SMITHSONIAN CONTRIBUTIONS TO

ZOOLOGY

NUMBER 12

R. E. Crabiii, jr. Tracheotaxy as a

Generic Criterion in

Himantariidae, with

Proposal of Two New

Bothriogastrine Genera

(Chilopoda:

Geophilomorpha)

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ABSTRACT

Two new Bothriogastrine genera are proposed and based upon three species, two of

them new, thereby recording for the first time the presence of the subfamily in the

New World. The characterizations utilize a new intertaxic criterion, tracheotaxy, the

study of tracheal configurations and patterns, which is introduced herein and

described. Some evolutionary inferences are suggested. Additionally, other new criteria

are employed, e.g., the nature of the pretarsal parungues. A key to all known

Bothriogastrine genera is appended.

Introduction

The Himantariidae, a family of persisting ancient

habitus, is properly divisible into two subfamilies:

Himantariinae, occurring widely but almost exclusively

in the northern hemisphere; Bothriogastrinae,1 heretofore known only from Greece and the adjacent Near

East, including northern Africa and Asia. The most

useful and reliable (but not the only) feature distinguishing the two is the second maxillary isthmus, anteromedially strongly diastemate in the nominate

group, but somewhat protracted and not at all diastemate in the Bothriogastrinae. In terms of genera and

species the Himantariinae is much the larger.

To date several dozen species and more than a

dozen himantariid genera have been described from

North America including Mexico, chiefly by R. V.

Chamberlin.2 Over a period of many years and until

recently all of the specimens that I had examined from

North America were plainly referable to Himantariinae. Chamberlin, for his part, neither in publication nor in letters to me ever suggested any reason for

supposing that his forms were not similarly assignable.

So it was with some surprise that I discovered the presence recently of unquestioned bothriogastrines in Mexico and California. My surprise was not total, however,

for considering chilopod distributional patterns between eastern Asia and the North American west coast,

it would not have been unreasonable to predict the

1

 As used here, exactly equal to Attems' Bothriogastrini

(1929, p. 46) and to Verhoeff's Bothriogastrinae and Mesocanthinae (1925, p. 548).

2

 For a detailed systematic summary, see Crabill (1959,

pp. 117-126, 154-159).

R. E. Crabill, Jr., Department of Entomology, Smithsonian

Institution, Washington, D.C. 20560.

presence of bothriogastrines in this part of the New

World.

The Mexican Bothriogastrinae that I have examined

so far are divisible into two new genera and three

species, two of them new. All are described below, and

a revised key to all of the genera of Bothriogastrinae is

appended.

One of the chief generic characteristics that I rely

upon in this study makes use of what I have termed

tracheotaxy,3 utilizing kinds of criteria that have

never before been employed diagnostically below the

rank of Order. Because it is a new and unexplored

departure, a few words of explanatory orientation are

needed. The reader's attention is directed to Figures

22 and 23, which show semischematically, respectively

for Geoballus and Straberax, three typical segments

with their dorsal tracheal divisions and spiracles. Both

are in dorsal aspect, cephalic ends uppermost.

In Figure 22 the segments' spiracles (A) are indicated; from each there arises a single tracheal trunk

that soon bifurcates to give rise to a posterior ramus

( B) and an anterior ramus (c). Each posterior ramus

passes directly to a nodus (E) which receives nearly

all of the various tracheal components. Note that there

is one nodus posteriorly in each segment. In all Geophilomorpha known to me the connections of the posterior ramus, which directly connects the spiracular

trunk with the nodus, are unvarying; as we shall see, it

is the anterior ramus that is variable.

In Geoballus the anterior ramus (c) does not pass

directly to the next segment's nodus, although in most

non-himantariids it does pass directly to it. Rather, it

connects in the preceding segment with an internodal

8

 Tracheotaxy, a new term: the study of tracheae as systematic criteria.

1
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

connective (D) at a point just anterior to the nodus

of that segment. The internodal connectives pass from

one nodus to the next down the full length of the body

where they may function as a direct intersegmental

carrier of air, as an air storage reservoir, or as both.

Comparing Figure 23 (of Straberax) with the foregoing (of Geoballus), it may be seen that in the former

the posterior ramus has exactly the same connections as it has in the latter; the internodal connectives,

however, are very different. In Straberax, in effect,

each anterior ramus terminates not in the preceding

segment, but in the pre-antecedent segment at the

nodus. Therefore, this type of connection might be

described as saltate, whereas the former kind could be

termed seriosaltate. In most Geophilomorpha the anterior ramus connects directly with the nodus of the

antecedent segment, and internodal connectives are

absent; this simplest of all arrangements I term serial.

The rear terminus of each internodal connective has

detached, but the interior terminus has remained intact, so that the anterior division of the system is in fact

composite, consisting of the continuous anterior ramus

and internodal connective. It seems clear that the dorsal tracheal system of the Straberax-type represents a

derivative of that of the Geoballus-type.

I have encountered both tracheal types or their proximate variations throughout the family; of the two

the Geoballus-type is by far the more common and in

both subfamilies. But the uncommon Straberax-type

has also been discovered in both subfamilies, where

the anterior ramus' extension may skip up to five segments before terminating at a nodus. From these

studies, whose elaboration transcends the scope of this

paper, I have drawn several hypotheses. (1) In himantariid evolution there was no single point-of-departure

at which the saltate Straberax-type of tracheation arose

once and only once from the seriosaltate Geoballustype. I believe that the saltate type has arisen many

times in different evolving lines (independently in the

two subfamilies) from the common and antecedent

seriosaltate type. (2) A given tracheal configuration

seems constant at least throughout a genus, and it

seems often, though not always, constant throughout a

group of related genera.

This study was undertaken with the aid of a grant

from the National Science Foundation. I would like

to thank Dr. G. E. Ball, Jr., and Mr. D. R. Whitehead

of the University of Alberta, Dr. W. T. Keeton of

Cornell University, and Dr. B. D. Valentine of the

Ohio State University for the material that they

have so generously placed at my disposal.

Geoballus, new genus4

Comparable with both Bothriogaster and Straberex,

this genus appears to resemble Bothriogaster more

closely in what I take to be the more fundamental features. They differ conspicuously as follows: In Bothriogaster—(1) Certain sternites, usually anterior to midbody length, with deep fovea-like sternobothria. (2)

The legs at least of the foregoing segments with hypertrophied parungues that equal or exceed their claw

lengths. In Geoballus—(1) No sternites with sternobothria. (2) No parunguis equal to or exceeding its

claw; the former always much shorter.

Antennae: The more proximal articles slightly dorsoventrally flattened, those remaining terete, not flattened; distally slightly attenuate. Bucca: Mesal margins not sclerotized and pigmented.5

 Labral embayment

shallow; labral sides not completely divisive. First maxillary coxosternal lappets absent; telopodite lappets

present or absent. Second maxillary claw without basal

bristles or filaments; isthmus anteromedially without

diastema, protracted. Tarsungula without basal denticle, other articles unarmed; presentation dorsomesal.

Prosternal pleurograms present and complete to condyles. Tergites without paramedian sulci. Major paratergites absent. Tracheation seriosaltate type. Sternites

without sternobothria; anterior and posterior body

sternites wider than long. Porefields strongly transverse,

very wide, subreniform; present on 1 through penult

without interruption. Ultimate pretergite bilaterally

fissate, the pleurites discrete; presternite divided;

sternite wider than long, posteriorly embayed. Each

coxopleuron with one dorsal and one ventral glandular

fossa bordering the tergite and sternite. Telopodite relatively short, in the male slightly clavate. Anal pores

absent.

Two species are referable to the genus.

TYPE-SPECIES.—Geoballus caputalbus, new species.

Original designation.

4

 Geoballus, dedicated with gratitude to George Ball, carabid connoisseur and esteemed old friend, who for more than

20 years has generously sent me wonderful Chilopoda from

wherever his coleopterous safaris took him.

* Throughout the Himantariinae and in some Bothriogastrine genera, these margins are heavily sclerotized and pigmented : a new character.
NUMBER 12 3

Geoballus caputalbus, new species °

The type-species is, of course, most like the genus'

only other species, bebelus, new species, differing notably as follows: In caputalbus—(1) Leg pairs 91. (2)

Telopodite lappets absent. (3) Final porefields straight,

slightly shorter. In bebelus—(1) Leg pairs 61. (2)

Telopodite lappets present. (3) Final porefields somewhat bent, slightly longer.

HOLOTYPE.—Male. Mexico, Oaxaca, 21.8 miles

north of Tuchatengo. 18 August 1966; G. E. Ball, Jr.,

and D. R. Whitehead, legg. Deposited in the United

States National Museum.

GENERAL CHARACTERS.—Length 45 mm. Leg pairs

91. Shape: Dorsoventrally flattened; of essentially uniform width. Color sordid yellow. Vestiture extremely

sparse and short.

DETAILED DESCRIPTION.—Antennae: Not attenuate;

dorsoventrally only slightly flattened. All articles except 14th wider than long. Articles (4-5)-14 finely

densely setose; l-(3-4) with fewer and longer setae.

14th article setosensoria ectally and mesally, located

slightly distad of midlength. Cephalic plate: Greatest

width to length=ll:8. Shape: anteriorly rostrate;

sides slightly excurved; rear straight. Cephalic suture

not apparent. Clypeus: Without plagulae or anterocentral fenestra. Paraclypeal sutures nearly straight,

complete to outer ends of fulcra. With 30-40 short

weak setae on anterior half; posterior half glabrous.

Labrum: Deeply embayed, teeth 6 + 7; entirely demarcated from clypeus by hyaline suture. Mandible: Shaft

long, flat, proximally curved. Corpus with 7 pectinate

lamellae and 1 dentate lamella having 10 teeth. First

maxillae: Coxosternum medially deeply incised; lappets absent. Medial lobes relatively long, subtriangular.

Telopodite lappets absent; unipartite, not demarcated

from coxosternum. Second maxillae: Isthmus deep

anteroposteriorly, medially protracted, not diastemate.

Weak postmaxillary sclerites present. Telopodite: Both

basal condyles present, strong; articles 1 and 2 wider

than long; claw robust, without basal bristles and filaments, shallowly excavate within.

Forcipular segment: Tarsungula relatively long and

relatively little curved; edges smooth, not serrulate;

presentation dorsomesal. Poison calyx linear, not subspherical. Prostemal pleurograms present but weak,

complete to condyles. Anterocentral denticles absent.

9

 Named in honor of D. R. Whitehead, who with G. E. Ball

collected these and plethorous other excellent specimens.

Tergites: Essentially glabrous. Paramedial sulci absent. Pleurites: Paratergites absent. Spiracles subcircular. Legs: Sparsely shortly setose, robust. Pretarsi:

Robust and very long, not excavate beneath; parungues equal, about one-fourth as long as claws. Sternttes:

All wider than long. Very shortly, sparsely setose. Without carpophagus-stmctures, foveae, stigmoid, or virguliform structures. Porefields: On 1 through penult

without interruption, on anterior body subcentral in

position, on rear body posterior in position; transversely

wide and straight to exceedingly wide, anteroposteriorly short, each posteriorly slightly embayed or

subreniform.

Ultimate pedal segment: Pretergite bilaterally fissate, the pleurites discrete. Tergite much wider than

long, 17:8. Presternite medially separated. Sternite:

Greatest width to length, 12:7; sides straight and convergent, rear slightly incised. Coxopleuron: Moderately inflated, encroaching slightly upon penult; gland

canals opening into fossae along edges of stemite and

tergite, hence glands not in subsurface pockets. Telopodite: Slightly longer than penult; dorsoventrally

slightly flattened and distally slightly clavate; with two

tarsalia; pretarsus absent.

Postpedal segments: Syntergum densely hirsute. Female gonopods long, well separated, each biarticular.

Anal pores absent.

Geoballus bebelus, new species

Geoballus bebelus most resembles caputalbus; their

diagnostic differences are summarized under the latter's description.

HOLOTYPE.—Male. Mexico, Veracruz, San Andres

Tuxtla, Vulcan San Martin, 3,500'. 1 August 1959;

W. T. Keeton and B. D. Valentine, legg. Deposited

in the United States National Museum.

GENERAL CHARACTERS.—Length 40 mm. Leg pairs

61. Shape: Dorsoventrally flattened; body essentially

uniform in width front to rear. Color sordid yellow.

DETAILED DESCRIPTION.—Antennae: Not attenuate,

only slightly flattened dorsoventrally. Each article except 14th very slightly wider than long. Articles 6-14

finely densely setose, 1-5 with fewer but longer setae.

14th article setosensoria ectally and mesally, located

far distad of midlength. Cephalic plate: Greatest width

to length 19:17. Shape: anteriorly rostrate; sides excurved, rear straight. Cephalic suture not apparent.

Clypeus: Without plagula or fenestra. Paraclypeal
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

sutures nearly straight, complete to outer ends of fulcra.

Setae about 30, on anterior third of clypeus, the rest

glabrous. Labrum: Relatively shallowly embayed.

Teeth 6+7. Demarcated from clypeus by a hyaline

suture. Mandible: Shaft long and flat, proximally

curved. Corpus with 7 pectinate and 1 lamellate lamellae, the latter with 9 teeth. First maxillae: Coxosternum medially deeply incised; lappets absent; medial

lobes relatively long and subtriangular. Telopodites

small, lappets present; unipartite, not discernibly demarcated from coxosternum. Second maxillae: Isthmus

deep anteroposteriorly, medially protracted, not diastemate. With weak postmaxillary sclerites. Telopodite: Dorsal and ventral condyles present; claw

robust, without basal bristles or filaments, shallowly

cochlear. Forcipular segment: Tarsungula relatively

long, relatively little curved; edges smooth, not serrulate; poison calyx linear; presentation mesodorsal.

Prosternal pleurograms present, complete to condyles,

weak; anteromesal denticles absent. Tergites: Essentially glabrous. Paramedial sutures absent. Pleurites:

Paratergites absent. Spiracles subcircular. Legs:

Sparsely, shortly setose, robust. Pretarsi: Robust and

very long, not excavate beneath; parungues equal,

about one-fifth as long as claw. Sternites: All much

wider than long. Very shortly, sparsely setose. Without

carpophagus-structures, foveae, stigmoid or virguliform

structures. Porefields: On 1 through penult without

interruption; on anterior body subcentral in position, on

rear body posterior in position; tranversely wide to exceedingly wide and slightly arched, anteroposteriorly

short, somewhat reniform.

Ultimate pedal segment: Pretergite bilaterally fissate, the pleurites discrete. Tergite: Much wider than

long, 15:10. Presternite medially separated. Sternite:

Greatest width to length 18:13; sides straight and convergent, rear strongly incised. Coxopleuron: Moderately inflated; encroaching slightly upon penult; gland

canals opening into fossae along edges of sternite and

tergite, hence not in subsurface pockets. Telopodite:

Slightly longer than penult; dorsoventrally flattened,

not notably clavate; densely finely hirsute; with two

tarsalia; pretarsus absent.

Postpedal segments: Syntergum densely hirsute.

Male gonopods long, biarticular. Anal pores absent.

Straberax, new genus

 7

Straberax seems most like Geoballus; the two differ

most conspicuously as follows: In Geoballus—(1) Tracheaton is, like that of Bothriogaster, seriosaltate as in

Figure 22. (2) Coxopleural glands in deep, subcircular

crypts. (3) Second maxillary telopodite lappets, if

present, normal, not amalgamated with telopodites. In

Straberax—(1) Tracheation is saltate as in Figure 23.

(2) Coxopleural glands in shallow elongate fossae. (3)

Second maxillary telopodite lappets present but

curiously amalgamated with telopodites.

Antennal proximal articles dorsoventrally slightly

flattened, the more distal articles not flattened, terete;

distally slightly attenuate. Buccae: Mesal margins not

sclerotized and pigmented. Labral embayment deep,

completely divisive. First maxillary coxostemal lappets

absent; telopodite lappets present but amalgamated

with telopodites. Second maxillary claw without basal

bristles or filaments; isthmus anteromedially without

diastema, protracted. Tarsungula without basal denticles, all articles unarmed; presentation dorosomesal;

article 1 basomesally with a prominent pigmented

macula. Prosternal pleurograms strong and complete to

condyles. Tergites without paramedian sulci. Major

paratergites absent. Tracheation saltate. Sternobothria

absent. Anterior body sternites wider than long;

posterior sternites longer than wide or rectilinear.

Porefields transversely not notably wide; present on 2

through penult without interruption. Ultimate pretergite bilaterally fissate, pleurites discrete; presternite

divided, sternite wider than long, posteriorly embayed.

Each coxopleuron with one dorsal and one ventral

glandular crypt, these deep, subcircular, heterogeneous. Telopodite relatively short, clavate in male, not

so in female, dorsoventrally slightly flattened; tarsalia

2, without pretarsus. Anal pores absent.

The genus is known to contain one species.

TYPE-SPECIES.—Gosiphilus morelos Chamberlin

(= Straberax morelos (Chamberlin)). Monotypic and

original designation.

Straberax morelos (Chamberlin), new combination

Gosiphilus morelos Chamberlin, 1943, p. 11.

GENERAL CHARACTERS.—Length to 56 mm. Leg

pairs 77-97. Shape: Dorsoventrally flattened; essentially uniform in width front to rear. Color sordid yellow.

DETAILED DESCRIPTION.—Antennae: Length to head

length (2—2.5) : 1. Proximal 3-4 articles slightly

flattened, the remainder terete; distally slightly attenuate; all articles except 14th wider than long. Articles 6-

14 more setose than 1-7. Article 14 ectally and mesally

with a setosensorium immediately proximad of mid7

 A name of arbitrary coinage.
NUMBER 12

length. Head: Greatest width to length, 9:7. Anteriorly

rostrate, sides excurved, rear straight. Cephalic suture

weakly indicated. Clypeus: Fenestra and plagula absent. Paraclypeal sutures straight, slightly oblique, complete, ending at ectal third of the fulcra. Setae

numerous, short, about 40 on each side of the glabrous

midline; prelabrals absent. Buccae without deeply pigmented sclerotic margins. Labrum: Protuberant and

dependent; deeply medially embayed, there with some

14—18 teeth. Mandible: Proximal half of shaft slightly

curved, expanded. Corpus with 4 pectinate and 1

dentate lamellae, the latter with 10—11 teeth. First

maxillae: Coxosternum essentially divided medially

into right and left halves, i.e., deeply, widely cleft,

lappets absent. Medial lobes short. Telopodites:

Weakly demarcated from coxosternum, not discernibly

biarticular; evidently broadly amalgamated with lappets. Second maxillae: Isthmus anteroposteriorly deep;

anteromedially protracted, not diastemate. Postmaxillary sclerites present, atrophied. Telopodite: Articles

1 and 2 wider than long; both basal condyles present;

terminal claw short, robust, evidently cochlear, without

basal bristles or filaments.

Forcipular segment: Prehensors: Claw relatively

short and but slightly curved; smooth, not serrulate,

presentation mesodorsal; poison calyx linear,

elongate; all articles without denticles; article 1

posteromesally with a prominent, dark macula.

Prosternum: Without anterior denticles; pleurograms

strong, complete to condyles.

Tergites: Paramedial sulci not discernible.

Pleurites: Paratergites absent. Spiracles subelliptical,

vertical. Legs: Robust, essentially glabrous; each

pretarsus with two equal parungues, these about onethird as long as claw. Sternites: On anterior half of

body wider than long; on rear of body essentially rectilinear. Porefields: On 2 through penult without

interruption; on anterior body subcentral in position,

transversely subelliptical, thereafter becoming

smaller in size and more posterior in position. Without foveae, stigmoid, and virguiliform structures.

Ultimate pedal segment: Pretergite laterally fissate, the pleurites discrete. Tergite much wider than

long. Presternite medially separated. Sternite: Greatest width exceeds length, 8:7; sides essentially

straight, rear slightly embayed. Each coxopleuron:

Inflated but not encroaching upon penult; with one

dorsal and one ventral heterogeneous, multicanaliculate glandular cavity, opening respectively along

tergite and sternite margins. Telopodite: In the

female as long as penult and no more robust, in the

male as long as penult but notably more robust and

slightly clavate; with two tarsalia; pretarsus absent.

Postpedal segments: Female gonopods medially

contiguous but not fused, each distinctly biarticular.

Anal pores absent.

SPECIMENS STUDIED.—I have studied seven specimens from three Mexican states; all agree closely

with each other. All are deposited in the collection of

the U.S. National Musuem. Their leg-pair dispersion

is as follows: 77 leg-pairs, 1; 89, 5; 95, 1. Locality

data is as follows: Morelos: 9.1 miles east of Cuernavaca, Canon de Lobos at 4,300*, 24.VII.66, Ball and

Whitehead; Cuatla, 24.IV.63, Whitehead. Oaxaca:

18.4 miles south of Suchixtepec at 4,500*, 20.111.66,

Ball and Whitehead. Veracruz: 5.8 miles west of

Ojazarco at 5,400', 10.VII.59, Keeton and Valentine.

Key to BOTHRIOGASTRINAE

1. Coxopleura entirely without glands and their associated cavities, fossae, canals, and pores

(Africa, Asia) Mesocanthus Meinert

Coxopleura with glands in cavities or fossae, with associated canals and pores 2

2. Each coxopleuron with only one dorsal gland cavity, this exiting along tergite; ventral glands

absent. Second maxillary claw with robust basal bristles. (Africa, Asia)

Polyporogaster Verhoeff

Each coxopleuron with dorsal and ventral glands, these exiting via one dorsal and one ventral

gland cavity or fossa. Second maxillary claw basally unarmed 3

3. Tracheal anterior rami saltate as in Figure 23. Second maxillary telopodite lappets present

but amalgamated with telopodites (Mexico) Straberax, new genus

Tracheal anterior rami seriosaltate as in Figure 22. Second maxillary telopodite lappets, if

present, normal, not amalgamated with telopodite 4

1. Certain sternites (usually anterior to midbody length) with prominent, deep sternobothria.

Legs, at least of foregoing segments, with hypertrophied parungues equal to or exceeding

their claw length (Africa, Asia) Bothriogaster Selivanoff

Sternites without sternobothria. Parungues not hypertrophied, always much shorter than their

claws (Mexico) Gtoballus, new genus
6 SMITHSONIAN CONTMBUTIONS TO ZOOLOGY

Literature Cited CrabiU, R. E.

1959. A Synonymical List of American Himantariidae,

Attems, C. with a Generic Key and Description of a New

1929. Geophilomorpha. Lieferung 52 in Das Tierreich. Genus. Entomological News, 70(5-6): 117-126,

154-159.

Chamberlin, R. V. Verhoeff, K. W.

1943. On Mexican Centipeds. University of Utah 1925. Chilopoda. In Bronn, Klassen und Ordnungen,

Biological Series, 7 (3): 11. 5:548.
NUMBER 12

8

FIGURES 1-11.—Geoballus caputalbus, holotype: 1, 5th sternite (a=porefield); 2, ultimate

sternite minus 8; 3, penult sternite; 4, ultimate pedal and postpedal sternites (a=pores in coxopleural fossa). Geoballus bebelus, holotype: 5, 8th sternite; 6, ultimate sternite minus 8; 7, penult

sternite. Straberax morelos (Chamberlin): 8, 10th sternite; 9, ultimate sternite minus 10;

10, penult stemite; 11, ultimate pedal and postpedal sternites (a=concealed gland canals in

crypt, b=crypt opening).
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 12-21.—Geoballus caputalbus, holotype: 12, right side of prostemum with prehensor

(a=pleuroprosternal suture, b=pleurogram) ; 13, labrum with left fulcrum and paraclypeal

suture, 14, ultimate antennal article (a=setosensorium); 15, first and second maxillae (a=

protracted medial portion of isthmus). Geoballus bebelus, holotype: 16, first maxillae, left

telopodite with medial lobe (a= lappet); 17, ultimate antennal article showing both setosensoria.

Straberax morelos (Chamberlin): 18, right side of prostemum with prehensor (a=poison calyx,

b=macula); 19, ultimate antennal article showing both setosensoria; 20, labrum with both fulcra

and adjacent areas (a=right, paraclypeal suture, b=right fulcrum, c=clypeolabral suture, d =

labral embayment); 21, first and second maxillae (alright lappet, b=postmaxillary sclerite,

c=salivary pore, d=protracted portion of isthmus).
NUMBER 12

Geoballus

FIGURE 22.—Geoballus caputalbus, holotype, semischematic FIGURE 23.—Straberax morelos (Chamberlin) (A=spiracle;

(A=spiracle; B=posterior ramus; C=anterior ramus; D = B=posterior ramus; C=anterior ramus; D=internodal coninternodal connective; E:=nodus). nective; E=nodus).

US. GOVERNMENT PRINTING OmCE: 1969 O—333-340

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Copy must be typewritten, double-spaced, on one side of standard white bond paper, with

1

 l/t" top and left margins, submitted in ribbon copy with a carbon or duplicate, and accompanied

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.