New cavernicolous millipeds of the family Cambalidae (Cambalidea: Spirostreptida) from Texas (U.S.A.) and Mexico
.Millipeds SpirostrepLida) ancl Mexico
Family Texas 0
0 1) Department of Zoology and Physiology , Balon Houge, Louisiana , U. S. A
Millipeds of the genus Cambala arc abundant in Texas caves. In three widely separated cave areas, there are two allopatric species, of which one is represented by two subspecies. Cilmbala speobia (Chamberlin), a troglobite, surely will also be found in Coahuila. Of the two subspecies of C.reddelli n. sp., the nominate subspecies is a troglophile that also OCCllI'S in epigean sites in l\'ew Mexico, and inornatlls is probably a troglobite. The last is of special interest because of the very weak development of the dOl'sal crests and pore crests. Mexicambala !'llssf:lli, n. gen. and sp., a troglobite from San Luis Potosi, is more highly modified by cave life than any other cambalid. These are the only cambalids that are known from the caves of western North America and }dexico. In the states east of Texas, Cambala minor (Bollman) and occasionally other congeners are collected in the entrance and twilight zone of ca ves. Cambala loomisi (HofTman) occurs in a cave in south-central Alabama. ~lost of the specimens that were studied in the preparation of this paper were collected by the biological division of the Texas Speleological SllI'vey, of which i\Ir. ,James Heddell is editor. The remaining specimens were collected by Dr. Hichard O. Albert, Mr. James K. Baker, Dr. T. C. Barr, Jr., and DI'. C. C. Holl'. I am grateful to all of these collectors. The deposition is mentioned in the section on each species.
This genus was summarized by Loomis in 1938. A needed revision
is contemplated by HolTman (
) to bring together the species that
have been described since then.
Carnbala diITers from Jll exicarnbala chiefly in the longer, slightly
salient ventral margins of the collum, the fewer setae on the anal
valves, and the bilobed coxite of the anterior gonopods. The species
of Cambala diITer chiefly in body size, number of ocelli, amount of
body pigment, prominence of dorsal crests and pore crests, segment
on which crests begin, size and distribution of lobes on legs of matme
males, presence or absence of tarsal claw of legpair 1 of males, and
details of both pairs of gonopods. Dorsal crests begin on segments 2,
3, or 4. Pores begin on segment 5. A troglobite, C. speobia, lacks ocelli;
all other species have several ocelli in 1 or 2 series. Two setae are
on the mesial margin of each anal val ve.
Article 2 of legpair 1 of males is about twice the width of article 3.
One or 2 articles of 2 or more pairs of legs of mature males are
enlarged. The articles that are modified are It and 5, 01' only article 4,
of the following legpairs: 6 and 7, 4 Lhrough 7, and Lhe first 24 or so
Sternum of anLerior gonopods is somewhat rectangular and longer
than it is wide. DisLal margin of coxite is bilobed; anteriOl' lobe bears
macro setae ; flagellum is long. TelopodiLe is seLose on disLal sUl'face;
distal margin is divided inLo two shallow lobes.
Anterior .process of coxa of posterior gonopods is either simple 01'
bilobed; iL bears cUl'ved macroseLae. Ectal process of coxa is smaller
than anteriOl' process and eiLher simple or divided. A vertical series
of minute spllrs is on mesial smface of coxa. TelopodiLe bears shorL,
thick SPUl'S on apex. I am 11llcerLain wheLher the telopodite always
consists of 2 articles; possibly Lhe nllmber is noL constanL.
DistribuLion. The UniLed StaLes from western WashingLon and
noLhern Idaho Lo the AtlanLic Coast. MosL abundanL in Lhe
S p e c i e s. The numbel' is lIncerLain. Of Lhe to thaL were listed by
Chamberlin and 1IofIman (
), some are probably synonyms or
subspecies of C. minor. Epigean, Lroglophilic, and Lrogiobitic species
OF CA.\IBALA IN TEXAS
'1. Without ocelli. Maximum body width '1.9 mm. Central and
southwest Texas. . . . . . . . . . . . . . . . . . . . . . . . .. C.speobia (Chamberlin)
With ocelli. Maximum body width '1.2 mm. Distribution otherwise.
(C.reddelli, n. sp.) 2
2. With distinct dorsal crests and pore crests. Ocelli usually in 2 series.
""VestTexas C.reddelli reddelli, n. subsp.
Lacking distinct dorsal crests and pore crests. Ocelli in 1 series.
Northwest Texas C.r. inornatus, n.subsp.
The presence of a tarsal claw on legpair 1 of the male distinguishes
the speeies of Cambala that are treated in this paper from the more
eastern eongeners, including those in cast Texas.
Diagnosis. A polytypic species near C.washingtonensis Causey in
the form of the gonopods; diners in the smaller body size, fewer
segments, the smaller lobes on the pregonopodallegs, and the absence
of lobes on the postgonopodal legs. Characterized especially by a pair
of setae on the apex of the caudal tergite.
Length 16 to 24 mm. Width 0.9 to '1.2 mm. 41 to 56 segments.
From 9 to 2 ocelli are arranged in either 1 or 2 irregular rows.
Dis t ri but ion. Ca ves and epigean sites in west and northwest Texas
and I\'ew Mexico. It possibly occnrs also in Colorado.
Subspecies. Two subspecies are descr.ibed here. They diner in the
prominence of the crests and in the number of ocelli. An epigean
collection from northern I\'ew Mexico is intermediate between them.
This species is named for Mr . .James L. Heddell in recognition of his
outstanding collections of troglobitic millipeds from Texas and
Diagnosis. A small-bodied troglophile that is like C. r. inornatlls
in the form of the gonopods and lobation of Lhe legs; differs in the
greater development of the dorsal crests and pore cresLs and the
al'l'angemenL of the ocelli in 2 irregular series.
Male holotype. Length 16 mm. WidLh 0.9 mm. 40 segmenLs, of
which the last 2 are legless. Flesh-colored and wiLh red lateral glands
when freshly preserved; became red-brown in alcohol. lIead is as in
congeners. Antennae reach back to margin of segment 3; raLio of
length of articles 'I thr01lgh 7 and width of arLicles 5 and 6 (in
parentheses) is: 19:26:30:21:23(22):25(20):4. Ocelli, which are partly
covered by collum, are in rows of abouL 3, 5.
Body is not markedly narrowed behind head or swollen at segment 7.
Collum (fig. 1) is a little longer than segments 2 and 3, the
anteroventral margin is lifted slightly to allow the antenna to be carried
between it and mandible, but caudo-ventral angle is not lifted. Dorsal
surface of first 3 and last 2 segments is entirely smooth. Six weak
crests are on segment!1 and the third and fourth segments preceding the
last. Dorsal crests of typical body segments are a little less prominent
than in C. washingtonensis Causey and much less prominent than in
C.speobia (Chamberlin); pore crests have the usual pear shape, but are
flattened and no more prominent than the !I dorsal crests; surface
between crests is smooth; below the pore crests there are about
16 horizontal striae, of which the upper are the most prominent.
Horizontal striae on the first 3 segments are indistinct. Horizontal
striae are distinct on lateral and ventral surfaces and along segmental
sulcus of dorsal surface of prozonites. Caudal tergite is a little shorter
than anal valves; caudal margin is rounded; there is a pair of setae
at the apex and another pair at the intersections with the anal scale.
Two pairs of setae are on mesial margins of anal valves, and 1 pair
is on anal scale. Caudal margin of anal scale is weakly convex.
Legpair' 1 consists of 6 articles and a tCl'minal claw; as in the
congeners, article 2 is about twice as wide as article 3. Articles !I and 5 of
legpnirs 6 (fig. 2) and 7 are moderately enlarged for the genus. No
postgonopodal legs are enlarged.
FigUl'e 3 is an anterior view of the anterior gonopods, and figUl'e 4
is an ectal view of the right posterior gonopod of a paratype. There
is very little difference between the gonopods of C. reddelli and
C. washingtonensis. The laUe!' dillers in the larger' body size and in the
lobation of article 5 (fig. 6) of the first 24 or so legpairs immediately
behind the gonopods.
Female para type. Length 19 mm. Width 1.0 mm. 46 segments,
of which the last 2 are legless.
Variations. Summary of 9 specimens: Length 16 to 20 mm.
Width 0.9 to 1.1 mm. 41 to 49 segments, of which 2 or 3 are legless.
Ocelli are unequal in size; there are from 7 to 11 in 2 irregular rows
that are partly covered by the collulll.
Dis tri but ion. Ca \'ernicolous and epigean in west Texas and
Type locality and specimens. 'I'exas. Culbertson Co.: BordCl'
Can). 30 Sept. 1962, 200, 3~~; 10 Oct. 1962, 900, including the
holotype, 24W, immatUl'e specimens; .J. K. Baker, collector. IIolotype and
paratype of both sexes are in the American ~lus. Nat. !list. (NewYOI'k).
Hemaining paratypes of both sexes are in the U.S. Nat. Mus. and
Other record. New .Mexico. San Miguel Co.: west of Cowles, end
of road to Lake Catherina, sifting in oak litter, 8,400 ft. alt., 26 Aug.
1952,200; C. C. HolT, collector. Amer. Mus Nat. 1.list.
This milliped has not appeared in any of the several collections
that I have from the caves of Eddy County, New Mexico.
Hecord of population intermediate between C. r. reddelli and C.I'.
inornatus: New l\Iexico. Colfax Co: Philmont Scout Hanch, Porcupine
Trail, 9,600 ft. alt., about 15 miles southwest of Cirnmaron, Aug. 12 to
28 Aug. 1962, many specimens; H. C. Albert, collector. ArneI'. lvlus. Nat.
I-list. and author's collection. Summary of 4. specimens: Length 20 to
24 mm. Width LO to L2 mrn. 51 to 56 segments. Ocelli are unequal
in size and arranged in 2 (or rarely 1) uneven rows of 7-2,7-1,6-1,7-0.
Dorsal crests and pore crests are low, wide, and much nearer the
condition in inomatlls than in the nominate subspecies. Horizontal
striae are well developed, as in the nominate subspecies. Lobes on
legpairs 6 and 7 are a little larger than in other populations.
Diagnosis. A small-bodied cavernicole that is like C.r.rcddelli in
the form of the gonopods and lobation of the legs; dilTers from it and
all other congeners in the presence of only very slight traces of dorsal
crests and pore crests; also dilTers from the nominate subspecies in that
the ocelli, of which there are from 2 to 7, are arranged in a single series.
Male holotype. Length about 21 mm. Width LO mm. 51 segments,
of which the last 2 are legless. Ocelli are small, round, and contiguous;
3 are on one side and 2 on other side. Pores begin on segment 5; slight
traces of large, circular pore crests remain. Below the pores,
metazonites and posterior half of prozonites are horizontally striated;
across the dorsum, prozonites lack striae and a series of pits is in
segmental sulcus. Figure 5 is a lateral view of a typical body segment.
I f the segments are viewed from behind, slight traces of 4 dorsal
crests and the upper horizontal striae can be seen.
Gonopods and characters that are not mentioned are as in the
Variations. Length 18 to 24 mm. Width 0.9 to LO mm. 44 to
55 segments. 2 to 7 ocelli are in one row; some may be covered by
The two populations in the caves in Childress and Wheeler Counties
dilTer in body proportions, number of segments, and number of ocelli.
Specimens from Big Mouth Cave and Small Mouth Caves (Wheeler
County) are about 24 times as long as wide, have from 51 to 54 body
segments, and 2 to 4 round ocelli. Specimens from Windmill Crack
Cave (Childress County) are about 20 times as long as wide, have from
44 to 51 body segments, and 3 to 7 ocelli.
Ecology. The records suggest that C.r.inornatus is an obligate
cavernicole. The reduced number of ocelli is the only morphological
evidence of modification by cave life. The loss of the crests may have
occmred before the cave habitat was adopted. The ecological evidence,
as reported by one of the collectors, Mr . .James H. Heddell, suggests
that inornatus is a troglobite. He wrote (letter to author 22 May 1963):
"We recently investigated about 40 gypsum caves in Northwest Texas.
Most of the caves examined are quite recent in origin, despite their
large volume, and most are subject to violent and complete flooding.
I found almost no indications of troglobites in the caves, Big and
Small Mouths in Wheeler County and Bateman Cave in King County
being the only notable exceptions. In these large caves and Windmill
Crack I found the only small, pale spiders ... The area is definitely not
a good collecting area." Later (letter 12 .June 1963 to author) he
added: "I have thought at length about the question of its being
troglobitic, and would be tempted to say that it is. The only active
forms that we saw were in total darkness. Intensive searching in
daylight at the entrance to Small Mouth Cave, where it was quite
moist and food was abundant, yielded no animals. The same held true
at Big Mouth Cave, where the most moist part of the cave was in
subdued daylight. The only animals found, however, in that cave
were in a very dim twilight, under rocks, and quite inactive. Another
matter of interest is that since the discovery of Pleistocene fossils, the
caves appear to be older than we had postulated at fil'St, at least some
of them. What is apparently a troglobitic isopod was also found in
another cave. None of these are conclusive, but I do think that they
suggest that [inornatlls] is a troglobite."
Caves in northwest
Type locality and specimens. 'l'exas. Wheeler Co.: Big Mouth
Cave, 2 miles north of Shamrock, 19 Oct., 1957, ~, T.C.13arr, .Jr.;
May, 1963, 9c3'c3i'n,cluding the holotype, 5W, J. Heddell and W. Hussell.
Small Mouth Cave, 100 yards east of Big Mouth Cave, May, 1963, ~,
Heddell and Hussell. Holotypes and paratypes of both sexes are in
the American Mus. Nat. I-list. (New York); remaining specimens are in
the author's collection.
a th e I' re co rd. Childress Co.: Windmill Crack, 20 miles north of
Childress, May, 1963, 9c3'3',12~~, Heddell and Hussell. Amer. Mus. Nat.
Hist. and author's collection.
Nomenclatorial note. I have examined the many collections of
millipeds made by the Texas Speleological Survey and am convinced
that only one cambaloid species has been collected in the caves of
central and southwest Texas. The members of the Survey made large,
careful collections in Wyatt Cave (the type locality of E.speobills and
C. caeca) and nearby caves in a special elTort to determine whether'
there are two cambaloid species there. They found only one,
confirming Loomis' (1953) conjecture that serious errors were introduced into
the original description of Eclomus speobius and that Cambala caeca
is a synonym of E.speobills.
D i a gn 0 sis. A medium-sized troglobite that is related to the eastern
congeners in the form of the anterior gonopods. Distinguished from
them by the absence of ocelli and by the presence of a tarsal claw on
legpair 1 of the male.
Maximum width and length are 1.9 and 33 mm., respectively. 37 to
50 segments, of which the last 1 to 3 are legless. Body is flesh-colored
and pore glands are red in life; turns dark red-brown in alcohol. With
no conspicuous cave modifications other than absence of ocelli; an
occasional immature specimen has 1 or 2 ocelli. In the mature male,
large, rounded lobe is on the ventral surface of article 4 of legpairs
G and 7, and article 5 of the same legs is swollen. Dorsal crests are
slightly indicated on segment 3; on segment 4 there are G subeqllal
slender crests which are slightly lower than crests on typical segments;
pore crests and pores begin on segment 5. Crests are narrower and
higher than in C. r. reddelli. Apex of caudal tergite is unusually thick
for the genus, rounded, a little longer than anal valves, and asetose.
The following corrections should be made to the figure of the
posterior gonopod, as drawn by Causey (1959): the telopodite consists
of 2 articles, with the length of the longer terminal article
approximately twice as great as the width; the posterior coxite is bilobed,
with the mesial lobe flattened and asetose; a vertical series of minute,
cun'ed spurs is on the mesial surface of the coxa.
Ecology. Found on moist guano, silt, and walls and under organic
debris and rocks in caves from the twilight zone back several hundred
feet in total darkness. Frequently is collected with members of the
polydesmoid genus Speodesmus.
Distrihution. Caves in the following central and southwest Texas
counties: Bandera, Bell, Bexar, Burnet, Comal, Coryell, Edwards,
Hays, Irion, Kendall, Kinney, Lampasas, Medina, Menard, Heal, San
Saba, Sutton, Terrell, Travis, Uvalde, Val Verde, and Williamson.
Type locality and specimens. Eclomus speobills Chamberlin.
'fcxas. Edwards (not Sutton) Co.: Wyatt Cave. Many cotypes and
paratypes (Felton Cave) are in the Chamberlin Collection. I have not
examined them. Topotypes, including 69 mature and immature
specimens collected by .James Reddell and Dill McKenzie 21 Sept.
1963, have been divided among the U.S. Nat. Mus., Amer. Mus. Nat.
lEst. and the author's collection.
Cambala caeca Loomis. 'l'cxas. Edwards Co.: Wyatt Cave, 6
h010type, U.S. Nat. Mus., no. 2087. Felton Cave, 12W paratypes.
Cambala captiosa Causey. 'I'cxas. Williamson Co.: Beck's Hanch Cave,
61 mi. west of Round Rock. 6 holotype, ~ paratype, ArneI'. Mus. Nat.
Hist. Paratypes and many topotypes are in the author's collection.
Genus iUcxic:unhala, new
Type species. Mexicambala russelli, new species. Monobasic.
Diagnosis. Near Cambala; differs in that the sides of the collum
are shorter and closer to the head, the setae on the anal valves and
anal scale are numerous, and the distal margin of the coxite of the
anterior gonopods is not divided.
BALA R U SSELLJ,
D i agn 0 sis. A depigmented, eyeless troglobite that is characterized
by 4. conspicuous, serrated dorsal crests, prominent pore crests, and
elongated tarsal claws and antennae.
Male holotype. Length 21 mm. Width 1.3 mrn. 39 segments, of
which the last 2 are legless. Flesh-colored except for the darker pore
crests when freshly preserved; later became medium gray in alcohol. .
Head resembles species of Cambala. 16 labral setae. 4 widely
separated clypeal setae. Antennae reach back to margin of segment 4;
ratio of length of articles 1through 7 and width of articles 5 and 6 (in
parentheses) is: 30: 40: 57: 46: 45 (29) : 51 (30) 10; the usual sensory
areas are on segments 5 and 6; 4 terminal cones.
Body is slightly nanowed behind head. Segment 7 is not swollen.
Prozonites are markedly narrower than metazonites. Collum (fig. 7)
is length of segments 2 and 3; its lateral margin is short and close
to the head, and the thin margin of the head capsule fits into a
shallow emargination. Dorsal surface of collum is smooth. Segments 2
and 3 have slight traces of dorsal crests and distinct but uneven
horizontal striae. 4 dorsal crests, lacking serrations, are on segment 4.
By segment 9 dorsal crests reach their maximum size and typical shape
(fig. 8); they are thin, high, and divided into 3 angular or rounded
serrations. Pores and pore crests begin on segment 5; distal two-thirds
of each pore crest is thin and sharp, resembling the dorsal crests;
anterior one-third of pore crest is high, rounded in front and angular
behind; pore opens on flat outer surface of crest. Crests are lower on
2 segments preceding the last. Last segment lacks crests and lateral
striae. Surface between crests is smooth and shining. Below crests
there is one weak horizontal crest and a double series of uneven
rectangular areas that are continuous with the single series of
rectangular areas on posterior half of prozonite; anterior half of prozonite
is minutely honeycombed. Caudal tergite is a little longer than penult
segment; caudal margin is rounded, entirely covering anal valves
from above. Distal margin of anal scale is straight. Anal valves are
not margined. Setae are scattered over anal valves and anal scale;
anal tergite is asetose (fig. 9).
Legpair 1 (fig. 10) consists of 6 articles and a claw; article 2 is a
little wider than article 3, but not as wide as in species of Cambala. No
legs are lobed or swollen. Tarsal claws are about two-thirds the length
of the tarsi. Penes are short, contiguous, and 2 macrosetae are on the
apex of each one. As in species of Cambala, ventral margins of segment
6 overlap, ventral margins of segment 7 are contiguous, and opening
of gonopodal cavity is semicircular.
Gonopods closely resemble species of Cambala. Anterior gonopods
difTer in that sternum is shorter and distal margin of coxite and
telopodite are not divided (fig. 11). Posterior gonopods difTer (figs. 12,
13) in that the mesial row of nodules is larger and the teIopodite does
not have a distinct transverse septum.
Note. The holotype may lack a molt of maturity. If so, some of the
legs of the male may be found to be enlarged after the final molt.
Female paratype. Length about 25 mm. Width 1.6 mm. 43
segments, of which the last 2 are legless.
Variations. Mature specimens have from 39 to 43 segments, are
21 to 26 mm. long, and 1.3 to 1.6 mm. wide. Immature specimens are
known with 38, 36, and 32 segments.
Distribution. Type locality in eastcentral Mexico.
Type locality and specimens. Mexico. San Luis Potosi. Cueva
de la Parra, 5 km. north of Xilitla, in total darkness, 26 Oct. 1963.
This species is !lamed
for the collector,
The cavernicoles include: (I) Cambala speobia (Chamberlin), troglobitic
in central and southwest Texas; (2) C.reddelli reddelli n. sp. and subsp.,
troglophilic in west Texas and epigean in New Mexico; (3) C. reddelli
inornatus n. subsp., troglobitic in northwest Texas; and (t,) l1le.r.ieambala russelli
n. gen. anrl sp., troglobitic in southern San Luis Potosi. They are described
and figured, and a key is given.
IU~S UM l~
Les Diplopodes Cambalides actuellemen t connus des parties cen trale et
occidentale du Texas (U.S.A.) et de la partie du Nord-Est du "lexique
sont les formes suivantes: 1° - Cambala speobia (Chamber'lin), troglobie du
Texas, dans les parties cen trale et sud-occiden lale; 2° - C. reddelli reddelli sp.
et subsp. nov., lroglophile dans Ie Texas occidental et epig6 dans Ie
Nouveaul\Jexique; 3° - C. reddelli inol'llatus subsp. nov., troglobie dans Ie Nord-Ouest
du Texas; et f,o - jlTexieambala russelli gen. et sp. nov., lroglobie dans Ie
Sud de l'Etat de San Luis Potosi (Mexique). L'aute11l' donne ici les
descriptions et dessins de ces especes, etun tableau de leur d6termination.
57 (1)-58 (2)
Figs.l-flo Cambala r. reddelli, n. subsp.,
body; 2. Sixth leg.; 3. Anterior gonopods,
gonopod, ectal view.
male paratype. 'l.Anterior end of
an tm'ior view; flo High t posterior
Cambala r. inol'llatus, n. subsp.
Figs. 7-13.l1lexieambalarusselli, n.gen. and sp., male paratype. 7.Anterior
end of body; 8.l\Jiddle body segment; 9. Posterior end of body; 10. First leg.;
I1.Anterior gonopods, anterior view; 12. Hight posterior gonopod, ectal
view; 13. Right posterior gonopod, posterior view.
EclytllS (not Holmgren 1855 ) speobills Chamberlin, 1952 , Ent. News 63 : 10 .
Eclomlls speobills , Chamberlin, 1952 , EnL News 63: 7'1. Chamberlin and HolTman , 1958 , U.S. NaLl\Ius. Bull. 212 : 175 .
Cambalaeaeea Loomis , 1953 , Jour. Washington Acad. Sci., 1 , 3 ( 10 ): 43 ( 12 ): !d 7, figs. 1- 3 . New SYllonomy.
Cambala eaptiosa Causey , 1959 , Proc. BioI. Soc. \Vashington , 72 : 69 - 71 , figs. 1- 3 . Now SYllonomy.
CAUSEY , NELL B . - 1959: Two new troglobyric millipeds from Texas. 1'roc . BioI. Soc. \Vashington 72 : 69 - 76 , 6 figs.
CIIAMBERLIl '(, H. V. , and HOFF~JAi\', H. L. : -1958 Checklist of the l\lillipeds of North America . U.S. Nat. "Ius. Bull . 2 \ 2 : 1 - 2a6 .
HOFnIAN , ILL. -1956: New genera and species of cavernicolous diplopods from Alabama . Geol. Sur. Alabama, Mus. Paper 35 , pp. 1 - 11 , 1\ figs.
- 1958: Appalachian Cambalidae: Taxonomy and distribution (Diplopoda: Spirostreptida) .. Jour. Washington Acad. Sci. f , 8 (il) : 90 - 9f " I map.
HEDDELL , JAMES L., and RUSSELL , \VJI. LIA~I H . -1962: The caves of Northwesl Texas . Texas Spel. Sur . 1 ( 8 ) : I- 56 . 1Ilimeographed .