A review of the silverfish (Lepismatidae, Thysanura) of the United States and the Caribbean area

Amencan Museum

Novitates

PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY
CENTRAL PARK WEST AT 7QTH STREET, NEW YORK, N. Y. T0024

NUMBER 2481 FEBRUARY 16, 1972

A Review of the Silverfish
(Lepismatidae, Thysanura)

of the United States and the Caribbean Area

By PEDRO WyGopZINsky!

ABSTRACT

An analysis of the composition of the lepismatid fauna of the United States and
the Caribbean area is made, with special emphasis on the geographic origin of the
species. Among the 13 species found in the United States, only three are autoch-
thonous; among the 13 species reported from the Caribbean area, six seem to be
endemic. All other species probably have been introduced by man. Keys for the
determination of the lepismatids of the United States and of the Caribbean area
are followed by an enumeration of all species concerned, with distributional data
and taxonomic comments. Four species are reported for the first time from the
United States, viz., Acrotelsa collaris, Ctenolepisma diversisquamis, C’. targionii, and
Stylifera gigantea. Lepisma mucronata Packard is synonymized with Acrotelsa collaris
(F.), and Ctenolepisma reducta Folsom with C. diversisquamis Silvestri; Ctenolepisma
campbell Barnhart is transferred to Thermobia. The female of Ctenolepisma hum-
melincki Wygodzinsky is described for the first time. Peliolepisma calva, until now
only known from Ceylon, is reported from Guyana and Cuba, where it is
supposedly a common house lepismatid.

RESUMEN

Este trabajo constituye una puesta al dia de nuestros conocimientos sobre la
fauna de lepismatidos (trazas) de los Estados Unidos de Norte America y de la
regién del Caribe. Se hace una analisis de la composicién faunistica de las dos
areas, con consideracién especial de las origenes geogr4ficas de las especies. Entre

‘Curator, Department of Entomology, the American Museum of Natural History.

2 AMERICAN MUSEUM NOVITATES NO. 2481

las 13 especies encontradas en los Estados Unidos, solamente tres son autéctonas;
y de las 13 halladas en el area del Caribe, seis parecen ser endémicas. Todas las
otras especies probablemente han sido introducidas por el hombre. Se presentan
claves separadas para la determinacidén de todos los géneros y especies, seguidas
de una enumeracién comentada de las especies, con su sinonimia primaria y datos
sobre su distribucién. Se establece la sinonimia de Lepisma mucronata Packard con
Acrotelsa collaris (F.), y la de Ctenolepisma reducta Folsom con Ctenolepisma diver-
sisquamis Silvestri. La hembra de Ctenolepisma hummelincki se describe por la primera
vez. Ctenolepisma targionit, conocida de la subregion mediterranea, se sefiala por
primera vez del hemisferio occidental (Carolina del Sur). Peliolepisma calva Ritter,
antes solamente conocida de Ceylan, se senala de la Guyana y de Cuba, donde se
informe es comun en las habitaciones. Ctenolepisma campbelli Barnhart, especie
sinantropa de Ohio y Pennsylvania, se transfiere al género T7hermobia.

INTRODUCTION

Silverfish, firebrats, and related species are frequently commensals of
man, in whose habitations and storage buildings they have established
themselves, and where they may cause considerable damage. Several of
these synanthropic species have been used in the laboratory in research on
physiology. The literature on economic importance and physiology of
these insects is voluminous and is not reviewed here.

The present paper is concerned with the classification, nomenclature,
origin, and distribution of lepismatids found in North America and the
Caribbean. The last key to synanthropic lepismatids of the United States,
published by Slabaugh (1940) is out of date; a new key is now provided.
No distinction is here being made between synanthropic and free-living
species; the latter may locally invade human habitations, and synan-
thropic species may secondarily become free-living again.

A second key allows the determination of the lepismatids of the Carib-
bean area, including the Caribbean coast of Venezuela and Colombia. It
is quite possible that certain species included in this key and not known
from the United States will later be found in such regions as southern
Florida and adjacent islands, a possibility that should be taken into
account when identifying material from the extreme south.

ACKNOWLEDGMENTS

For the present study I have used the collections of the American
Museum of Natural History (AMNH) as well as material received from
other public institutions and individual scientists. I owe thanks to the
following for loan of material and other assistance: Dr. J. N. Belkin,
University of California at Los Angeles (UCLA); Dr. P. J. Darlington,
Museum of Comparative Zoology, Harvard University; Dr. P. Hurd,
formerly of the California Insect Survey (CIS), University of California,

1972 WYGODZINSKY: SILVERFISH 3

Berkeley; Dr. K. Krishna, the American Museum of Natural History;
Dr. J. E. Luke, Department of Entomology, Pennsylvania State Uni-
versity, College Park; Dr. J. A. Payne, formerly at Clemson University,
Clemson, South Carolina; Dr. E. H. Slifer, Glenside, Pennsylvania;
Dr. C. A. Triplehorn, Ohio State University, Columbus; Dr. H. V.
Weems, Florida State Collection of Arthropods (FSCA), Gainesville; and
Ing. Fernando de Zayas, Instituto de Biologia, La Habana, Cuba.
All photographs and drawings were made by the author.

GEOGRAPHICAL DISTRIBUTION

The geographical distribution of Neotropical Thysanura has been
analyzed by Wygodzinsky (1967); that paper should be consulted to
supplement data contained in the present study.

Any interpretation of distribution patterns of lepismatids is encumbered
by the frequency with which species of this family are introduced by man
into areas and regions different from those where the species are native.
This is facilitated by the ease with which domestic or peridomestic
populations of silverfish originate from free-living ones, and, inversely, the
frequent derivation of secondarily free-living populations from domestic
ones. Not all free-living populations of lepismatids found in areas or
regions that are not their original ones have necessarily arisen from
domestic populations. Lepisma wasmanni Moniez, for instance, a circum-
Mediterranean species also found in the coastal deserts of Peru has never
been reported as associated with man; it probably was carried to Peru
with soil or plant material.

Table 1 shows the species now known to occur in the United States and
the Caribbean area. Species marked with an italic a are believed to be
native, or at least have not been reported from anywhere else. All other
species are considered to have been introduced.

It is remarkable that although the total number of species found in the
United States and the Caribbean area is identical (13), the faunistic
composition of these adjacent areas, which even slightly overlap in
southern Florida, is quite different. Only five species share the two areas.
Three of these (A. collaris, C. longicaudata, and L. saccharina) are common
in households, and the two others (C. diversisquamis and S. gigantea) are
common free-living, wide-ranging species reaching their northern limit in
southern Florida.

The species not shared comprehend precinctive and introduced forms;
the latter are examined first.

Ctenolepisma ciliata and C. lineata pilifera, originating from the Mediter-
ranean area, may not be able to tolerate the comparatively hot and

4 AMERICAN MUSEUM NOVITATES NO. 2481

humid conditions of the Caribbean area; the second species is very
common in the Southwest.

The only record of C. targionit from the United States is based on a
sample of what apparently is a localized population in South Carolina;
its absence from the Caribbean is thus not surprising. On the other hand,

TABLE 1
DisTRIBUTION OF LEPISMATIDAE IN THE UNITED STATES AND CARIBBEAN AREA

United States Caribbean Area

Acrotelsa collaris Acrotelsa collaris
Allacrotelsa spinulata®
Ctenolepisma ciliata
Ctenolepisma diversisquamis Ctenolepisma diversisquamis
Ctenolepisma dubitalis®
Ctenolepisma hummelincki*
Ctenolepisma lineata pilifera
Ctenolepisma longicaudata Ctenolepisma longicaudata
Ctenolepisma targioniana
Ctenolepisma targronit
Ctenolepisma versluyst®
Heterolepisma horni*
Heterolepisma sp. (Bimini)
Lepisma saccharina Lepisma saccharina
Leucolepisma arenaria®
Murolepisma deserticola®
Peliolepisma calva
Stylifera gigantea Stylifera gigantea
Stylifera impudica®
Thermobia campbell
Thermobia domestica

"Believed to be native.

C. targioniana ranges widely over the tropics, and might imaginably find a
foothold in southern Florida.

Peliolepisma calva is also a tropical species, free-living in Ceylon but
apparently domestic in Guyana and Cuba; this adaptability makes the
species a candidate for accidental introduction and possible establishment
in the warmer parts of the United States.

Of the two species of Thermobia, domestica is well established in the
United States, not only in human habitations but also with secondarily
free-living populations in the West and Southwest. Its apparent absence
from the Caribbean may be attributable, as in Ctenolepisma ciliata and C.

1972 WYGODZINSKY: SILVERFISH 5

lineata pilifera, to the preference for a drier climate by the species. Ther-
mobia campbelli is only known from two domestic populations in the eastern
United States, and no domestic or free-living populations are known from
anywhere else. The time of arrival and place of origin of the species are

Fics. 1, 2. 1. Allacrotelsa spinulata. 2. Thermobia domestica.

unknown, but as the remaining species of Thermobia occur naturally in the
south of the Palearctic and in the Ethiopian region, it may be hypothe-
sized that 7. campbell: will eventually be found in that area.

The precinctives are of special interest; all are free-living.

Allacrotelsa spinulata is very common in the western and southern United
States. The species has a considerable ecological range, but it does not
occur in extremely hot and arid or hot and moist conditions. Signifi-
cantly, it seems to be absent from southern Florida and has not been
reported from the Caribbean. A second species of the genus is Mediter-

6 AMERICAN MUSEUM NOVITATES NO. 2481

4

Fics. 3-5. 3. Lepisma saccharina. 4. Ctenolepisma lineata pilifera. 5. Thermobia
campbell.

ranean in distribution, and the third is known only from Baltic amber
(Wygodzinsky, 1961).

Leucolepisma arenaria and Maiurolepisma deserticola are both desert forms
found only in the Southwest. Both genera are monotypic, and probably
are specialized derivatives, the first of Ctenolepisma-like, the second of
Prolepismina-like ancestors. |

None of the Caribbean precinctives belongs to an endemic genus, and as
the world fauna of the Lepismatidae becomes better known, some of these
species now considered precinctive may turn out not to be endemic at all.

The great majority of the species of the large genus Ctenolepisma are
native to the Old World (Africa, southern Europe, and Asia), and the
presence of three supposedly native species in the Caribbean is surprising.
Ctenolepisma hummelincki and C. versluysi are not different in any significant
way from most other species of the genus; only C. dubitalis is highly
aberrant morphologically compared with the rest of the genus. Wygod-
zinsky (1967) thought that the fact that dubitalis is marginal taxonomically
as well as geographically in relation to the main stock of Ctenolepisma
suggests the hypothesis that it or its ancestors arrived on the scene much
earlier than the rest of the species of Ctenolepisma, viz., that it is truly
autochthonous.

1972 WYGODZINSKY: SILVERFISH 7

Heterolepisma is an old genus with a basically amphinotic distribution
pattern, with several native species found in South America. Although
there is no prima facie evidence to suggest that the Caribbean species are
not autochthonous, it should be mentioned that at least one species of the
genus, trisetosa (Escherich), is known to occur in the Brazilian state of
Pernambuco, in East and West Africa, on Lombok, and in Japan; this
clearly indicates that passive dispersal is a possibility in this genus.

The case of the two species of Stylzfera is far from clear. S. impudica, listed
as an endemic, seems to be basically restricted to the Caribbean. A record
of a specimen caught on a boat in Oceania (Paclt, 1966) is highly doubtful
(see Systematics, p. 22). Paclt (1966) also reported specimens from
Hamburg (Germany), introduced from Venezuela or Colombia; even if
correctly identified, these finds do not change the basic picture.

Stylifera gigantea is very common in the Caribbean area where it ranges
from the mainland of South America to southern Florida, but it is also
exceedingly common in the coastal desert of Peru; a closely related form of
perhaps only subspecific difference occurs on the Galapagos Islands. It is
possible that the areas mentioned represent the natural range of the species,
a common distribution pattern in other animals and in plants. On the
other hand, all the 13 remaining species of Stylifera are Pacific in distribu-
tion (Wygodzinsky, 1967), and it is possible that the two Caribbean
species are not autochthonous.

SYSTEMATICS

KEY TO THE FAMILIES OF THYSANURA IN NorTH AMERICA
AND THE CARIBBEAN

1. Eyes and ocelliabsent . . . . . . . . . +. . . Nicoletiidae
EYesPprcsenta!. 27.8 50 3a wom OF te Bie oe Be pot lx ben
2. Ocelli present; scales absent. . . . . . . . +. . Lepidotrichidae
Ocelli absent; scales present. . . . . . . . +. +. . Lepismatidae

Nicoletiidae are subterranean or cavernicolous, sometimes myrmeco-
philous or termitophilous; they are never pigmented, and their scales are
colorless. The North American species are not well known.

Lepidotrichidae are found in leaf litter and under bark; their pigment
is reddish. Only one genus and species is known, precinctive to a small
area in northern California.

Lepismatidae comprehend the most commonly encountered species;
they are found free-living, in birds’ nests, or associated with humans; a few
are myrmecophilous or termitophilous. They frequently possess hypo-
dermal pigment, and the scales of the dorsal surface may form complex
patterns (fig. 2) or not (fig. 1). Although most species are tropical or sub-

8 AMERICAN MUSEUM NOVITATES NO. 2481

tropical, man has carried some to temperate climates where they survive
in shelter provided by their host.

The taxonomy of lepismatids is based to a large extent on the structure
and distribution of large, regularly arranged setae, the macrochaetae
(figs. 7C, E; 10G). These may not be extant in poorly preserved specimens,
but sockets indicate the place of their insertion (fig. 10D, F). Some
experience and adequate optical equipment are necessary to analyze the
arrangement of the macrochaetae; in case of doubt, a slide mount of the
whole insect or of critical structures must be prepared.

I would like to point out that although the present survey includes all
species now known from the areas under consideration, further species
will probably be found in the future. Caution is thus required when
determinations are made.

KEY TO THE LEPISMATIDAE OF THE UNITED States, ExcLupiInc Hawa

1. Last tergum triangular, sharply pointed and with more than 1+1 bristle-

combs (fig.6D) . . 2
Last tergum not sharply pointed (igs. TF; 8C, E, H, K; 10F) or, hee tri-
angular, with only 1+-1 bristle-combs (fig. 7D) wees a)

2. Submedian fields of macrochaetae of head capsule narrowly elongate, not
attaining anterior border of head capsule (fig. 6A); prosternum covered by
fore coxae from below, and with a tuft of macrochaetae centrally (fig. 6C);
ovipositor with fossorial spines oe a on posterior gonapophyses (fig. 6G);
male with parameres. . . . . Aecrotelsa collaris

Submedian fields of macrochaetae on “head capsule broadly triangular,
attaining anterior border of head capsule (fig. 6B); prosternum covering
fore coxae from below, without central tuft of macrochaetae (fig. 6E);

ovipositor without fossorial spines .. . . Stylifera gigantea
3. Macrochaetae Sine often bifid or trifid apically (fig. 7E); males with
parameres. 4

Macrochaetae barbed or feathered (fig. 10G); males lacking Derathercse eS
4, Last tergum truncate or faintly rounded apically (fig. 7F); apical segment of
labial palp wider than long (fig. 7H); dorsum of well-preserved living
specimens uniformly slate gray (fig. 3) . . . . . Lepisma saccharina
Last tergum triangular (fig. 7D); apical segment of labial palp longer than
wide (fig. 7G); dorsum of Serre living specimens dark brown
(fig.1) .. . . . Allacrotelsa spinulata
5. Insects golden yellow, not more than 6 mm. when mature, generally myrme-
cophilous; their shape as shown in figure 7A; caudal appendages very short,

not longer than maximum width of abdomen (fig. 7A) .
. Murolepisma deceriola
Color different; when mature, ‘length 61 mm. or more; shape more elongate
(figs. 2, 4, 5); caudal appendages elongate, much longer than maximum
width of abdomen (figs. 2,4,5) . . . 6
6. Lateral claws at least as long as second tarsomere (fig. OF, G); median claw
with conspicuous cuticular projections (fig. 9E-G); ovipositor apically with

1972 WYGODZINSKY: SILVERFISH 9

10.

FL:

12.

fossorial spines (fig. 9H, I) . . . . . Leucolepisma arenaria
Claws shorter than second tarsomere (fig. 8D); median claw glabrous;
ovipositor lacking fossorial spines. 7

. Several abdominal terga with 3+3 bristle- combs (fig. “8F); maxillary palps
8

with five segments
Abdominal terga at most with 242 bristle- combs; maxillary palps with five
(fig. 10B) or six (fig. 10A) apparent segments. . rele

. Abdominal sterna each with one median bristle-comb (similar to fig. 10D);

3+3 bristle-combs on terga II-V . . «S.C. Ctenolepisma targionit
Abdominal sterna without median bristle-combs; 3+3 bristle-combs on
abdominal terga II-V, II-VI or II-VII . . . . . Yd

. Only abdominal terga II-V with 3-++3 bristle-combs

Ce Ctenolepisma doersisquaris

Abdominal terga II-VI or II-VII with 3-43 bistle-combs . . . . . 10
Only abdominal terga II-VI with 3++3 bristle-combs; tenth abdominal tergum
truncate or faintly emarginate posteriorly (fig. 8E) ee so ve tt
Abdominal terga II-VII with 3+3 bristle-combs; tenth abdominal tergum
subtriangular (fig. 8C) . . . . . Ctenolepisma lineata pilifera
Maximum length about 10 mm.; antennae and caudal filaments not longer
than body; distinct violaceous hypodermal pigment on head, body and
appendages... . . Ctenolepisma ciliata
Length 13-15 mm.; ; antennae and caudal appendages longer than body; head,
body, and appendages practically without hypodermal pigment :

a ae Ctenolepisma longicaudata

Maxillary palp with five segments (fig. 10B); only one pair of stylets present;
scales of living specimen, when intact, uniformly gray dorsally (fig. 5). .

. Thermobia campbell:

Maxillary palp with six - apparent segments (fig. 10A); two or three pairs of
stylets present; scales of living specimen, when intact, forming complex
pattern (fig.2) . . . . . . . . «3...» Thermobia domestica

KEY TO THE LEPISMATIDAE OF THE CARIBBEAN AREA

. Last abdominal tergum triangular, sharply pointed, and with more than 1+1

bristle-combs (fig.6D) . . 2
Last tergum not sharply pointed; if triangular, then with ‘only 141 bristle-
combs (fig. 7D) . . . . 4

. Submedian fields of macrochaetae on head Capmile narrowly elongate, not

attaining anterior border of head capsule (fig. 6A); prosternum covered
below by fore coxae, and with tuft of macrochaetae centrally (fig. 6C);
ovipositor with fossorial spines a” on posterior gonapophyses (fig. 6G);
male with parameres. .. . . Aerotelsa collaris
Submedian fields of macrochaetae on head capsule broadly triangular, attain-
ing anterior border of head capsule (fig. 6B); prosternum covering fore
coxae from below, and without central tuft of macrochaetae (fig. 6E);
ovipositor with or without fossorial spines; male without parameres . 3

. Abdominal sterna with 2+2 multiseriate bristle-combs at hind border, one at

each side of insertion of stylets (fig. 6F), the latter present in more than two
pairs; ovipositor of female lacking fossorial spines . . Stylifera gigantea

10 AMERICAN MUSEUM NOVITATES NO. 2481

Abdominal sterna with 1++1 uniseriate bristle-combs at hind border, mesad of
insertion of stylets, the latter only in two pairs; ovipositor of female with

fossorial spines. . . Stylifera tmpudica
4. Large setae bare, often bifid or -trifid apically (fig. TE) . a Ip
Large setae feathered or barbed (fig. 7C) . . “hes

5. Abdominal sterna with median bristle-combs in addition: to sublateral ones
(similar to fig. 10D); tenth abdominal tergum rather elongate, distinctly
longer than wide (fig. 7F) . . . . . . Lepisma saccharina

Abdominal sterna lacking median bristle- combs: tenth abdominal tergum
rather short, subsemicircular, shorter than wide (similar to fig. 10F) . 6
6. Stylets present in only one pair . . . .. . . Heterolepisma horm
Stylets present in two pairs. ors Heterolepisma sp. (Bimini)
7. Thoracic nota at hind margin with only 1+1 single macrochaetae (fig. 7B)
. Peliolefisma calva
Thoracic ots at hind margin ‘with 141 bristle- combs composed each of at
least two large setae (fig. 8A) . . . 8

8. Last abdominal tergum subtriangular, strongly salient apically (fig. 8K).
Ctenolepisma hummelinckt
Last abdominal tergum subtrapezoidal or broadly rounded apically (fig.
9

8E,H). .
9. 343 bristle- combs present on abdominal terga II- VII (very rarely II-VIID);
ovipositor of female with fossorial spines apically . . Ctenolepisma dubitalis
3+3 bristle-combs present on fewer abdominal terga; ae of female
lacking fossorial spines... se a AD
10. 3+3 bristle-combs only present on | abdominal terga IV . Cell
3+3 bristle-combs present on abdominal terga II-VI. . . . . . 12

11. Stylets present in only two pairs; size under 10 mm. ne BS ase Oe:

ee ede ee ee ee’ Ctenolepisma diversisquamis

Stylets present in three pairs; size over 10 mm. . . Ctenolepisma targioniana

12, Size over 10 mm.; last abdominal tergum distinctly truncate or slightly
emarginate apically (fig. 8E); stylets present in two pairs

. . Ctenolepisma longicandata

Size less than 10 mm.; last abdominal tergum subsemicircular (fig. 8H); sty-

lets present in three pairs . . . . . . . . Ctenolepisma versluysi

Acrotelsa collaris (Fabricius)
Figure 6A, C, G

Lepisma collaris Fasricius, 1793, p. 64.
Lepisma mucronata Packarb, 1873, p. 49 (new synonymy).

MATERIAL EXAMINED: USA: Florida: Levy County: Seahorse Key,
Aug. 24, 1955 (C. H. Wharton; FSCA), one male. Monroe County:
Loggerhead Key, Dry Tortugas Islands, in house, Jan. 9, 1962 (H. A.
Denmark; FSCA), one male, one female. Bahamas: Nassau, summer 1945
(A. S. Vernay; AMNH), one male, one female.

I have examined the type of Lepisma mucronata Packard, kept in the
Museum of Comparative Zoology, Harvard Universtiy, Cambridge. The

1972 WYGODZINSKY: SILVERFISH 1]

type is a moderately well-preserved male, which agrees fully with Acrotelsa
collaris; hence the above synonymy, already suspected by Paclt (1966,
1967).

This is a tropicopolitan synanthropic species, often of economic im-
portance; it has not been reported from the United States before.

In addition to the characters given in the key, this species can also be
recognized by the macrochaetae arranged in distinct whirls on the legs
and cerci; in other lepismatids, such setae are irregularly distributed on
these appendages. The descriptions given by Escherich (1905) and Stach
(1935) should be consulted for further taxonomic data.

Allacrotelsa spinulata (Packard)
Figures 1, 7D, G

Lepisma spinulata PACKARD, 1873, p. 48.
Stachisma mexicana WyGODZINSKY, 1949, p. 224.

MaTERIAL ExaAmineD: USA: Florida: Martin County: Hobe Sound,
Pinus clausa debris, Feb. 1, 1959 (H. A. Denmark; FSCA), one female;
8 miles NNW of Stuart, Dec. 12, 1962 (W. Ivie; AMNH), one male, one
female. Saint Lucie County: 9.4 mi. N of Saint Lucie River, Pinus clausa
debris, Feb. 1, 1959 (H. A. Denmark; FSCA), one female. Highlands
County: Sebring, in leaf litter in dry, sandy scrub oak-live oak association,
Dec. 26, 1961 (H. V. Weems, Jr.; FSCA), several specimens; ibid., under
board in dry white sandy area, Feb. 1, 1960 (FSCA), one female; id:d.,
between layers of weathered cardboard in dry white sandy field, March 1,
1960 (H. V. Weems, Jr.; FSCA), four males, four females; ibid., in Pinus
clausa association, Dec. 23, 1961 (H. V. Weems, Jr.; FSCA), several
specimens. Orlando County: 3 miles NW of Maitland, Dec. 10, 1962
(W. Ivie; AMNH), one female. Putnam County: near Red Water Lake,
under bark of dead Quercus virginiana, March 22, 1962 (H. V. Weems, Jr.;
FSCA), two specimens. Alachua County: Gainesville, in sawdust mound,
Dec. 20, 1959 (H. V. Weems, Jr.; AMNH, FSCA), numerous males and
females; ibid., in malt traps, Aug. 7, 1962 (R. E. Woodruff and B. Benesh;
FSCA), one male, one female; ibzd., near Newman’s Lake, Nov. 2, 1959
(R. E. Woodruff; FSCA); one female. South Carolina: Clemson, under
pine bark, Jan. 9-10, 1966, June-July, 1966 (J. A. Payne; AMNH), 14
females, seven juveniles; Pendleton, in cardboard box (J. A. Payne;
AMNHB), two females; zbid., under bark of elm, Sept. 26, 1961 (J. A.
Payne; AMNH), two females. Tennessee: Oak Ridge, beneath pile of
boards in old building, Feb. 9, 1965 (J. A. Payne; AMNH), five females.
Arkansas: Lafayette County, Apr. 27, 1967 (I. Brown; AMNH), one fe-

12 AMERICAN MUSEUM NOVITATES NO. 2481

Fic. 6. A. Acrotelsa collaris, head capsule with fields of macrochaetae, schematic.
B. Stylifera gigantea, head capsule with fields of macrochaetae, schematic. C. Acro-
telsa collaris, anterior portion of body, seen from below. D-F. Stylifera gigantea.
D. Apex of abdomen, seen from above and slightly lateral. E. Anterior portion of
body, seen from below. F. Lateral portion of fifth abdominal sternum, with stylet
and two multiseriate bristle-combs. G. Acrotelsa collaris, apex of posterior
gonapophysis.

t

1972 WYGODZINSKY: SILVERFISH 13

Fic. 7. A. Murolepisma deserticola, general aspect. B, C. Peltolepisma calva.
B. Pronotum. C. Macrochaeta of posterior border of pronotum, with sensory seta
and sockets of additional smaller hairs. D. Allacrotelsa spinulata, apex of abdomen,
seen from above. E, F. Lepisma saccharina. E. Macrochaeta. F. Apex of abdomen,
seen from above. G. Allacrotelsa spinulata, labial palp. H. Lepisma saccharina,
labial palp.

male; Ashley County (I. Brown; AMNH), one female. Oklahoma:
Arrowhead State Park, July 17, 1966 (J. and W. Ivie; AMNH), one
female. Texas: Montopolis, near Austin, in sand, June 5, 1903 (AMNH),
one male, three females; Somerset, Nov. 15, 1942 (E. S. Ross; CAS), one
female, identified by Wall; Lockhurst State Park, April 13, 1963 (W. J.
Gertsch and W. Ivie; AMNH), four males, one female.

This is a free-living species, originally described from Texas. It is very
common in the western United States, from the state of Washington to
Arizona; it has also been found in Mexico. The above records widen its
known range considerably.

The best description of the species (as Acrotelsa spinulata) is given by
Silvestri (1948).

Ctenolepisma ciliata (Dufour)
Lepisma ciliata DuFour, 1831, p. 420.
MaTERIAL ExamineD: USA: California: Contra Costa County:

Lafayette, July 6, 1952 (R. A. Stirton; CIS), one male. Los Angeles
County: Pasadena, April 1, 1953 (UCLA), one male. Mexico: Chihuahua:

14 AMERICAN MUSEUM NOVITATES NO. 2481

Salaices, Cueva del Diablo, mouth of cave, July 23, 1947 (W. J. Gertsch;
AMNB), one female.

This species apparently is not common in North America; the above are
all the specimens I have seen, as opposed to hundreds of specimens of the
very successful Ctenolepisma lineata pilifera.

The best descriptions of this species are given by Escherich (1905) and
Stach (1935).

Ctenolepisma diversisquamis Silvestri
Figure 8B

Ctenolepisma diversisquamis SILVESTRI, 1908b, p. 153.
Ctenolepisma reducta Fotsom, 1923, p. 170 (new synonymy).

MATERIAL EXAMINED: USA: Florida: Monroe County: Loggerhead
Key, Dry Tortugas Islands, in leaf litter of Casuarina sp., Sept. 2-3, 1961
(H. V. Weems, Jr.; FSCA, AMNH), very numerous males and females;
Garden Key, Dry Tortugas Islands, in sand and debris back of beach,
May 9, 1961 (H. V. Weems, Jr.; FSCA), two females.

The above synonymy was tentatively suggested by Paclt (1967), and is
here formally established. The main difference between diversisquamis and
reducta was supposed to be the presence in the first species of scales of two
different kinds, those with numerous and those with few rays, and the
absence of this scale polymorphism in reducta. | have now examined
specimens of reducta from Florida, and have found the dorsal scales to be of
two types, just as illustrated in the original description of diversisquamis by
Silvestri (1908). Another difference seemed to reside in the shape of the
terminal segment of the labial palp; according to the original descriptions,
this segment is wider than long in diversisquamis but longer than wide in
reducta. In immature specimens from Florida now examined, the terminal
segment of the labial palp (fig. 8B) is about as wide as long, but in mature
specimens it is slightly wider than long, as in diversisquamis. Paclt (1967)
also synonymized Ctenolepisma brachyura Silvestri under diversisquamis.

The species has been reported from the Cape Verde Islands from where
it was described originally, from East Africa (as brachyura) and from
Puerto Rico, Aruba, the Caribbean coast of Colombia, and the Marquesas
(as reducta) ; it is now also known from the United States. This is obviously
a species dispersed by man, although it is not necessarily synanthropic.

Ctenolepisma dubitalis Wygodzinsky

Ctenolepisma dubitalis WyGopzinsky, 1959, p. 35,

1972 WYGODZINSKY: SILVERFISH 15

This species has been collected only on the Leeward Group of the
Lesser Antilles.

Ctenolepisma hummelincki Wygodzinsky
Figure 8G, I-N
Ctenolepisma hummelinckt WyGopzinsky, 1959, p. 31.

MATERIAL EXAMINED: Bahamas: Leaf Cay, Allen Cays, Exuma Island,
Jan. 6, 1953 (Hayden and Giovannoli; AMNH), one female.

This species was described from Barbuda and St. Kitts, in the Windward
Group; only males were known. The female now examined agrees with
the males in such characters as shape, pigmentation, structure of mouth-
parts, abdominal chaetotaxy, chaetotaxy of stylets, and shape of last
tergum. The size of the female examined is 8.5 mm.; the maximum size of
the males was 9 mm. The main difference between the males and the
female is the number of bristle-combs on the prosternum: 5-6+5-6
bristle-combs in the males, and only 2++2 in the female (fig. 81).

The eighth and ninth coxites are as illustrated (fig. 8J, L). Each coxite
VIII bears a bristle-comb composed of approximately 14 macrochaetae.
The inner projections of coxites [IX are unusually long and slender, viz.,
slightly more than three times as long as wide at their base. The ovipositor
is long and slender, surpassing the apex of the process of coxite [IX by
about twice the length of the latter. The gonapophyses consist of approxi-
mately 35 articles. The terminal article of the anterior gonapophyses
(fig. 8N) is pointed, that of the posterior gonapophyses (fig. 8M) strongly
flattened dorsoventrally and somewhat scoop-shaped. The chaetotaxy of
the apical segments of the gonapophyses is as illustrated (fig. 8M, N).

Ctenolepisma lineata pilifera (Lucas)
Figures 4, 8C

Lepisma pilifera Lucas, 1840, p. 560.
Lepisma quadriseriata PACKARD, 1873, p. 47.
Lepisma rubro-violacea ScHOTT, 1897, p. 190.
Lepisma reticulata ScHOTT, 1897, p. 192.

MATERIAL EXAMINED: USA: South Carolina: Clemson, on desk, Dec.
15, 1965 (AMNH), one female. Pennsylvania: Glenside, in house, April
14, 1969 (E. H. Slifer; AMNH), numerous males and females.

This introduced, often synanthropic, species is exceedingly common in
the western states of Oregon, Nevada, California, and Arizona and also in
Mexico; lineata pilifera is generally secondarily free-living in the southern

16 AMERICAN MUSEUM NOVITATES NO. 2481

portion of this area. In the East, the species is also common, according to
Sweetman and Kulash (1944) who listed it as Ctenolepisma quadriseriata.
These authors report the species ranging from south-central Canada in
the north to Georgia and Tennessee in the south, and at least to Missouri
westward.

This species is very variable, especially as to pigment distribution,
although individual populations tend to be rather uniform. This vari-
ability might explain the large synonymy of this species.

Acceptable descriptions of lineata pilifera may be found in Escherich
(1905), Stach (1935) and, as Ctenolepisma rubroviolacea, in Wall (1954).

Ctenolepisma longicaudata Escherich
Figure 8A, D-F

Ctenolepisma longicaudata EscHERIcH, 1905, p. 83.
Ctenolepisma urbana SLABAUGH, 1940, p. 95.

MATERIAL ExamInED: USA: Florida: Putnam County: near Red Water
Lake, July 18, 1959 (Pam Weems; FSCA), one female. Alachua County:
Gainesville, in buildings, various dates (H. V. Weems, Jr., H. A. Denmark,
and others; FSCA), numerous males and females. South Carolina:
Clemson, under old book, Sept. 22, 1965 (H. G. Young; AMNH), one
male. Louisiana: Baton Rouge, June 23, 1960, in bookcase (H. Daly;
AMNH), one male.

This species is common in the warmer parts of the world. In the United
States it is strictly synanthropic and has been reported, generally as
Ctenolepisma urbana, from North Carolina, Missouri, Louisiana, Illinois, and
southern California (Mallis, 1941; Slabaugh, 1940; and Sweetman and
Kulash, 1944). Ctenolepisma urbana has been correctly synonymized with
longicaudata by Paclt (1966).

The original description is sufficient to recognize the species.

Ctenolepisma targionit (Grassi and Rovelli)
Lepisma targionit GRASSI AND RovELtI, 1889, p. 7.

MatTERIAL EXAMINED: USA: South Carolina: Anderson, Nov. 12, 1965
(R. M. O’Neal; AMNH), two males.

This is the first record of the species from the Western Hemisphere;
the original range is circum-Mediterranean.

Acceptable descriptions of this species can be found in Escherich (1905)
and Stach (1935).

1972 WYGODZINSKY: SILVERFISH 17

Fic. 8. Species of Ctenolepisma <A. C. longicaudata, left half of pronotum. B. C.
diversisquamis, immature female from Florida, labial palp. C. C. lineata pilifera,
apex of abdomen, dorsal view. D-F. C. longicaudata. D. Tarsus of hind leg.
E. Apex of abdomen, dorsal view. F. Abdominal tergum V, right lobe folded
under. G. C. hummelincki, labial palp. H. C. versluysi, apex of abdomen, dorsal
aspect. I-N. C. hummelincki, female from Leaf Cay, Bahamas. I. Prosternum.
J. Coxite IX, with stylet and base of posterior gonapophysis. K. Tergum X.
L. Coxite VIII, with base of anterior gonapophysis. M. Apical article of posterior
gonapophysis; one sensory rod with high magnification. N. Apical articles of
anterior gonapophysis.

18 AMERICAN MUSEUM NOVITATES NO. 2481

Ctenolepisma targioniana Silvestri
Cienolepisma targioniana SILVESTRI, 1908a, p. 11. —

This species, originally described from East Africa, was later reported
from Brazil, the Caribbean coast of Venezuela, and the Leeward Group
of the Lesser Antilles.

The original description is the only one available.

Ctenolepisma versluyst Escherich
Figure 8H

Ctenolepisma versluyst ESCHERICH, 1905, p. 84.

This species is known from the Lesser Antilles, the Caribbean coast of
Colombia, and Acapulco, on the Pacific coast of Mexico.
Escherich’s description is the only one available.

Heterolepisma horn Stach
Heterolepisma horni Stacn, 1933, p. 341.

This species, originally described from the coast of Ecuador, was later
found in Jamaica and the Leeward Group of the Lesser Antilles.
The original description is sufficient to identify this species.

Heterolepisma sp.

MATERIAL EXAMINED: Bahamas: Bimini, East Bimini, Jan. 31, 1964
(K. Krishna; AMNH), one female; Bimini, South Bimini, Jan. 30, 1964
(K. Krishna; AMNH), two males, one female.

All specimens were collected on the beach under dead vegetation.

This species keys out to Heterolepisma pampanea Silvestri, found in the
colder areas of Argentina, but is not identical. Not enough material is
available to describe this species, member of a notoriously difficult genus.

Lepisma saccharina L.
Figures 3, 7E, F, H
Lepisma saccharina Linn£, 1758, p. 608.
MATERIAL EXAMINED: USA: Florida: Monroe County: Loggerhead
Key, Dry Tortugas Island, in leaf litter of Casuarina sp., with large numbers
of Ctenolepisma reducta, Sept. 3, 1961 (H. V. Weems, Jr.; FSCA), one male.

The above is the only record of free-living Lepisma saccharina in the
United States. The species is normally domestic and probably widely

1972 WYGODZINSKY: SILVERFISH 19

distributed over the country, although older identifications must be open
to doubt.

This is the only Lepisma recorded from North America, but other species
may well be found in the future; as mentioned above, Lepisma wasmanni is
commonly found free-living in coastal deserts of Peru, and I have seen a
further, as yet unidentified, species from Cuba.

Good illustrated redescriptions of Lepisma saccharina are found in Uchida
(1943) and Wygodzinsky (1941).

Leucolepisma arenaria Wall
Figure 9
Leucolepisma arenaria WALL, 1954, p. 74.

MATERIAL ExAMINED: USA: California: San Diego County: Borrego
Valley, on sand dunes at night, April 1955 (P. Wygodzinsky; AMNH),
numerous males and females; Borrego Valley, sand dune area, April 28,
1961 (J. Rozen, Jr. and R. Schrammel; AMNH), several males and
females. Riverside County: 3 mi. W of Indio, June 10 and 16, 1956 (M.
Wasbauer; CIS), several males and females; Hopkins Well, 18 mi. W of
Blythe, sand dunes, April 12, 1958 (J. Powell; CIS), several specimens;
Palm Desert, June 23, 1956 (M. Wasbauer; CIS), several specimens. San
Bernardino County: Cronise Station, April 28, 1956 (P. D. Hurd and J.
Powell; CIS), several males and females; Saratoga Springs, Death Valley,
May 30, 1953 (MacDonald; UCLA), one male, one female. Arizona:
Ocotillo, Maricopa County, April 26, 1961 (W. J. Gertsch; AMNH), one
female. Nevada: Nelson, Clark County, April 27, 1952 (H. Washburn;
UCLA), four males, one female.

Leucolepisma arenaria was described from San Diego and Riverside
counties in southern California; additional records from California and
the finding of the species in Arizona and Nevada widen its range con-
siderably.

A few illustrations of the coxites IX and genital appendages of the
female supplement those given by the author of the species (Wall, 1954).
Figure 9E shows the microtrichia of the median claw, a character men-
tioned by Paclt (1967) as characteristic for the species. Illustrations of the
mouthparts and hind tarsi of specimens from Borrego, southern California,
and from Nelson, Nevada show certain features distinguishing the Nevada
population from all other material seen. In the Nevada specimens, the
maxillary palp (fig. 9C) is comparatively more elongate, the distal segment
of the labial palp (fig. 9D) is less strongly widened, the tarsi (fig. 9F) are
relatively more slender, and the claws comparatively much shorter, than

20 AMERICAN MUSEUM NOVITATES NO. 2481

Fic. 9, Leucolepisma arenaria. A, B. Specimen from Borrego, California.
A. Maxillary palp. B. Labial palp. C, D. Specimen from Nelson, Nevada.
C. Maxillary palp. D. Labial palp. E. Median claw of hind leg, California
specimen. F. Hind tarsus, Nevada specimen. G—J. California specimen. G. Hind
tarsus. H. Apex of anterior gonapophysis. I. Apex of posterior gonapophysis.
J. Coxite X, with stylet and posterior gonapophysis.

in other specimens. These meristic differences are constant for all specimens
observed, but as the populations examined agree in all other structural
characters, they are considered to be conspecific.

Murolepisma deserticola Silvestri
Figure 7A
Mirolepisma deserticola SILVESTRI, 1938, p. 353.

This is the only myrmecophilous lepismatid found in North America.
It is known only from Arizona and California.
The original description is very detailed.

1972 WYGODZINSKY: SILVERFISH 21

Peliolepisma calva Ritter
Figure 7B, C
Peliolepisma calva RitTER, 1910, p. 380.

MatTERIAL EXAMINED: Guyana [former British Guiana]: Demerara, in
book in library, 1964 (C. P. Kennard; CIE, AMNH), three males, one
female. Cuba: La Habana (F. de Zayas; AMNH), one male, two females.

Paclt (1967) synonymized Peliolepisma Ritter with Ctenolepisma Esche-
rich, and listed the present species under Ctenolepisma. No reasons were
given for this synonymy.

Peliolepisma calva was described from Ceylon and had not been reported
from anywhere else. Its discovery in the Western Hemisphere adds to the
considerable list of introduced lepismatids already discovered. The
collector of the Cuban specimens considers the species to be the ‘‘common
household silverfish”’ in La Habana.

The original and only description is quite short; a redescription is
desirable.

Stylifera gigantea (Escherich)
Figure 6B, D-F
Acrotelsa gigantea EscHERICH, 1905, p. 109.

MATERIAL EXAMINED: Mexico: Perez Island, Yucatan, Sept. 2, 1952
(J. Pallister; AMNH), one female. Cuba: Oriente: Moa, May, 1954
(F. de Zayas; AMNH), one male; Matanzas: Itabo, Apr. 1962 (AMNH),
two males, two females. Bahamas: Long Cay, Feb. 10 and 25, 1953 (E. B.
Hayden; AMNH), one male, four females; New Portsmouth, Eleuthera
Island, March 29, 1953 (G. Rabb; AMNH), one female; South Bimini,
June, 1951 (M. A. Cazier, C. and P. Vaurie; AMNH), one female. USA:
Florida: Monroe County: Garden Key, Dry Tortugas Islands, collected by
beating a dead sea-purslane (Suriana maritima), Jan. 12, 1962 (H. A.
Denmark; FSCA); ibid., under fallen coconuts, May 8, 1965 (H. A.
Denmark; FSCA), numerous males and females; zbéd., under bark of
Casuarina sp., Sept. 2-3, 1961 (H. V. Weems, Jr.; FSCA), several males
and females; Big Pine Key, Nov. 20, 1958 (R. E. Woodruff; FSCA), one
male; Key Largo, under bark of dead tree, Aug. 29, 1961 (H. V. Weems,
Jr.; FSCA), one male; South Matecumbe Key, Nov. 30, 1961 (W. J.
Gertsch; AMNH), one female; Little Torch Key, under bark of dead tree,
May 7, 1961 (H. V. Weems, Jr.; FSCA), numerous males and females;
Middle Torch Key, May 7, 1961 (R. E. Woodruff; FSCA), one male,

22 AMERICAN MUSEUM NOVITATES NO. 2481

one female; Torch Key, June.8, 1960, under bark of small dead trees
(H. V. Weems, Jr.; FSCA), numerous males and females. Dade County:
Homestead, under bark of Casuarina equisetifolia, May 7, 1962 (J. Knowles;
FSCA), one female; Miami, Feb. 10, 1961 (R. E. Woodruff; FSCA), two
males, three juveniles.

This species was originally described from the Antilles, but was later
collected also in the coastal deserts of Peru, and in arid regions of Colombia
and Venezuela, on the shores of the Caribbean.

The original description is sufficient to identify this species, the largest
of all lepismatids of the area under consideration.

Stylifera impudica (Escherich)
Acrotelsa impudica EscHERICH, 1905, p. 112.

This species is known from the Caribbean coastal regions of Colombia
and Venezuela, and from various islands of the Leeward Group of the
Lesser Antilles.

The rather summary original description (Escherich, 1905) was
recently supplemented by a detailed redescription (Wygodzinsky, 1959).

The chaetotaxy of the posterior gonapophyses of a female from Oceania
as shown by Paclt (1966) is very different from that illustrated in Wygod-
zinsky’s (1959) redescription and obviously does not represent conditions
found in impudica as understood by Wygodzinsky (loc. cit.).

Thermobia campbelli (Barnhart), new combination
Figures 5, 10B—]
Ctenolepisma campbelli BARNHART, 1951, p. 184.

MarTERIAL ExaAMINED: USA: Pennsylvania: University Park, campus of
Pennsylvania State University, in buildings (J. F. Luke; AMNH)
numerous males and females.

The above specimens agree well with the data given in the description of
Ctenolepisma campbelli by Barnhart (1961), who found specimens in build-
ings in Ohio. Barnhart (loc. cit.) described his specimens as not having more
than 2+2 bristle-combs on any abdominal tergum, which places the
species in 7 hermobia and not in Ctenolepisma, but none of the authors who
have published on this species (Barnhart, 1961; Theron, 1963; and Paclt,
1967) placed campbelli correctly. An examination of the types of Cteno-
lepisma campbelli confirmed this identification.

Barnhart’s description and drawings are here supplemented by some
additional illustrations. Although Barnhart mentioned that the terminal

1972 WYGODZINSKY: SILVERFISH 23

segment of the labial palp bears five sensory papillae, his figure shows a
larger number, obviously in error; the five sensory papillae are shown
here (fig. 10E) in detail. The chaetotaxy of the apical articles of the
ovipositor, of some importance in the taxonomy of the Lepismatidae,
is also illustrated here (fig. 10I, J).

Fic. 10. Species of Thermobia. A. T. domestica, maxillary palp. B-J. T. campbellt.
B. Maxillary palp. C. Labial palp. D. Abdominal sternum V. E. Apex of labial
palp, high magnification. F. Tergum X, macrochaetae shown only on one side.
G. Bristle-comb of abdominal tergum, high magnification. H. Coxite [X of female,
with stylet and posterior gonapophyses. I. Apex of anterior gonapophysis.
J. Apex of posterior gonapophysis.

The description of the chaetotaxy of the abdomen as given by Barnhart
(1961) is somewhat confusing and not quite correct. Thermobia campbelli
has abdominal terga I-VIII and X with 1+1 lateral bristle-combs, and
terga II-VII with 1+-1 sublateral bristle-combs. Ventrally, sterna I and
II lack bristle-combs; there are 1+1 sublateral bristle-combs on sterna
IV-VIII, and median bristle-combs on sterna IJI-IV.

Thermobia campbelli adults especially when preserved in alcohol, can
easily be mistaken for juvenile Thermobia domestica. I am indebted to
Mr. J. F. Luke for having called my attention to the fact that living

24 AMERICAN MUSEUM NOVITATES NO. 2481

campbelli can easily be distinguished from domestica by their dorsal scales,
which are uniformly gray (fig. 5); in domestica, the dorsal scales form a
complex color pattern (fig. 2). Thermobia campbelli differs from domestica
and the three other named species of Thermobia, aegyptiaca (Lucas),
infelix Silvestri, and longimana Escherich by the presence of only one pair
of stylets; there are two or three pairs in the remaining species. 7 hermobia
campbell: differs, furthermore, from aegyptiaca by its smaller size, and the
comparatively short and stout inner processes of the coxite [X in the
female; longimana is distinguished by its extremely elongate appendages,
and infelix by the strongly widened apical segment of its labial palp.
Thermobia campbell: is only known from two domestic populations in the
eastern United States; it is probably Mediterranean or northern Ethiopian
in origin, to judge from the distribution of the other species of the genus.

Thermobia domestica (Packard)
Figures 2, 10A
Lepisma domestica PACKARD, 1873, p. 48.

The firebrat, a normally synanthropic insect and introduced in the
Western Hemisphere, is rare in South America but rather common in the
United States. In colder climates the species is generally restricted to
basements where permanent heat sources, such as boilers, exist, whereas
in warmer areas it is found in any part of a given building. In the west and
southwest of the United States secondarily free-living populations have
become established in semiarid areas.

LITERATURE CITED

BARNHART, C. S.
1951. A new silverfish of economic importance found in the United States
(Thysanura: Lepismatidae). Ohio Jour. Sci., vol. 51, pp. 184-186.
Durour, L.
1831. Description et figures de deux espéces nouvelles du genre Lepisma. Ann.
Sci. Nat. (Zool.), vol. 22, pp. 419-421.
Escuericu, K.
1905. Das System der Lepismatiden. Zoologica [Stuttgart], vol. 43, pp. 1-164.
Fasricius, J. C.
1793. Entomologia systematica. Copenhagen, vol. 2, 519 pp.

Fotsom, J. W.
1923. A new lepismid from Porto Rico. Proc. Ent. Soc. Washington, vol. 25,
pp. 170-171.

Grass!, B. AND G. ROVELLI
1889. Tavola analitica dei Tisanuri italiani da noi finora riscontrati. Bull.
Soc. Ent. Italiana, vol. 21, pp. 3-8.

1972 WYGODZINSKY: SILVERFISH 20

LinnE, C.

1758. Systema naturae. Editio decima, reformata. Stockholm, vol. 1, IV+

823-++1 pp.
Lucas, H.

1840. Histoire naturelle des crustacés, des arachnides et des myriapodes.

Paris, 600 pp.
MALLIs, A.

1941. Preliminary experiments on the silverfish Ctenolepisma urbani Slabaugh.

Jour. Econ. Ent., vol. 34, pp. 787-791.
PAcKARD, A. S.

1873. Synopsis of the Thysanura of Essex County, Mass., with descriptions of a

few extralimital forms. Fifth Ann. Rept. Peabody Acad. Sci., pp. 23-51.
Pactt, J.

1966. Neue Beitraege zur Kenntnis der Apterygoten-Sammlung des Zoo-
logischen Staatsinstitutes und Zoologischen Museums Hamburg. II.
Lepismatidae und Maindroniidae (Thysanura). Ent. Mitt. Zool.
Staatsinst. Zool. Mus. Hamburg, vol. 3, pp. 147-162.

1967. Thysanura. Fam. Lepidotrichidae, Maindroniidae, Lepismatidae. In
Wytsman, P., Genera Insectorum, fasc. 218, pp. 3-86.

Ritter, W.

1910. Neue Thysanuren und Collembolen aus Ceylon und Bombay, gesammelt

von Dr. Uzel. Ann. K. K. Naturhist. Hofmus., vol. 24, pp. 379-398.
Scu6rt, H.

1897. North American Apterygogenea. Proc. California Acad. Sci., ser. 2,

vol. 6, pp. 169-196.
SILVESTRI, F.

1908a. Thysanura. Jn Sjéstedts Kilimandjaro-Meru Expedition, Upsala, vol.
18, pp. 11-14.

1908b. Tisanuri raccolti da L. Fea alle isole del Capo Verde, alla Guinea
Portoghese e alle isole S. Thomé, Principe e Fernando Poo. Ann. Mus.
Civ. Stor. Nat. Genova, ser. 3, vol. 4, pp. 133-187.

1938. Due novi generi deserticoli di Lepismatidae (Insecta: Thysanura). Boll.
Lab. Ent. Agr. Portici, vol. 1, pp. 340-353.

1948. Descrizione di una specie nota e di una nuova di Lepismatidae (Insecta,
Thysanura). Jbid., vol. 8, pp. 100-106.

SLABAUGH, R. E. A.

1940. A new Thysanuran, and a key to the domestic species of Lepismatidae

(Thysanura) found in the United States. Ent. News, vol. 51, pp. 95-98.
STACH, J.

1933. Heterolepisma horni und die Zusammenstellung der Arten der Gattung
Heterolepisma (Thysanura). Ann. Mus. Zool. Polonici, vol. 9, pp.
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1935. Die Lepismatidenfauna Aegyptens. Jbid., vol. 11, pp. 27-111.

SWEETMAN, H. L. anp, W. M. KutasH

1944. The distribution of Ctenolepisma urbana Slabaugh and certain other

Lepismatidae. Jour. Econ. Ent., vol. 37, p. 444.
THERON, J. G.

1963. The domestic fish moths of South Africa (Thysanura: Lepismatidae).

South African Jour. Sci., vol. 6, pp. 125-130, 6 figs.

26 AMERICAN MUSEUM NOVITATES NO. 2481

Ucuiwa, H.
1943. Hompo ni okeru Seiyo-shimi (Lepisma saccharina L.) no saikento. Bol.
Zool. Tokyo, vol. 11, pp. 8-10.
WALL, W. J.
1954. A redescribed species and a new genus and species of the family Lepis-
matidae in California (Thysanura). Pan-Pacific Ent., vol. 30, pp. 72-76.
Wycopzinsky, P.
1941. Zur Kenntnis einiger europaeischen Dipluren and Thysanuren. Verh.
Naturf. Gesell. Basel, vol. 52, pp. 63-100.
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matidae, Thysanura). Acta Zool. Lilloana, vol. 6, pp. 215-227,
1959. Thysanura and Machilida of the Lesser Antilles and Northern South
America. Jn Wagenaar Hummelinck, P., Studies on the Fauna of
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