MILLIPEDS OF THE OKDEK COLOBOGNATHA, WITH
DESCRIPTIONS OF SIX NEW GENERA AND TYPE
SPECIES, FROM ARIZONA AND CALIFORNIA
By
O. F.Cook and H.
F.Loomis
OftheBureauofPlantIndustry, United StatesDepartment ofAgriculture
INTRODUCTION
A
special interestmay
be claimed formillipeds of the order Colo-bognatha
as examples of interrupted or residual distribution, in widely separated regionswhich
could not be reachedby any method
of transportationnow
at the disposal of these animals.The
expla- nation forsuch factsof distribution istobefound by
tracingback
toformer
ages of theworld when
vegetationand
surface conditionsmust
have been widelydifferentfrom
those ofthe present time.The Oolobognatha have
greater environmental limitations than themem-
bers of
most
of the other orders,though
the habitsand
living requirements areremarkably uniform through
thewhole
class of millipeds.From
the general uniformity of habits at the present time itmay
be inferred that a similar uniformity existed in the past,
and
that therequirements for existenceand
survivalhave
beenmuch
thesame during
thewhole
period of biological history of the group, or since theirworld-wide distributionwas
attained.In
comparison withmost
of the other millipeds, theColobognatha
aredelicate, fragile, slow-moving creatures, unable toburrow
in the soil or to withstand surface exposure.The
legsand
other appen- dagesareveryshortand
unspecialized,and
restricted food habits are indicatedby
minute,rudimentarymouth
parts.The
outstanding requirement for the existence of delicatehumus
animals is a continuous supply of moisture, notnecessarily a regular supply, but one that is never completelyinterrupted, or the creatures at that place are exterminated.Thus
the existence ofhumus
faunais
an
evidence of the moisture continuity ofany
locality,and may
afford a better assurance regarding local conditions in the
humus
layer than is obtainable in
any
other way.^1Cook, O. F. Notes on the distribution of millipeds in southern Texas, etc. Proc.
U. S. Nat. Mus., vol. 40, pp. 147-167, 1911.
No. 2714.—Proceedings U. S. National Museum, Vol. 72, Art. 18
58237—28 1 1
2 PROCEEDINGS
OFTHE
NATIOISTAIjMUSEUM
VOL.72 Species that lack the protection of a well-chitinized exoskeleton aremore
definitely confinedto areaswhere
thehumus
layer inwhich
they live remains moist at all timesand
so furnishes foodand
con- tinuous protection.Such
areas aremost commonly found
in the Tropicswhere
the rainfall is usuallymore abundant
than in tem- perate regions.The
greater differences betweensummer and
winter climate in the temperate regions doubtlessmay
also be considered as limiting factors of distribution,though
the soil conditions appear to bemuch more
important.Humus
faunasmay
be limitedby
the nature of the soil, as well asby
the need of continuous moisture.The
drying of the surface does not endanger the existence of ahumus
faunawhere
the condi- tions are such that its creatures can take refuge in a moist subsoil, but clays or colloidal soils do not afford such protectionand
are not adaptedtotheneedsof ahumus
fauna. If thesoil cracks in drying, thehumus
animalsmay
enter,and may
seek deeper levels as the season advances,butwhen
the raincomesand
waterfillsthecrevices, the soil dissolves into softmud and
creaturesburied in ithave littlechanceto escape.
Whether
forthis cause or others still to be recog- nized, the rule seems to be that the colloidal soilshave
little in theway
ofhumus
faunas,and
oftenno
animal life, evenunder
condi- tions that in other respectsmay
appear quite favorable.While humus
faunas exist inmany
regions of colloidal soils, it isfound
in such regions that the animals live in thehumus
blanket,and
are confined to the placeswhere
the blanket is thickenough
to holdits moisture through the dry season.
Under
such conditions the animalsmay
be said to live in thehumus
rather than in the soil,except as the character of the soil
may
be modifiedunder
ahumus
blanket.
As
with othercosmopolitanorders ofmillipeds,most
of the species of Colobognatha are tropical, but thefew
thathave
beenfound
intemperateregions have beenrecognized as distinct genera or even as distinct families, not represented in the Tropics.
From
the United States only five species of Colobognatha have beenknown and
in recent years thisnumber
has notbeen increased; three of the species were describedfrom
the Eastern Statesand two from
the Pacificslope.
One
of the eastern species has been reportedfrom
several localitiesand
belongs to the genus Polyzonium^ that occurs also in Europe.The
other eastern species, representingtwo
very distinct genera, Brachycy'beand Andrognathus,
apparently are rareand
local,
no two
species being reportedfrom
thesame
place.With
the addition of thenew
forms described here, the preponderance in Colobognathamust now go
to theWestern
States,where
eightgeneraAKT.18
NEW
GE15TEEAOF MILLIPEDS COOK AFD LOOMIS 6
are representedfrom
six separate districts.Only
one genus in California is shared with the Eastern States.^A
genus HypozoniuTn, closely related to Pol'j/Boniwm,was
de- scribed in1904from
Seattle,Wash, One
of thenew
generadescribedon
a laterpage was found by
the writers in the fall of 1919,on Mount
Tamalpais, nearSan
Francisco, Calif.Three
other generawere
distinguished in a remarkable collection of millipedsfrom
a small area inPlumas
County, Calif., obtained in the winter of 1922-23,by H.
S. Barber, of theBureau
ofEntomology. Although
this collection included only five species, four of
them
belonged to the Colobognatha.Three
of the specieshad
tobetreated as distinct genera, while thefourth isreferable tothe genus Brachycybe,known
also
from
the Eastern States.In
October, 1924, anothermember
of this orderwas
collectedinthe PinalMountains
of southern Arizona, the first specimen being securedby
L. K.Lytton and
othersfrom
thesame
localityby
thewriters. This animal belongsto adistinctly tropical family, Siphonophoridae, not previouslyknown
to exist in the United States.^The
speciesfrom Mount Tamalpais was found
in loose stony soilunder
a layer ofoak
leaveson
thewestern slope of themountain
not farfrom
theredwood
forestknown
asMuir Woods. Equable
condi- tions are maintained in the coast districtsby
the breezesand
fogsfrom
the ocean.The
survival of theredwood and
giant sequoias in the coast belt of Californiashow
that climatic conditionshave remained
nearly thesame
for long periods,though
the trees are reckoned as survivals of a former age of wider extension of forests,when
thevegetation generallywas more
luxuriantifnotmore
tropical in character.The
forest firesand
the periodical burnings of brush- landand
grasslands undoubtedlyhave
been the chief restricting fac- tor during the period ofhuman
occupation.Changes
of climatehave
been alleged, but changes of vegetationmay have
beensufficient to localizesuch types as
redwood
treesand
millipeds in thefew
places that have afforded continuous protection through the centuries.Whether
thehumus
faunain Californiawas
derivedfrom
tropicalAmerica
orfrom
other land masses ofthe ancientworld
is hardly to be conjectured at present, but it isworthy
of note that the nearest relatives of the millipedfrom Mount Tamalpais
are in the Mediter- raneanregion,not inother parts ofAmerica.A
southernorigin oralliancemay
be claimedforthe milliped froni Arizona,which
belongs to a family widely distributed in the Tropics'-An additional genus, GosodesmusChamberlin, related toBrachycybe, was described in 1922fromsouthernCalifornia. Itscharacters aregiven at theendof thispaper..
^Another memberof this familywas found still farther northin California, in Novem-
ber, 1926, while the paper was awaiting publication, so that two new genera ol Siphonophoridae are included.
4 PEOCEEDINGS OF THE NATIONAL MUSEUM
vol.72but not
known
previously in theUnited
StatesThe
locationmight
appearmost
unlikely for the survival of atropicaltype— an
elevated,exposed place
where
in cold periods theground
probably freezes to a depth of several inches.The
vegetation is sparse, with scattered small junipers, shrubby oaks,and
bushes.Probably
themost
essen- tial feature of the environment is the very coarse,open
soilformed
ofdecomposing
granite.The
creatureswere found under
stoneswith clean surfaces underneath.In no
casewere
millipedsfound under
stoneswhere
the surfaceswere
coated withdark
colloidal material,which
also discolors the graveland
doubtless represents a different condition of the soil.The
placewas
ina
divide afew
milesfrom Miami,
in the direction of Superior.Though
surroundedby
very broken country, the spotwas
nearly leveland
withno
through drainage or flooding of the surface. It seems thatthehumus
faunas of colloidal regions are limited to spotswhere
ahumus
blankethas accumulated above the soil, orwhere
the surface is not flooded even inheavy
rains.Thus
the restriction of thenew
tropicalmillipedto theverysmall area inwhich
itwas found
could be considered as a specialand
rather striking illustration of a general relation thathad
beeii observed in the study of thehumus
fauna.
From
the distribution of thehumus fauna
itmay
be possible tothrow
lighton
the question of the natural conditions in the south- western area before the period ofhuman
activity, or even further back.There
can be little doubt that the surface conditions have beengreatlychanged
duringthehuman
period,and
the archeologicai indications of extensiveand
long-standinghuman
occupation in this region are accumulating.Whatever
the causes of thechanged
conditions, widerand more
continuous distribution of thehumus fauna
in former times ishardlyto bedoubted.CHARACTERS OF THE COLOBOGNATHA
The
Colobognatha, as thename
indicates, are millipeds with a restricted development of themouth
partsand
other cephalic organs, except that the antennae are relatively large.The mouth
parts especially are reduced or
much
less developed than in the other orders,and have
been described as suctorial rather thanman-
ducatory, after the analogy of insects.
The
labral region of thehead
inmost
of the genera isnarrow
or isproduced
into a sharp beak.A
remarkably divergent genus described in this paper has thehead
broadened below instead of narrowed.The
structure of the segments is primitive rather than specialized, with aswide
a diversity ofform
as inany
other order of millipeds.The
segments ofmost
of the genera are definitely flattened, in con-AET.18
NEW GENEEA OF MILLIPEDS — COOK AND LOOMIS 5
trast with other long-bodied millipeds.
But
afew
of the genera are rather short-bodied,and
thebroad lateral expansions, or carinae of the segments, are similar to those ofsome members
of the order Merocheta.The
skeleton isweakly
chitinizedand
the segmental sclerites aredistinct or not so
much
fused as inmost
of the other orders.In
all of the Colobognatha the pedigerous laminae are free
and
the pleurae alsomay
be freefrom
the tergites or unitedby
a lateral suture.A median
suture appears in one of the subordersand
is doubtless to be considered as a primitive character.Median
sutures arefound
in several other orders,notably theMonocheta, Merocheta,and
Anocheta, suggesting a fusion oftwo
primitive sclerites toform
the dorsal scutes of millipeds.The Colobognatha
differfrom
all other millipeds inhaving
eight pairs ofnormallyformed
legsin front of thetwo
pairs ofspecialized copulatory legsof the males. Also the copulatorylegs, or gonopods, aremuch
less specialized than in other ordersand have
the jointsand
claws distinct, as in the ambulatory legs.The
posterior pair ofgonopods
is themore
specializedand
does nothave
thesame
position as themore
specializedgonopods
in the other orders,which
are locatedon
the seventhsegment
of the body.Where
only one pair of legs is modified for copulatory purposes, as in the order Merocheta, they are the anterior legsof the seventh segment.In
all of the orderswhere
both pairs of legson
the seventhsegment
are modified, the specialization of the anterior pair is alwaysmuch
greater than that of the posterior pair.
But
in the Colobognatha themore
specialized posteriorgonopods
have been developedfrom
the anterior pairoflegs, notofthe seventh segment, but of the eighth segment,where no
similar modification occurs inany
of the other orders.With
this positional differenceadded
to the structural differences, it is plain that thegonopods
of theColobognatha
are not reallyhomologous
with those of the other orders of millipeds, butgo back
as a separate line of development to a remote ancestral stagewhen
theappendages and
segments of thebody were
unspecialized.The
different position of thegonopods
affordsan
explanation of the fact that theColobognatha have
eight pairs of legs in front of thegonopods
while other orders have only seven pairs. Before the different attachment of thegonopods was
recognized itwas
cus-tomary
to describe the Colobognatha ashaving
an additional pair of legson some
of the anterior segmentswhich
in other orders are footless or are provided with only one pair.Some
ordershave no
legs
on segment
3and some
havetwo
pairs of legson segment
5,but in the Colobognatha each of the first 5 segments has a single pair of legs, while the subsequent segments
have
2 pairs.Thus
theQ PEOCEEDINGS
OFTHE NATIONAL MUSEUM
vol.72attachment of legs in the order Colobognatha is the
same
as the order Anocheta, a largegroup
of millipeds traditionally referred to the genus Spirobolus.As
a further consequence of the different position of thegonopods
in the Colobognatha, it follows that an
uneven number
of pairs of unmodified legs will befound on
the posterior part of the body, behind the gonopods, supposing that each of the segments hastwo
pairs, except thelast
two
segments,which
arefootless.The
posterior pair of segment8 renders thenumber
uneven.Pocock, writing in the Biologia Centrali Americana, reports an even
number
of post-copulatory legs ina species of Platydesmus,and on
that accountwas
unwilling toadmit
a different position for the gonopods.While Pocock
undoubtedlywas
a very careful observer, accurate countingof the largenumbers
of legs on specimens of Colo- bognathaisnoteasy.The
ventral platesarenotunitedtothe pleuraeand
the attachment of the legs to a particular segmentoften is diffi- cult to determine. Since it ismore
feasible to count the legs accu- ratelyon
photographs thanon
the specimens, photographs are sub- mitted as evidence of the actual occurrence ofuneven numbers
of post-copulatory legs.In
addition to accidental abnormalities of thelegs,thereisa further possibility, as recognizedby
Pocock, that the last of the leg-bearing segmentsmight have
only one pair. Thiswould
be anotherway
of explainingan
evennumber
of pairs behind the gonopods, but should not betaken for granted without proof.Two
suborders of Colobognatha are represented in America.In
the suborder Polyzonoidea thehead
isnarrowed
in frontand
insome
genera isproduced into a slender beak.The mouth
parts have been described as suctorial rather than manductatory,though
themanner
offeeding seems nottohave beenobserved.
The
segments are convex or flattened, sharply angled at the junction of the scuta with the pleurae, but not produced into lateral carinaeand
lacking amedian
suture. In thesuborder Platydesmoidea the
head
isnotnarrowed
or produced, the segments have lateral carinae,and
themedian
line of thesegments ismarked
withadistinctgrooveor suture.Three
families areknown
to represent this order north of theMexican
boundary.The
Siphonophoridae have thehead
suddenly constricted in front into a slender, sharp-pointed beak; thedorsum
ispilose
and
tuberculatebut lacksan
impressedmedian
sulcus.The
Polyzoniidaehave
thedorsum
without hairs, tubercles, oran
im- pressedmedian
sulcus,and
are lacking in flattened or projecting lateral carinae but are provided with eyes.In
theAndrognathidae
thedorsum
is either hairy or tuberculate or both,and
there is a well-markedmedian
sulcus, the lateral carinae are flattened or pro- jectingand
all of the species are without eyes.AET.18 ISTEW
GENEEA OF MILLIPEDS COOK AND LOOMIS 7 Family SIPHONOPHORIDAE
The
genusSiphonophora
has been the only representative of this family in America.There
arenumerous
tropical species, but until recentlyno member
of this family has been reported in or near the United States.In
1923 a representative of thefamilywas
describedfrom
Sonora, Mexico,and
in theautumn
of 1924 anotherwas
dis- coveredinthemountains borderingthenorthern edge of the southernArizona
desert. ThisArizona
species offers sufficiently distinctive characterstocallfor ageneric recognitiondistinctfrom
Siphonophora.A
second genus of Siphonophoridaewas found
inSan
Benito County, Calif., inNovember,
1926. This genus is remarkable be- cause themouth
parts are not produced into a beak, because of the extreme slenderness of the body,and
because of the very largenum-
bers of segments, in this respect probably exceeding
any
other milliped.SIPHONACME,
new genusType.
— Siphonacme
lyttoni, anew
speciesfrom
Arizona.Diagnosis.
—
Closely related toSiphonophora
but differing in hav- ing the posteriorgonopods
very slenderand
the distal joint of both anteriorand
posteriorgonopods
nearly straight, as long or longer than the preceding joints taken together. Also thehead
is broadly ovaland
the beakrelatively short, slender,and
abrupt.Description.
— Body
slender,from
twenty to thirty times as long as broad, rather strongly convex.Head
subglobose-pyriform, abruptly contracted to a short slender beak.Antennae
short, crassate, subclavate, a sense organ present near the margin, on the outer face of the fifth joint.First
segment
oblong with anteriormargin
transverse or but slightly emarginate.Segments
rather strongly convex; anteriorand
posterior subsegments tuberculate, the latter hirsute, with the hairs risingfrom
between the tubercles.Repugnatorial poresopening
from
a slight but abrupt prominence on the dorsal surface close to the lateralmargin;
the orifice of the pore surroundedby
a finerim
borderedby
a series of short, erect, closely placed hairs.On
afew
of the anterior segments the pore isnear the front
margin
of the posterior subsegment, buton
the other segments it is near the posteriormargin and
almost in the angle.Pleurae scarcelyexceeded
by
the dorsal plates at the lateral margin.Last
segment
with pleuraland
ventral sutures fused; the ventral sutureopen on
the penultimate segment.Preanal scale distinct.
8 PROCEEDINGS OF THE NATIONAL MUSEUM
vol.72Both
pairs ofgonopods
simple, longand
slender, the terminal joint of each pair nearly straight; anteriorgonopods
directed for- ward,with thelast joint as longas all the precedingjoints together;posterior
gonopods
longerand more
slenderand
reachingforward
between the anteriorgonopods
to their tips, the last joint longer than all the otherjoints together.The
type ofSiphonophora
is S. portoricensis Brandt.The
orig- inalspecimensare in the BerlinMuseum and show
a tapering conicalhead
equaled or exceeded in lengthby
a long slender beak.The
antennae are moderately crassate with the joints gradually thickerfrom
the base, joint 2 slightly longer thanjoint 3, joints 3 to 5 sub- equal in length, joint 6 about twice as long as joint 5,and
joint 7 very shortand
scarcely projectingbeyond
joint 6; the first segmentmore
than twice as long as the secondand
distinctly emarginate in the middle.The
repugnatorial pores are borneon
largerounded
prominenceswhich
give the segments the appearance of being dis- tinctly shouldered.A
larger female specimen with YO segmentshas the beak
much
longer than the head,and
thehead
is smallerand more
abruptlynarrowed
than in themale
specimen,which
has a shorter beak, scarcely longer than the head.SIPHONACME LYTTONI, newspecies
Description.
— Body
small, very longand
slender, twenty to thirty times as long as broad, scarcely tapering at the ends; the segments rather strongly convex. Motions in crawling very slow.Male
with 78 segments, females with 88and
121 segments.Length
of male, 16mm.; width
0.7mm.;
largest female 29mm.
longand
0.9mm.
broad.
Living colors.
—
^Head, antennae,and
firstsegment
white; segments 2 to 4 with a longitudinalmedian band
of dull pinkish abruptly broadenedand
divided atsegment
5,becoming
gradually paler to the middle of thebody and
finally disappearing so that the last seg-ments
also are white.The beak
is a palewaxy
yellow,and
the large lateral sense organs of the antennae,on
joint 5, appear as distinct orange-yellow spots.Head
subglobose-pyriform, abruptly contracted to a short, slenderbeak
piliferous to the tip, less than one-third as long as themass
of the head; surface ofhead
tuberculateand
pilose, the beaksmooth
with scattered largeand
small hairs.Gnathochilarium distinct, narrowly triangular, a
group
of large bristles atthe apex aslong asthe slender apical portion of thebeak;
median
groove present,basal anglesrounded, very close to the inser- tion of the first pair of legs.AET.18
NEW GENERA OF MILLIPEDS — COOK AND LOOMIS 9
Antennae
crassate, ratherstronglyclavate, joints 5and
6 distinctly thickerthanthe others; joint 6 about twice aslong as joint 5; joints 2and
4subequal,distinctlyexceededby
joint 5; joint7 distinctlypro- jecting; a large, orange-yellow transversely-oval sense organ near therim
of joint 5on
the lateral face.In
crawlingthe antennae are carried directly forward.Fig. 1.
—
SiPHONACMB lyttoni. a. Lateral tibw of head. 6. Antenna ofFEMALE, VENTRAL VIEW. C. GONOPODSj VENTRAL VIEW
First
segment
large, subelliptic, the anteriormargin
transverse or with a slight angular emargination; a transverse crescentic area in front of the middle of the segment slightly depressed or less convex than the surface behind it,forming
a thin collar over the base of the head.Segments
rather stronglyconvex; the surface of the posterior divi- sion tuberculateand
pilose, the hairs inserted between the tubercles, the posteriormargin
minutelyand
regularly serrate, supplementarymargin
not apparent; anterior subsegments beset with circular or ovalflattenedorbroadlyrounded
tubercleslikethose ofthe posterior subsegments, but usually without hairs or with only afew
near the posterior subsegment; pleural sutures at the lateralmargins
of the segments, the dorsal plates scarcely projectingbeyond
the pleurae.Last
two
segments longer thanany
of the preceding, without legs or ventral plates; last segment with the pleuraland
ventral sutures completely fused; penultimatesegment
with the sutures open.Last segment distinctly longer than the penultimate, gradually
narrowed
at the sidesand
rather broadlyrounded
at the apex; pre- anal scale distinct, themargin
transverse, nearly straight; anal valves prominent, evenly convex,somewhat
exceeding the posteriormargin
ofthesegment,the surfacepilosebut notdistinctlysculptured.58237—28 2,
10 PKOCEEDINGS
OPTHE NATIONAL MUSEUM
vol.72Gonopods
very long; the anterior pair extendingforward beyond
the basal joints of thetwo
precedingpairs of legs, the terminal seg-ment
quite slender, slightly tapering, nearly straight,and
as long or slightlylonger than the other joints taken together; posterior gono- pods longer than the anterior pairand more
slender, the last joint simple, very longand
slenderand
reaching the apex of the anterior gonopods.Type.—Q^i.
No. 975,U.S.N.M.
Three
specimens, amale and two
females,were found by
L. R.Lytton, in
company
with O. F. Cook, October 27, 1924,on
slightly moistsoillargelycomposed
of loosedecomposed
granite, under large stones inan open dry
place near thesummit
of a pass, \y^ miles north of themonument marking
theboundary
of Gilaand
Pinal Counties,on
the road between Superiorand Miami,
Ariz.The
con- ditions appeared quiteunfavorable for ahumus
faunaand
even withmuch
diggingno
other millipedsand no
thread centipeds (Geophili- daeand
related families)were
found.Only
afew
Scolopendridaeand
Lithobidaeand
afew
specimens of Ja'pyxwere
seen.On January
28, 1925, after the regionhad
been coveredby snow and
ice, alongand
diligent searchof thesame
localityby
the writerswas rewarded
with buttwo
living female specimensand two
dead specimens, the sex ofwhich was
not determined.Other humus
ani-mals were much more
plentiful than in Octoberand
four other speciesof millipedswere
collected.The
localitywas
visited againby Mr. and
Mrs.Loomis on March
1, 1925,when
fourmore
specimens were found.Two
of thesewere
males with 98and
108 segments, respectively.SIPHON
ACME
PSEUSTES (Chamberlin)Siphonophora pseustesChambeblin,Proc. Cal.Acad. Sci.,ser. 4, vol. 12, 1923, pp. 389-402.
From
thedrawings and
description ofSiphonophora
pseustesChamberlin
* itseems probable that this species should be placed in SiphonacTne, butSiphonacme
pseustes apparently is distinctfrom Siphonacme
lyttoni inhaving
the beakand
antennae longer, in themore
deeply emarginatefirst segment,and
thebody
with fewer seg- ments, especially inthe males. Possibly an examination of the gono-pods
ofSiphonophora
glohicepsPocock would show
itshould be in- cluded inSiphoTUicme.ILLACME, new
genusType.
—
Illacme plenipes^ anew
speciesfrom
California.Diagnosis.
—
SimilartoSiphonophora and Siphonacme
but readily distinguishedby
the triangular-cordate head,narrowed
gradually*0nchilopodsand diplopods from islands in the Gulf of California. Proc. Cal. Acad.
Sci., ser. 4, vol. 12, pp.389-407,1923.
ART.18
NEW GENERA OF MILLIPEDS COOK AND LOOMIS 11
to
an
acute angle below, butshowing no
tendency toform
a pro- jecting beak.Description,
— Body extremely slender, monilifoirm, about fifty
times as long as broad; composed
of a very large number
of seg-
ments, in the longest specimens approaching 200; dorsum
strongly
convex, nearly semicylindrical; the surface shining, moderately
hirsute.
Head
triangular-cordate inoutline, resemblingthat ofPolyzoniv/mand
withoutany
suggestion of a beak, the sides continuousand
sharply converging to a rather blunt angle, but notproduced
into a snout. Position of thehead
nearly vertical to the body, not reflexedunder
the body.Antennae
inserted at the sides of the head, abruptly capitate- clavate, subgeniculate; joints 1 to 4 very small, joints 5
and
6 abruptly thickerand
larger, held parallel to the sides of the head;joint 6 the longest, but scarcely as broad as joint 5; joint 7 broad
and
short.First segment slightly narrowed, about half again as long as the second; oblong, the lateral
margins
evenly rounded, anteriormargin
nearly parallel with the posterior; the surface nearly even, lacking the abrupt convexity of the posterior subsegmentson
the rest of the body; segments 2 to 4 shorter than the following, the trans- verse constriction withan
irregularrow
ofrounded
tubercleson
the anterior slope; surface of anterior subsegments rather coarsely re- ticulate^ surface of posterior subsegments finely hirsute, each of the hairs subtendedby
a short sublunate transverse groove, giving the general effect ofan
indistinct network; supplementarymargin
dis- tinct, with regular truncate divisions.Eepugnatorial pores difficult to detect, located
on
sliglit promi- nences near the posteriormargins
of the segments, not close to the lateral margins, pores of segment 5 in thesame
position as the others; lateral sutures open, borderedby
slight parallel ridges;surface of pleurae with a rather
open
reticulation, sparsely hirsute along the posterior margins.Last three segments
somewhat
longer than the preceding; the analsegment
nearly as long as the penultimate, the sides converg- ing to a broadlyrounded
apex, evenwith the valves, not projecting;
surface of the anal valves rather strongly
and
evenly convex, themargins
not prominent; preanal scale not distinct.Legs
rather short, scarcely exceeding the sides of the body, the basal joints prominentand
swollenon
the inner side, sometimes withan
extrudedmembrane
or exudate, nearly in contacton
themedian
line.This genus separates at once
from
Siphonacineand Siphonophora
by
the 9,bsence of a beak.The
antennae are differentfrom
those of12
PEOCEEDIISrGSOF THE NATIONAL MUSEUM
vol.72SiphonacTne in
having
joints 1 to 4much
narrowerand more
slen- der, while joint 5 is abruptly largerand more
nearly equal to joint 6. Also thebody
ismuch more
slenderand
the nimiber of segments ismuch
greater, approaching twice asmany, and
perhaps exceedingany
other milliped.A
diplopod with 192 segments, as countedon
one of these specimens, has a totalequipment
of 750 legs,and may
be the nearest approach to a literal "thousand legs."ILLACME PLENIPES, new species
Body
veryslenderand
flexible,filiform,stronglyand
evenly convex, moderately pubescenton
all of the exposed surfaces;number
of seg-ments
attaining192, with a length of 36mm. and
awidth
of0.7mm.
in the largest female specimen; other individuals 26 to 29
mm.
long, 0.5to 0.6mm.
wide, with 136to 152 segments.Head
rather narrowly triangular-cordate, the vertexmore
densely hirsute, the hairs rather short, the clypeus with longerand
sparser hairs, nearlynaked
above the rather blunt-pointed labium; position ofhead
nearly vertical, not strongly recurvedunder
the body.Antennae
inserted at the sides of the head, moderately hirsute, abruptly capitate-clavate, subgeniculate, the terminal joints carried at the sides of the head, the secondand
third joints at the lateralmargin
ofthefirstsegment; joints2 to4 graduallythicker butmuch
smaller
and
narrower than joints 5and
6; joint 2somewhat
longer than joints 3and
4,which
are subequaland
nearly asbroad as long;joint 5 also about as broad as long, but
much
thicker than joint 4;joint 6 slightly narrower than joint 5,
and
distinctly longer, about oneand
one-half times as long as broad, cylindric-oval, slightlynarrowed toward
the end; joint 7 projecting as a rather broad frustum about one-sixth of the length of joint 6; olfactory conesnot prominent.First segment with the lateral
margins
evenlyrounded and
the anteriormargin
nearly parallel with the posterior; the surfacemore
even thanon
other segments,which
are abruptly convex behind the transverse constriction.Penultimate segment withoutlegs, the large pleura meeting in the middle
and
apparently united, but the sutures indicatedby
a finemedian
groove; last segment converging to a broadlyrounded
apex, not projectingbeyond
themargins
of the valves, scarcely equal to themargin when
viewedfrom
the side.Type.—C^it
No. 976,U.S.N.M.
Numerous
specimenswere
collectedby
O. F.Cook
inSan
Benito County,Calif.,November
27, 1926, ashort distance after crossing the divide between Salinasand San Juan
Bautista.Only
one colonywas
found, in a small valley of a northern slopewooded
with oaks, under a rather large stone.The
living animalswere
nearly white,moved
very slowly,and
rolled themselves into regular, close spiral coilswhen
disturbed, thecoils with three or four turns.ART.18
NEW
GEISTEEAOF MILLIPEDS COOK AND LOOMIS 13 Family POLYZONIIDAE
ANALYTICAL KEY TO GENEEA OF POLYZONIIDAE
Segmentsstrongly depressed, the dorsum nearlyhorizontal; segments 20.
Platyzonium/
Segmentsstronglyconvex, 30to50 innumber.
Sterna with abroadmediangroove separating thebasal joints of thelegs; head broadly rounded below; antennae and first two pairs of legs distinctly
crassate Buzoniiiim, new genus.
Sterna narrow, the basal joints of the legs nearly in contact; head nar- rowed below; antennae and first legs slender.
Bodystronglynarrowed infrontfrom thefifth segment; first segmentlessthan half aswide as the body; antennae long, extending beyond the sides of the body, joints 5 and 6 distinctly larger than the others.
Bdellozonium,
new
genus.Body moderately narrowed in front, the first segment more than half the body width; antennae short, the joints subequal.
Last segment covered and exceeded by the large, broadly rounded penultimate
segment; segments 30 Hypozonium,
Last segmentexposedandprojecting beyondthe distinctlyemarginateposterior edgeof thepenultimate; segments variable innumber, adults commonly with
40 to 50 Polyzonium.
BUZONIUM, new
genusType.
— Buzomum
crmsipes^ anew
speciesfrom
California.Diagnosis.
— Head
subquadrate, the clypeuswidened
belowand
thelabrum
broadly rounded; antennae robust, clavate, uncolored;first
two
pairs oflegs ofboth sexes distinctly crassate.Description.
— Body small, moderately convex; body
cavity about
one and
three-quartertimes as broad as high.
Head
small, subquadrate,narrowed
between the antennae but the clypeus broadand
convexand
thelabrum
broadly rounded, themargin
denticulate.Eyes,
two on
each side, notprominent
nor heavily pigmented, reddish or brownish.Antennae
not colored, robust, distinctly clavate, joints 5and
6 longerand
thickerthan the others.First segment scarcely half as
wide
as the broadest diameter of the body, crossed in frontabove thelateral anglesby
adistinctridgefrom
the lower edge ofwhich
the thin anteriormargin
is produced.Subsequent segments
somewhat
polished,marked
with fine longi- tudinal striations;dorsum margined by
the extended pluraewhich
are visiblefrom
above; pore of the fifthsegment
lower thanon
the following segmentsand
locatedon
the exposed posterior corneron
the anteriorsubsegment.^Platyzonium is a European genus known only from Spain, originally described as a species of Cryptodesmus. See Cook, O. F., On Cryptodesmus Getschmannil Karsch.
Zoologiseher Anzeiger, No. 488. 1895.
14 PROCEEDINGS
OFTHE NATIONAL MUSEUM
VOL.72 Last segment subtriangular,exposed above.Legs
shortand
with large coxal joints, separatedby
the broad sternawhich
arenarrowerand more
elevatedbehind;firstand
second pairs oflegs conspicuously smallerand more
crassate than those fol- lowing, with theclaws also reduced.BUZONIUM CRASSIPES, new species
Number
of segmentsfrom
52 to 72; length of largest specimen 21mm.,
width2.8mm. Dorsum
stronglyconvex;body
cavity hardly twice aswide
as high;body
very abruptlyrounded
behind,more
gradually in front.Alcoholic specimens with the
dorsum
pinkish brown, the sides lighter, ventral surface pale.The
semitransparency of the exoskele- ton allows the internal organs to affect the color of the dorsum.d Fig. 2.
—
Buzonidm ckassipes. a. Head. Antennae and first segment, an- terior VIEW. b. First legs of male, posterior view. c. Segments4, 5, 6,
AND7,LATERAL VIEW. d.LASTTHREESEGMENTS, DORSALVIEW
Head
with the clypeusand labrum
greatlywidened and rounded
and
withnumerous
veryshort,stifflyerecthairs;labrum
considerably broader than the distance between the base of the antennae, themargin
coarselyserrateon
thesides,more
finely inthemiddle; vertexand
frontroughened by
rather deepand somewhat
irregular length- wise impressions; eyesvery widely separated, not hiddenby
thefirst segment,composed
oftwo
ocelli each, placed quite obliquely; anten-nae
subclavateand
aslong asthewidth
ofthefirst segment,firstand
second joints together equal in length to the sixth,which
is the longest; sense cones 4.ABT.18
XEW GENERA OF MILLIPEDS
—
COOK AND LOOMIS 15
First segment subreniform, slightly emarginate in frontand
behind; thelateralanteriormargin
thick,rounded,and
carriedup
as a highridge across the dorsal portionbehind athin,expanded margin
producedfrom
thelower edgeof theridge.Subsequent segments slightlyshining; posterior divisionimpressed with
many
fine, short longitudinal striationsmuch
stronger at themiddle
of thedorsum
near the anterior edge of the subsegment;plurae projecting slightly,
forming
anarrow
raised lateralmargin
ofthe segments; pores of thefifth segment locatedlow down on
the enlarged caudal corners of the anterior subsegment, poreson
subse- quent segments higherand on
the posterior subsegment.Last segment exposed above, subtriangular; segments immediately preceding it sharply
and
deeply emarginate.Legs
not reaching the sides of the body; sterna broad, separating the legs, narrowerand much more
elevated behind; coxae of all legs at leasthalfagainasbroadasthenextjoint,withtheexception of the firsttwo
pairs of legsthe coxaehave a large oval pit at apexon
the innerside; firstand
second pair oflegs reduced in sizeand
stronglycrassate,
more
noticeable inthemale,and
with the claws shorterand
less acutethan
on
the other legs.Type.—Cdii. No. 977,
U.S.N.M.
Six
mature
specimens collectedbeneath bark ofoak
logs near Sen- eca,Plumas
County, Calif,December,
1922,and
January, 1923,by H. S
Barber,and two
otherspecimensfrom same
locality collected inMarch,
1924,by
F. J. Silor.BDELLOZONIUM, new
genusType.
—
BdellozoniwTh cerviculatum, anew
speciesfrom
California.Diagnosis.
—
RelatedtoPolyzoniuTnand
Hypozoniurm^ but withthe clypeal portion of thehead
longer, the labral extremity broaderand more
rounded, the antennaemuch
longerand
the jointsmore
equal, the anterior segments relatively smaller.Description.
— Body
small,moderatelyconvex,with thebody
cavity about three times as broad as high.Head
very small, subconic, the clypeus gradually narrowed, the apex obtusely angled in front; anterior surface pigmented.Eyes
of four ocelli on each side inan
oblique row, prominent, heavily pigmented.Antennae
deeply colored, scarcely clavate, long, projectingbeyond
the sides of the body, joint 6 the longest; joints 3and
5 subequal, longerthan joints2and
4; joints 5and
6 longerand
slightly thicker than joints 2to 4.First segment very small, scarcely longer than segment 2
and
about half aswide
; anteriormargin
with a fine raised rim.16 PEOCEEDINGS OF THE NATIONAL MUSEUM
VOL.72Segments
flat underneath, moderately convex above, with a dis- tinct transverse groove separating the subsegments; anterior sub- segmentsmooth and
even,more
exposed than inPolyzonium;
pos- terior subsegmentsomewhat
inflated behind the groove, the surface minutely wrinkled, punctate or striate, especially in the groove, Re- pugnatorial poreson
slightprominencesin front of the middleof the posterior subsegmentand
rather remotefrom
themargins
on an- terior segments, closer to themargin on
posterior segments; pore of segment 5much
closer to the lateralmargin
thanon
the other seg-ments, located
on
the anterior subsegment, withthe transverse suture of the segment sinuate behind the pore,forming
a small lateral elevationaround
the pore.Penultimate segment slightly longer than the preceding, with a
wide
posterior sinus exposing the last segment.Last segment small, with a broadly
rounded apex
concealing the anal valves.Basal jointsof the legs nearly in contact atmiddle, while inPoly- BomuTTh they are distinctly separated, especially
on
the anterior segments.BDELLOZONIUM CERVICULATUM, new species
Number
of segments 39 to 46.The
largest specimen 16mm.
longby
3.3mm.
broad.Body
rather stronglyconvex above, the posteriorend more
abruptlyrounded
than the anterior;body
cavity about twiceaswide
ashigh.Fig. 3.
—
Bdhllozonium ceeticulatum. a. Lateral margins and kepugna- TORIAL PORES OP SEGMENTS 4, 5, AND 6. &. LAST THREE SEGMENTS OF BODY,DORSALVIEW
Color of the
dorsum salmon
pink in living specimens,more
brownish in alcohol, theside of the segmentsand
the ventral surface pale;head and
antennaelight,mottledwith purplish.Antennae
about oneand
a half times as long as the first segmentis
wide and
reaching wellbeyond
the sides of the body; joints 3and
6 distinctly longer than the others. Ocelli 3 to 4 on each side in aART. 18
NEW GENERA OP
MILLIPEDSCOOK AND LOOMIS 17
very oblique row, 1 or 2 in each series usually being coveredby
thefirst segment.
A low
tubercle bearing a long hair above the lower ocellus of each series.Below
the base of the antennae is a broad transverse depression in front ofwhich
the clypeus issomewhat
inflated; labral areagraduallynarrowed,the apex
somewhat
rounded.First segment very small, only one-third as
wide
as the full-sized segments,almost squarely truncate in frontand
with anarrow
raisedrim
; posteriormargin
broadlyand
evenly rounded.Body
shining; anterior subsegments almost smooth, afew
faint impressed longitudinal lines near the caudalmargin;
posterior sub- segmentsmore
conspicuously impressed with similar lines, rather deepon
the anterior third, faintertoward
the middle, thehind mar-
gin smooth; exposed portion of the anterior subsegment usually about one-third the length of the posterior subsegment, thehind margin
of the middle segments of thebody
slightlyand
broadly emarginate at middle; pores of the fifthsegment on
the anterior subsegment slightly above the posterior cornerand
near the trans- verse sulcus; other poreson
the posterior subsegments, closer to the transverse sulcus thanto the lateralmargin
; dorsal encroachment of the pleurae very slight butforming
anarrow
raised lateralmargin
to each segment.
Legs
almostreachingthesidesof thebody,slightly separatedfrom
each otherby
the sternawhich
are elevated behindand on
the sidesand form
a depressionopen
in front; basal joints of the,legs with rounded-oval pits.Females
with the first pair of legs very smalland
slender,much more
reduced than those of the males,which
also are smaller than the following pair.Type.—Cat
No. 978,U.S.N.M.
Several specimens
from
Belden,Plumas
County, Calif., collectedby H.
S. Barber,January
2, 1923. "It isnumerous, in several sizesfrom
about 4mm.
up,under bark and
in splintered crevices ofoak
lognear themouth
ofChipps
Creek."Genus
HYPOZONIUM
CookHYPOZONIUM
ANURUM
CookEypozonium anurum Cook, Myriapoda of Northwestern America, Harriman Expedition, p. 63.
This species
was
describedfrom
Seattle,Wash.
It has been reported alsofrom
Bremerton,Wash.^
8Chamberlin, R.V., Can. Ent., p. 262, August,1911.
18
PEOCBEDIlSrGSOP THE NATIONAL MUSEUM
vol.72 GenusPOLYZONIUM
BrandtPOLYZONIUM BIVIRGATUM (Wood)
Octoglena bivirgata Wood, Proc. Plaila. Acad. Nat. Sci., p. 186, 1864.
Petaserpes rosalhusCope, Trans.Amer. Ent. Soc,vol. 3, p. 65, 1870.
HexaglenacrpptocephalaMcNeill, Proc. U. S.Nat.Mus., vol. 10, p. 328, 1887.
Polyzonium rosaWum Bollman, and others.
This specieshas long beencalledPolyzoniv/mrosalbum (Cope) but that it resembled Octoglena tivirgata
Wood
has beenknown
formany
yearsand
the likelihood of itsbeing a
synonym
of Mvirgatawas
also consideredon
the basis ofWood's
draw- ings being poorlymade and
failing toshow
the true relationand
shape of thehead and
anterior segments. Sinceno
material has ever beenfound
to provePIG. 4.-P0LYZ0NIUM BIVIRGATUM. ^^^ distiuctncss of the
two
spccics, Itlatekal mabgins and eepug- seems best to assume that rosaTbwm is a
IT^lto
r^^^""^ segments
synomyn
of hivirgatum,which
is theolder name.
The
specieshasbeen reportedfrom
Georgia, Tennessee,New
York, Indiana,and Michigan and may
be expectedfrom many
other of the Eastern States.A
peculiarity ofPolyzordwm
is that the repugna- torial secretion has a camphor-like odor, but whether this is true of othermembers
of the family isnotknown.
^
Family ANDROGNATHIDAE
The
separation of this familyfrom
the tropical Platydesmidae isbased
on
the structure of the sterna.The Andrognathidae
have the basal jointsof the legsalmost in contact, but separatedby
a peculiar fungiformstructure, whilethePlatydesmidaehave
thelegsseparatedby
broadsternaand
arewithouttheintercoxalprojections.analytical key to genekaopandrognathidae
Repugnatorial pores elevated on a short stalk or pedicel; fifth segment with
lateralcarinae deeply emarginate, bilobed; anal scaleabsent Andrognathns..
Pores not elevated on pedicels; fifth segment not different from adjacent seg- ments; anal scalepresent.
Body very slender, attaining a length more than twenty times the width;
number ofsegments verylarge, sometimesmore than 100; dorsum not tuber-
culate, evenly convex Mitocybe, new genus.
Body broader, less than fifteen times as long as wide; segments less than 70;
dorsum tuberculate; lateral carinae longitudinal or depressed.
Bodyabout twelve times as long as broad; loosely jointed, with anterior sub- segments partly exposed; first segment not tuberculate, without lateral
^Cook, O. P. Camphor secreted byan animal. Science, new ser., vol. 12, pp. 516-521, 1900.
AET.18
NEW
GEITERAOF MILLIPEDS COOK AND LOOMIS 19
earinae, about twice as broad aslong, scarcelywider than the head,following segments with lateral earinae rather narrow, nearly horizontal, not tuber-
culate Ischnocyhe, new genus.
Body abouteighttimes as long asbroad,very compact, theanteriorsubsegments concealed; first segment with distinct tubercles and lateral earinae,
much
wider than the head; following segments with lateral earinaebroad, sloping slightly, with rows of dorsal tubercles extending to near the lateral
margin Brachycyhe.
Genus
ANDROGNATHUS
CopeANDKOGNATHUS
CORTICARIUS CopeAndrognathus corticariusCope, Proc.Amer. Philos. Soc,p. 182, 1869.
Known from
Virginiaand
Tennessee.MITOCYBE, new
genusType.
—
Mitocyhe cmriportcce, anew
speciesfrom
California.Diagnosis.
— From
theform and
sculpture of the body,thelamina-tion of the segments
and
the situation of the repugnational pores, this genus appears to be closely related toEuropean
DoUstenus,from which
it differs at least in having a distinct preanal scale.The
presence of a preanal scale, the location of the poreson
the surface of the segments,and
thenormal
formation of segment 5 distinguishMiiocyhe from
its closestAmerican
relative,Andro-
gnathivs.
Description.
— Body
slender,from
fifteen to twenty-five times as long as broad; segments very numerous, in females exceeding 100.Head
cordate, notproduced
into a pointed snout nor coveredby
the first segment, turnedunder
the body; antennae shortand
stout, widely separated at base; eyes lacking.First segment subreniform, emarginate in front, the angles ab- ruptly
rounded and
the anteriormargin
raised;median
sulcus absent;
surface not tuberculate but covered with fine, erect hairs.
Subsequent segments hirsute, not tuberculate;
dorsum
evenly con- vex, the lateral earinae thickened instead of being flattened or depressed; the repugnatorial poreson
the caudal angles; sulcus presenton
all segmentsfrom
the second to the penultimate.Last
segment
hoodlike, rather longand
with the apex truncate;no
impressedmedian
sulcus.Anal
valves visiblebeyond
the last segment.Preanal scalepresent, comparatively large
and
pilose.Sterna
produced
into lobeswhich
separate the coxae.Legs
shortand
slender, not reaching the sides of thebody
; coxae twiceas thick as the otherjoints; in addition to the coxal joint there appear to be six other joints.Males
smallerand
with fewer segments than the females.The
genericname
alludes to the long, attenuatebody
contrasting withBrachycyhe and
associated genera.20 PEOCEEDINGS OF THE NATIONAL MUSEUM
vol.72MITOCYBE AUKIPORTAE, newspecies
Segments
variable innumber, two
maleshave
52and
64 segmentsand
four females have 80, 84, 108,and
110 segments; the specimens rangingin lengthfrom
14mm.
to44mm. and
inwidth from
0.8mm.
to 1.6
mm. Dorsum
evenly convex without flattened or depressed lateral carinae.Color
creamy
white in living animals, suggesting a resemblance tosome
speciesofSiphonophora.Head
turnedback under thebody
butnot coveredby
the first seg- ment, cordate; the labral region bluntly rounded; vertex slightly inflatedon
the sides above the base of the antennae; surface pilose, with longer hairson
the sidesofthe front; antennae shortand
thick,clavate, pilose;bases remote;jointsincreasing inthicknessto joint6;
joint 2 longest; joint 3 equal in length to joint 7; joint 5 broader
thanlong. i
First segment subreniform, slightly emarginate in front; anterior
margin
raised; angles abruptly rounded;no
impressedmedian
line;surface evenly covered with fine, erect hairs.
Following segments with the anterior subdivisions glabrous; with a very fine raised
median
linefrom which
four or five annual foldsbend
sharplyback and
then proceedaround
the segment, the folds intersectedby many
fine impressed lines. Posterior subsegments evenly convex;thelateralcarinaethickenedinstead ofbeingflattened or depressed;median
sulcus present, broaderand
deeper in front than behind; surface with fine, erect short hairs risingfrom minute
punctationswhich
give thedorsum
a granular appearance,thepubes- cencemore abundant on
the lateral carinae. Lateralmargins
of carinae parallel; front angles abruptly rounded; anteriormargin
of the subsegment with a broad, shallow emargination between the an- gles;hindanglesrounded on
several of the anterior segments, slightly producedbackward
;more
stronglyand
sharply producedon
the lastfew
segments; hindmargin
of all segmentsrounded
excepton
the lastfew
segmentswhich
are broadly emarginate behind.Eepugna-
torial pores
on
the caudal angle of the carinae, surroundedby
anarrow
margin.Last segment rather long, hood-like, the hind