Database of Hymenoptera in America north of Mexico

This database is originally based on the
Catalog of Hymenoptera in America North of Mexico, Volumes 1-3,
prepared cooperatively by specialists under the direction of
Karl V. Krombein, Paul D. Hurd, Jr., David R. Smith and B. D. Burks
published by the Smithsonian Institution Press, Washington, D.C., 1979,
and digitized by the Biodiversity Heritage Library, 2009.

Draft-1, contains unformated sections and lots of OCR errors!
Please cite as follows:
Pickering, J. 2009. Database of Hymenoptera in America north of Mexico.
Proceedings of Life http://www.discoverlife.org/proceedings/0000/6
Updated: 26 February, 2014

Index to Taxa of Hymenoptera • Highertaxa

(supplement to catalog of species north of Mexico). — Berland, 1958. Atlas des
Hymenopteres de France. I, Tenthredes, Parasites, Porte-aiguillon (Bethyloides, pars), 155
pp., 14 pis. II, Porte-aiguillon (Bethyloides (fin), Scolioides, Formicoides, Pompiloides,
Vespoides, Sphecoides, Apoides), 184 pp., 14 pis. — Oeser, 1961. Zool. Mus. Berlin, Mitt. 37:
75-85, figs. 106-107 (phylogeny based on ovipositor). — Krombein, Burks et al., 1967. U. S.
Dept. Agr., Agr. Monog. 2, Sup. 2, 584 pp. (supplement to catalog of species north of
Mexico). — Malyshev, 1968. Genesis of Hymenoptera and phases of their evolution 319 pp.
(English translation by Richards and Uvarov). — Borror and DeLong, 1970. An
Introduction to the Study of Insects, ed. 3, pp. 537-616, figs. 495-577. — Riek et ai, 1970.
/)( The Insects of Australia, C. S. I. R. 0., pp. 867-959, 40 figs. -Richards, 1972 (1971).
Ent. Essays to Commemorate Retirement of Prof. K. Yasumatsu, pp. 1-13, 10 figs,
(thoracic spiracles in classification). — Rasnitsyn, 1971. XIII Internatl. Congr. Ent., Proc. 1:
289 (origin). -Koenigsmann, 1976. Deut. Ent. Ztschr. (n.f. 23) 4-5: 253-279, 4 figs,
(phylogeny).

Biology: Bishoff, 1927. Biologie de Hymenopteren, 598 pp., 224 figs. — Fulmek, 1957.
Naturhist. Mus. Wien, Ann. 61: 110-227 (aphid parasites and predators). — iWata, 1972
(1971). Evolution of Instinct: comparative Studies of Hymenoptera Behavior, 503 pp.
(Japanese edition). — Farish, 1972. Anim. Behavior 20: 662-676 (grooming behavior).
— Iwata, 1976. Evolution of Instinct: Comparative Ethology of Hymenoptera, 535 pp.,
frontisp., 50 figs. (English translation, Natl. Tech. Inform. Serv., PB 257052).

Morphology: Snodgrass, 1911. U. S. Natl. Mus., Proc. 39: 37-91, 16 pis., 19 text figs, (thorax).
— rohwer and Gahan, 1916. Ent. Soc. Wash., Proc. 18: 20-76, 3 pis. (wing venation).
—Comstock, 1918. The Wings of Insects, pp. 362-381, pi. 10, text figs. 380-405.
—Snodgrass, 1935. Principles of Insect Morphology, 667 pp., 319 figs. —Ross, 1936. Ent.
Soc. Amer., Ann. 29: 99-111, 2 pis. (wing venation). —Snodgrass, 1941. Smithsn. Inst., Misc.
Collect. 99 (14): 1-86, 33 pis., 6 text figs, (male genitalia). — Reid, 1941. Roy. Ent. Soc.
London, Trans. 91: 367-446, 81 figs, (thorax of wingless and brachypterous spp.).
—Richards, 1956. Roy Ent. Soc. London, Proc. (A) 31: 99-104, 7 figs, (interpretation of
thoracic venter). -Short, 1959. Roy. Ent. Soc, London, Trans. Ill: 175-203, 11 figs,
(abdominal musculature). —Oeser, 1961. Zool. Mus. Berlin, Mitt. 37: 3-119, 107 figs, (female
ovipositor). —Daly, 1963. Ent. Soc. Amer., Ann. 56: 295-306 (thoracic musculature).
—Smith, 1970. Ent. Soc. Amer., Ann. 63: 1-27 (evolutionary morphology of external
genitalia). — Eady, 1974. Jour. Ent., Ser. B, 48: 63-72 (wing venation).


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drous in the Macroxyelinae and strophandrous in the Xyelinae.

All members of this suborder are phytophagous except for the family Orussidae which is
parasitic on other wood-boring insects. Host plants and habits are various. Many larvae are ex-
ternal feeders on the foliage of the host, but others are leaf miners, gall formers on twigs,
petioles, leaves, or buds, leaf-edge rollers, stem borers, wood borers, or petiole miners. Because
of their plant feeding habits, many sawflies are pests of ornamental plants, agricultural crops, or
forest stands. Their occurrence is normally local, but outbreaks in forests may be extensive,
covering many thousands of acres and resulting in considerable growth loss or death of trees.
Appearance in epidemic proportions is commonly sporadic, suddenly appearing for a year or
more then practically disappearing for several years, probably being kept under control by their
natural parasite-predator-disease complex. Some of the most destructive species are aliens
which were accidentally introduced from abroad and found a vacuum for development without
natural enemies. Many parasites have been introduced to help combat these pests. Some of the
more important sawflies are those that feed on timber species, roses, apples, pears, peaches, and
wheat and other grain crops. On the other hand, a few sawflies have been used beneficially, or at
least have been attempted for use in the biological control of weeds. One species, Ucona acaenae
Smith (=Antholcus varinervis of authors), was introduced into New Zealand from Chile in the
early 1930's to help control the spread of Acaena. This attempt was only partially successful.
Another species, Priophorus morio (Lepeletier), was released in Hawaii from the western
United States to help control the spread of Rubus on the islands, but this attempt failed.

The life cycle of many sawflies follows a similar pattern, though with many variations. Adults
of most species fly in spring and early summer and are very short-lived. It is difficult to keep in-
dividual adults alive in captivity for much more than a few days although the actual flight of a
species may extend over several weeks or months. Some adults may not feed but some may feed
on moisture, nectar and pollen of flowers, leaf pubescence, or other insects. The most productive
areas for collecting are in vegetation on edges of rivers or streams, marshes, open and scrubby
woodland, or undisturbed meadows. The catkins of willows and alders are especially productive
for many groups. Oviposition is in the foliage, stems, twigs, or wood, and the eggs are most al-
ways inserted in the plant tissue though some Pamphiliidae may glue their eggs to the leaf sur-
face. Larval feeding time varies, but usually lasts about two weeks. Many larvae, especially
Diprionidae, feed gregariously at first but later disperse to other parts of the host. After feed-
ing is completed the larva molts into a non-feeding stage called the prepupa or resting stage.
The prepupa normally leaves the host in search of a site for pupation. This may be in a cell in the
ground or litter, in a papery cocoon in the ground, in some other substance such as wood, stems
of the host or nearby plants, or fruits of other plants. The prepupal stage differs morphologically
from the feeding stages, especially in the shape of the mandibles and sometimes color pattern.
Some larvae with spines loose these in the final molt. If there is a single generation, the prepupa
will remain in its cocoon or cell the rest of the summer and pupate the following spring. How-
ever, depending on the species, latitude, or diapause requirements, there may be several genera-
tions a year, or it may take several years to complete the cycle.

The following references are to general articles on the Symphyta. Some, such as Maxwell,
1955, in morphology, pertain to many species and the reference is not repeated under each spe-
cies. Under some species, such as Pristiphora erichsonii (Hartig), a listing of all biological
references would take many more pages, and the references are selected or pertain to some
recent synopsis which contains a good bibliography. Some of the new distribution records, hosts,
and taxonomic changes are taken from unpublished notes. Only those taxonomic changes cited as
being new have not before appeared in the literature.

I am indebted to several colleagues for reviewing parts of this section and or providing some
of the information contained herein: H. E. Milliron, formerly with the Biosystematics Research
Institute, Agriculture Canada, Ottawa and other authorities of the Institute for allowing study
of their sawfly collection; H. R. Wong, Northern Forest Research Centre, Canadian Forestry
Service Edmonton, Alberta, Canada; and H. Greenbaum, Department of Entomology, University
of Arkansas Fayetteville.

Taxonomy: Norton, 1867-1869. Amer. Ent. Soc, Trans. 1: 31-84, 193-280; 2: 211-242, 321-368
(N. Amer. catalog). — Cresson, 1880. Amer. Ent. Soc, Trans. 8: 53-68 (N. Amer. catalog).
— Dalla Torre, 1894. Cat. Hym., v. 1, 459 pp. (world catalog). — Ashmead, 1898. Canad. Ent.
30: 141-145, 177-183, 205-213, 225-232, 249-257, 281-287, 305-316 (classification, keys to


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genera). — Konow, 1905. In Wytsman, Gen. Ins., fasc. 27, 27 pp.; fasc. 28, 14 pp.; fasc. 29,
176 pp. (world catalog). -Dyar, 1893-1895. Canad. Ent. 25: 244-248; 26: 42-45, 185-189; 27:
191-196, 208-212 (larvae). —Dyar, 1895 Amer. Ent. Soc, Trans. 22: 301-312 (larvae) -Dyar,
1897. N. Y. Ent. Soc, Jour. 5: 18-30, 190-201 (larvae) -Dyar, 1898. Canad. Ent. 30: 173-176
(larvae) -Dyar, 1898. N. Y. Ent. Soc, Jour. 6: 121-138 (larvae) -Dyar, 1900. N. Y. Ent.
Soc, Jour. 8: 26-31 (larvae). -Rohwer, 1911. Ent. Soc. Wash., Proc. 13: 215-224
(classification). -Rohwer, 1911. U. S. Dept. Agr., Bur. Ent., Tech. Ser. 20, pt. 2, pp. 69-109
(genotypes and work of Ashmead). —Rohwer, 1911 Ent. News 22: 218-219 (additions and
correction to "genotypes"). — Enslin, 1912-1918. Deut. Ent. Ztschr., Beih. 790 pp.
(Tenthredinoidea of middle Europe). — MacGillivray, 1913. Ent. Soc Ont., Ann. Rpt. 44:
54-75 (larvae). —MacGillivray, 1916. Conn. State Geol. and Nat. Hist. Survey Bui. 22:
25-175. -Yuasa, 1923. 111. Biol. Monog. 7(4): 1-168 (N. Amer. larvae). —Ross, 1937. 111. Biol.
Monog. 15(2): 1-173 (generic revision, N. Amer.). —Benson, 1938. Roy. Ent. Soc. London,
Trans. 87: 353-384 (world classification). —Benson, 1951. Handb. for Ident. of Brit. Ins., v.
6, pt. 2(a): 1-49; 1952, v. 6, pt. 2(b): 51-137; 1958, v. 6, pt. 2(C): 139-252. -Lorenz and
Kraus, 1957. Die Larvalsystematik der Blattwespen, 339 pp. (larvae of European species).
—Ross, 1960. Amer. Ent. Soc, Trans. 85: 315-321 (early history of sawfly study in N.
Amer.). —Benson, 1962. Brit. Mus. (Nat. Hist.) Ent., Bui. 12: 381-409 (Holarctic sawflies).
— Rasnitsyn, 1969. Origin and evolution of the lower Hymenoptera, 187 pp. (fossil).

Biology: Rohwer, 1915. Ent. Soc. Wash., Proc 17: 195-198 (mating habits). —Benson, 1950.
Soc. Brit. Ent., Trans. 10: 45-142 (natural history of Brit, sawflies). —Wong, 1954. Canad.
Ent. 86: 154-158 (sawflies on white birch in Man., Sask.). — Neilson, 1958. Canad. Ent. 90:
229-234 (life histories of sawflies in low-bush blueberry fields in N. B. ). — Raizenne, 1957.
Canada Dept. Agr. Pub. 1009, 45 pp. (forest sawflies of s. Ont. and their parasites).
— Lindquist, 1959. Canad. Ent. 91: 625-627 (leaf mining sawflies on birch). — Lindquist and
Miller, 1970. Ent. Soc. Ont., Proc. 100: 117-123 (free-feeding sawflies on birch and alder in
Ont.). -Lindquist and Miller, 1971. Ent. Soc. Ont., Proc. 102: 118-122 (larvae feeding on
spruce and balsam fir in Ont.).

Morphology: Marlatt, 1891. Ent. Soc. Wash., Proc. 2: 115-117 (final molting of larvae).
— Marlatt, 1894. Ent. Soc. Wash., Proc. 3: 78-82 (neuration of wings). —MacGillivray, 1906.
U. S. Natl. Mus., Proc 29: 569-654 (wings). —Van Dine, 1906. Hawaii. Ent. Soc, Proc. 1:
19-22 (comparative anatomical study of mouthparts). — Crampton, 1919. Ent. Soc. Wash.,
Proc 21: 129-155 (genitaHa and terminal abdominal structure of males and larvae).
— Middleton, 1921. Ent. Soc. Wash., Proc. 23: 139-144 (terminal abdominal structures).
— Middleton, 1921. Ent. Soc. Wash., Proc 23: 173-192 (suggested homologies between
larvae and adults). — Boulange, 1924. Mem. et Travaux des Facultes Catholiques de Lille
28: 1-444 (genitalia). —Taylor, 1931. Roy. Phys. Soc, Proc. 22: 41-70 (morphology of the
tenthredinid head). -Taylor, 1931. Ent. Soc. Amer., Ann. 24: 451-466 ("Dyar's Rule", its
application to sawfly larvae). — Malloch, 1936. Inaugural- Dissertation, Erlangung der
Doktorwurde, Philosophischen Fakultat der Friedrich- Wilhelms-Universitat zu Berlin, 55
pp. (thorax and cenchri). —Parker, 1934. Bol. Lab. Zool. Gen. e Agr. Portici 28: 159-191
(anatomy of larvae, with special reference to head). —Ross, 1936 Ent. Soc. Amer., Ann. 29:
99-111 (ancestry of wing venation). —Ross, 1945. Ent. News 56: 261-268 (genitalia:
terminology and study techniques). —Maxwell, 1955. Canad. Ent. 87 (sup. 1): 1-132
(comparative internal larval anatomy). — Arora, 1956. Panjab Univ., Res. Bui., Zool. 90:
85-119 (relationship of Symphyta to other insect orders based on adult external
morphology). — Togashi, 1965. Kontyu 33: 230-234 (rectal papillae). — Togashi, 1970. Mushi
43 (sup.): 1-114 (comparative morphology of internal reproductive organs). — Kenchington,
1972. Jour. Ent. 46: 111-116 (variations in silk gland morphology among larvae).


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Superfamily Megalodontoidea

• Family Xyelidae
• Family Pamphiliidae

Superfamily Tenthredinoidea

• Family Pergidae
• Family Argidae
• Family Cimbicidae
• Family Diprionidae
• Family Tenthredinidae

Superfamily Siricoidea

• Family Syntexidae
• Family Siricidae
• Family Xiphydriidae
• Family Orussidae

Superfamily Cephoidea

• Family Cephidae

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Superfamily Ichneumonoidea

By Paul M. Marsh and Robert W. Carlson

Present data on numbers of species ranks the Ichneumonoidea as the largest superfamily of
Hymenoptera. It includes the bulk of the larger Parasitica, most ichneumonoids being over 5 mm
in length and a few being longer than 50 mm. A greater percentage of Ichneumonoidea than
other Parasitica have the ovipositor conspicuously exposed, but, on the other hand, a very sig-
nificant portion of Ichneumonoidea have ovipositors that protrude scarcely or not at all beyond
the median dorsal extremeties of the apical tergites. Townes (1975) discussed the Hymenoptera
(mostly Ichneumonoidea) with the longest ovipositors; he cited a species of IphiaidaxV
(Braconidae) with an ovipositor 14 times the length of the body.

The distinguishing characters of Ichneumonoidea are: the usual fusion of the costal and sub-
costal veins of the fore wing; the long antennae, which are usually more than 14-segmented; and
the 2-segmented hind trochanters (the other trochanters usually also being 2-segmented). The
characters are for the most part shared by the Braconidae, Aphidiidae, Hybrizontidae, Ichneu-
monidae, and Stephanidae, the five families here considered to comprise the Ichneumonoidea. In
the case of the Stephanidae, however, there is some question about the correctness of placement
in the Ichneumonoidea (see Townes, 1969, p. 3). The larval head capsule of stephanids is con-
siderably different from those of other ichneumonoid families (personal commun., J. R. T. Short,
1976), and in stephanids the costa and subcosta are more distinctly separated than in other
ichneumonoid families. Nevertheless, we believe it best to leave the Stephanidae in the Ichneu-
monoidea until further studies are made.

The Aphidiidae and Hybrizontidae are sometimes treated as subfamilies of Braconidae (see
van Achterberg, 1976). Some early 19th Century authors referred to the combination of the
latter three groups as the "Ichneumonidum adscitorum" (?unauthentic Ichneumonidae) which
they distinguished from the "Ichneumonidum genuinorum" (genuine Ichneumonidae). The
distinction largely resulted from Jurine's (1807) classification of the veins and cells of hymenop-
terous fore wings and his provision of terms for some of the veins and cells (e.g. "nervi recur-
rentes"). Eady (1974) provided an excellent discussion of the way in which the Jurinean system
of wing vein and cell nomenclature was modified and expanded by those who adopted Jurine's
ideas. He reviewed the systems of wing vein nomenclature which are currently used for
Braconidae and compared them with usages for Aphidiidae, Ichneumonidae, and other
Hymenoptera. He proposed an "interim method [in order] to overcome the more frequently vo-
iced objections to ... [the Comstock-Needham] system without adding to the confusion or ob-
structing progress toward uniformity."

It was apparently by mutual agreement that Gravenhorst (1819) and Nees ab Esenbeck (1819)
decided to specialize on the Ichneumonidorum genuinorum and Ichneumonidorum adscitorum,
respectively. In the papers referred to, they simultaneously outlined their plans for the mono-
graphs which are here cited in the sections to which they pertain. Thunberg (1822, 1824) chose to




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ignore the revolutionary advances in classification made possible by the work of Jurine and
reverted to lumping all of the Ichneumonoidea under the generic name Ichneumon (see in-
troduction to Ichneumonidae). Consequently, Thunberg's work was largely ignored prior to
Roman's (1912) study of his type specimens.

The actual number of species in the Ichneumonoidea can only be estimated. The Braconidae
contains about 2,000 described species in North America and about 10,000 worldwide; the Ichne-
umonidae about 3,000 in North America and about 15,000 worldwide. However, the total number
of species is estimated to be 60,000 worldwide in the Ichnuemonidae (Townes, 1969, p. 7) and
40,000 in the Braconidae.

Except for Hybrizontidae and Stephanidae, the families of Ichneumonoidea occur in all zoo-
graphical regions and in all terrestrial habitats. In the Ichnuemonidae the Western Palearctic
fauna is best known followed by the Nearctic, whereas the reverse seems to be true for the
Braconidae. As in the case of Chalcidoidea, most of our knowledge of the Ichneumonoidea has
been derived from species of economic importance to agriculture. For the vast majority of spe-
cies, there is little or no knowledge of biology.

The ichneumonoids are parasitic on nearly all groups of insects as well as on spiders, and all
stages of these hosts are attacked. Aphidiidae, many Braconidae, and possibly Hybrizontidae
(hosts of latter unknown) attack paurometabolous insects, while no paurometabolous hosts are
known for Ichneumonidae or Stephanidae. The only ichneumonoids which attack adults of
holometabolous insects are certain euphorine and blacine Braconidae. Aside from the limitations
which have been mentioned, large numbers of Ichneumonoidea are polyphagous and the limits of
the host range seem to be related more to the host habitat than to the taxonomy of the host.

The only families of Ichneumonoidea which are known to include hyperparasitic species are
Ichneumonidae and Braconidae, but only a very few Braconidae could be regarded as hyper-
parasitic (i.e. a few Euphorinae which attack adult Ichneumonidae). The fact that hyper-
parasitism is much more prevalent in the Ichneumonidae than in the Braconidae is explained lar-
gely (but not in the case of mesochorine and eucerotine Ichneumonidae) by the fact that certain
Ichneumonidae have the habit of attacking hosts which are confined within silken cocoons (e.g.
sawfly prepupae, ichneumonoid prepupae, spider eggs, chrysopid eggs, etc.), while this habit has
not developed among Braconidae. Further discussion of host relations is deferred to the in-
troductions to the various taxa.

Taxonomy: Jurine, 1807. Nouv. Meth. Class. Hym. Dipt. v. 1, 324 p. and 14 pi. — Gravenhorst,
1819 (1818). Nova Acta Leopoldina 9: 281-298. — Nees ab Esenbeck, 1819 (1818). Nova
Acta Leopoldina 9: 299-310. — Thunberg, 1822; 1824. Acad. Imp. des Sci. St. Petersburg,
Mem. 8: 249-281 (key); 9: 285-368. -Ashmead, 1900. U. S. Natl. Mus., Proc. 23: viii and 220
p. (classification of Ichneumonoidea and Evanioidea). —Roman, 1912. Zool. Bidr. Uppsala
1: 231-293. -Viereck, 1914. U. S. Natl. Mus., Bui. 83: v and 186 p. (type-species of
Ichneumonoidea and Evanioidea). —Viereck, 1921. U. S. Natl. Mus., Proc. 59: 129-150 (sup.
to Viereck, 1914). —Townes, 1969. Amer. Ent. Inst., Mem. 11: 2-7. — Eady, 1974. Jour. Ent.
(B) 43: 63-72. —Townes, 1975. Ent. News 86: 123-127. —van Achterberg, 1976. Tijdschr. v.
Ent. 119: 33-78.

• Family Braconidae
• Family Aphidiidae
• Family Hybrizontidae
• Family Ichneumonidae
• Family Stephanidae

Superfamily Chalcidoidea

By Gordon Gordh

The Chalcidoidea are among the most difficult groups of Hymenoptera to identify because of
their small size and the lack of adequate keys to the North American species. Several species vie
for the distinction of being the smallest insect (about 0.2 mm long), and most species are less
than 3-4 mm long. The characters used to distinguish chalcidoids from other Hymenoptera in-
clude the presence of a prepectus, failure of the pronotum to meet the tegula, 13 or fewer seg-
ments comprising the geniculate antenna, and drastically reduced wing venation.

Ashmead (1904) recognized 14 families and provided the first comprehensive modern classifi-
cation of the Chalcidoidea. Despite its many errors, this was a prodigious work and remarkable
considering the primitive optical equipment and state of knowledge about the Chalcidoidea at
that time. Nikol'skaya (1952) elevated the number of families to 24, and Boucek and Hoffer
(1957) (subsequently translated by Peck, 1964) recognized 18 families. In this catalog Burks has
reduced the number of families to eleven. Casual thought may lead one to wonder why there is
so much inconsistency among workers regarding higher classification of chalcidoids. These clas-
sifications are based on external morphology, and the chalcidoids are exceedingly plastic
morphologically. This plasticity generates differences of opinion over the limits of higher taxa
because workers weight characters differently.

Chalcidoids are found in all zoogeographical regions, in all terrestrial habitats, and all families
are found in each zoogeographic region. Despite their omnipresence, chalcidoids remain one of
the poorest known superfamilies. Taxonomically, the western Palearctic fauna is best known,
followed by the Nearctic. The remainder of the zoogeographical regions (Neotropical, Ethiopian,
Australian, and Oriental) are almost completely unknown with respect to their endemic faunas.
Much of our knowledge of chalcidoids stems from species which are associated with agriculture.

The body size and searching habits of chalcidoids make them suitable for fossilization in
resinous amber, but fossil records of the Chalcidoidea are incomplete. Fewer than 50 species are
known, and these belong to less than half of the chalcidoid families. The most comprehensive ac-
counts of fossil chalcidoids are by Brues (1910), Doutt (1973), and Yoshimoto (1975). The last
study provides a summary of knowledge about fossil chalcidoids. Yoshimoto (1975) reports that
mymarids, trichogrammatids, and tetracampids are referable to the Cretaceous Period (70-90
million years before present).

Owing to the paucity of knowledge about fossil chalcidoids and their morphological plasticity,
the relationship of this superfamily to other parasitic Hymenoptera has not been conclusively
established. We are not certain that the Chalcidoidea are monophyletic in the Hennigian sense,
although some investigators have that opinion. An interpretation of chalcidoid phylogeny based
on the known fossil record is provided by Yoshimoto (1975).

The actual numerical dimension of the Chalcidoidea can only be speculated. The ichneumonid
specialist Henry Townes (1969) has estimated that there are 60,00() species of Ichneumonidae.




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DeBach (1974) has estimated that somewhere between 70-90 percent of the parasitic Hymenop-
tera remain to be described. I believe that the Chalcidoidea wiir ultimately be recognized as
larger than the Ichneumonidae. There are some who would disagree with this estimate, but their
estimates are based on impressions developed from examining species that repeatedly have
been submitted for identification. These species mostly are associated with the agroecosystem
and represent only a portion of the total chalcidoid fauna. My primary interest in making these
assertions is to stimulate research on the Chalcidoidea because they are a fertile area for in-
vestigations in biology, behavior, ecology, and systematics.

Chalcidoids have diverse and frequently specialized feeding habits. Most species of chalcidoids
are parasitic, but phytophagy probably has evolved several times in the Chalcidoidea because it
is found in several distantly related taxa and many unrelated species of plants serve as hosts.
Phytophagy is found most frequently in association with gall-forming habits, but the evolutiona-
ry significance of this observation remains unknown.

Agaonids demonstrate the most intimate expression of phytophagy in the Chalcidoidea. This
group is poorly represented in North America because all agaonids develop in fig seeds (Ficus
spp.), and these plants occur naturally only in tropical and subtropical climates. All figs are de-
pendent on agaonids for pollination, and agaonids can only develop within the receptacles of
Ficus. Host specificity seems to be the trend in agaonids with each species of fig having its own
agaonid for pollination (Ramirez, 1970 a,b; Grandi, 1961). Numerous other chalcidoids are as-
sociated with Ficus as inquilines (Hill, 1967 a,b).

Other taxa of chalcidoids with phytophagous species include the Eurytomidae, Torymidae,
brachyscelidiphagine Pteromalidae, and Tanaostigmatidae.

Some ecologists prefer to use the term parasitoid to characterize parasitic insects. Protelean
parasite is a phrase often used to distinguish between typical parasites and insects that are
parasitic in the larval stage only (Askew, 1971).

One definition of parasitism for all parasitic organisms is impractical because animal species
are parasitic in many different ways. The parasitological definition of parasitism in the sense of
parasitic worms and protozoa is unsuitable in the present context because parasitic chalcidoids
do not behave in a manner consistent with that definition. Therefore it seems more appropriate
to list some of the biological attributes of parasitic chalcidoids. Parasitic chalcidoids are charac-
terized as follows: (1) they are obligate parasites in the larval stage only; (2) they require only
one host to complete development; (3) they attack related taxa (other arthropods and usually in-
sects); (4) if the parasite completes development, the host invariably dies; (5) the ratio of size
between the parasite and host approximates unity (except in some cases where the parasitic lar-
vae are gregarious or polyembryonic delevopment occurs).

Adults of some species host feed, but the significance of this behavior is not always clear.
Host feeding may provide nutrients necessary for ovary or egg development, or it may be a con-
venient source of nutrients necessary for sustaining life (Flanders, 1953; Doutt, 1964; Quezada et
ai, 1973).

Parasitism by insects reaches its most elaborate development in the Chalcidoidea. Primary
parasitism (larval development on a phytophagous host) is the most common type of parasitism
by chalcidoids. Hyperaparasitism (a parasite attacking another species of parasite) is found al-
most exclusively in the Hymenoptera, and reaches its most extensive development in the Chalci-
doidea as indicated by the fact that most families have hyperparasitic species. Further evidence
of the extensiveness of hyperparasitism in this superfamily is found in the fact that several
types of hyperparasitism have evolved in the group. These include secondary (a parasite at-
tacking a primary parasite), tertiary (a parasite attacking a secondary parasite) and quaternary
(a parasite attacking a tertiary parasite). Hyperparasitism probably evolves out of primary
parasitism in situations involving strong interspecific competition.

An unusual type of hyperparasitism occurs in Coccopkagoides utilis Doutt and various related
genera such as Coccophagus, Eyicarsia, and Prospaltella. Female larvae develop as primary
parasites of armored-scale insects, and the male larvae develop as hyperparasites of their own
females (Broodryk and Doutt, 1966). This phenomenon is called adelphoparasitism or au-
toparasitism and appears restricted to the aphelinines (Zinna, 1961; Flanders, 1959, 1967).

Parasitic chalcidoids can be categorized on the basis of where the egg is deposited and how
the larva feeds. Most species attack the host directly, but adult female eucharitids and perilam-
pine pteromalids oviposit on vegetation and the first-instar larva (planidium) searches for the
host (Smith, 1912; Clausen, 1940 a,b). Species in which the adult female directly attacks the host







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lay their eggs on the host's body and the larvae develop externally, or deposit their eggs inside
the host's body and the larvae develop internally. There is a tendency for parasites that attack
exposed hosts to develop internally (exception: elachertine Eulophidae), and parasites that at-
tack concealed hosts to develop externally.

The intra- and interspecific relationships among parasitic chalcidoids vary. Some species are
solitary (one parasite per host), and others are gregarious (several parasites per host). When
more than one parasite species develops on a host simultaneously, the condition is termed multi-
ple parasitism. When more eggs of one parasite species are laid on a host than can develop to
maturity, the condition is termed superparasitism. Supernumerary individuals are eliminated
through larval combat or physiological suppression (Salt, 1961).

The distinction between parasitism and predation sometimes fails, and some chalcidoids could
be called predators. A prime distinction between parasites and predators is that predators
frequently consume several prey, but parasites consume only one host per individual. The eu-
notine pteromalids and some mymarids could be regarded as egg predators because their larvae
feed externally on scale-insect eggs in the "brood chamber" after they are oviposited by the
female scale-insect (Clausen, 1940 a).

Parasitic species that attack many species of hosts are called polyphagous; parasitic species
that attack only a few species of hosts are called stenophagous; and parasitic species that attack
only one species of host are monophagous. Complete host specificity is difficult to establish
because it is based essentially on negative evidence. The fact that a parasite will not attack a
host under some conditions does not constitute proof that it will not parasitize that species.
Nevertheless, there is a tendency towards specialization in the Chalcidoidea, and this is reflected
by: (1) repeated recovery of a parasite from a host species over a large area, but not from re-
lated host species that occur sympatrically; (2) demonstrated preference for a host species when
a choice is available; (3) superior reproductive capability on a host species; and (4) physical
limitations that prevent a parasite from attacking a potential host.

Some polyphagous chalcidoids appear to prefer habitats rather than a taxonomically cohesive
group of hosts. For example, Zagravimosovia species parasitize leaf-mining insects whether
they are Lepidoptera, Diptera, or perhaps Hymenoptera. In contrast, related Diglyphus species
parasitize only leaf-mining agromyzid Diptera. Other chalcidoids are extremely polyphagous.
Dibrachys cavus (Walker) is an example. This species, like several others, has an exceedingly
long host list that includes representatives of several orders. It usually develops as a primary
parasite, but frequently also acts as a facultative hyperparasite (Graham, 1969). No explanation
has been provided as to why one species should be so polyphagous and a closely related,
morphologically similar species should be stenophagous or even monophagous.

Likewise, there are associations between host stage attacked and the taxonomic assignment of
the parasite. For instance, the Trichogrammatidae and Mymaridae exclusively develop on the
egg stage of other insects and the spalangine pteromalids are pupal parasites (Annecke and
Doutt, 1961; Boucek, 1963; Doutt and Viggiani, 1968).

Chalcidoids parasitize more hosts in more different taxonomic categories than any other
group of parasitic insects. This spectrum extends from spider eggs (Desantisca) to aculeate
Hymenoptera {Melittobia, Leucospidae). A detailed account of the biology of chalcidoids
requires more space than is available here. However, a short summary of some interesting host
relationships is provided.

A bizarre host association is found in Ixodiphagus and Hunterellus (Encyrtidae) whose spe-
cies are primary, internal parasites of tick larvae and nymphs. These genera are cosmopolitan
and may prove to be beneficial insects in tick control (Cooley and Kohls, 1934; Cole, 1965; Doube
and Heath, 1975).

The mymarid Caraphractus cinctus Walker is unusual in that it parasitizes dytiscid beetle
eggs that are submerged beneath the surface of the water. The female parasite swims in the
water by vibrating her wings and oviposits in the host's eggs. Females have considerable dis-
criminative ability, and can detect eggs that have been parasitized (Jackson, 1958, 1966).

The Eucharitidae are parasitic on Formicidae. The association apparently is an old one, and
eucharitids oviposit on vegetation visited by worker ants. The eggs hatch, and the triungulin lar-
vae are phoretically transported to the ant nest. Inside the nest the triungulin larvae eventually
move into the brood chamber where they parasitize immature ants (Clausen, 1923; 1940 b,c).

Other information about host association of chalcidoids is limited by a lack of knowledge about
the immature stages of many groups of potential hosts. However, the higher taxonomic catego-




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ries that include the most host species for chalcidoids include Lepidoptera, Homoptera, Diptera,
Coleoptera, and Hymenoptera. Chalcidoids generally have failed to adapt to the nymphal stage
of paurometabolous insects. The host spectrum of chalcidoids is being expanded constantly by
more comprehensive biological studies of other insects. Given the diversity of habits, host as-
sociations, and stages attacked, it seems reasonable to conclude that any insect potentially in-
cludes several niches where a chalcidoid can develop.

All known Hymenoptera develop parthenogenetically and chalcidoids demonstrate three
types: arrhenotoky, thelytoky, and deuterotoky. Arrhenotoky is the most common type of
parthenogenesis among chalcidoids. Uninseminated arrhenotokous females deposit haploid eggs
that develop into hemizygous males. Inseminated arrhenotokous females produce female off-
spring from fertilized eggs and males from unfertilized eggs. Arrhenotoky is a mechanism
whereby lethal and deliterious genes can be relatively rapidly eliminated from a population and
superior genotypes can be relatively rapidly selected.

Thelytoky is parthenogenesis in which males are unknown or rare and females produce
females by various asexual mechanisms. Cytologically, diploidy is maintained by apomixsis and
automixsis. Apomixsis (ameiotic thelytoky) is characterized by an absence of meiosis, and
chromosome number is not reduced. Automixis (meiotic thelytoky) has reduction divisions, and
diploidy is maintained in several ways. Rossler and DeBach (1973) review the methods of main-
taining a constant chromosome number.

Thelytoky is common among parasitic Hymenoptera, but the extent of thelytoky in the Chalci-
doidea is not known because our knowledge of their biology is limited. Many species are known
from the original description only, and many species have been described from the female sex
only. Thelytoky may be more common than now realized. In the rather well known genus
Aphytis, DeBach (1969) records that about 30% of the species are thelytokous.

The evolutionary significance of thelytoky is an issue of debate. Traditional views hold that
thelytoky is an "evolutionary blind alley". However, Rossler and DeBach (1972) have shown that
at least one species of thelytokous chalcidoid has females that are capable of sexual reproduc-
tion.

Deuterotoky is parthenogenesis in which unfertilized eggs develop into both sexes. The
cytological mechanism of deuterotoky has not been examined in chalcidoids. This form of
parthenogenesis is common in some other animals, and has been reported in some species of
chalcidoids (Doutt, 1959).

The cytogenetics of the Hymenoptera have been reviewed by Crozier (1975). That paper
points to a lack of knowledge developed about chalcidoid karyotypes and cytological phenomena.

Hymenoptera are haplodiploid and this has been confused with sex determination. The cor-
relation between males being haploid and females being diploid is positive and strong, but
haploidy and diploidy in themselves do not determine sex. Diploid males are known to occur
(Whiting, 1945). Several theories have been advanced to explain sex determination in the
Hymenoptera, but in no instance has one theory proven adequate to explain determination in all
groups (Whiting, 1940, 1943; daCunha and Kerr, 1957; Slobodchikoff and Daly, 1971). Crozier
(1975) suggests that any general theory should accommodate the multiple allele case with as lit-
tle modification as possible.

Polyembryony is a cytological phenomenon in which a single egg develops into many in-
dividual progeny. Among Hymenoptera the process occurs in the Platygastridae
(Proctotrupoidea) and copidosomatine Encyrtidae (Silvestri, 1906; Leiby, 1922, 1926).

Sex ratio in many species of animals approximates unity. In arrhenotokous chalcidoids the sex
ratio usually is female biased and fluctuates between 60 and 80 percent female. Numerous fac-
tors have been implicated in the determination of sex ratio including size, stage, or species of
host (Abdelrahman, 1974 a,b; Avidov and Podoler, 1968; Clausen, 1940 a), rate of oviposition
(Abdelrahman, 1974 b), egg orientation (King, 1961), genetic factors (Wilkes, 1964), differential
mortality (Roberts, 1933; Flanders, 1937; Abdelrahman, 1974 a), density fluctuations (Flanders,
1956), nutrition (Flanders, 1965; Moran et al., 1969), and many others. This Hst could be
lengthened substantially and its only limitation now is lack of research.

Mayr (1969) defines sibling species as "pairs or groups of closely related species which are
reproductively isolated but morphologically identical or nearly so." Recent studies of chalcidoids
have demonstrated that this group has many sibling species complexes (Hafez and Doutt, 1954;
Claridge and Askew, 1960; DeBach, 1959, 1960, 1969; Khasimuddin and DeBach, 1976, a,b,c; Rao
and DeBach, 1969 a,b,c). These complexes suggest that chalcidoids are in an active state of




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evolution and speciating rapidly. Factors of chalcidoid biology that promote rapid speciation in-
clude: (1) short generation time; (2) several generations per season; (3) intensive inbreeding via
sib mating; (4) microgeographic isolation; and (5) host preference.

Chalcidoids are the most important group in applied biological control. Other taxa (Tachinidae,
Ichneumonidae, Braconidae, Proctotrupoidea) are used extensively in biological control, but spe-
cies-for-species chalcidoids have been used more successfully. DeBach (1964) lists 25 pest species
with which complete biological control was achieved. Chalcidoids are responsible for control in
13 of these programs, a number far greater than any other taxonomic group. Agricultural pests
in these control programs include many Homoptera, but some Coleoptera have also been con-
trolled by chalcidoids (Taylor, 1937; Tooke, 1953; Williams et a/., 1951).

It is a pleasure to acknowledge the comments and suggestions on the preceding account made
by the following individuals: Kenneth Cooper, Paul DeBach, Eric Grissell, Peter Price, and
David Rosen.

Taxonomy: Ashmead, 1904. Carnegie Mus., Mem. 1 (4): 225-555. — Brues, 1910. N. Y. Ent.
Soc, Jour. 18 (1): 1-22 (Fossils). — Nikol'skaya, 1952. Akad. Nauk SSSR 44, 593 pp.
—Boucek and Hoffer, 1957. Klic Zuireny CRS 2: 208-288. —DeBach, 1959. Ent. Soc. Amer.,
Ann. 52 (4): 354-362. -DeBach, 1960. Ent. Soc. Amer., Ann. 53 (6): 701-705. -Annecke and
Doutt, 1961. Rep. So. Africa Dept. Agr. Tech. Serv., Ent. Mem. 5: 1-71. —Boucek, 1963.
Mus. Natl. Pragae, Acta Ent. 35: 429-512. —Peck, Boucek, and Hoffer, 1964. Canad. Ent.
Soc, Mem. 34: 120 pp. —Doutt and Viggiani, 1968. Calif. Acad. Sci., Proc. (4) 35 (20):
477-586. — Townes, 1969. Amer. Ent. Inst., Mem. 11: 1-300. —Graham, 1969. Brit. Mus.
(Nat. Hist.) Ent., Bui. Sup. London, 16: 908 pp. —Doutt, 1973. Pan-Pacific Ent. 49 (3):
221-228 (fossils). -Yoshimoto, 1975. Canad. Ent. 107 (5): 499-528 (fossils).

Biology: Silvestri, 1906. Lab. Zool. Gen. e Agr. Portici, Bol. 1: 17-64. -Smith, 1912. U. S.
Dept. Agr., Tech. Ser. 19 (4): 33-69. -Leiby, 1922. Jour. Morph. 37: 195-285. —Clausen,
1923. Ent. Soc. Amer., Ann. 16 (3): 195-219. -Leiby, 1926. Ent. Soc. Amer., Ann. 19 (3):
290-299. —Roberts, 1933. U. S. Dept. Agr., Tech. Bui. 365: 1-21. — Cooley and Kohls, 1934.
5th Pacific Sci. Congr., Proc. 5: 3375-3381. —Flanders, 1937. Calif. Univ. Pubs., Ent. 6 (15):
401-422. —Taylor, 1937. Biol. Cont. Ins. Fiji 239 pp. —Clausen, 1940 a. Entomoph. Insects,
688 pp. -Clausen, 1940 b. Ent. Soc. Wash., Proc. 42 (8): 161-170. -Clausen, 1940 c. Wash.
Acad. Sci., Jour. 30: 504-516. -Whiting, 1940. Jour. Morph. 66 (2): 323-355. -Whiting, 1943.
Genetics 28: 365-382. —Whiting, 1945. Quart. Rev. Biol. 20: 231-260. -Williams et ai,
1951. Bui. Ent. Res. 42 (1): 23-28. -Tooke, 1953. Union So. Africa Dept. Agr., Ent. Mem. 3,
282 pp. —Flanders, 1953. Jour. Econ. Ent. 46 (4): 541-544. —Hafez and Doutt, 1954. Canad.
Ent. 86 (2): 90-96. —Flanders, 1956. Insectes Sociaux 3 (2): 322-334. — daCunha and Kerr,
1957. Forma et Functios 1 (1): 33-36. —Jackson, 1958. Roy. Ent. Soc. Lond., Trans. 110
(17): 533-556. -Doutt, 1959. Ann. Rev. Ent. 4: 161-182. -Flanders, 1959. Ent. Expt. et
Apl. 2 (2): 125-142. — Claridge and Askew, 1960. Entomophaga 5 (2): 141-153. — Grandi,
1961. Univ. Bologna Inst. Ent., Bol. 26: 1-13. -King, 1961. Nature 189 (4761): 330-331.
—Salt, 1961. Soc. Expt. Biol., Symp. 15: 96-119. — Zinna, 1961. Lab. Ent. Agr. "Filippo
Silvestri", Portici, Bol. 19: 301-358. —DeBach, 1964. in: Biol. Control Ins. Pests Weeds, pp.
673-713. —Doutt, 1964. in: Biol. Control Ins. Pests Weeds, pp. 145-167. —Wilkes, 1964.
Science 144 (3616): 305-307. —Cole, 1965. WHO-EBL 43: 65. —Flanders, 1965. Amer. Nat.
99: 489-494. — Broodryk and Doutt, 1966. Hilgardia 37 (9): 233-254. -Jackson, 1966. Roy.
Ent. Soc. Lond., Trans. 118 (2): 23-49. -Flanders, 1967. Entomophaga 12 (5): 415-427.
—Hill, 1967 a. Figs (Ficus spp.) of Hong. Kong, 130 pp. —Hill, 1967 b. Jour. Nat. Hist. 1:
413-434. — Avidov and Podoler, 1968. Israel Jour. Ent. 3 (1): 1-16. — Mayr, 1969. Principles
Syst. Zool., 428 pp. — Moran et al., 1969. Roy. Ent. Soc. Lond., Trans. 121 (2): 41-58.
—DeBach, 1969. Israel Jour. Ent. 4: 11-28. —Rao and DeBach, 1969 a. Hilgardia 39 (19):
515-553. -Rao and DeBach, 1969 b. Hilgardia 39 (19): 555-567. -Rao and DeBach, 1969 c.
Evolution 23: 525-533. —Ramirez, 1970 a. Univ. Kans., Sci. Bui. 49 (1): 1-44. —Ramirez,
1970 b. Evolution 24 (4): 680-691. —Askew, 1971. Parasitic Insects, 316 pp. — Slobodchikoff
and Daly, 1971. Amer. Zool. 11: 273-282. — Rossler and DeBach, 1972. Entomophaga 17 (4):
391-423. —Rossler and DeBach, 1973. Hilgardia 42 (5): 149-176. — Quezada, DeBach and
Rosen, 1973. Hilgardia 41 (18): 543-604. — Abdelrahman, 1974 a. Austral. Jour. Zool. 22 (2):
213-230. —Abdelrahman, 1974 b. Austral. Jour. Zool. 22 (2): 231-247. —DeBach, 1974. Biol.
Control by Nat. Enemies, 323 pp. — Crozier, 1975. Hymen. Anim. Cytogen. 3: Ins. 7, 95 pp.




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—Doube and Heath, 1975. Jour. Med. Ent. 12 (4): 443-447. — Khasimuddin and DeBach,
1976 a. Entomophaga 21 (1): 81-92. —Khasimuddin and DeBach, 1976 b. Entomophaga 21
(1): 113-122. —Khasimuddin and DeBach, 1976 c. Ent. Soc. Amer., Ann. 69 (1): 15-20.

• Family Torymidae
• Family Pteromalidae
• Family Eurytomidae
• Family Chalcididae
• Family Leucospididae
• Family Eucharitidae
• Family Eupelmidae
• Family Encyrtidae
• Family Eulophidae
• Family Mymaridae
• Family Trichogrammatidae

Superfamily Cynipoidea

By B. D. Burks

The Cynipoidea have gone through an evolutionary development that approximately parallels
that of the Chalcidoidea. Both superfamilies have parasitic and phytophagous forms, although
the Cynipoidea does not include known genera that have phytophagous as well as parasitic spe-
cies. In the Chalcidoidea several genera include phytophagous and parasitic species; in the Cyni-
poidea no such mixtures occur below the family level. It should be borne in mind, however, that
the present classification of the Cynipoidea may have given unjustifiable weight to the habits of
the forms placed in the various higher categories. In any case, in each superfamily it is debatable
whether the phytophagous or parasitic habit is the more primitive. Although all authorities
agree that the members of each superfamily came originally from phytophagous ancestors, there
is disagreement as to whether the first Hymenoptera recognizable as representing either super-
family were parasitic or phytophagous. If they were parasitic, as many authorities believe, the
present-day phytophagous species are secondarily evolved from parasitic ancestors. The close
agreement in essential morphological characters among all members of each superfamily ex-
cludes the possibility of polyphyletic origin for the parasitic and phytophagous forms.

Revision: Dalla Torre and Kieffer, 1910. Das Tierreich, 24, Hymenoptera, Cynipidae, 891 pp.
(world monograph, literature through 1905). —Weld, 1952. Cynipoidea 1905-1950. Ann
Arbor, Mich., Privately Printed, 351 pp. (supplement to Dalla Torre and Kieffer;
illustrated keys).

Taxonomy: Felt, 1940. Plant Galls and Gall Makers, pp. 78-228; 254-269; 329-338 (illustrated
keys to galls). —Weld, 1957. Cynipid Galls of Pacific Slope. Ann Arbor, Mich., Privately
Printed, 80 pp. (illustrated keys to galls). —Weld, 1959. Cynipid Galls of the Eastern
United States, Ann Arbor, Mich., Privately Printed, 158 pp. (illustrated keys to galls).
—Weld, 1960. Cynipid Galls of the Southwest. Ann Arbor, Mich., Privately Printed, 51 pp.
(illustrated keys to galls).

Biology: Mani, 1964. Ecology of Plant Galls, 434 pp.

• Family Ibaliidae
• Family Liopteridae
• Family Figitidae
• Family Eucoilidae
• Family Alloxystidae
• Family Cynipidae

Superfamily Evanioidea

By Robert W. Carlson

This superfamily includes tiiree families, Evaniidae, Aulacidae, and Gasteruptiidae. There ap-
pears to be little evidence to indicate that the Aulacidae and Gasteruptiidae are actually related
to the Evaniidae. There does appear to be some evidence to indicate that the Aulacidae and
Gasteruptiidae are distantly related. Because there is no evidence to suggest that a common an-
cestor of Aulacidae and Gasteruptiidae would have been less closely related to ancestral
Evaniidae than to the ancestor of any other group, there is little reason for not maintaining
placement of the Aulacidae and Gasteruptiidae in the Evanioidea.

Revision: Kieffer, 1912. Das Tierreich, v. 30, XIX and 431 p. (spp. of world).

Ta.xonomy: Crosskey, 1951. Roy. Ent. Soc. London, Trans. 102: 282-301. — Muesebeck and
Walkley, 1956. U. S. Natl. Mus., Proc. 105: 319-419 (type-species).

Morphology: Crosskey, 1951. Roy. Ent. Soc. London, Trans. 102: 247-281.

• Family Evaniidae
• Family Aulacidae
• Family Gasteruptiidae

Superfamily Pelecinoidea

By Carl F. W. Muesebeck
Taxonomy: Handlirsch, 1933. Handbuch der Zoologie, v. 4 (2), Insecta 2, pp. 981-982.

• Family Pelecinidae

Superfamily Proctotrupoidea

By Carl F. W. Muesebeck

This is a large and diverse group of parasitic Hymenoptera, but the North American
proctotrupoid fauna has had little critical study with the result that apparently most of our spe-
cies remain undescribed. The only comprehensive treatments of the North American Proctotru-
poidea are that by Ashmead, in 1893, and that included by Kieffer in his monograph of the world
fauna, which comprises three volumes that were published between 1914 and 1926. Kieffer used
the name Serphoidea, based on Serphus Schrank, 1780, for this superfamily and that usage was
followed generally in the literature of the next thirty years; but Serphus Schrank was set aside
by the International Commission on Zoological Nomenclature in 1946 (Opinion 178), under
suspension of the Rules, and its synonym Proctotnipes Latreille, 1796, with P. brevipennis
Latreille as type-species, was added to the Official List of Generic Names in Zoology as Name
No. 616. As a result of that action the name Proctotrupoidea has come into general use again.

I am indebted to Dr. Lubomir Masner, Biosytematics Research Institute, Canada Department
of Agriculture, for much helpful advice and for new records, in connection with the preparation
of this section of the Catalog.

Revision: Ashmead, 1893. U. S. Natl. Mus., Bui. 45: 5-472 (the Bethylidae and Dryinidae were
included). -Kieffer, 1907-1914. In Andre, Spec. Hym. Eur. Alg., v. 10, pp. 1-1014 and v. 11,
pp. 1-447. -Kieffer, 1914-1926. Das Tierreich, Lief. 42 (1914), pp. VI-XVII, 1-254; Lief. 44
(1916), pp. VI-XXX, 1-627; Lief. 48 (1926), pp. VII-XXXVI, 1-885.

Taxonomy: Ashmead, 1887. Ent. Amer. 3: 73-76, 97-100, 117-119. —Harrington, 1899. Roy.
Soc. Canada, Trans. (2) 5: 117-206. -Ashmead, 1902-1903. N. Y. Ent. Soc, Jour. 10: 240-247
(1902); 11: 28-35 and 86-99 (1903). — Mani, 1941. Cat. Indian Ins., part 26, pp. 1-60.
— Risbec, 1950. Trav. Lab. d'Ent. du Sect. Soudanais de Recherches Agron., v. 2, pp.
511-637. -Muesebeck and Walkley, 1956. U. S. Natl. Mus., Proc. 105: 319-419 (Types of the
genera and subgenera). —Masner, 1965. Brit. Mus. (Nat. Hist.), Bui., Ent, Sup. 1, pp. 3-154.
—Masner, 1965. Psyche 72: 295-304. —Masner and Muesebeck, 1968. U. S. Natl. Mus. Bui.
270: 1-143. -Masner, 1969. Nat. Canad. 96: 775-784. -Kozlov, 1970. Ent. Obozr. 49:
203-226 (English transl.: Ent. Rev. 49: 115-127). —Kozlov, 1971. Soc. Ent. Unionis Sovet.,
Horae 54: 7-67.

Biology: Clausen, 1962. Entomophagous Insects, pp. 239-270.

Morphology: Snodgrass, 1941. Smithsn. Misc. Collect. 99 (14): 38-41. — Reid, 1941. Roy. Ent.
Soc. London, Trans. 91: 428-430. — Pschorn-Walcher, 1955. Schweiz. Ent. Gesell., Mitt. 28:
214-219.


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• Family Vanhorniidae
• Family Roproniidae
• Family Heloridae
• Family Proctotrupidae
• Family Diapriidae
• Family Scelionidae
• Family Platygastridae

Superfamily Ceraphronoidea

By Carl F. W. Muesebeck

Superfamily rank for this group was first proposed by Masner in 1956 (Acta Faun. Ent. Mus.
Nat. Pragae 1: 101), although without indication of basic differences from the Proctotrupoidea.
In 1967, however, Masner and Dessart (Inst. Roy. Sci. Nat. de Belg., Bui. 43: 1-33) defined the
superfamily in detail and recognized it as comprising two families, Ceraphronidae and
Megaspilidae, the latter being divided into the subfamilies Megaspilinae and Lagynodinae. This
classification appears to be sound. The members of this superfamily are remarkable for the pos-
session of two apical spurs on the anterior tibiae.

In his 1914 revision of this group Kieffer used the name Calliceratidae, based on Calliceras
Nees, 1834, since Ceraplivon Jurine, 1807, which had been considered the type-genus of the fami-
ly, was a junior homonym of Cemphrou Panzer, 1805. Kieffer was generally followed in the
literature of the succeeding thirty years. However, in its Opinion 174, issued in 1946, the Inter-
national Commission on Zoological Nomenclature suppressed Ceraphron Panzer, under suspen-
sion of the Rules, and validated Ceraphron Jurine with C. sulcatus Jurine, 1807, as type-species;
since then the names Ceraphron and Ceraphronidae have again come into general use.

Apparently species of this superfamily are largely hyperparasites, developing especially on
larvae of Aphidiidae, Braconidae, Ichneumonidae, Bethylidae, Dryinidae and Tachinidae. Some,
however, are evidently primary parasites of Aleyrodidae, various small Diptera (i. e., Cecido-
myiidae, Phoridae), certain neuropteroids and some cynipoid gall-makers; others occur in ant
nests where they presumably develop as parasites of certain myrmecophilous Diptera.

Revision: Ashmead, 1893. U. S. Natl. Mus., Bui. 45: 102-136 (North American species).
—Kieffer, 1907. In Andre, Spec. Hym. Eur. Alg., v. 10, pp. 5-261, 10 pis. (Old World
species). —Kieffer, 1909. In Wytsman, Gen. Ins., fasc. 94, 24 pp., 2 pis. (generic key and
assignment of species). —Kieffer, 1914. Das Tierreich, Lief. 42, pp. 63-238 (World fauna).
—Dodd, 1914. Roy. Soc. So. Austral., Trans. 38: 85-118 (Australian fauna). — Hellen, 1966.
Fauna Fennica 20: 3-45 (Finnish forms).

Taxonomy: Haliday, 1834. Ent. Mag. 1: 272. — HaHday, 1839. Hymenoptera Britannica, Hym.
Synopsis, p. ii. — Foerster, 1856. Hym. Stud., v. 2, pp. 27, 97-99. —Ashmead, 1903. N. Y.
Ent. Soc, Jour. 11: 33-35. —Masner, 1956. Acta Faun. Ent. Mus. Nat. Pragae 1: 101.
— Dessart, 1962. Soc. Roy. d'Ent. de Belg., Bui. et Ann. 98: 305-309 (key to genera).
—Masner and Dessart, 1967. Inst. Roy. Sci. Nat. de Belg., Bui. 43: 1-33.

Morphology: Reid, 1941. Roy. Ent. Soc. London, Trans. 91: 429-430. —Masner and Dessart,
1967. Inst. Roy. Sci. Nat. de Belg., Bui. 43: 6-25.

• Family Ceraphronidae
• Family Megaspilidae

Superfamily Trigonaloidea

By Robert W. Carlson

• Family Trigonalidae

pendently a number of times in all aculeate superfamilies. However, these biological distinctions
break down in most of the more primitive wasps belonging to the Bethyloidea and Scolioidea.
Many of these behave as true parasitoids in that the larval prey may be only temporarily para-
lyzed, occasionally several eggs may be laid on a single prey larva, and frequently no nest what-
ever is made, the prey being left in situ, or at most a crude cell may be constructed around the
subterranean prey as in most Tiphiidae and Scoliidae. Parasitism of the egg stage of the host is
known only among the Amiseginae (Chrysididae). Polyembryony is unknown, but parthenogene-
sis occurs in some Aculeata. Usually this is of the facultative kind as is found in social insects
such as some ants, vespid wasps and honeybees. It may be obligate in some aculeates, such as
the tiphiid wasp Methocha and some species of the bee genus Ceratina, where males are rare or
unknown.

The simplest kind of nest among the aculeates is made by the wasp dragging the paralyzed
prey into a crevice in or above ground or back into the prey's burrow; the opening is usually
sealed off by particles of the substrate to make a crude cell. A second type of nest is also made
in a pre-existing cavity, such as borings of beetle larvae in wood or twigs, or in old insect galls or
abandoned mud cells. The nest in this second type may be unicellular as in the first kind of nest,
or it may consist of a linear series of cells, each cell separated from its neighbor by a partition of
mud, wood chips, resin, masticated plant leaves, or other substances. A third kind of nest is ex-
cavated by the wasp or bee in the ground, in rotten wood, or in the soft pith of such shrubs as
sumac and elderberry. The subterranean nests are frequently unicellular but multicellular nests
in the ground, rotten wood or pith may have the individual cells arranged in a linear series or in
clusters with the individual cells sealed by a partition or closing plug of the substrate. Occa-
sionally a mud turret may be constructed over the entrance of subterranean nests. Next, there
are the nests constructed entirely from foreign materials. Usually these are above ground
although some Vespidae and Bombinae have subterranean nests. The nests of solitary species
may be made of mud, or a mixture of resin and pebbles, with the cells arranged in parallel tubes,
or in clusters or with separate but adjacent cells; some exotic social Vespidae make a mud en-
velope around combs of hexagonal cells. A number of social species make paper or carton nests
in which the nesting material consists of masticated wood fibers, bark or rotten wood. Finally,
there is the complex nest of the honeybees constructed from wax secreted from glands in the
abdomen of workers.

Aculeate larvae are normally cannibalistic if they come in contact accidentally. This tendency
is prevented in multicellular nests by the existence of partitions separating adjacent larvae.
However, some species make brood cells in which several larvae develop amicably without the
occurrence of cannibalism. Such nests have been reported for a few North American Isodontia
(Sphecidae) and Megachile (Megachilidae), and for many exotic Allodapini (Anthophoridae).

True sociality (eusociality) has arisen independently several times in the higher aculeates, the
Formicoidea, Vespoidea, Sphecoidea and Apoidea. All of the ants are eusocial or are social
parasites of other ants, but the majority of wasps and bees are solitary species. Wilson (1971)
considers that eusocial insects must possess three traits: "individuals of the same species
cooperate in caring for the young; there is a reproductive division of labor, with more or less
sterile individuals working on behalf of fecund individuals; and there is an overlap of at least
two generations in life stages capable of contributing to colony labor, so that offspring assist
parents during some period of their life." Presocial insects exhibit one or two of the above traits.
Solitary species have none of these traits.

Most solitary aculeates practice mass provisioning, that is, the egg is laid and a store of food is
placed in the cell with it, then the cell is closed; in many species the store of food is provided be-
fore oviposition. Some sphecoid wasps, such as many Bembicinae (Nyssonidae), have taken the
first step toward subsocial status by adopting progressive provisioning. This behavior is charac-
terized by placing the egg on a single prey specimen and not furnishing additional prey until the
egg has hatched or by the hatching of the egg before any food is provided; after hatching the
larva is fed daily or at intervals as required. Other aculeates, e.g., Moniaecera (Crabronidae),
some Andrena (Andrenidae) and Exomalopsis (Anthophoridae), have achieved the higher level
of communal status, in which several females use a common burrow entrance but presumably
maintain separate cells. A higher level of presocial behavior (quasisocial) is found rarely in some
exotic bees where two or more gravid females of the same generation cooperatively construct
and provision the cells. Some of our Halictidae have attained the semisocial stage which is
similar to the quasisocial except that unmated females of the same generation associate with a




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gravid female or females and care for the larvae of the latter. A semisocial colony may evolve
into a primitive eusocial colony as happens later in the season in some Angochlorella nests when
the colony becomes monogynous and only the workers forage for pollen. Another stage toward
the eusocial is the subsocial in which one female cares for her own larvae as in many Allodapini
(Anthophoridae) and young nests of Boynhus (Apidae); such a colony may later become truly
eusocial as happens when the first brood of Bombus workers ecloses and takes over the foraging
activities previously performed by the queen.

Taxonomy: Lanham, 1960. Ent. News 71: 85-86 (significance of hind tibial strigil in
classification). —Richards, 1972 (1971). Ent. Essays to Commemorate Retirement of Prof.
K. Yasumatsu, pp. 1-13, 10 figs, (thoracic spiracles in classification). —Brothers, 1975.
Kans. Univ. Sci. Bui. 50: 483-648, 101 figs., 7 tabs, (phylogeny, especially Mutillidae).

Biology: Walsh and Riley, 1869. Amer. Ent. 1: 122-143, figs. 96-112 (habits of wasps).
— Ashmead, 1894. Psyche 7: 19-26, 39-46, 59-66, 75-79 (habits of wasps). — Peckham and
Peckham, 1898. Wis. Geol. Nat. Hist. Survey, Bui. 2: 1-245, 14 pis. (instincts and habits of
solitary wasps in Wis.). — Hartman, 1905. Tex. Acad. Sci., Trans. 7: 15-85, 24 figs., also
published as Univ. Tex., Bui. 65, Sci. Series 6: 8-73, 24 figs, (habits of some Texan solitary
wasps). —Peckham and Peckham, 1905. Wasps social and solitary, 311 pp. (prey, nests, life
history in Wis.). — Rau and Rau, 1918. Wasp studies afield, 372 pp., 68 figs, (nests, prey,
life history in Mo.). —Wheeler, 1919. Amer. Phil. Soc, Proc. 58: 1-40 (evolution of parasitic
Aculeata). —Rau, 1922. Acad. Sci. St. Louis, Trans. 24: 1-44 (prey, nests, ecology of Mo.
wasps, bees, ants). —Wheeler, 1923. Social Life Among the Insects, 375 pp., 113 figs.
—Rau, 1926. Acad. Sci. St. Louis, Trans. 25: 157-277, 8 pis. (ecology of wasps and bees
nesting in clay bank in Mo.). —Williams, 1928. Hawaii. Sugar Planters' Assoc. Expt. Sta.,
Bui. Ent. Ser. 19: 30-60, 112-174 (tropical wasps and bees). —Wheeler, 1928. Social Insects,
378 pp., 79 figs. —Rau, 1928. Acad. Sci. St. Louis, Trans. 35: 325-489, 68 figs, (behavior
non-social wasps in Mo.). — Reinhard, 1929. The witchery of wasps, 291 pp., 14 pis., 10 text
figs, (prey, nests, life history in Md.). —Rau, 1933. Jungle bees and wasps of Barro
Colorado Island, 324 pp. (nests, prey, life history). — Iwata, 1942. Tenthredo 4: 1-46, 5 pis.,
1 fig., 2 pp. unnumbered figs, (compar. studies behavior of solitary wasps). — Hurd, 1955.
Century of Progress in Natural Sciences, pp. 573-575, Calif. Acad. Sci. (history of wasp
taxonomy). —Cooper, 1957. Jour. Expt. Zool. 134: 469-514, 26 figs, (functions of cell
partitions in preventing parasitism, predation and cannibalism, and in orienting larva for
pupation). —Evans, 1958 (1956). Tenth Internatl. Congr. Ent., Proc. 2: 449-457 (evolution
of social life in wasps). — Olberg, 1959. Das Verhalten der solitaeren Wespen
Mitteleuropas, 402 pp., 779 photos, (prey, nests). —Evans and Linsley, 1960. South. Calif.
Acad. Sci., Bui. 59: 30-37 (sleeping aggregations of aculeates). — Michener, 1961. Roy. Ent.
Soc. London, Symp. 1: 43-56 (aspects of social polymorphism). — Krombein, 1962. Ent. Soc.
Wash., Proc. 64: 11-19 (parasitism of several wasps and bees by acarid mites). —Linsley,

1962. Ent. Soc. Amer., Ann. 55: 148-164, 9 figs, (sleeping aggregations). —Evans, 1962.
Evolution 16: 468-483, 6 figs, (evolution of prey-carrying mechanisms in wasps). —Evans,

1963. Wasp farm, 178 pp., 25 pis., 16 text-figs, (popular account). —Hamilton, 1964. Jour.
Theoret. Biol. 7: 1-52 (genetic evolution of social behavior). —Evans, 1966. Ann. Rev. Ent.
11: 123-154, 2 figs, (behavior patterns of solitary wasps). —Krombein, 1967. Trap-nesting
wasps and bees: Life histories, nests and associates, 570 pp., 29 pis., 2 text-figs. Cited in
text as Trap-nesting wasps and bees. — Andrewes, 1969. The lives of wasps and bees, 204
pp., 16 pis., 15 text-figs, (popular account). —Evans and Eberhard, 1970. The wasps, 265
pp., 122 figs, (synthesis of data on life history, behavior, ecology). —Flanders, 1970. Canad.
Ent. 102: 898-905 (cannibalistic infanticide in social Hymenoptera). —Wilson, 1971. Insect
societies, 548 pp., figs, (synthesis of insect sociology). —Iwata, 1972 (1971). Evolution of
instinct-comparative studies of Hymenoptera behavior, 503 pp., 50 figs, (in Japanese).
—Michener and Lin, 1972. Quart. Rev. Biol. 47: 131-159 (evolution of sociality).

— Spradbery, 1973. Wasps: an account of the biology and natural history of solitary and
social wasps, 408 pp., 28 pis., 131 text figs. —Hamilton, 1973. Ann. Rev. Syst. Ecol. 3:
193-232 (altruism in social insects). — Cazier and Linsley, 1974. Amer. Mus. Novitates 2546:
1-20, 6 figs, (foraging behavior of bees and wasps on Kallstroemia). —Schmidt et al,
1974. Sozialpolymorphismus bei Insekten, 974 pp. — Trivers and Hare, 1976. Science 191:
249-263, 7 figs., 6 tabs, (haplodiploidy and evolution of social insects). —Iwata, 1976.




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Evolution of Instinct: Comparative Ethology of Hymenoptera, 535 pp., frontisp., 50 figs.
(English translation, Natl. Tech. Inform. Serv., PB 257052).




Page 1203 • Top • -1 • +1 • Original • Database of Hymenoptera in America north of Mexico • Highertaxa

Superfamily Bethyloidea

By Karl V. Krombein

• Family Bethylidae
• Family Sclerogibbidae
• Family Chrysididae
• Family Dryinidae
• Family Embolemidae

Superfamily Scolioidea

By Karl V. Krombein

Taxonomy: Ashmead, 1903-1904. Canad. Ent. 35: 4-8, 39-46, 95-107, 155-158, 199-205, 303-310,
323-332; 36: 5-9 (keys to genera).

• Family Tiphiidae
• Family Sierolomorphidae
• Family Mutillidae
• Family Scoliidae
• Family Sapygidae

Superfamily Formicoidea

By David R. Smith

• Family Formicidae

Superfamily Vespoidea

By Karl V. Krombein

Included in this superfamily are several groups with quite diverse habits. The primitive fami-
ly Masaridae includes the only solitary wasps which store pollen and nectar as food for their lar-
vae rather than paralyzed or dismembered Arthropoda. The solitary eumenid wasps store para-
lyzed lepidopterous or coleopterous, or rarely hymenopterous, larvae as food for their young. All
of the truly social wasps belong to the family Vespidae.

Taxonomy: Ashmead, 1902. Canad. Ent. 34: 163-166, 203-210, 219-221 (keys to families and
genera). — Dalla Torre, 1904. In Wytsman, Gen. Ins., fasc. 19, pp. 1-108, 6 pis. (keys and
species catalog). — Bequaert, 1928. Ann. and Mag. Nat. Hist. (10)2: 138-176 (notes on
vespoid types in British Museum). —Bequaert, 1928. Brooklyn Ent. Soc, Bui. 23: 53-63
(Fabricius types in Banks coll.). — Reid, 1942. Roy. Ent. Soc. London, Trans. 92: 285-331,
137 figs, (larval classification). —Richards, 1962. Revisional study of masarid wasps, pp.
3-27 (reclassification, key to subfamilies, phylogeny). — Charnley, 1973. Buffalo Soc. Nat.
Sci., Bui. 26: 1-79 (value of propodeal orifice and male genitalia in higher classification).

Biology: Spradbery, 1973. Wasps, 408 pp., 28 pis., 131 text figs, (natural history of British
Vespoidea).

• Family Masaridae
• Family Eumenidae
• Family Vespidae

Superfamily Pompiloidea

By Karl V. Krombein

This superfamily includes two families, the very large and abundant Pompilidae and the very
small and rare Rhopalosomatidae.

Members of the Pompilidae are commonly called spider wasps because they prey exclusively
on spiders. Only a single larger spider is stored per cell, rather than a number of smaller spiders
as in the spider-preying sphecoid genera such as Trypoxylon, Trypargilum and Sceliphron. The
majority of species, including the more primitive forms, capture the spider before preparing a
burrow in the soil. Some Pepsinae make multicellular nests in pre-existing cavities in twigs or in
the ground, and some build individual mud cells which may be joined in a series; in both types
the cell is constructed before the spider is captured. Some genera, such as Pepsis, may use the
burrow of the prey spider as a nesting site. The peculiar genus Minagenia is a spider ec-
toparasite.

The Rhopalosomatidae are ectoparasites of nymphal crickets.

• Family Pompilidae
• Family Rhopalosomatidae

Superfamily Sphecoidea

By Karl V. Krombein

For nearly a century most specialists in this group, influenced by Kohl's ultra-conservative
views, considered that the superfamily contained a single family, the Sphecidae. The monumen-
tal generic reclassification by Bohart and Menke (1976) embraces this opinion. However,
Brothers (1975) demonstrates convincingly that each of the aculeate superfamilies should com-
prise several families if the family-level groups are to represent categories of equal phylogenetic
value. The classification used elsewhere in this catalog supports the latter conviction. Ac-
cordingly the major subfamilies recognized by Bohart and Menke are restored to family rank, a
position accorded them by most specialists of the previous century.

Some authors believe that the sphecoid wasps and the bees belong to a single superfamily, the
Sphecoidea. For example, Brothers divides the Sphecoidea into two informal groups, the
Spheciformes and Apiformes, with eight and nine famiHes respectively. However, on the basis of
the presence or absence of a hind tibial strigil, Boerner (1919) divides the Aculeata into two sub-
sections, the sphecoids, pompiloids and vespoids, and the formicoids, scolioids and apoids. The
phylogeny of the sphecoid wasps and bees requires more intensive investigation than they have
had hitherto, for the possibility exists that the bees may not be so closely related to the sphecoid
wasps as supposed by some workers. At this time the Sphecoidea and Apoidea are maintained as
separate superfamiHes.

The behavior and Hfe history of this diverse assemblage of wasps has attracted a host of ob-
servers both in the United States and abroad. Many species are ground nesters and are known
popularly as digger wasps; most of them dig their own nests but some species appropriate
pre-existing burrows of other arthropods and modify them as needed. Numerous species nest
above ground in pre-existing cavities such as abandoned borings of beetle larvae in wood, old in-
sect galls and old mud-dauber nests; many of these species can be induced to nest in borings in
wood called trap-nests. Some of our species excavate their own nests in soft pith of shrubs such
as sumac and elderberry, or in rotten wood. Relatively few North American species are mason
wasps, building various kinds of mud cells. Several genera are cleptoparasites of other
ground-nesting sphecoids. So far as known the North American species are all solitary wasps,
but apparent eusociality has been discovered in the Jvleotropical genus Microstigynus Ducke.

Members of the Sphecoidea prey upon a great variety of terrestrial insect orders as well as
upon spiders. Varying degrees of host specificity are found among the several families and
lesser categories. In general the more primitive sphecoids prey upon the more primitive and an-
cient groups of Hemimetabola while the more advanced groups prey upon the higher groups of
Holometabola.

The pre- 1920 references listed below under the side-head Taxonomy are not reliable for
generic or specific discrimination.

Revision: Bohart and Menke, 1976. Sphecid wasps of world, 695 pp., 190 figs, (reclassification
of world genera, lists of species-level taxa).




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Taxonomy: Ashmead, 1899. Canad. Ent. 31: 145-155, 161-174, 212-225, 238-251, 291-300,
322-330, 345-357 (keys to families and genera, and lists of No. Amer. spp.). —Smith, 1908.
Nebr. Univ., Studies 8: 323-410, 1 pi. (keys to Nebr. spp.). — Mickel, 1918 (1917). Nebr.
Univ., Studies 17: 342-456, 2 figs, (keys to Nebr. spp.). —Pate, 1937. Amer. Ent. Soc, Mem.
9: 1-103 (generic names and type-species). —Evans and Lin, 1956. Amer. Ent. Soc, Trans.
81: 131-166, 13 pis. (larvae of Sphecinae). —Evans and Lin, 1956. Amer. Ent. Soc, Trans.
82: 35-66, 13 pis. (larvae of Nyssoninae). —Evans, 1957. Amer. Ent. Soc, Trans. 83: 79-117,
12 pis. (larvae of Philanthinae, Trypoxyloninae and Crabroninae). —Evans, 1959. Amer.
Ent. Soc, Trans. 85: 137-191, 7 pis. (addendum to larvae, keys to subfamilies and genera
based on larval characters, and remarks on evolution and classification based on larval
characters). —Evans, 1964. Amer. Ent. Soc, Trans. 90: 235-299, 12 pis. (larvae,
supplement). —Evans, 1964. Ent. News 75: 225-237, 3 figs, (classification and evolution of
digger wasps based on larvae). —Brothers, 1975. Kans. Univ. Sci. BuL 50: 586-587, 640-641
(phylogeny).

Biology: Evans, 1963. Sci. Amer. 208 (4): 145-154 (evolution as evidenced by predatory
behavior). — Evans, 1966. Science 152: 465-471, 6 figs, (accessory burrows of digger wasps).
— Kurczewski and Snyder, 1968. Conservationist 23 (2): 28-31, 11 figs, (evolution of
cliff-nesting in digger wasps). —Miller and Kurczewski, 1973. hi Dindal, Proc. First Soil
Microcommunities Conf., USAEC, CONF-711076; Natl. Tech. Inform. Serv., USDC, pp.
204-217 (ecology of digger wasps). —Evans, 1975 (1974). N. Y. Ent. Soc, Jour. 82: 259-267,
4 figs, (digger wasps as colonizers of new habitats). — Alcock, 1975. Anim. Behaviour 23:
893-894 (male behavior and territoriality).

• Family Ampulicidae
• Family Sphecidae
• Family Pemphredonidae
• Family Astatidae
• Family Larridae
• Family Crabronidae
• Family Mellinidae
• Family Nyssonidae
• Family Philanthidae

Superfamily Apoidea

By Paul D. Hurd, Jr.

This superfamily contains the bees which, like many other aculeates, visit flowers for nectar.
However, unhke nearly all other aculeates, the bees and most wasps of the Masaridae provide or
feed their larvae with a mixture of pollen and nectar or, in some bees, the mixture is converted
into glandular substances which are then fed to their larvae as well as to certain adults. It is
presumed by many specialists that bees as a group evolved from flower-visiting wasps, most
likely the sphecoid wasps (but possibly also other groups of aculeate wasps), by developing a de-
pendence for food upon pollen and nectar (and sometimes other substances, such as plant oils)
and thereby have abandoned the habit of provisioning their nests with insect or spider prey. It
is not known when this dependency arose, but it could not have occurred before the advent of
Angiosperms which did not begin to flourish until the latter half of the Cretaceous period and
which then became the dominant flora of the earth by the Tertiary. In general, the evolution of
the entomophilous flower has resulted in the replacement of a shallow, flat or bowl-shaped
flower by a corolla-tube. The progressive increase in the depth of the corolla-tube conceivably
has resulted from coevolutionary interactions between the flowers and entomophilous insects,
especially bees, with progressively more elongated mouthparts. Although the earliest known
fossils of bees (Tertiary) are insufficient to establish ancestral relationships with other aculeates
or to demonstrate that the evolution of mouthparts proceeded from the short to the
long-tongued condition, it is probably significant that this sequence is being corroborated by stu-
dies on the systematics, morphology, biology, behavior and biogeography of the contemporary
bee fauna of the world. This fauna is at present considered to consist of eight famihes which in
most current classifications are usually arranged phylogenetically as follows: Colletidae, Ox-
aeidae, Andrenidae, Hahctidae, Melittidae, Megachihdae (including the Fideliinae),
Anthophoridae and Apidae. Even though there is some and sometimes much variation in the
length of the glossa within each of these families, the first five families listed above contain the
so-called short-tongued bees while the remaining families consist of the so-called long-tongued
bees. Although the short-tongued bees of only three families (Colletidae, Melittidae and Halic-
tidae) are present on all continents, only the Colletidae are exceptionally diverse and
well-represented on the southern continents, especially in Australia where other presumably an-
cient groups of plants and animals survive today. The glossa of the Colletidae, in addition to
being normally short and bilobed, is also usually truncate and bifid and therefore is more
wasp-like in structure than is the glossa of any other family of bees. Thus it is believed that the
initial evolution of bees resulted in the development of short-tongued bees which radiated
throughout the earth and this event was subsequently followed by the coevolution of the corol-
la-tube and the long-tongued bees which also have spread throughout the world. Consequently,
the Colletidae are regarded by most speciaHsts as representing the most primitive group of liv-
ing bees. Most, but not all, authors believe that the Apoidea represent a monophyletic assem-




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blage which evolved from a sphecoid ancestor. Brothers (1975) has concluded that the sphecoid
wasps and bees belong to a single superfamily, the Sphecoidea, which according to him consists
of two informal groups, the Spheciformes (8 families) and the Apiformes (9 families). However,
Lanham (1960) has commented upon the similarities and differences of the Sphecoidea and
Apoidea and points out that the presence or absence of the strigil on the hind leg long known to
European hymenopterists may be of as much phylogenetic significance as other characters of
presumably phylogenetic importance. The value of the strigil in these matters was employed by
Boerner (1919) who divides the aculeate Hymenoptera into the Haplocnemata (ants, scolioid
wasps, and bees) and the Diplocnemata (sphecoids, pompilids, and vespids) thereby indicating
that bees are more closely related to the scolioids than to the sphecoid wasps. The phylogeny of
the Aculeata deserves a more thorough study and reevaluation before we can dismiss the possi-
bility that the Apoidea is a polyphyletic assemblage having been derived from both scolioid and
sphecoid ancestors.

Most species of bees construct their nests in the ground usually excavating their own tunnels
and cells, although many others appropriate preexisting burrows or other cavities in the ground
and sometimes modify these to accompHsh their needs. Still many others make their nests above
ground. Among these are species that gnaw out their nesting tunnels in wood substrates of vari-
ous kinds including hollow stemmed plants, while others appropriate a wide variety of preexist-
ing cavities, such as abandoned beetle burrows, hollow trees, old mason wasp nests, old bird
nests, empty snail shells and old insect galls, while still others make their nests of wax and other
materials such as mud, resin, pebbles, pieces of leaves or petals, plant down, etc., and place their
nests either in the open attaching them to branches and so forth or place their nests under
eaves, bridges, rocks, cow chips and so on. As a consequence of these habits, many species of the
families Megachilidae and Apidae readily accept artificial nesting devices (hives, trap-nests, etc.)
which not only has made possible detailed studies of their biologies, but also has made possible
the manipulation and management of several species, including the common honeybee, for use in
the poUination of agricultural crops or for the production of honey and other useful products of
value and benefit to mankind.

In nesting behavior the vast majority of bees are solitary including all members of the fami-
lies Colletidae, Oxaeidae and Melittidae. Except for a few communal species, all members of the
family Andrenidae also exhibit sohtary nesting behavior. Similarly most species of the family
Megachilidae are solitary in habit, although some nest communally (parasocial behavior) while
some are quasisocial and possibly a few are even semisocial in their nesting behavior. Among the
remaining three families (Halictidae, Anthophoridae and Apidae), the Halictidae and
Anthophoridae, each represented by many solitary species, contain some parasocial and eusocial
(primitively social) species as well as some subsocial species in the anthophorid tribe Ceratinini.
While most species of the family Apidae live in perennial, highly eusocial colonies, others exhibit
solitary, parasocial or primitively social nesting behavior.

Bees are exceptionally important pollinators of both native vegetation and many agricultural
crops. Unquestionably bee-plant (bee-flower) relationships reflect various strategies on the part
of both sets of participants and two of the most commonly recognized behavioral modes of pollen
collection by bees are oligolecty and polylecty (see reviews by Grant, 1950; Linsley, 1958; and
Baker and Hurd, 1968). The intrafloral relationships of bees not only involves their own survival
and evolution, but also through coevolutionary interactions with flowers insures the main-
tenance and evolution of much of the earth's flora that depends upon entomophilous polHnation
for reproduction. These interrelationships are of unus'ial significance to us because we in turn
depend upon the earth's flora for our own livelihood and welfare.

In America north of Mexico there are about 3,500 species of bees. This area represents nearly
one-sixth of the earth's land surface and, disregarding ecological diversity, this means that the
world fauna of bees approximates 21,000 species. Somewhat more than 2,700 of our species are
pollen-collecting bees while slightly more than 700 species (or about 21%) are cleptoparasitic spe-
cies. Only about 800 species of bees occur east of the Mississippi River and thus the apifauna of
the larger and more ecologically diverse western portion of America north of Mexico is more
than three times richer in species. Since it is well established that the apifaunas of arid regions
are consistently much richer in species than any other climatic regions, it is not surprising that
most of the North American species of bees are to be found in the southwestern United States
and adjacent northern Mexico. All eight recognized famihes of bees are present in America
north of Mexico and are represented by the following numbers of species: Colletidae (153), Ox-




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aeidae (4), Andrenidae (1,199), Halictidae (506), Melittidae (30), Megachilidae (607),
Anthophoridae (919), and Apidae (47). Among the largest genera in our fauna are: Andrena
(511), Perdita (502), Nomada (286), Dialicius (189), Osmia (133), Megachile (114), Triepeolus
(102), Melissodes (99), and Colletes (96). In all, our fauna consists of 121 genera of which 92 con-
tain the pollen-collecting species. Several of our species are Holarctic in distribution and eight
species, including the European honeybee, are of introduced origin.

The biology of the Apoidea including their behavior, intrafloral ecology, nesting habits, life
histories, communication and the like has always fascinated and attracted much interest and
study by the laity, the beekeeper and the specialist. An immense literature on these and related
subjects has already accumulated and continues to develop so that it is becoming almost an im-
possibility to accomplish an in-depth bibliography of the Apoidea. For example, there are now
more than 100,000 references to the European honeybee alone. Fortunately, the task is made
easier by several literature information retrieval systems and especially by the International
Bee Research Association, headquartered in England, which publishes the key abstracting jour-
nal on bees, Apicultuml Abstracts, two key research journals in English, Bee World and the
Journal of Apicultural Research, as well as comprehensive bibliographies (/. B. R. A. Bibliogra-
phies) on selected subjects pertaining to bees.

Revision: Dalla Torre, 1896. Cat. Hym., v. 10: viii and 643 pp. (classification, catalog of world
spp.; lists 6,165 spp. in 136 genera distributed among 14 subfamilies and all placed in one
family, the Apidae). — Boerner, 1919. Biol. Zentrbl. 39: 145-186, 6 figs, (classification).
— Friese, 1923. Die Europaischen Bienen (Apidae), vii and 456 pp., W. de Gruyter and Co.,
Berlin and Leipzig (classification, life histories). — Grutte, 1935. Arch. Naturgesch. (n, f.) 4:
449-534 (classification of parasitic spp.). — Sandhouse, 1943. U. S. Natl. Mus., Proc. 92:
519-619 (type-species of genera and subgenera). — Michener, 1944. Amer. Mus. Nat. Hist.,
Bui. 82: 151-326, text figs. 1-246, diagrams 1-13 (morphology, phylogeny and classification).
— Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bui. 141: 1-538, 134 figs., 16 tables (east. U. S.
spp. of Colletidae, Andrenidae, Halictidae and Melittidae). —Mitchell, 1962. N. C. Agr.
Expt. Sta. Tech. Bui. 152: 1-557, 134 figs., 18 tables (east. U. S. spp. of Megachilidae,
Anthophoridae, Xylocopidae, Apidae). —Michener, 1965. Amer. Mus. Nat. Hist., Bui. 130:
1-362, 789 text figs., 15 pis., 4 tables (classification). —Weber, 1965. Colo. Univ. Studies,
Series in Bibliography 1: 1-124, 1 frontis. (bibliography of T. D. A. Cockerell). —Stephen,
Bohart and Torchio, 1969. Oreg. State Univ. Agr. Expt. Sta., pp. 1-140, 320 figs,
(classification, morphology, phylogeny and biology of northwest. U. S. spp.). —Michener,
1974. The social behavior of the bees, chapter 3: 25-29. Cambridge, Mass. The Belknap
Press of Harvard Univ. Press (classification).

Taxonomy: Cockerell, 1898. Sci. Lab. Denison Univ., Bui. 11: 41-73 (N. Mex. spp.).
—Cockerell, 1898. N. Mex. Univ., Bui. 1: 43-73 (N. Mex. spp.). — Ashmead, 1899. Amer.
Ent. Soc, Trans. 26: 49-100 (classification). —Fowler, 1902. Calif. Agr. Expt. Sta., Rpts.
1899-1901, pt. 2, pp. 316-330 (long-tongued Calif, spp.). —Cockerell, 1903. Psyche 10: 74-78
(Calif, spp.). —Robertson, 1903. Amer. Ent. Soc, Trans. 29: 163-189 (synopsis of
Megachilidae and Bombinae). —Cockerell and Robbins, 1910. Colo. Univ. Studies 7:
179-195, 8 pi?. (Rocky Mts. spp.). — Graenicher, 1911. Wis. Nat. Hist. Soc, Bui. 1: 221-249
(north. Wis. spp.). — Lutz and Cockerell, 1920. Amer. Mus. Nat. Hist., Bui. 42: 491-641
(notes on distribution, bibliography and floral records of N. Amer. spp. of Anthophoridae
and Apidae). —Cockerell, 1924. Ent. Soc. Wash., Proc. 26: 77-85 (tax. characters).
—Cockerell, 1928. Colo. Univ. Studies 16: 99-126 (Colo. spp.). —Michener, 1941. Sixth
Pacific Sci. Congr., Proc. 4: 297-303 (distributional history of N. Amer. fauna). —Michener,
1947. Amer. Midland Nat. 38: 443-455 (south. Miss. spp.). —Stevens, 1948. N. Dak. Agr.
Expt. Sta. Bimonthly Bui. 10: 187-194 (N. Dak. spp.). —Stevens, 1948. N. Dak. Agr. Expt.
Sta., Bimonthly Bui. 11: 49-54, 2 figs. (N. Dak. spp.). —Stevens, 1949. N. Dak. Agr. Expt.
Sta., Bimonthly Bui. 11: 130-135, 210-225, 4 figs. (N. Dak. spp.). —Richards, 1949. Linn. Soc.
London, Proc. 161: 40-41 (evolution of cuckoo spp.). —Stevens, 1949. N. Dak. Agr. Expt.
Sta. Bimonthly Bui. 12: 14-22 (N. Dak. spp.). —Stevens, 1950. N. Dak. Agr. Expt. Sta.,
Bimonthly Bui. 12: 90-98, 3 figs. (N. Dak. spp.). —Stevens, 1950. N. Dak. Agr. Expt. Sta.,
Bimonthly Bui. 13: 72-80, 4 figs. (N. Dak. spp.). —Bohart and Knowlton, 1950. Utah State
Agr. Expt. Sta. mimeo series, 371: 1-5 (Utah spp.). — Buckell, 1951. Ent. Soc Brit.
Columbia, Proc. 47: 7-24 (B. C. spp.). —Stevens, 1951. N. Dak. Agr. Expt. Sta., Bimonthly




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Bui. 13: 199-205, 2 figs. (N. Dak. spp.). —Stevens, 1951. N. Dak. Agr. Expt. Sta., Bimonthly
Bui. 14: 27-31, 59-64, 2 figs. (N. Dak. spp.). —Stevens, 1952. N. Dak. Agr. Expt. Sta.,
Bimonthly Bui. 14: 105-112, 2 figs, (collecting, mounting, labeling, identifying, study, life
histories). -Michener, 1953. Kans. Univ. Sci. Bui. 35: 987-1102, figs. 1-287 (larvae).

— Michener, 1954. Pan-Pacific Ent. 30: 63-70, fig. 1, table 1 (pupae). —Michener, 1954.
Amer. Mus. Nat. Hist., Bui. 104: 1-176, figs. 1-55, tables 1-3 (classification). —Michener,
1955. A century of progress in the natural sciences, pp. 575-579, Calif. Acad. Sci., San
Francisco (Apoidea). — LaBerge, 1956. Kans. Univ. Sci. Bui. 38: 501-531 (types in Snow
Entomological Museum). — Krunic, 1959. Zbornik Matice Srpske (Novi Sad) 17: 102-111
(transitional forms between solitary and social spp.). — Lanham, 1960. Ent. News 71: 85-86
(diagnostic characters). —Moure, 1960. Studia Ent. 3: 97-160 (Fabricius types of
Neotropical spp.). — Hurd, 1966. Ent. Soc. Amer., Bui. 12: 110-111 (distributional patterns
in west. N. Amer.). —Nielsen and Bohart, 1967. Ent. Soc. Amer., Ann. 60: 414-419, 18 figs,
(larval sex characters). — Covell, 1972. Ent. Soc. Wash., Proc. 74: 10-18 (Lovell types).

— Kerr and da Silveira, 1972. Evolution 26: 197-202 (karyotypic evolution and tax.
implications). —Bohart and Knowlton, 1973. Utah Acad. Sci. Arts and Letters, Proc. 50:
1-9 (spp. of Curlew Valley of Utah and Idaho). —Brothers, 1975. Kans. Univ. Sci. Bui. 50:
586-587, 640-641 (phylogeny). — Moldenke, 1977 (1976). Wasmann Jour. Biol. 34: 147-178, 1
fig., 6 tables (evolutionary history and diversity of faunas of Chile and Pacific North
America).

Biology: Loew, 1884. Jahrb. Bot. Garten 3: 69-118 (floral relationships, oligotrophy and
poly trophy). —Robertson, 1888. Bot. Gazette 13: 33-34 (effect of wind on behavior).
— Verhoeff, 1892. Zool. Anz. 15: 41-43 (relationships between host and parasitic bee
larvae). — Bulman, 1892. Sci. Gossip 329: 98-99 (floral constancy). —Robertson, 1899. Bot.
Gazette 28: 215 (oligotrophy). — Friese, 1899. Zool. Jahrb., Abt. Syst. 3: 847-870 (parasitic
bees and their hosts). —Bulman, 1902. Zoologist 6: 220-222 (floral constancy).
— Graenicher, 1905. Wis. Nat. Hist. Soc, Bui. 3: 153-167 (life history and habits of parasitic
bees). —Latter, 1906. Nature 74: 200 (how inquiline bees find their hosts). —Lovell, 1913.
Ent. News 24: 104-112 (origin of oligotrophy). —Robertson, 1914. Ent. News 25: 67-73
(origin of oligotrophy). —Lovell, 1914. Ent. News 25: 314-321 (origin of oligotrophism).
— Gutbier, 1915. Soc. Ent. Ross., Horae 41: 1-57, 2 pis. (classification and evolution of
nests). —Robertson, 1918. Ent. News 29: 340-342 (proterandry and flight behavior).
— Betts, 1920. Bee World 2: 10-11 (floral constancy). —Clements and Long, 1923. Carnegie
Inst. Wash., Pub. 336: 1-274 (experimental pollination). — Lutz, 1924. N. Y. Acad. Sci., Ann.
29: 181-283 (u. v. floral patterns and flower visiting habits). —Robertson, 1924. Ecology 5:
393-407 (phenology of entomophilous flowers). —Robertson, 1925. Ecology 6: 412-436
(heterotrophy). —Robertson, 1926. Psyche 33: 116-120 (phenology of inquiline and
nest-making bees). — Rau, 1926. Acad. Sci. St. Louis, Trans. 25: 157-277, 8 pis. (life
histories). —Hicks, 1926. Colo. Univ. Studies 15: 217-310 (nesting habits and parasites of
certain Boulder County, Colo. spp.). —Robertson, 1926. Ecology 7: 378-380 (list of
oligolectic spp.). —Robertson, 1928. List of visitors of 453 flowers, 221 pp., Carlinville,
Illinois. —Robertson, 1929. Psyche 36: 112-118 (phenology of oligolectic spp.). —Robertson,
1929. Flowers and insects, 221 pp., Lancaster, Pa., Science Press. —Bromley, 1930. N. Y.
Ent. Soc, Jour. 38: 159-175 (bee-killing robber flies). —Graenicher, 1930. Ent. Soc. Amer.,
Ann. 23: 285-310 (bee-fauna and vegetation of Miami, Fla.). —Robertson, 1930. Ent. News
41: 154-157, 331-336 (proterandry and flight behavior). — Atwood, 1933. Canad. Jour.
Research 9: 443-457 (apple blossom visiting spp. in N. S.). — Cockerell, 1933. Amer. Nat.
67: 1-3 (excessive abundance). —Pearson, 1933. Ecol. Monog. 3: 374-441 (ecological
relationships of spp. in Chicago region). —Rau, 1933. Jungle bees and wasps of Barro
Colorado Island, 324 pp., 112 figs., Kirkwood, Mo. (life histories). —Hicks, 1934. Colo. Univ.
Studies 21: 265-271 (parasites). —Rau, 1934. Acad. Sci. St. Louis, Trans. 28: 219-224
(behavior of certain solitary and social spp.). —Betts, 1935. Bee World 16: 111-113 (floral
constancy). —Cockerell, 1935. Science 81: 458-459 (origin of higher flowering plants and
their insect visitors). —Graenicher, 1935. Ent. Soc. Amer., Ann. 38: 285-310 (bee-fauna and
vegetation of Wis., visitors). — Malyshev, 1935. Eos 11: 201-309, pis. III-XV (nesting
habits of solitary spp.). — Linsley and MacSwain, 1942. South. Calif. Acad. Sci., Bui. 40:
126-137 (nest drepredation by Pti7ius californicus Pic). —Linsley, 1942. Calif. Univ. Pubs.




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Ent. 7: 189-206, pis. 6-7, 1 text fig. (bionomics of Homia, a nest parasite). — Linsley and
MacSwain, 1944 (1943) Ent. Soc. Amer., Ann. 36: 589-601 (predation by Trichodes omatus
Say). —Linsley, 1944. Pan-Pacific Ent. 20: 67-68 (bee prey records of Callinicus calcaneus
Loew). —Linsley, 1944. Brooklyn Ent. Soc, Bui. 39: 54-55 (sapygid parasites). —Popov,
1945. Zhur. Obsch. Biol. 6: 183-203 (parasitism in bees). —Linsley, 1946. Econ. Ent., Jour.
39: 18-29 (alfalfa pollinating spp. in Calif.). —Mitchell, 1946. Research and Farming,
Raleigh, N. C. 4: 1-2, 11 (DDT as threat to bees). -Peck and Bolton, 1946. Scij»Agr. 26:
338-418 (alfalfa pollinating spp. in Sask.). — Bohart, 1947. Farm and Home Sci.'Utah Agr.
Expt. Sta. 8: 13-14 (alfalfa pollinating spp. in Utah). —Linsley and MacSwain, 1947. Econ.
Ent., Jour. 40: 349-357 (factors influencing effectiveness of alfalfa pollinating spp. in
Calif.). —Richards, 1949. Linn. Soc. London, Proc. 161: 40-41 (evolution of cuckoo bees and
wasps). —Linsley, MacSwain and Smith, 1950. Econ. Ent., Jour. 43: 59-62 (DDT
susceptibility). —Grant, 1950. Bot. Rev. 16: 379-398 (flower constancy). —Linsley and
MacSwain, 1951. South. Calif. Acad. Sci. Bui. 50: 92-95 (parasitism by Tricrania stansburyi
Hald.). —Larking, 1952. Agron. Jour. 44: 216-218 (alfalfa pollinating spp.). —Linsley and
MacSwain, 1952. Wasmann Jour. Biol. 10: 91-102 (parasitism by Nemognatha spp.).
—Linsley, MacSwain and Smith, 1952. Ecology 33: 558-567 (outline for study of life
histories of solitary and semisocial spp.). — Pengelly, 1953. 84th Ann. Rpt. Ent. Soc. Ont.,
pp. 101-118 (alfalfa pollinating spp. in Ont.). — Michener, 1953. Century of Progress in the
natural sciences, pp. 575-578, Calif. Acad. Sci., San Francisco (Apoidea). —Michener, Cross,
Daly, Rettenmeyer and Wille, 1955. Ins. Sociaux 2: 237-246 (techniques for studying
behavior). —Stephen, 1955. Econ. Ent., Jour. 48: 543-548 (alfalfa pollinating spp. in Man.).
—Bohart and Nye, 1956. Gleanings in Bee Culture 84: 265-268, 317, 331-333, 337, 400-405,
468-472, 508, 602-606, 639 (place of bees in the world of insects). —Kerr and Laidlaw, 1956.
Advances in Genetics 8: 109-153 (genetics of bees). —Manning, 1956. Behaviour 9: 114-139
(honey-guides). —Manning, 1956. Royal Physiol. Soc, Proc. 25: 67-71 (floral constancy).
— Leppik, 1957. Evolution 11: 466-481 (coevolution of entomophilous plants and
anthophilous insects). —Michener and Lange, 1957. Kans. Ent. Soc, Jour. 30: 71-80
(ethology of colletid spp.). — Hobbs, 1957. Canad. Ent. 89: 230-235 (alfalfa and red clover as
sources of nectar and pollen). —Linsley and MacSwain, 1957. Calif. Univ. Pubs. Ent. 11:
395-430 (stylopization). —Bohart, 1958. Internatl. Congr. Ent., Proc. 10: 929-937 (alfalfa
pollinating spp.). — Koerber and Medler, 1958. Wis. Acad. Sci. Arts and Letters 47: 58-63
(trap-nest survey of solitary spp. in Wis.). —Linsley, 1958. Hilgardia 27: 543-599, 3 figs., 8
tables (ecology of solitary Apoidea). —Linsley and MacSwain, 1958. Evolution, 12: 219-223
(significance of floral constancy). —Medler, 1958. Ent. News 69: 21-24 (parasitism by
Lencospis affinis Say of trap-nesting spp.). — Michener, 1958. Xth Internatl. Congr. Ent.,
Proc. 2: 441-448 (evolution of social behavior). —Michener, Lange, Bigarella and Salamuni,

1958. Ecology 39: 207-217 (factors influencing distribution of nests in earth banks).
—Michener and Lange, 1958. Science 127: 1046-1047 (primitive social behavior). —Hobbs,
1958 (1956). Tenth Internatl. Congr. Ent. 4: 939-942 (factors affecting value of bees as
pollinators of alfalfa and red clover.). —Evans and Lin, 1959. Wasmann Jour. Biol. 17:
115-132 (predation by Philanthus spp.). —Linsley and Hurd, 1959. Ent. News 70: 63-68
(ethological observations on spp. in Ariz, and N. Mex.). —Linsley and MacSwain, 1959.
Kans. Ent. Soc, Jour. 32: 8 (sound production in nocturnal spp.). —Linsley and MacSwain,

1959. Calif. Univ. Pubs. Ent. 16: 1-46 (ethology of Ranunculus visiting spp.). —Powell and
Chemsak, 1959. Kans. Ent. Soc, Jour. 32: 115-120 (predation by Philanthus spp.).
—-Linsley, 1960. N. Y. Ent. Soc, Jour. 68: 13-20 (matinal bees at flowers of Cucurbifa,
Ipomoea and Datura). —Evans and Linsley, 1960. South. Calif. Acad. Sci., Bui. 59: 30-37, 1
pi. (sleeping aggregations). —Linsley, 1960. Calif. Univ. Pubs. Ent. 16: 357-392, pis. 48-55
(ethology of bee- and wasp-killing robber flies in Ariz, and N. Mex.). —Hobbs, Nummi and
Virostek, 1961. Canad. Ent. 93: 409-419 (food gathering behavior of honey-, bumble-, and
leaf-cutter bees in Alta.). —Bohart, 1962. 1st Internatl. Symp. Pollination, August 1960.
Swedish Seed Assoc. Copenhagen Publ. Comm., Proc. 7: 181-188 (introduction of foreign
polHnating spp.). —Linsley, 1962. Ent. Soc. Amer., Ann. 55: 148-164 (sleeping
aggregations). —Linsley, 1962. Sartyrck ur Meddelande nr 7 fran Sveriges
Froodlareforbund, pp. 189-197 (ethological adaptations of solitary spp. for pollination of
desert plants). —Michener, 1962. Rev. Biol. Tropical 10: 167-175, 2 figs, (pollen collection
from flowers with tubular anthers). —Linsley and Cazier, 1963. Pan-Pacific Ent. 39: 1-18, 6




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figs., 2 tables (spp. which take pollen from Solaniivi flowers). — Linsley, MacSwain and
Raven, 1963. Calif. Univ. Pubs. Ent. 33: 1-58, 6 pis., 6 text figs, (comparative behavior of
Camissonia and Oenothera spp. of the Colorado Desert and the Great Basin). —Linsley,
MacSwain and Raven, 1963. Calif. Univ. Pubs. Ent. 33: 59-98, 3 pis. (comparative behavior
of Camissonia and Oenothera spp. of the Mojave Desert). — Michener, 1963. Science 141:
434-435 (division of labor among primitively social spp.). — Wille, 1963. Rev. Biol. Tropical
11: 205-210 (behavioral adaptations of pollen-collecting spp. at flowers of Cassia).
—Michener, 1964. Ins. Sociaux 11: 317-342 (reproductive efficiency in relation to colony
size). —Michener, 1964. Amer. Zool. 4: 227-239 (evolution of nests). — Armitage, 1965.
Kans. Ent. Soc, Jour. 38: 89-100, 4 figs., 4 tables (predation by Philaiithus spp.).
—Matthews and Fischer, 1965. North Central Branch, Ent. Soc. Amer., Proc. 19: 79-81, 1
fig. (modified trap-nest). — Fye, 1965. Econ. Ent., Jour. 58: 803-804, 4 figs, (trap-nesting
methods). —Fye, 1965. Canad. Ent. 97: 863-877, 6 figs., 4 tables (ethology of spp. taken in
trap-nests in northwest Ont.). —Parker and Bohart, 1966. Pan-Pacific Ent. 42: 91-98
(host-parasite relationships as determined by use of trap-nests). —Butler, Werner and
Levin, 1966. Kans. Ent. Soc, Jour. 39: 434-436 (safflower visiting spp.). —Levin and
Butler, 1966. Econ Ent., Jour. 59: 654-657, 3 tables (safflower pollinating spp.).
— Krombein, 1967. Trap-nesting wasps and bees, vi and 570 pp., 29 pis., Smithsn. Press
(life histories, nest architecture, nest associates). — Kerfoot, 1967. Amer. Nat. 101: 65-70
(correlation between ocellar size and foraging activities). — Parker and Bohart, 1968.
Pan-Pacific Ent. 44: 1-6 (host-parasite relationships as determined by use of trap-nests).
—Baker and Hurd, 1968. Ann. Rev. Ent. 13: 385-414 (intrafloral ecology). — Batra and
Torchio, 1968. Mycologia 60: 189-190 (N. Amer. records of Ascocphaera apis L.). —May
and Stockhammer, 1968. Kans. Ent. Soc, Jour. 41: 339-341, 1 fig. (mass colonization by use
of artificial substrate). —Kerr, 1969. Ecol. Biol. 3: 119-175 (evolution of social bees).
—Michener, 1969. Ann. Rev. Ent. 14: 299-342, 3 tables (comparative behavior of social
bees). —Bohart, 1970. Ent. Soc. Amer., Bui. 16: 8-9 (management of native spp. for
commercial crop production). — Schlissing, 1970. Ecology 51: 1061-1067 (foraging behavior
in flowers of Ipomoea and A^iiseia). — Gerber and Klostermeyer, 1970. Science 167: 82-84
(sex control). — Darchen, 1970. Gaz. Apicult. 754: 48-51 (division of labor in social spp.).
— Mickel, 1970. Minn. Univ. Agr. Expt. Sta. Tech. Bui. 27: 1-77 (references to literature
pertaining to Mutillidae parasitic on Apoidea). —Bohart, 1970. Utah State Univ. 41st
Faculty Honor Lecture, 33 pp. (evolution of parasitism). — Macior, 1970. Amer. Jour. Bot.
57: 716-728 (pollinating spp. of Pedicularis in Colo.). —Hurd, Linsley and Whitaker, 1971.
Evolution 25: 218-234, 4 figs., 3 tables (squash and gourd bees and origin of cultivated
Cucurbita). —Bohart, 1971. Tall timbers Conf. Ecol. Animal Contrib. Habitat
Management, Proc, pp. 253-266, 9 figs, (management of habitats for native spp.).
—Michener and Brothers, 1971. Kans. Ent. Soc, Jour. 44: 236-239, 4 figs, (observation nest
for burrowing spp.). —Batra, 1972. Kans. Ent. Soc, Jour. 45: 208-218 (nest-building
secretions). —Osgood, 1972. Maine life Sci. Agr. Expt. Sta. Tech. Bui. 59: 1-8 (nesting sites
of native spp. associated with low-bush blueberries in Maine). —Bohart, 1972. Ann. Rev.
Ent. 17: 287-312, 4 tables (management of native spp. for crop pollination). —Michener,
1972. Kans. Ent. Soc, Jour. 45: 373-376 (direct food transferring behavior). — Cruden, 1972.
Evolution 26: 363-389 (pollination biology oiNemophila menziesii with comments on
evolution of oligolectic bees). —Cruden, 1973. Amer. Jour. Botany 60: 802-809 (pollination
of Mirabilis). —Thorp, 1973. Pan-Pacific Ent. 49: 89 (robber fly predation). —Batra, Batra
and Bohart, 1973. Mycopath. Mycol. Appl. 49: 13-44 (mycoflora of domesticated and wild
bees). -MacSwain, Raven and Thorp, 1973. Calif. Univ. Pubs. Ent. 70: 1-80, 3 pis., 10 figs.,
18 tables (comparative behavior of Clarkia visiting spp.). —Linsley, MacSwain, Raven and
Thorp, 1973. Calif. Univ. Pubs. Ent. 71: 1-68, 6 pis., 15 figs., 10 tables (comparative
behavior of Caynissonia and Oenothera visiting spp. in cismontane Calif, and Baja
Cahfornia). -Vogel, 1973. Umschau 73 (22): 701-702 (collection of fatty oil from plants and
its incorporation into larval food.). —Macior, 1973. Amer. Jour. Bot. 60: 863-871 (pollinating
spp. of Pedicularis on Mt. Rainier). —Torchio, 1973. Kans. Ent. Soc, Jour. 46: 446-453, 1
fig., 4 tables (relative toxicity of insecticides to honeybee, alkah bee and alfalfa leafcutting
bee). — Frankie, 1973. Ent. Soc. Amer., Ann. 66: 690-691, 1 fig. (field technique for marking
bees). —Macior, 1974. Melanderia 15: 1-59 (pollinating spp. of Front Range of Colorado
Rocky Mts.). — Cazier and Linsley, 1974. Amer. Mus. Novitates 2546: 1-20, figs. 1-6, tables




Page 1747 • Top • -1 • +1 • Original • Database of Hymenoptera in America north of Mexico • Highertaxa

1-2 (foraging behavior of spp. visiting flowers of Kallstroemia grandiflom). —Jones and
Buchman, 1974. Anim. Behaviour 22: 481-485, 2 tables (u. v. floral patterns as orientation
guides). -Estes and Thorp, 1974. Torrey Bot. Club, Bui. 101: 272-276 (poHinating spp.
visiting flowers of Ludwigia peploides). — Michener, 1974. The social behavior of the bees,
xii and 404 pp., The Belknap Press of Harvard Univ. Press. —Michener and Brothers,
1974. Natl. Acad. Sci. U. S. A., Proc. 71: 671-674 (queen inhibitory behavior). — Macior,

1974. Missouri Bot. Garden, Ann. 61: 760-769, 21 figs, (coadaptation between flowers and
pollinating spp.). — Moldenke and Neff, 1974. The bees of California: A catalogue with
special relevance to pollination and ecological research. Origin and Structure of
Ecosystems (IBP) Technical Rpts. 74-1 to 74-6, Calif. Univ. Santa Cruz. —Austin and
OHver, 1974. Arnold Arboreteum Harvard Univ., Jour. 55: 291-299 (pollinators of
Sisyrinchiiim solstitiale, a Fla. endemic). — Bottema, 1975. Paleohistoria 17: 17-35, 9 figs.,
7 tables (contamination of pollen spectra by burrowing bees in prehistoric settlements).
— Cazier and Linsley, 1975. Pan-Pacific Ent. 51: 248-253, 6 figs., 2 tables (bee and wasp
visitors to the flowers of Kallstroe))iia grandiflora after two years drought). —Thorp,
Briggs, Estes and Erickson, 1975. Science 189: 476-478, 1 fig. (nectar flourescense and
foraging efficiency of bees). —Estes and Thorp, 1975. Amer. Jour. Bot. 62: 148-159
(pollination oi Pyrrhopappns carolinianus). —Thorp and Estes, 1975. Kans. Ent. Soc,
Jour. 48: 175-184, 6 figs, (intrafloral behavior of bees on flowers of Cassia fasciculata).

— Heinrich, 1975. Ann. Rev. Ecol. Syst. 6: 139-170 (energetics of pollination). — Heinrich,

1975. Evolution 29: 325-334, 4 figs., 2 tables (bee flowers). —Barrows, Bell and Michener,
1975. Natl. Acad. Sci. USA, Proc. 72: 2824-2828 (odor differences and their social function). '

— Hurd and Linsley, 1975. Smithsn. Contrib. Zool. 193: iii and 74 pp., 18 figs., 15 tables
(principal Larrea visiting spp. of southwest. U. S.). — Kevan, 1975. Biol. Conservation 7:
301-309, 1 fig., 4 tables (effect of fenitrothion on pollinators of lowbush blueberries).

— Macior, 1975. Amer. Jour. Bot. 62: 1009-1016, 19 figs., 17 tables (pollination of
Delphinium tricome). —Macior, 1975. Amer. Jour. Bot. 62: 1065-1072, 21 figs., 7 tables
(pollination ecology of Pedicularis in the Yukon Territory). —Evans, 1975. Ent. Soc.
Amer., Ann. 68: 888-892, 6 figs., 3 tables (predation by Pliilanthus albopilosus Cress.).

— Linsley, 1976. Pan-Pacific Ent. 52: 177-178 (defensive behavior of males about plants not
visited by their females). — Bouseman, 1976. Pan-Pacific Ent. 52: 178-179 (predation by
Apiomenis crassipes). — Iwata, 1976. Evolution of instinct, comparative ethology of
Hymenoptera, 535 pp., Washington, D. C, Smithsonian Institution (behavior). —Erickson,
Enns and Werner, 1976. Ent. Soc. Amer., Ann. 69: 959-970, 4 tables (bee-associated
Meloidae). — Jander, 1976. Physiol. Ent. 1: 179-194, 8 figs., 2 tables (grooming and pollen
manipulation). —McGregor, 1976. U. S. Dept. Agr., Agr. Handbook 496: 1-411, 196 figs,
(pollination of cultivated crop plants). — Frankie, Opler and Bawa, 1976. Jour. Ecol. 64:
1049-1057, 1 fig., 4 tables (foraging behavior). —Rust and Clement, 1977. Kans. Ent. Soc,
Jour. 50: 37-48, 5 figs., 4 tables (role in pollination of Colli7isia sparsiflora).

Morphology: Braue, 1913. Jenaische Ztschr. Naturwiss. 50: 1-96 (pollen-collecting apparatus).

— Stoeckhert, 1924. Arch. Naturgesch. (A) 90 (2): 109-131 (gynandromorphism). — Kuhn,
1927. Ztschr. vergleich. Physiol. 5: 762-800 (color vision). — Pessotzkaya, 1929. Soc. Nat.
Leningrad. Trav. 59: 21-46 (gland apparatus in instinctive behavior). —Beck, 1933. Utah
Acad. Sci., Proc. 10: 89-137, 8 pis. (male genitalia). —Michener, 1943. Pan-Pacific Ent. 19:
96-100 (homologies between male and female appendages). —Michener, 1944. Ent. Soc.
Amer., Ann. 37: 336-351 (appendages of eighth and ninth abdominal segments).

— Michener, 1944. Amer. Mus. Nat. Hist., Bui. 82: 158-225 (comparative external
morphology). — Auclair and Jamieson, 1948. Science 108: 357-358 (amino acids in pollen
collected by bees). — Wille, 1956. Kans. Univ. Sci. Bui. 38: 439-499 (thoracic musculature).
— Wille, 1958. Ent. Soc. Amer., Ann. 51: 538-546, 24 figs, (dorsal vessel). — Altenkirch,
1962. Zool. Beitrag (n. f.) 7: 161-238 (abdomen). —Michener, 1965. Amer. Mus. Nat. Hist.,
Bui. 130: 27-32, figs. 2-17 (morphological terminology). — Cruz-Landim, 1967. Arq. Zool. S.
Paulo 15: 177-290 (glands). — Rothenbuhler, Kulincevic and Kerr, 1968. Ann. Rev. Genetics
2: 413-438 (genetics). —Graf, 1968. Bol. Univ. Federal Parana, Zool. 3: 65-78 (salivary
gland). —Stephen, Bohart and Torchio, 1969. Oreg. State Univ. Agr. Expt. Sta., pp. 3-31,
figs. 1-112 (external morphology). — Tanabe, Tamaki and Nakamo, 1970. Japanese Jour.
Genetics 45: 425-428 (variation in esterase isozymes). — Tulloch, 1970. Lipids 5: 247-258




Page 1748 • Top • -1 • +1 • Original • Database of Hymenoptera in America north of Mexico • Highertaxa

(composition of beeswax). -Lello, 1971. Kans. Ent. Soc, Jour. 44: 5-20, 21 figs, (adnexal
glands of sting apparatus). —Lello 1971. Ciencia e Cultura 23: 253-258 (adnexal glands of
sting apparatus). — Wille, 1971. Rev. Biol. Tropical 18: 33-51 (musculature of salivary
syringe and neck region). —Kerr, 1972. Kans. Ent. Soc, Jour. 45: 111-122, 20 figs,
(chromosome numbers). —Almeida Correia, 1973. Faculdade Cienc. Porto Univ., An. 56:
67-175 (morphological and morphometric study of mouthparts of principal genera).
—Almeida Correia, 1973. Inst. Zool. "Dr. Augusto Nobre" Facul. Cienc. Porto 118: 1-117, 8
pis. (mouthparts). — Cruz-Landim, 1973. Studia Ent. 16: 209-215, 2 figs., 1 table (thoracic
saHvary glands). -luga, 1973. Mus. Hist. Nat. "Grigore Antipa", Trav. 13: 203-226, 26 figs,
(apical abdominal appendages). — Michener, 1974. The social behavior of the bees. Chapter
1: 3-19 (development, structure and function). —Snyder, 1975. Evolution 28: 687-689
(allozymic variability). — Pasteels and Pasteels, 1975. Arch. Biol, Bruxelles 86: 453-466, 13
figs, (stereoscan studies of pollen collecting scopae of Fideliidae). —Lello, 1976. Kans. Ent.
Soc, Jour. 49: 85-99, 22 figs., 3 tables (adnexal glands of sting apparatus). —Pasteels and
Pasteels, 1976. Arch. Biol., Bruxelles 87: 79-102, 25 figs, (stereoscan studies of pollen
collecting scopae of Colletidae and Oxaeidae).

• Family Colletidae
• Family Oxaeidae
• Family Andrenidae
• Family Halictidae
• Family Melittidae
• Family Megachilidae
• Family Anthophoridae
• Family Apidae