B. D. BUR Species of Spalangia

Latreille in the

United States

W National Museum

Collection

(Hymenoptera: Pteromalidae)

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • 1969 NUMBER 2

SMITHSONIAN CONTRIBUTIONS TO

ZOOLOGY

NUMBER 2

B. D. Burks Species of

Spalangia Latreille

in the United States

National Museum

Collection

(Hymenoptera:

Pteromalidae)

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1969
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ABSTRACT

A reappraisal of species of Spalangia in the USNM collection. The types of nine species

described by Ashmead, Girault, Richardson, and Howard are all redescribed, their

present condition given, and lectotypes designated, if necessary. Spalangia brasiliensis

Ashmead is synonymized under chontalensis Cameron; muscidarum var. texensis

Girault is synonymized under cameroni Fullaway. Spalangia attae, from the nest of an

Atta ant in El Salvador, and dozieri, parasite of the sarcophagid fly Sarcodexia

sternodontis Townsend in Puerto Rico, are described as new.

Z. Bouc'ek (1963, 1965) has published a comprehensive study of the difficult genus Spalangia Latreille for

the world. His papers are based on a study of the types

of Foerster, Latreille, Curtis, and Walker, all of which

are located in European museums, as well as on material identified as species described by Ashmead, Girault,

and Perkins, types of which are located in North

American, Australian, and Hawaiian museums. The

latter specimens were supplied to Bouc'ek by several

entomologists, principally those at the Canadian National Collection and the United States National

Museum (USNM). The papers by Bouc'ek have made

it possible for the first time to place with confidence

most specimens of Spalangia, the bulk of which belong

to species worldwide in distribution, but which were

described first from the Palearctic region.

The USNM collection contains the types of (Spalangia brasiliensis Ashmead) =S. chontalensis Cameron,

S. drosophilae Ashmead, S. haematobiae Ashmead,

(S. rugosicollis Ashmead) =S. nigra Latreille; (S.

muscarum Girault) =S. nigripes Curtis, (S. muscidarum var. stomoxysiae Girault) =S. endius Walker,

(S. muscidarum var. texensis Girault) =S. cameroni

Fullaway; the only known remaining types of (S. muscidarum Richardson) = S. nigroaenea Curtis; and the

type of (S. muscae Howard) —S. nigra Latreille, which

may have been described through inadvertence. In

addition, in the USNM a large amount of material

that accumulated over the years has been identified by

various taxonomists as the same as these types or as

other species, based on literature. When Bouc"ek's paper

was published, a reworking of all this material was

necessary. In the course of this restudy of the USNM

B. D. Burks, Systematic Entomology Laboratory (United

States Department of Agriculture), Smithsonian Institution,

Washington, D.C. 20560.

material, I found two undescribed species of Spalangia,

and these are described in this paper. It also developed

that it would be desirable to clarify the status of all

the types of Spalangia now in the collection. In some

it was necessary to designate lectotypes, and for all it

was advantageous to give the present condition of the

specimens. For each I have given a short summary of

the diagnostic characteristics, in case the types should

be damaged later.

Bouc'ek (1963, p. 498) considered S. brasiliensis

Ashmead to be a valid species, but I am here

synonymizing it under chontalensis Cameron. He

(p. 458) placed muscidarum var. texensis Girault as

a synonym of endius Walker, but it is a synonym of

cameroni Fullaway instead. Otherwise, I agree with

his placement of the species whose types are in the

USNM collection.

Of the nine species represented by types in the

USNM collection, only two names survive as valid.

The other seven specific names have fallen into

synonymy, despite the fact that all these species were

described more than 45 years ago, and four of them

more than 60 years ago. In all but one species, they

have fallen into synonymy under Palearctic species that

were described a century or more ago; the one exception is a synonym of a worldwide species that was first

described from Hawaii in 1910.

Spalangia attae, new species

FIGURE 2

This species agrees with Spalangia drosophilae Ashmead in being small, less than 2.0 mm long, with the

thorax depressed and flat, in lacking the crossline of

punctures near the posterior margin of the pronotum,

in having the frenal crossline of the scutellum indicated

only at the lateral margins, and in having closely set,

longitudinal striae on the mesopleuron. The two differ

1
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

in that attae has the head and pronotum shagreened

and mat rather than smooth and shining with scattered punctures; the petiole is as long as the hind coxa,

rather than being shorter than the hind coxa; and

the foretarsal segments are markedly shorter in attae

than they are in drosophilae (Figures 2, 3).

FEMALE.—Length 1.5 mm. Black, with basal 4 segments of each tarsus white, apical segment brown; tegulae brown, wing veins tan. Surface of head minutely

shagreened and mat, except for ocellar triangle, which

is smooth, antennal scape closely and irregularly sculptured, dull; pronotum, except for narrow, smooth crossband at posterior margin, minutely shagreened and

mat; scapulae shining; praescutum shining at anterior

margin, elsewhere with minute, contiguous and shallow pits in transverse, closely set rows, these becoming

irregular near posterior margin of sclerite; axillae, scutellum, and postscutellum smooth and shining; metepisternum minutely shagreened and mat; propodeum shining; petiole with numerous, closely set,

longitudinal carinae, interstices with minute cross

septa; surface of gaster shining. Anterior coxae subshining, with minute, closely set, parallel striae; middle coxae reticulated at bases, smooth and shining

apically; hind coxae smooth and shining. Prepectus

heavily and irregularly sculptured. Mesopleuron with

lineolate sculpture dorsally and near base of midcoxa,

pitted sculpture in anterior oblique impressions and

in 2 oval impressions posteriorly, elsewhere smooth.

Eyes and head hairy, but head hairs not arising from

obvious punctures (these obscured by the shagreenate sculpture) ; distinct groove extending from anterior ocellus into apex of scrobe cavity; malar furrow almost hidden by head sculpture; length of malar space

equal to eye height. Relative lengths of parts of antenna, scape 55; pedicel, 20; first funicular segment, 8;

second, 5; third, 6; fourth, 7; fifth, 7; sixth, 8; seventh,

8; club, 25.

Dorsum of thorax depressed and flat, as in drosophilae Ashmead. Pronotum bearing numerous bristles,

but punctures from which they arise obscured

by shagreened surface sculpture; crossline of punctures wanting near posterior margin. Notaulices and

lateral margins of scutellum with crenulate sculpture;

margin of scutellar frenum indicated only by 2 punctures at each lateral margin; posterior margin of scutellum crenulate, setting off postscutellum. Foretarsal

segments compact, first segment as long as following

three combined (Figure 2); middle and hind tarsi

slightly more elongate.

Propodeum with crenulate anterior and lateral margins, a double row of coarse, rectangular punctures on

meson. Petiole stout, half as wide as long, and as long

as hind coxa. Gaster as long as thorax and propodeum,

apical sternite (subgenital plate of Snodgrass) produced as a minute point on meson, not indented, as in

drosophilae; apices of ovipositor sheaths slightly

exserted.

MALE.—Unknown.

TYPE-LOCALITY.—Near San Salvador, El Salvador.

HOLOTYPE.—USNM 69871.

Described from a single specimen from debris of a

nest of the leaf-cutter ant, Atta mexicana (F. Smith),

25 June 1958, by O. L. Cartwright. The host of this

Spalangia is not known, but it should be some small

dipteron. It was taken along with a large assortment

of beetles, including staphylinids, scarabaeids, tenebrionids, and others (the collector is a Coleopterist).

I have another specimen of this Spalangia, unfortunately badly broken, that was taken from a nest of

Atta texana (Buckley) at San Antonio, Texas,

16 January 1935, by E. V. Walter. It was collected in

the ant nest at a point seven feet below the surface of

the ground. I have not designated this specimen a paratype because it is fragmentary.

Spalangia chontalensis Cameron

Spalangia brasiliensis Ashmead, 1904, p. 502 [female].—Bou2ek, 1963, p. 498. [New synonymy.]

Spalangia brasiliensis was described from one specimen. It is labeled: "Santarem, H. H. Smith Coll. Spalangia brasiliensis $ Type Ashm., Type No. 60553

U.S.N.M."

Boucek (1963, p. 498) treated brasiliensis as a valid

species, but he found it difficult to place with certainty from only a reading of Ashmead's description.

Actually a study of the type of brasiliensis shows it to

be a synonym of S. chontalensis Cameron. Bouc"ek has

redescribed and figured the type of chontalensis, and

the type of brasiliensis differs from it in no significant

way.

The type of brasiliensis shows the following diagnostic characters:

Head with dense umbilicate punctation, with smooth

interstices, setae noticeably long; pronotum with a

prominent, slightly irregular, anterior carinate margin,

dorsal meson of pronotum smooth, lateral areas densely

umbilicate-punctate, crossrow of closely set, deep punctures located just anterior to posterior margin of
NUMBER 2

pronotum. Anterior half mesopraescutum smooth,

posterior half densely covered with rugose pits, a

narrow, longitudinal carina present on meson; each

notaulix indicated by a row of five large, rectangular

pits; scutellum with frenal crossrow of punctures complete, deep, closely set, scutellum otherwise smooth

except for three pits in each posterolateral corner.

Propodeum coarsely punctured laterally and on meson;

petiole half as wide as long, with coarse, longitudinal

rugae and prominent lateral bristles, interstices of

petiole between longitudinal rugae minutely roughened; posterior margin of second gastral tergum

broadly emarginate; gaster as long as thorax and

petiole combined.

There are specimens of chontalensis from St. Vincent, W. I., labeled as Spalangia nigra Latr. by Ashmead, in the USNM collection. These were the basis

for the St. Vincent record of nigra that was published

by Riley (1894, p. 59). Other published West Indian

records of nigra are undoubtedly based on this reference. Spalangia nigra is not known to occur in the

Neotropical region.

Spalangia dozieri, new species

FIGURE 1

This species agrees with drosophilae Ashmead in

lacking a crossline of punctures near the posterior

margin of the pronotum, in having the thorax depressed and flattened, and in being small, less than 2.0

mm long. The two differ in that dozieri has the parallel, longitudinal striae of the mesopleuron, characteristic of drosophilae, much less well developed, and has

a complete frenal crossrow of punctures on the scutellum, which is lacking in drosophilae; dozieri also has a

prominent, median, dorsal, longitudinal groove on the

pronotum, and this is lacking in drosophilae.

FEMALE.—Length 1.65-1.75 mm. Black, with base

of each tibia and basal 4 segments of each tarsus pale

tan or white; tegula and wing veins dark brown. Head

smooth and shining, sparsely punctate; antennal scape

minutely shagreened, mat; pronotum smooth, shining,

sparsely punctate, scapulae shining; praescutum lightly

and minutely sculptured on anterior three-fifths, this

sculptured area bounded posteriorly by an irregular,

transverse row of punctures, and surface of praescutum

posterior to this crossline smooth; axillae, scutellum,

and postscutellum shining; metepisternum strongly

shagreened; propodeum smooth on each side of me3

dian, longitudinal sculptured area; petiole with strong

and irregular, alveolate sculpture; first gastral tergum

smooth, shining, following terga shining, but surface

faintly sculptured; apices of ovipositor sheaths sculptured and mat. All coxae shining. Prepectus heavily

and irregularly sculptured. Mesopleuron dorsally with

minute, parallel and closely set, lineolate striae; another similar, oblique area of striae in median region

and near posterior margin of mesopleuron; area near

anterior and ventral margins of this sclerite smooth.

Eyes and head sparsely hairy; distinct groove extending from anterior ocellus into scrobe cavity; malar

furrow distinct, its length two-thirds the eye height.

Relative lengths of parts of antenna, scape, 42; pedicel,

12; first funicular segment, 5; second, 6; third to

seventh, each 5; club, 20. All funicular segments wider

than long, first only slightly wider than long, but segments gradually widening, so that seventh is twice as

wide as long.

Dorsum of thorax depressed and flat, as in drosophilae Ashmead. Pronotum bearing scattered bristles and

a median longitudinal groove; notaulices and lateral

margins of scutellum with crenulate sculpture; margin

of scutellar frenum set off by a complete crossrow of

closely set punctures; posterior margin of scutellum

incised, crenulate only laterally. First foretarsal segment as long as following three combined, middle and

hind tarsi more elongate.

Propodeum with lateral and posterior margins irregular (Figure 1); a double arcuate row of coarse,

rectangular punctures at base, these fusing before

reaching apex. Petiole only slightly longer than wide,

shorter than hind coxa. Gaster depressed dorsally,

broader than thorax and shorter than thorax and propodeum combined; apices of ovipositor sheaths slightly

exserted.

MALE.—Unknown.

TYPE-LOCALITY.—Mayaguez, Puerto Rico.

HOLOTYPE.—USNM 69872.

Described from 19 female specimens as follows:

Type and 16 paratypes, Mayaguez, Puerto Rico,

reared 26 August 1936 from the pupa of Sarcodexia

sternodontis Townsend (Diptera, Sarcophagidae) by

H. L. Dozier; 1 paratype, Punta Arenas, P.R., 12

March 1963, taken in bait trap, E. F. Legner; 1 paratype, Central Portugalata, Cuba, collected February

1928 by C. F. Stahl. All specimens in the USNM

collection.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIOURES 1-3.—1, Spalangia dozieri, new species. 2, Spalangia attae, new species, foretarsus.

3, Spalangia drosophilae Ashmead, foretarsus.

Spalangia drosophilae Ashmead

FIGURE 3

Spalangia drosophilae Ashmead, 1887, p. 199 [female].

This species was described from a single female

specimen. The unique type is labeled: "Jacksonville,

Fla., Type, Spalangia drosophilae Ashm., Type No.

26039 U.S.N.M." It is preserved in fair condition. It

seems to have been remounted at some time, but it

clearly shows the diagnostic characters of this valid

species. Body small and flattened, antenna! scape short,

all funicular segments short, with second segment the

shortest; pronotum lacking punctate crossrow near

posterior margin; frenal line of scutellum indicated

only by a few punctures at each lateral margin; mesopleuron with numerous, closely set, parallel, longitudinal striae; gaster broader than thorax.
NUMBER 2

Spalangia haematobiae Ashmead

Spalangia haematobiae Ashmead, 1894, p. 37 [female].

This species was described from one female specimen. The type is labeled: "4285/24, Sept. 13. '9. Spalangia haematobiae Ashm., Type No. 2169 U.S.N.M."

The petiole and gaster are now missing from the type

specimen.

The following description is drawn from the type:

Black; legs and wingveins dark brown. Front and

vertex smooth, shining, with few scattered hair punctures; height of compound eye greater than length of

malar space, 33:25. Surface of antennal scape irregularly and minutely shagreened; relative lengths of

parts of antenna, scape, 50; pedicel, 18; first funicular

segment, 10; second to seventh, each 8; club, 22. Distinct groove extending from anterior ocellus into top

of scrobe depression. Thorax not markedly flattened,

as in drosophilae Ashmead, but moderately rounded.

Pronotum ecarinate anteriorly; lacking punctate crossline near posterior margin; surface smooth, shining,

with relatively sparse hair punctures laterally, these

wanting on dorsal meson; scapulae shining; praescutum smooth and shining anteriorly, faintly sculptured

on posterior half, with a thin, median, longitudinal

carina, this bordered on each side by large, shallow and

slightly irregular punctures; notaulices with large,

crenulate punctures; axillae smooth, shining; scutellum smooth, shining, without frenal crossline of punctures. (Specimen mounted in such a way that pleura

cannot be seen.) Propodeum smooth, with crenulate

basal, apical, and lateral margins and a double row of

large, shallow pits on meson, a median carina present

between the rows in basal half.

I consider this to be a valid species of Spalangia,

as does Boudek (1963, p. 475). There are specimens

in the USNM collection from New Hampshire, Maryland, Texas, California, Mexico, Costa Rica, and St.

Croix in the Virgin Islands. The only known host is

Haematobia irritans (L.).

Spalangia nigra Latreille

Spalangia muscae Girault.—Howard, 1911, p. 90.

Spalangia muscae has been thought to be a nomen

nudum, a manuscript name, and a described species.

Bouc"ek (1963) considered it a nomen nudum;

Richardson (1913) and Girault (1920) treated it as a

manuscript name; while Bishopp (1913) and Riihl

(1916) listed it as a valid, described species. Crawford

(in Richardson 1913) took a decided position that

muscae was a valid species and Peck (1951, 1963)

listed it twice as a valid species.

Howard (1911), in discussing a parasite that had

emerged from a puparium of Musca domestica when

it was opened by a laboratory worker, made the following statement:

a fully formed and active adult black Spalangia crawled

immediately from the opening made by his dissecting needle.

This will be described by Girault as Spalangia muscae. A

certain proportion of the housefly puparia are affected by

this parasite in precisely the same way as are the puparia in

Europe by Spalangia nigra as described by Bouche\

Girault never published a description of muscae, and

it is extremely doubtful that the passage quoted from

Howard can stand by itself as the description of a

species. Howard himself evidently did not consider

that he had described the species, because he did not

designate a type specimen. In 1923, however, S. A.

Rohwer located the specimen of Spalangia that

Howard had discussed in his book and designated it

as Howard's type. Rohwer assigned to it USNM Type

No. 26329 and labeled it: "Spalangia muscae How.

Type, labeled by Rohwer."

As was pointed out by Boucek (1963, p. 447),

muscae, whatever its standing, is the same as the holarctic species nigra Latreille, 1805.

Spalangia nigripes Curtis

Spalangia muscarum Girault, 1920, p. 213 [female].

This species was described from one female specimen. The type is labeled: "Urbana, 111., 39589, Spalangia muscarum Girault. 9 Type. Type No. 20877

U.S.N.M." The specimen is intact.

This specimen agrees in all diagnostic characters

with reliably determined European specimens in the

USNM of Spalangia nigripes Curtis, the species with

which muscarum has been synonymized by Bouc"ek

(1963, p. 468). Five apical funicular segments of

antenna broader than long; head and pronotum with

relatively sparse, umbilicate punctation and smooth

interstices; pronotum lacks an isolated crossline of

punctures just anterior to posterior margin; scutellum

with a complete frenal crossrow of punctures, anterior

scutellar area sparsely punctate laterally, smooth on

meson; and second gastral tergum with posterior margin broadly emarginate.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Spalangia nigroaenea Curtis

Spalangia muscidarum Richardson, 1913, p. 38 [male,

female].

This species was described from one male type and

an undesignated number of paratypes of both sexes.

The type has been lost, but there are 3 male and 1

female paratypes in the USNM collection. They are

labeled: "Forest Hills, Mass., IX. 1912, Paratype No.

40219 U.S.N.M., Spalangia muscidarum Richardson

Paratype."

These specimens are identical with European specimens in the USNM of Spalangia nigroaenea Curtis,

the species with which muscidarum has been synonymized by Bou^ek (1963, p. 449).

Pronotum with distinct anterior margin and crossline of punctures just anterior to posterior margin;

median area of pronotum smooth, lateral areas sparsely

punctate; scutellum with complete frenal crossline,

otherwise impunctate or with one or two weak punctures in posterolateral angles.

Spalangia nigra Latreille

Spalangia rugosicollis Ashmead, 1894, p. 36 [female].

This species was described from one female specimen. The type is labeled: "Mo., Spalangia rugosicollis

Ashm., Type No. 2168 U.S.N.M." The pedicel and

flagellum of each antenna and the right forewing are

missing; otherwise the type specimen is intact.

This type agrees in all diagnostic characters with

European specimens in the USNM of Spalangia nigra

Latreille, the species with which rugosicollis has been

synonymized by Boucek (1963, p. 446). Antennal

scape with longitudinal aciculations on inner face;

head and pronotum with umbilicate punctation and

smooth interstices; pronotum without anterior carina

but with isolated crossline of punctures just anterior

to posterior margin; pronotum with a smooth, median

triangle at posterior margin; scutellum with complete

frenal crossrow of punctures; petiole twice as long as

broad, with strong longitudinal rugae; gaster no wider

than thorax.

Spalangia endius Walker

Spalangia muscidarum var. stomoxysiae Girault, 1916, p. 57

[female].

Spalangia stomoxysiae Girault, Peck in Muesebeck et al.,

1951, p. 535.

This was described from two female cotypes. Lectotype labeled: "Hunter No. 2970, B. 18, Dallas, Tx.,

XI.24.12, Par. of Stomoxys calcitrans, Spalangia

muscidarum stomoxyiae (!) Girault, Lectotype No.

19674 U.S.N.M.," present designation of lectotype.

(The other cotype is a specimen of S. earneroni

Perkins.)

The lectotype of this species agrees in diagnostic

characters with European specimens in the USNM of

S. endius Walker, the species with which stomoxysiae

has been synonymized by Bou£ek (1963, p. 458). Pronotum lacking anterior carina, but with an isolated

crossrow of punctures just anterior to posterior margin,

this crossrow subangularly produced anteriorly on the

meson; median area of pronotum bare, lateral areas

sparsely punctate; scutellum with complete frenal

crossline of punctures, area anterior to this bare except

for two or three minute punctures.

Spalangia cameroni Fullaway

Spalangia muscidarum var. texensis Girault, 1920, p. 213

[male; new synonymy].

This form was described from two male specimens,

only one of which can now be found. Lectotype labeled:

"Hunter No. 2970, B.I8. Dallas, Tex., XI.24.12, Par.

of Stomoxys calcitrans, Spalangia muscidarum texensis Girault 3 type, Lectotype No. 20876 U.S.N.M."

The lectotype is a specimen of S. cameroni Fullaway, not S. endius Walker, the species with which it

has been synonymized by Bouc"ek (1963, p. 458).

Earlier, Peck (1951) placed it as a synonym of stomoxysiae Girault. Pronotum lacking anterior carinate

margin, but with an isolated crossline of punctures just

anterior to posterior margin; pronotum with lateral

punctures crowded and rugose, a median, longitudinal

dorsal groove extending from anterior border almost

to punctate crossline; notaulices terminating in a pair

of large pits; scutellum with a complete frenal crossline of punctures, scutellum smooth anterior to this.

Literature Cited

Ashmead, W. H.

1887. Studies on the North American Chalcididae, with

Descriptions of New Species, Chiefly from Florida.

Transactions of the American Entomological Society, 14:183-202.

1894. A Synopsis of the Spalangiinae of North America.

Proceedings of the Entomological Society of Washington,?,^) :27-37.

1904. Classification of the Chalcid Flies. Memoirs of the

Carnegie Museum, 1 (4) :xi+225-55l.
NUMBER 2

Bishopp, F. C.

1913. The Stable Fly (Stomoxys calcitrans L.), an Important Livestock Pest. Journal of Economic Entomology, 6:112-126.

Boucek, Z.

1963. A Taxonomic Study of Spalangia Latr. (Hymenoptera, Chalcidoidea). Ada Entomologica Musei

Nationalis Prague, 35:429-512.

1965. Descriptions of New Species of Spalangia Latr.

(Hym., Chalc.) from Africa and South America,

with Notes on Some Known Species. Ada Entomologica Musei Nationalis Pragae, 36: 593—602.

Girault, A. A.

1916. Descriptions of Four New Species of North American

Pteromalidae. Societas Entomologica, 31:56—58.

1920. New Serphoid, Cynipoid, and Chalcidoid Hymenoptera. Proceedings of the United States National

Museum, 58:177-216.

Howard, L. O.

1911. The House Fly. 312 pages. New York: Stokes.

Peck, O.

1951. In Muesebeck et al., Hymenoptera of America North

of Mexico, Synoptic Catalog, Superfamily Chalcidoidea. United States Department of Agriculture Monograph, 2:410-594.

1963. A Catalogue of the Nearctic Chalcidoidea (Insecta:

Hymenoptera). Canadian Entomologist, supplement

30, 1092 pages.

Richardson, C. H.

1913. An Undescribed Hymenopterous Parasite of the

Housefly. Psyche, 20(l):38-39.

Riley, C. V.

1894. In Riley, Ashmead, and Howard, Report upon the

Parasitic Hymenoptera of the Island of St. Vincent.

Journal of the Linnean Society of London,

25:56-254.

Ruhl, M.

1916. Liste neuerdings beschriebener oder gezogener

Parasiten und ihrer Wirte, V. Societas Entomologica, 31:60.

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by the original artwork. Duplicate copies of all material, including illustrations, should be retained

by the author. There may be several paragraphs to a page, but each page should begin with a new

paragraph. Number consecutively all pages, including title page, abstract, text, literature cited,

legends, and tables. The minimum length is 30 pages of typescript and illustrations.

The title should be complete and clear for easy indexing by abstracting services. Taxonomic

titles will carry a final line indicating the higher categories to which the taxon is referable:

"(Hymenoptera: Sphecidae)." Include an abstract as an introductory part of the text. Identify

the author on the first page of text with an unnumbered footnote that includes his professional

mailing address. A table of contents is optional. An index, if required, may be supplied by the

author when he returns page proof.

Two headings are used: (1) text heads (boldface in print) for major sections and chapters

and (2) paragraph sideheads (caps and small caps in print) for subdivisions. Further headings

may be worked out with the editor.

In taxonomic keys, number only the first item of each couplet; if there is only one couplet,

omit the number. For easy reference, number also the taxa and their corresponding headings

throughout the text; do not incorporate page references in the key.

In synonymy, use the short form (taxon, author, date, page) with a full reference at the end

of the paper under "Literature Cited." Begin each taxon at the left margin with subsequent

lines indented about three spaces. Within a taxon, use a period-dash (.—) to separate each

reference. Enclose with square brackets any annotation in or at the end of the taxon. For

references within the text, use the author-date system: "(Jones, 1910)" or "Jones (1910)." If

the reference is expanded, abbreviate the data: "Jones (1910, p. 122, pi. 20: fig. 1)."

Simple tabulations in the text (e.g., columns of data) may carry headings or not, but they

should not contain rules. Formal tables must be submitted as pages separate from the text, and

each table, no matter how large, should be pasted up as a single sheet of copy.

Illustrations (line drawings, maps, photographs, shaded drawings) can be intermixed

throughout the printed text. They will be termed Figures and should be numbered consecutively;

however, if a group of figures is treated as a single figure, the individual components should be

indicated by lowercase italic letters on the illustration, in the legend, and in text references:

"Figure 96." If illustrations (usually tone photographs) are printed separately from the text as

full pages on a different stock of paper, they will be termed Plates, and individual components

should be lettered (Plate 9b) but may be numbered (Plate 9: figure 2). Never combine the

numbering system of text illustrations with that of plate illustrations. Submit all legends on pages

separate from the text and not attached to the artwork.

In the bibliography (usually called "Literature Cited"), spell out book, journal, and article

titles, using initial caps with all words except minor terms such as "and, of, the." (For capitalization of titles in foreign languages, follow the national practice of each language.) Underscore

(for italics) book and journal titles. Use the colon-parentheses system for volume, number, and

page citations: "10(2) :5-9." Spell out such words as "figures" and "plates" (or "pages" when

used alone).

For free copies of his own paper, a Smithsonian author should indicate his requirements on

"Form 36" (submitted to the Press with the manuscript). A non-Smithsonian author will receive

50 free copies; order forms for quantities above this amount with instructions for payment will be

supplied when page proof is forwarded.