Thomas C. Barr, Jr.
The Status and Affinities of Duvaliopsis Jeannel (Coleoptera: Carabidae).
Psyche 71(2):57-64, 1964.

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THE STATUS AND AFFINITIES OF
DUVALIOPSIS JEANNEL (COLEOPTERA: CARABIDAE)m BY THOMAS C. BARR, JR.
Department of Zoology, University of Kentucky The genus Duvaliopsis was established by Jeannel (1928) for a small group of endogenous, anophthalmous trechines from the Car- pathian Mountains and the Transylvanian Alps of Romania, Czecho- slovakia, and Poland. Although earlier authors had classified them with Anophthalm~us Sturm, Trechus Clairville, or Duvalius Delarou- zie (formerly considered a subgenus of Trechus) , Jeannel ( 1928) clearly demonstrated their morphological similarity to Trechoblemus Ganglbauer and to North American cavernicole trechines of the genera Pseudanophthalmus and Neaphaenops. Trechoblemus, Duvaliopsis, Pseudanophthalmus, and Neaphaenops were placed in a "s&e phyl6- tique de Trecf~oblemus"', united by the common possession of certain characters: ( I) the mentum is fused to the prementum; (2) the re- current portion of the apical groove of the elytron is usually connected to or directed toward the 3rd longitudinal stria; (3) the copulatory sclerites (of which there are one or two) are placed laterally (aniso- topic), rather than ventrally (isotopic), in the internal sac; and (4) the anterior tibiae are pubescent on the outer side. Subsequent to 1928, additional genera in North America and Japan have been described which should probably be allied with this series (Valentine 1952, Yoshida and Namura 1952, Uino 1956 and 1958, Barr 1960).
In the eastern United States, the largest and most widely distributed genus of cave beetles is Pseudanophthalmus, species of which are now known from Indiana, Kentucky, Tennessee, Alabama, Georgia, Ohio, Virginia, West Virginia, and Pennsylvania. Although found only in caves up to the present time, a few Virginia species have rudimentary eyespots, suggesting comparative recency of adopting a wholly sub- terranean mode of life. The absence of epigean trechines from North America which seem to share a relatively recent ancestry with Pseu- danophthalmus and other cave genera has provoked considerable specu- lation on the history and evolution of the group. Trechoides fasciatus Motschulsky, from the Oligocene Baltic amber, could belong either to Laswtrechus or Trechoblemus (Jeannel, 1928). This fossil demon- 'This investigation was supported in part by a grant from the National Science Foundation, no. G-18765.
Manuscript received by the editor February 25,1964. Pwhe 7157-64 (196-1). hup Wpsycht cnlclub orgi71/71-057.htw.l



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58 Psyche [June
strates the presence of the series in Europe in the mid-Tertiary, but reveals nothing about the North American representatives. Morpho- logically, the closest known relative of Pseudanophthalmus is Duva- liopsis.
The Museum of Comparative Zoology, Harvard University, re- cently obtained part of the collection of Dr. Eduard Knirsch, Kolin, Czechoslovakia, which contains 44 specimens of Duvaliops'is, including all 8 forms treated by Jeannel in his monumental Monographie des Trechinae (1928). I am indebted to Dr. Philip J. Darlington, Jr., curator of entomology at the Museum of Comparative Zoology, for permission to undertake a study of these beetles. When Jeannel (1928) established the genus Duvaliopsis, only 10 species of the North American Pse.udanophthahus were known, pre- senting a far narrower conception of the limits of the latter genus than is held today. The chief diagnostic characters of Duvaliopsis were said to be (Jeanne1 1928) : ( I ) punctures 3 and 4 of the margi- nal series ("fouets humeraux de la sirie ombiliquie") are closely applied to the marginal gutter; and (2) the transfer apparatus con- sists of a single copulatory piece in the form of a very long, concave spoon, bifid at the apex, the convex side facing the right side of the internal sac.
Pse'udanophthalmus differed in having : ( I ) punctures 3 and 4 of the marginal series farther from the marginal gutter than punctures I and 2; and (2) the transfer apparatus consisting of 2 pieces, not bifid.
In examining all 8 forms of Duvaliopsis known to Jeannel when he established the genus and in comparing them with most of the known species of Pseudanophthalmus, I am unable to find any consistently significant difference in the chaetotaxy of the humeral marginal set. In larger species of Pseudanophthalmus, especially those with moder- ately convex elytra, the 3rd and 4th punctures do appear farther from the gutter than the 1st and 2nd. As Jeannel himself very clearly explained (Monographic, 111, p. 18), the relative positions of the umbilicate series are far from absolute, and are related to the hyper- trophic enlargement of the external interstriae. Thus the alleged
generic character would appear valid when a large, convex Pseuda- nophthalnzus (e.g. P. menetriesii Motsch., the generotype) is compared with a Duvaliopsis (all of which are small and rather depressed), but would break down when the comparison is made with a small species of Pseudanophthalmus with depressed elytra. The transfer apparatus of Duvaliopsis is indeed distinctive, but so are the many transfer apparatus types of the twenty-odd species groups of Pseudanophthalmus. The unusual length of the copulatory piece in



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19 641 Burr - Duvalio psis 59
itself is not diagnostic, since certain species of Pseudanophthalmus, e.g. the gracilis section of the hubbardi group, have equally long trans- fer apparatuses. Even the single copulatory piece is not peculiar, since P. cumberlandus Val. and its allies have but a single copulatory piece. In conclusion, there appears to be no reason why Duvaliopsis should be maintained as a genus distinct from Pseudanophthalmus. Its dis- tinctiveness is on the order of magnitude of the difference between various species groups of Pseudanophthalmus, and it is to this status which I propose it be relegated.
The study of the Knirsch material has suggested certain changes in the taxonomic arrangement proposed by Jeannel ( 1928). A revision is given below.
PseudanophthaZmus Jeannel
Jeannel 1920 : p. 154; type species: Anophthalmus menetriesii Mo,tschulsky. SYNONYM: Duvaliopsis Jeannel 1928 : p. 106 ; type species: Anophthalm~ bielzi Miller.
bielzi group
( = Duvdiops'is Jeannel )
Size small (3.3-4.0 mm.) , integument pubescent. Head rounded or slightly wider than long; eyes absent, their site indicated in some species by a small, oblique cicatrix; antennae about half the body length (except in rybinkskii). Pronotum transverse, 1/5 to 1/3 wider than long; margins arcuate in apical 1/2 to 2/3, then sloping evenly back to the hind angles with little or a very brief sinuosity; hind angles rather small, variable; disc with sparse, long pubescence. Elytra 3/5 as wide as long, subconvex or depressed; striae regular or irregular, deeply or shallowly impressed, finely punctulate (except in bielzi) ; first discal puncture at the level of the 4th marginal puncture, slightly anterior to it, or slightly posterior; apical recurrent groove highly variable, parallel or oblique to the suture, connecting with either the 3rd or 5th longitudinal stria, with or without a crochet, but always terminating well in advance of the apical puncture. Aedeagus usually arcuate and moderately slender, the basal bulb set off from the median lobe by a slight constriction, the apex attenuate with the tip reflexed (produced into the shape of a boot in meliki} ; transfer apparatus a single, elongate, spoon-shaped copulatory piece with its concave side facing the left side of the internal sac, its dorsal and ventral edges rolled and sclerotized and apically produced to give a bifid appearance ; internal sac with moderate armature of small, blunt scales; parameres with 3 or 4 setae at their apices. Type species: Anophthalmus bielzi Seidlitz.




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60 Psyche
I
Elytra with internal longitudinal striae deeply impressed, all striae more or less irregular; labrum with a rather deep, . .
...................................................... V-shaped emargination 2
Elytra with the internal longitudinal striae shallowly impressed and regular, finely punctulate; labrum shallowly emargi- nate .................................................................................... 3 2 ( I) Elytra subconvex, striae deep and fairly regular, with little or no punctulation ; humeri rounded, the prehumeral borders oblique to the median line (Romania : Transylvanian Alps) .................................................................... bielzi (Seidlitz) Elytra depressed, internal striae moderately impressed, slightly irregular, and finely punctulate; humeri more angular, the prehumeral border nearly perpendicular t,o the median line (Czechoslovakia: Carpathians) ............ pilosellus (Miller) 3 ( I ) Antennae half as long as the total body length ; segments VII-X ................................
thick, more than half as wide as long
4
Antennae 2/3 as long as total body length; segments VII-X slender, less than half as wide as long (P,oland: Carpathi- ...................................................... ans) rybinskii (Knirsch)
4(3) Humeral margin distinctly serrulate ; head as long as wide .... 5 Humeral margin with serrulations obsolete ; head slightly but distinctly wider than long (Romania: eastern Carpathians) calimanensis (Knirsch)
........................................................ 5 (4) Aedeagus long and slender, the apex reflexed then curved down- ward in the shape of a boot (Romania: eastern Carpathi- . . . .
.............................................................. ans) 7nelzkz ( Csiki )
Aedeagus robust, with a slender and briefly produced apex (Romania : Transylvanian Alps) .... transylvanicus ( Csiki ) Most of the species are figured by Knirs'ch ( 1924) or Jeannel (1928), who also list the precise localities from which each is known. Pseudanophthalnzus bielzi (Seidlitz) new combination Anophthalmus Bielzi Seidlitz 1867: p. 45. Trechus (Duvalius) Bielzi: Knirsch 1925: p. 90. Duvaliopsis Bielzi: Jeannel 1928 : p. 109, figs. 1349-1352. Pseudanophthalmus pilosellus (Miller) new combination Anophfhalmus pilosellus Miller 1868 : p. 11. Trechus (Duvalius) Bielzi pilosellus: Knirsch 1924: p. 65, figs. 3 and 8; Knirsch 1925: p. 91.
'Includes only the Knirsch material ; I have not seen D. p'-.losellus beskiden- sis Hliskowski 1942 (Ent. Listy 5 : 17).



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19 641
hrr - D UV&OPS~S
Duvaliopsis pilosellus: Jeannel 1928: p. 110, figs. 1354, 1346, and 1347. Anophthalmus Bielzi Stobieckii Csiki: 1907: p. 574. Trechus (Duvalius) Bieizi Stobieckii: Knirsch 1925: p. 91. Duvaliopsis pilosellus Stobieckii: Jeannel 1928 : p. 11 1. Jeannel's key to species (Jfonographie, 111, p. 108) is erroneous because bielzi is said to have angular humeri with the prehumeral border perpendicular to the median line. However, only in pilosellus are the humeri sharply angular and the prehumeral borders perpendicu- lar. This is amply confirmed by Knirsch's long series of both pilosellus and stobieckii. I can find no taxonomically significant differences be- tween these two supposed subspecies. The male genitalia are identical. Pse.udanophthalmus rybinskii (Knirsch) new combination Trechus (Duvalius) Bielzi Rybinskii Knirsch 1924: p. 63; figs. 5 and 6; Knirsch 1925: p. 91.
Duvaliopsis pilosellus Rybinskii: Jeannel 1928 : p. 111. Pseudanophthal7nus calimanensis (Knirsch) new combination Trechus (Duvalius) Blelzi calimanensis Knirsch 1924: p. 65, figs. 4and.7. Duvaliopsis pilosellus calimanensis: Jeannel 1928: p. Ill, fig. 1353. .
These two species, P. rybinskii and P. cali7nanensis, are quite distinct from pilosellus in characters given in the key as well as in genitalic differences. Judging from strictly morphological criteria, they are probably more closely related to nzeliki and transylvanicus than they are to either bielzi or pilosellus.
Pseudanophthalrnus meliki ( Csiki) new combination Anofhfhalmus Bielzi Meliki Csiki 1912: p. 537. Duvaliopsis Meliki: Jeannel 1928 : p. 114, figs. 1355, 1356, and 1357. Trechus (Duvalius) pauperculus Knirsch 1925 : p. 91. Duvaliopsis Meliki pauperculus: Jeannel 1928: p. 114. In the distinctive boot-shaped enlargement of the apex of the aedeagus, this species recalls a similar feature in P. valentinei Jeannel and P. vanburenensis Barr (Tennessee, U. S. A.). The other species of the bielzi group are possibly more closely related to each other than to P. nzeliki, but the transfer apparatus confirms their affinity with the latter species.
Pseudanophthal?nus transylvanicus (Csiki) new combination Anophthalmus Bielzi transylvanicus Csiki 1902 : p. 52. Trechus (Duvalius) transylvanicus: Knirsch 1925 : p. 91. Duvaliopsis transylvanicus: Jeannel 1928: p. 113.



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In this species the head is as long as wide; the labral emargination is very shallow; the pronotum is I 1/4 times as wide as long; the hind angles of the pronotum are acute, with a deep and very brief marginal sinuosity before them; the elytral humeri are clearly serrulate; the first discal puncture is at or behind the 4th humeral marginal puncture; and the apical recurrent groove is long and parallel to the suture, variably connected with the 5th or the 3rd longitudinal stria. The aedeagus of a topotype (Schuler Gebirge, Transylvanian Alps, Romania) in the Knirsch collection measures 0.86 mm. long, much larger and more robust than that of pilosellus. The apex narrows abruptly and is briefly produced. The copulatory piece measures about 1/3 of the total length of the aedeagus. DISCUSSION
The realization that the Pseudanophthabnus found in caves of the eastern United States have their endogenous counterparts in the moun- tains of eastern Europe is primarily of zoogeographic interest. Like many disjunct distributions, this one suggests an earlier, broader distri- bution followed by intermediate extinction. Certainly the geographic extent of compatible trechine microenvironments would have been considerably broadened under the influence of a periglacial climate. The species of the bielzi group are now as closely restricted to the higher elevations (1200 meters and above, according to Jeannel, op. cit.) of the Carpathians and Transylvanian Alps as the American Pseudanophthalmus are restricted to caves. Both American and Euro- pean species are presurnabl,~ descended from winged, Trechoblemus- like ancestors.
Although the species of the bielzi group are not primarily cavernico- lous, they are probably similar to forms which colonized North Ameri- can caves during the Pleistocene interglacials. Endogenous Pseuda- nophthalmus have not been discovered in the eastern United States. I made a careful search of the high mountains of North Carolina and Tennessee in the summer of 1960, finding many T'rechus (Barr 1962) but no Pseudanophthalmus. If we were to seek a close environmental parallel to the Carpathians and Transylvanian Alps in North America, however, we would have to look farther north, nearer to the terminal moraines of the Pleistocene glaciers. The few scattered peaks 4000 feet or higher in Virginia and West Virginia would bear careful search. A recent study of the Pseudanophthalmus of the Appalachian valley (Ban-, in press) suggests that the cave species of that area have descended, with slight modification, from a smaller number of endo- genous species. Each ancestral endogenous species is presumed to



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19641 Burr - Duvaliops'is 63
have occurred in the geographic area in which the caves are at present inhabited by closely similar, allopatric species or subspecies. Similar patterns of speciation occur among Pseudanophthalnzus of the horni group in the Bluegrass of Kentucky, where apparently two ancestral species colonized the caves. One species had a short aedeagus similar to that of P. horni Garman, while the other had a long, hooked aedeagus similar to that of P. inexpectatus Bars. A single ancestral species is postulated for the tiresias section of the engelhardti group, which occupies the Central Basin of Tennessee. Cave colonization and speciation does not seem to have been radically different in the Appala- chian valley, the Bluegrass, and the Central Basin. Patterns of trechine speciation are more difficult to explain in cave systems of the karst plains developed on Meramac and Chester lime- stones of the Interior Low Plateaus - specifically, the Mitchell plain of Indiana, the Pennyroyal plateau of Kentucky, and the Eastern Highland Rim of Tennessee. Here the networks of subterranean solu- tional openings are more extensive, and dispersal from one cave system to another takes place more readily.
Here it is possible for abundant,
mobile species to have (for cave trechines) fantastically extensive ranges, up to 75 miles long in Darlingtonea kentuckensis Valentine and 110 miles long in Neaphaenops tellkampfii Erichson. Here it is not uncommon for 3, 4, or even 5 species of troglobitic trechines to inhabit the same cave, a phenomenon best explained by multiple in- vasion.
But despite the special interest that American coleopterists may have in speculating that American Pseudanophthalnzus descended from preadapted, montane, endogenous species like those of eastern Europe, the bielzi group itself deserves further careful study. With the possible exception of P. pilosellus, all the species are quite rare, so that mor- phological variation cannot be adequately subjected to statistical analysis. No useful taxonomic purpose is served by naming each local population a different subspecies, as has been done for certain Euro- pean casabids (hundreds of names have been applied to Carabus granu- Zatus and C. cancellatus, for example).
It appears premature to apply
the polytypic species concept to the bielzi group. However, extensive collecting, especially in Romania, would make possible a sound study of alpine speciation in the b':elzi group, involving analysis of variation and comparison of existing geographic ranges with Pleistocene glacial patterns and inferred Pleistocene climatology. Few detailed studies of the flightless insects of the Carpathians have been made (Kaszab 1961 ) . Such an investigation, while increasing the store of informa- tion on the role of the Carpathians and Transylvanian Alps as a



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Psyche
Pleistocene refugium, would also significantly broaden the base of our knowledge of cave colonization and speciation among trechines. BARR, THOMAS C., JR.
1960. A new genus of cave beetle (Carabidae: Trechini) from south- western Virginia, with a key to the genera of the Trechini of North America north of Mexico. Coleopterists' Bull., 14(3) : 65- 70.
1962. The genus Trechus (Coleoptera : Carabidae : Trechini) in the southern Appalachians. Ibid., 16 (3) :65-92. CSIKI, E.
1902.
(Descr. A. Bielzi transylvanicus). Allat. Kozl., 1: 52. 1907. (Descr. A. Bielzi Stobieckii). Ann. Mus. Nat. Hung., 5 : 574. 1912. (Descr. A. Bielzi ,Meliki). Ibid., 10 : 537. JEANNEL, R.
1920.
Notes sur les Trechini. Bull. Soc. Entomol. France, 1920: 150-155. 1928. Monographie des Trechinae. Troisiime livraison. L'Abeille, 3 5 : 1-808.
KASZAB, ZOLTAN.
1961. Die zoogeographischen Beziehungen der Karpaten und seiner Becken. Rovart. Kozl.. 14 f 16) : 266-269. , > ,
KNIRSCH, E.
1924. Zwei neye Subspecies des Trechus (Duvalius) Bielzi Seidl. Casopis Ceskoslovenskk Spol. Entomol., 21 (3/4) : 63-66. 1925.
Ein neuer Trechus aus den Ostkarpathen, Trechus (Duv.) pauper- culus. Ibid., 21 : 89-91.
MILLER, L.
1868. Eine entomologische Reise in die ostgalizischen Karpathen. Abhandl. d. z001.-botan. Ges. Wien, 18: 3-34. SEIDLITZ, GEORG.
1867. Beitrag zur Kaferfauna Siebenburgens. Verhandl. u. Mitteil. d. ~iebenbur~ischen Vereins f. Naturw. in Hermannstadt, 18: 43-46. UENO, SHUN-ICHI.
1956. New cave-dwelling trechids of Kurasaivatrechus-group (Coleop- tera: Harpalidae). Mem. Coll. Sci. Unk. Kyoto (B) : 23 : 69-78. 1958.
Two new trechids of Kurasawatrechus-Group found in the lime- stone caves of Japan (Coleoptera: Harpalidae). Japanese J. Zool., 12 (2) : 123-131.
VALENTINE, J. M.
1952. New genera of anophthalmid beetles from Cumberland caves. Geol. Surv. Alabama Mus. Pap., 34: 1-41. YOSHIDA, A., AND S. NOMURA.
1952. A list of the Arthropoda in the limestone caves in Kant'6-Moun- tainland, with the descriptions of a new genus and three species. The Chuh& Tokyo, 6: 1-8.




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