/3 instead of 3/3/3. The pro- and mesotarsi are together as long as those of the males.

Genus TANAIA gen. nov.

Type species. Tanaia burmitica sp. nov.

Derivation of name.  After the village of Tanai in which the amber containing the type-species was found.

Diagnosis.  Closely similar to the extant genus Hypselosoma and the fossil genus Buzinia gen. nov. by the forewing venation defining four marginal cells and three narrow, elongate, median cells. Distinguished from Hypselosoma by the rather long partial fusion of veins Cu and M2, the first basal cross-vein between Cu and R+M instead of Cu and M, and the presence of only three macrosetae on the labrum instead of five; from Buzinia gen. nov. by the presence of a trapezoidal cell on the forewing, the long fusion of veins Cu and M2, four shorter and broader marginal cells, and longer vein R+M; from all other Hypselosomatinae by its forewing venation.

Tanaia burmitica sp. nov. Text-figures 5–6
image

Figure TEXT-FIG. 5.. Tanaia burmitica gen. et sp. nov. in Albian Burmese amber. A, male holotype, AMNH Bu-720; B, paratype AMNH Bu-965. Scale bars represent 0·5 mm.

Download figure to PowerPoint

image

Figure TEXT-FIG. 6.. Tanaia burmitica gen. et sp. nov. in Albian Burmese amber; male holotype AMNH Bu-720. A, dorsal habitus. B, forewing. C, genital appendages in dorsal view. Scale bars represent 0·25 mm in A and 0·1 mm in C.

Download figure to PowerPoint

Derivation of name.  After burmite, a name applied to amber from Myanmar (formerly Burma).

Types.  Holotype male AMNH Bu-720, paratype (female?) AMNH Bu-965, in mid-Cretaceous (Upper Albian) amber from Tanai Village in Kachin Province, Myanmar.

Diagnosis.  As for genus.

Description

Body length 1·18 mm (from head to wing apex). Head strongly declivous, 0·46 mm wide; a long trichobothrium near margin of right eye; no visible ocelli; antenna with numerous erect setae, first and second joints the shortest, length of joints: 0·06/0·06/0·33/0·4 mm; eyes very large, overlapping anterolateral margin of pronotum; three macrosetae on labrum; segmentation of labium not visible; frons not delimited.

Thorax.  Pronotum transverse, without longitudinal ridges, 0·18 mm long, 0·42 mm wide; scutellum short, 0·14 mm long, 0·25 mm wide.

Wings.  Forewing tegminal, 0·95 mm long; clavus large, 0·50 mm long, 0·15 mm wide; no costal fracture; embolium broad; veins R, M1, M2, Cu, 1V, 2V present, defining four rather short and broad marginal cells, three narrow, elongate, median cells and a quadrangular to pentagonal trapezoidal cell; cross-vein between R+M and Cu, rather long distal fusion between Cu and M2 separating again distally; fusion between Cu and 1V at base of most distal marginal cell; vein 2V distally evanescent. Hindwing membrane composed of a net of small cells and at least one longitudinal vein, but size and shape are undetermined.

Legs.  Tarsal formula 3/3/3, with mesotarsi trimerous but incrassate. Fore coxae extending below two-thirds of eyes; long macrosetae on fore femora; fore tarsi without ventral spines; arolia on fore- and mid-legs. Leg measurements (in mm): length of fore tibia 0·33, tarsi 0·03/0·04/0·04; mid femur and tibia 0·33, tarsi 0·03/0·04/0·07; hind femur 0·33, tibia 0·62, tarsi 0·03/0·05/0·07.

Abdomen.  As wide as long, length 0·40 mm, densely setose; four visible pairs of spiracles; male genital structures asymmetrical, with a coiled vesica making a curve, elongate right and left parameres and two short, lateral appendages (paraproctal appendages?).

Discussion

  1. Top of page
  2. Abstract
  3. Localities and stratigraphy
  4. Material
  5. Systematic palaeontology
  6. Discussion
  7. References

Recent Schizopteridae live in wet environments, with many species being ground- and litter-dwellers in rainforests, tropical palm swamps and grasslands where they are predators of small invertebrates (Schuh and Slater 1995). Hill (1984) indicated that Australian Hypselosomatinae are closely allied to rainforest refugia and elevated bogs. He recorded Hypselosoma species from a variety of habitats including rainforests, open forest leaf litter and long grasses (Hill 1987). In the mangroves of Singapore, schizopterids are part of the myriad fauna that shelter under dead wood during high tide and emerge during low water to feed on small invertebrates (e.g. springtails, copepods) deposited by the receding tide (Ng and Sivasothi 1999). Interestingly, all specimens of Buzinia are in two pieces of amber with numerous other arthropods that are clearly litter-inhabitants. The 76 synclusions fossilized with the holotype and paratype from Archingeay-Les Nouillers were listed and their palaeoecological implications noted by Perrichot (2004). Similarly, 16 litter-inhabiting arthropods are fossilized with the allotype and paratype in amber from La Buzinie: a chilopod, four acari, a springtail, nine dolichopodid flies and a tanaidacean crustacean. Tanaidaceans are small aquatic, mainly marine but also brackish-water forms with relatively few species being reported from estuaries. They are predominantly tube-dwellers and burrowers (Sieg 1983). These aquatic arthropods are thus not likely to be caught in resin exuded by trees on land, unless aggregated on the shore after death and trapped in a flow of resin from plants growing nearby. Therefore, it is tempting to speculate that Buzinia was a litter-dwelling scavenger living in mangroves or banks of estuaries where the resin-producing plants grew, feeding on small invertebrates deposited by the receding tide. It is reasonable to speculate that the Burmese genus Tanaia lived in similar habitats, given its close resemblance to Buzinia and the blattodean inclusion fossilized with the holotype specimen. This provides new evidence for mesohaline habitats in mid-Cretaceous amber forests in France and Myanmar. Although geographically distant, these contemporaneous ecosystems apparently shared similar estuarine and mangrove-like tropical environments. Previous studies have indicated tropical rainforests for Myanmar but more mesic forests for France (Grimaldi et al. 2002; Perrichot 2005), with different origins for the amber. The Burmese amber could have a taxodiaceous or an angiosperm origin (Grimaldi et al. 2002; Lambert and Poinar 2002); the French amber is thought to be araucariaceous or cheirolepidiaceous (Perrichot 2005).

The discovery of Early Cretaceous Hypselosomatinae, which are very similar to extant Hypselosoma, provides evidence for the great antiquity and morphological stability of the poorly known bug family Schizopteridae.

Acknowledgements.  We are grateful to David Grimaldi (AMNH) for the loan of the Burmese specimens; Alain Couillard and Ginette Lambert who found the amber deposit of La Buzinie and helped us in its excavation, together with Romain Vullo and Bernard Gomez (University of Rennes 1); Gilbert Hodebert (MNHN) for drawing of one of the French specimens studied; and to two anonymous reviewers for their helpful comments on the first version of the manuscript. This work was partly supported by the Alexander von Humboldt Foundation (to VP).

References

  1. Top of page
  2. Abstract
  3. Localities and stratigraphy
  4. Material
  5. Systematic palaeontology
  6. Discussion
  7. References
  • CRUICKSHANK, R. D. and KO, K. 2003. Geology of an amber locality in the Hukawng Valley, northern Myanmar. Journal of Asian Earth Sciences, 21, 441455.
  • EMSLEY, M. G. 1969. The Schizopteridae (Heteroptera) with the description of new species from Trinidad. Memoirs of the American Entomological Society, 25, 1154.
  • ENGEL, M. S. 2004. Arthropods in Mexican amber. In BOUSQUETS, J. L., MORRONE, J. J., ORDÓÑEZ, O. Y. and FERNÁNDEZ, I. V. (eds). Biodiversidad, taxonomía y biogeografía de artrópodos de México: hacia una síntesis de su conocimiento. Universidad Nacional Autónoma de México, Bulletin, 4, 175–186.
  • ESAKI, T. and MIYAMOTO, S. 1959. A new or little known Hypselosoma from Amami-Oshima and Japan, with the proposal of a new tribe for the genus (Hemiptera). Sieblodia, 2, 109120. [In Chinese, English summary].
  • GRIMALDI, D. A. and ENGEL, M. S. 2005. Evolution of the insects. Cambridge University Press, Cambridge, 755 pp.
  • GRIMALDI, D. A., ENGEL, M. S. and NASCIMBENE, P. C. 2002. Fossiliferous Cretaceous amber from Myanmar (Burma): its rediscovery, biotic diversity, and paleontological significance. American Museum Novitates, 3361, 171.
  • HEIDEMAN. 1906. A new genus and species of the hemipterous family Ceratocombidae from the United States. Proceedings of the Entomological Society of Washington, 7, 192194.
  • HILL, L. 1980. Tasmanian Dipsocoroidea (Hemiptera: Heteroptera). Journal of the Australian Entomological Society, 19, 107127.
  • HILL, L. 1984. New genera of Hypselosomatinae (Schizopteridae) from Australia. Australian Journal of Zoology (Supplementary Series), 32, 155.
  • HILL, L. 1987. Four new Australian species of Hypselosoma Reuter (Heteroptera: Schizopteridae). Journal of the Australian Entomological Society, 26, 265278.
  • HILL, L. 1989. A revision of Australian Pachyplagia Gross (Heteroptera: Schizopteridae). Invertebrate Taxonomy, 3, 605617.
  • HILL, L. 1990. Australian Ogeria Distant (Heteroptera: Schizopteridae). Invertebrate Taxonomy, 4, 697720.
  • HILL, L. 2004. Kaimon (Heteroptera: Schizopteridae), a new, speciose genus from Australia. Memoirs of the Queensland Museum, 49, 603647.
  • LAMARCK, J.-B.de 1816. Histoire naturelle des animaux sans vertèbres, Tome 2. Verdière, Paris, 568 pp.
  • LAMBERT, J. B. and POINAR, G. O. Jr 2002. Amber: the organic gemstone. Accounts of Chemical Research, 35, 628636.
  • NÉRAUDEAU, D., ALLAIN, R., PERRICHOT, V., VIDET, B., DE LAPPARENT DE BROIN, F., GUILLOCHEAU, F., PHILIPPE, M., RAGE, J.-C. and VULLO, R. 2003. Découverte d'un dépôt paralique à bois fossiles, ambre insectifère et restes d'Iguanodontidae (Dinosauria, Ornithopoda) dans le Cénomanien inférieur de Fouras (Charente-Maritime, Sud-Ouest de la France). Comptes Rendus Palevol, 2, 221230.
  • NÉRAUDEAU, D., PERRICHOT, V., DEJAX, J., MASURE, E., NEL, A., PHILIPPE, M., MOREAU, P., GUILLOCHEAU, F. and GUYOT, T. 2002. Un nouveau gisement à ambre insectifère et à végétaux (Albien terminal probable): Archingeay (Charente-Maritime, France). Geobios, 35, 233240.
  • NÉRAUDEAU, D., THIERRY, J. and MOREAU, P. 1997. Variation in echinoid biodiversity during the Cenomanian–early Turonian transgressive episode in Charentes (France). Bulletin de la Société Géologique de France, 168, 5161.
  • NG, P. K. L. and SIVASOTHI, N. 1999. A guide to the mangroves of Singapore II (animal diversity). Singapore Science Centre, Singapore, 168 pp.
  • PERRICHOT, V. 2004. Early Cretaceous amber from south-western France: insight into the Mesozoic litter fauna. Geologica Acta, 2, 922.
  • PERRICHOT, V. 2005. Environnements paraliques à ambre et à végétaux du Crétacé nord-aquitain (Charentes, Sud-Ouest de la France). Mémoires de Géosciences Rennes, 118, 1310.
  • POINAR, G. O. Jr 1992. Life in amber. Stanford University Press, Stanford, CA, 350 pp.
  • POPOV, Y. A. and HERCZEK, K. 1993. New data on Heteroptera in amber resins. Annals of the Upper Silesian Museum, Entomology, Supplement, 1, 712.
  • REUTER, O. M. 1891. Monographia Ceratocombidarum orbis terrestris. Acta Societatis Scientiarum Fennicae, 19, 1 pl.
  • SCHUH, R. T. and SLATER, J. A. 1995. True bugs of the world (Hemiptera: Heteroptera) classification and natural history. Cornell University Press, Ithaca, NY, 336 pp.
  • SHCHERBAKOV, D. E. and POPOV, Y. A. 2002. Superorder Cimicidea Laicharting, 1781. Order Hemiptera Linné, 1758. The bugs, cicadas, plantlice, scale insects, etc. (= Cimicida Laicharting, 1781, = Homoptera Leach, 1815 + Heteroptera Latreille, 1810). 143156. In RASNITSYN, A. P. and QUICKE, D. L. J. (eds). History of insects. Kluwer, Dordrecht, 517 pp.
  • SIEG, J. 1983. Tanaidomorpha (Crustacea: Tanaidacea) from the Ross Sea, Antarctica. Journal of the Royal Society of New Zealand, 13, 395418.
  • UHLER, P. R. 1894. A list of the Hemiptera-Heteroptera of the families Anthocoridae and Ceratocombidae collected by Mr. H. H. Smith in the Island of St. Vincent; with descriptions of new genera and species. Proceedings of the Zoological Society of London, 1894, 156160.
  • WHEELER, W., SCHUH, R. T. and BANG, R. 1993. Cladistic relationships among higher groups of Heteroptera: congruence between morphological and molecular data sets. Entomologica Scandinavica, 24, 121137.
  • ZHERIKHIN, V. V. and ROSS, A. J. 2000. A review of the history, geology and age of Burmese amber (Burmite). Bulletin of the Natural History Museum, London (Geology Series), 56, 310.