Abstract
External and internal head structures of Zorotypus hubbardi were examined. A detailed description is provided for the wingless morphs. The results are compared to the conditions found in alate specimens and in representatives of potentially related groups of insects. The pigmentation is strongly reduced in the wingless morphs and well-developed compound eyes, optic lobes and ocelli are absent. Kidney-shaped white spots between the paired ocelli and compound eyes were absent in all specimens examined. Most characters of the head (orthognathism, tentorium, musculature, mouthparts) are plesiomorphic. The wing and eye dimorphism, the strongly reduced head sutures, the reduced number of antennomeres, the presence of a prostheca on the left mandible and the presence of a transverse muscle of the head capsule and a retractor of the salivary sclerite are potential autapomorphies of Zoraptera. The unusual shape and large relative size of the brain and suboesophageal complex are also probably autapomorphic and a result of miniaturisation. Possible affinities with Dictyoptera were not supported by the results of this study. The only potential apomorphic feature of the head shared with Embioptera is the presence of a clearly delimited anteclypeus. However, this condition is also present in several groups of Acercaria. The very strongly developed M. clypeobuccalis, the medially divided prementum, and the slender, apically bifid laciniae without any mesally directed setae or spines are potential autapomorphies of Paraneoptera (incl. Zoraptera). If this group is indeed monophyletic, it is plausible to assume that the stem species was small like zorapterans and psocopterans and feeding on hyphae and spores of fungi, with mandibles with grinding molae and chisel-like laciniae suitable for loosening the food substrate. The feeding apparatus was then further modified within Acercaria, with a mortar-and-pistill apparatus in Psocoptera, and piercing–sucking mouthparts in Anoplura, Thysanoptera and Hemiptera.
Similar content being viewed by others
References
Badonnel A (1934) Recherche sur l’anatomie des Psoques. Bull Biol Fr Belge 18(Suppl):11–241
Beutel R, Gorb S (2001) Ultrastructure of attachment specializations of hexapods (Arthropoda): evolutionary patterns inferred from a revised ordinal phylogeny. J Zool Syst Evol Res 39:177–207
Beutel RG, Haas A (1998) Larval head of Hydroscapha natans LeConte, 1874 (Coleoptera, Myxophaga, Hydroscaphidae) with special reference to miniaturization. Zoomorphology 18:103–116
Beutel RG, Pohl H. Hünefeld F (2005) Strepsipteran brains and effects of miniaturization (Insecta). Arthr Str Dev (in press)
Bitsch J (1963) Morphologie céphalique des Machilides (Insecta Thysanura). Ann Sci Nat Zool Paris 5:501–706
Boudreaux HB (1979) Arthropod phylogeny, with special reference to insects. Wiley, New York
Buckup L (1957) Der Kopf von Myrsidea cornicis (De Geer) (Mallophaga-Amblycera). Zool Jb (Anat) 77:241–288
Carver M, Gross GF, Woodward TE (1991) Hemiptera (bugs, leafhoppers, cicadas, aphids, scale insects etc.). In: CSIRO (ed) The insects of Australia, vol 1. Cornell University Press, Ithaca, pp 429–509
Caudell AN (1918) Zorotypus hubbardi, a new species of the order Zoraptera from the United States. Can Entomol 50:375–381
Caudell AN (1920) Zoraptera not an apterous order. Proc Ent Soc Wash 22(5):85–97
Chaudonneret J (1950) La morphologie céphalique de Thermobia domestica (Packard) (Insecte Aptérygote Thysanoure). Ann Sci Nat 11:145–302
Choe JC (1989) Zorotypus gurneyi, new species, from Panama and redescription of Z. barberi Gurney (Zoraptera: Zorotypidae). Ann Entomol Soc Amer 82(2):149–155
Cope OB (1940) The morphology of Psocus confraternus Banks (Psocoptera, Psocidae). Microentomol 5(4):91–115
Deitz LL, Nalepa C, Klass K-D (2003) Phylogeny of the Dictyoptera re-examined (Insecta). Entomol Abh 61(1):69–91
Delamare-Deboutteville C (1951) Characteres sexuels cephaliques de type intermédiaires chez les males aptères de Zorotypus delamarei Paulian. Nat Malg 3(1):37–38
Dorsey CK (1943) The musculature of the labrum, labium and pharyngeal region of adult and immature Coleoptera. Smithson misc Coll 103(7):1–42, 24 pls
Engel MS (2003a) Phylogeny of the Zoraptera. Entomol Abh 61(2):147–148
Engel MS (2003b) Zoraptera. In: Resh VH, Cardé RT (eds) Encyclopedia of insects. Academic Press, New York, pp 1201–1203
Engel MS, Grimaldi DA (2000) A winged Zorotypus in Miocene amber from the Dominican Republic (Zoraptera: Zorotypidae), with discussion on relationships of and within the order. Acta Geol Hisp 35:149–164
Engel MS, Grimaldi DA (2002) The first Mesozioc Zoraptera (Insecta). Amer Mus Novitates 3362:1–20
Grebennikov VV, Beutel RG (2002) Morphology of the minute larva of Ptinella tenella, with special reference to effects of miniaturisation and the systematic position of Ptiliidae (Coleoptera: Staphylinoidea). Arthr Str Dev 31:157–172
Grimaldi D (2001) Insect evolutionary history from Handlirsch to Hennig, and beyond. J Paleontol 75:1152–1160
Grimaldi D, Engel MS (2005) Evolution of the insects. Cambridge University Press, Cambridge
Gurney AB (1938) A synopsis of the order Zoraptera, with notes on the biology of Zorotypus hubbardi Caudell. Proc Soc Wash 40:57–87
Hanken J, Wake DB (1993) Miniaturization of body size: organismal consequences and evolutionary constraints. Annu Rev Ecol Syst 24:501–519
Haub H (1967) Der Kopf von Pseudomenopon pilosum (Scopoli) (Mallophaga-Amblycera). Zool Jb (Anat) 84:493–558
Hennig W (1953) Kritische Bemerkungen zum phylogenetischen System der Insekten. Beitr Ent 3(Sonderheft):1–85
Hennig W (1969) Die Stammesgeschichte der Insekten. Waldemar Kramer, Frankfurt a M
Iturralde-Vinent M, MacPhee R (1996) Age and paleogeographic origin of Dominican Amber. Science 273:2750–2752
Kéler S v (1963) Entomologisches Wörterbuch. Akademieverlag, Berlin
Kéler S v (1966) Zur Mechanik der Nahrungsaufnahme bei Corrodentiern. Z Parasitenkde Berlin 27:64–79
Klass K-D (2003) Relationships among the principal lineages of Dictyoptera inferred from morphological data. Entomol Abh 61(2):134–137
Kristensen NP (1975) The phylogeny of hexapod “orders”. A critical review of recent accounts. Z Zool Syst Evol-forsch 13:1–44
Kristensen NP (1981) Phylogeny of insect orders. Annu Rev Entomol 26:135–157
Kristensen NP (1991) Phylogeny of extant hexapods. In: CSIRO (ed) The insects of Australia, vol 1. Cornell University Press, Ithaca, pp 125–140
Kukalová-Peck J, Haas F (2001) Dermaptera hindwing structure and folding: new evidence for familial, ordinal and subordinal relationships within Neoptera (Insecta). Eur J Entomol 98:445–509
Kukalová-Peck J, Peck SB (1993) Zoraptera wing structures: evidence for new genera and relationship with the blattoid orders (Insecta: Blattoneoptera). Syst Entomol 18(4):333–350
Leverault P (1938) The morphology of the Carolina Mantis. Section II: the musculature. Univ Kansas Sci Bull 25(21):557–633
Lewis T (1973) Thrips, their biology, ecology and economic importance. Academic, London
Lo N (2003) Molecular phylogenetics of Dictyoptera: insights into the evolution of termite eusociality and bacterial endosymiosis in cockroaches. Entomol Abh 61(2):137–138
Maki T (1935) A study of the musculature and of the phasmid Megacrania tsudai Shiraki. Mem Fac Sci Agric Taihoku Imp Univ 12:181–279
Maki T (1936) Studies on the skeletal structure, musculature and nervous system of the Alder Fly Chauliodes formosanus Petersen. Mem Fac Sci Agric Taihoku Imp Univ 16:117–243 + errata, 10 pls
Matsuda R (1965) Morphology and evolution of the insect head. Mem Amer Entomol Inst 4:1–334
Mickoleit E (1963) Untersuchungen zur Kopfmorphologie der Thysanopteren. Zool Jahrb Abt Anat Ontog Tiere 81:101–150
Minet J, Bourgoin T (1986) Phylogenie et classification des Hexapodes (Arthropoda). Cah Liaisons OPIE 20:23–28
Moulins M (1971) La cavité préorale de Blabera craniifer Burm. (Insecte, Dictyoptère) et son innervation: étude anatomo-histologique de l’épipharynx et l‘hypopharynx. Zool Jb (Anat) 88:527–586
Poinar Jr GO (1988) Zorotypus palaeus, new species, a fossil Zoraptera (Insecta) in Dominican amber. J NY Entomol Soc 96(3):253–259
Rähle W (1970) Untersuchungen an Kopf und Prothorax von Embia ramburi Rimsky-Korsakow 1906 (Embioptera, Embdiidae). Zool Jahrb Abt Anat Ontog Tiere 87(2):248–330
Rasnitzyn AP (1998) On the taxonomic position of the insect order Zorotypida = Zoraptera. Zool Anz 237:185–194
Riegel GT (1963) The distribution of Zorotypus hubbardi (Zoraptera). Ann Entomol Soc Amer 56:744–747
Riegel GT (1987) Order Zoraptera. In: Stehr FW (ed) Immature insects. Kendall/ Hunt Publishing Co., Dubuque, Iowa, pp 184–185
Risler H (1954) Der Kopf von Kopf Bovicola caprae (Gurlt) (Mallophaga). Zool Jb (Anat) 71:289–426
Silvestri F (1913) Descrizione di un nuovo ordine di insetti. Boll Lab Zool gen Agr R. Sc Sup Agric Portici 7:193–209
Slifer E, Sekhon SS (1978) Structures on the antennal flagellum of Zorotypus hubbardi (Insecta, Zoraptera). Not Nat 453:1–8
Staniczek AH (2000) The mandible of silverfish (Insecta: Zygentoma) and Mayflies (Ephemeroptera): its morphology and phylogenetic significance. Zool Anz 239:147–178
Strenger A (1953) Zur Kopfmorphologie der Ephemeridenlarven. Österr Zool Z 4:191–228
Tröster G (1990) Der Kopf von Hybophthirus notophallus (Neumann) (Phthiraptera: Anoplura). Eine funktionsmorphologishce und konsequent-phylogenetische Analyse. Stuttg Beitr Naturk A 442:1–89
Valentine BD (1986) Grooming behavior in Embioptera and Zoraptera (Insecta). Ohio J Sci 86:150–152
Vishnoi HS (1962) Musculature of a common Indian termite Odontotermes obesus (Rambur) (Isoptera, Termitidae). Bull Entomol (Loyola College, Madras) 3:13–20
Walker EM (1931) On the anatomy of Grylloblatta campodeiformis Walker. I. Exoskeleton and musculature of the head. Ann Entomol Soc Amer 24:519–536
Weber H (1929) Kopf und Thorax von Psylla mali (Hemiptera-Homoptera). Z Morph Ökol Tiere 14:59–165
Weber H (1938) Beiträge zur Kenntnis der der Überordnung Psocoidea. 4. Die Labialdrüsen der Copeognathen. Zool Jb (Anat) 66:131–346
Weidner H (1969) Die Ordnung Zoraptera oder Bodenläuse. Entomol Z 79(4):29–51
Weidner H (1970a) 14. Isoptera (Termiten). In: Helmcke J-G, Starck D, Wermuth H (eds) Handbuch der Zoologie. Bd. IV: Arthropoda—2. Hälfte: Insecta, 2. Teil: Spezielles. Walter de Gruyter, Berlin New York, pp 77–83
Weidner H (1970b) 15. Zoraptera (Bodenläuse). In: Helmcke J-G, Starck D, Wermuth H (eds) Handbuch der Zoologie. Bd. IV: Arthropoda—2. Hälfte: Insecta, 2. Teil: Spezielles. Walter de Gruyter, Berlin, pp 1–12
Weidner H (1972) 16. Copeognatha (Psocodea). In: Helmcke J-G, Starck D, Wermuth H (eds) Handbuch der Zoologie. Bd. IV: Arthropoda—2. Hälfte: Insecta, 2. Teil: Spezielles. Walter de Gruyter, Berlin, pp 1–94
Wheeler WC, Whiting M, Wheeler QD, Carpenter JM (2001) The phylogeny of extant hexapod orders. Cladistics 17:113–169
Willmann R (2003) Die phylogenetischen Beziehungen der Insecta: Offene Fragen und Probleme. Verh Westd Entom Tag 2001:1–64
Wilson EO (1959) Some ecological characteristics of ants in New Guinea rain forests. Ecology 40:437–447
Acknowledgements
We are very grateful to Dr. DC Carlton (Louisiana State University), who provided many well-preserved wingless specimens of Zorotypus hubbardi. We also wish to thank Dr. K-D Klass (Museum für Tierkunde Dresden) for the gift of mantodean specimens and to V Grebennikov (Agriculture Canada) for the gift of slide preparations of alate and wingless Zorotypus. The loan of alate specimens preserved in ethanol from the Smithsonian Institution is gratefully acknowledged. We are also grateful to Dr. MS Engel (Natural History Museum, University of Kansas) for reprints and valuable comments, to E Anton for technical support (slide preparations), to F Hünefeld for 3-D reconstructions (both Institut für Spezielle Zoologie und Evolutionsbiologie, FSU Jena) and to K-H Hellmer for technical assistance (SEM pictures, Abteilung Evolutionsbiologie der Invertebraten, EKU Tübingen).
Author information
Authors and Affiliations
Corresponding author
Rights and permissions
About this article
Cite this article
Beutel, R.G., Weide, D. Cephalic anatomy of Zorotypus hubbardi (Hexapoda: Zoraptera): new evidence for a relationship with Acercaria. Zoomorphology 124, 121–136 (2005). https://doi.org/10.1007/s00435-005-0117-z
Received:
Accepted:
Published:
Issue Date:
DOI: https://doi.org/10.1007/s00435-005-0117-z