Carl H. Lindroth and Richard Freitag.
North American Ground-Beetles (Coleoptera, Carabidae, excluding Cicindelinae) Described by Thomas Say: Designation of Lectotypes and Neotypes.
Psyche 76:326-361, 1969.

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NORTH AMERICAN GROUND-BEETLES
(COLEOPTERA, CARABIDAE, EXCLUDING
CICINDELINAE) DESCRIBED BY THOMAS SAY:
DESIGNATION OF LECTOTYPES AND NEOTYPES
INTRODUCTION
Thomas Say (1787-1834) was the founder of scientific entomology and conchology in North America. He described an immense number of insects of all orders, among the Carabidae (incl. Cicindelinae) no less than 165 species. They were included in the following eight papers :
1817. Descriptions of several new species of North American insects. Journ. Acad. Nat. Sci. I :2. Philadelphia. pp. 19-23. (Only Cicindela, among the Carabidae.)
1818. A monograph of North American insects, of the genus Cicindela. Trans. Amer. Philos. Soc. (N. S.) I, Philadelphia. pp. 401-426.
1823a. Descriptions of coleopterous insects collected in the late expedition to the Rocky Mountains, performed by order of Mr. Calhoun, Secretary of War, under the command of Major Long. Journ. Acad. Nat. Sci. ?:I. Philadelphia. pp. 139-216. 1823b.~ Descriptions of insects of the families of Carabici and Hydrocanthari of Latreille, inhabiting North America. Trans. Amer. Philos. Soc. (N. S.) 2 :I. Philadelphia. pp. 1-109. 1824. Appendix to the narrative of an expedition to the source of St. Peter's river, &c., under the command of Stephen H. Long, Major U. S. T. E. 2. Philadelphia. pp. 268-378. I 825. American entomology. 11. Philadelphia. Pis. I 9-36. 1828. American entomology. 111. Philadelphia. Pis. 37-54. 1834.~ Descriptions of new North American insects and observa- '2001. Inst., Lund, Sweden.
'Lakehead Univ., Port Arthur, Ontario, Canada. Manuscript received by the editor May 16, 1969 'The year of publication of this important paper is much discussed. It
is often given as 1825 (e. g. by Hagen 1862-63; Leng 1929; Horn & Schenk- ling 1928-29), but Leconte (1859b, p. 435) is obviously right in assuming the year to be 1823. This, among other things, gives certain Say names priority over names given by Germar (1824), as expressly stated by Say himself (1834, p. 421, Anchomenus cincticollz's}, *Part of Say's paper of 1834 had been printed in 1829-33 in a newspaper, "The Disseminator," and another part as a separate pamphlet, both at New Harmony, Ind. (see Leconte 1859b, p. 521). Neither can be regarded as valid publication.




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19691 Lindroth - Ground Beetles 327
tions on some already described. Trans. Arner. Philos. Soc. 4. Phil- adelphia. pp. 409-470.
In the present paper we have treated only the true ground-beetles. The I 6 Say species of tiger beetles (subfam. Cicindelinae) are omitted and also 12 species described from Mexico. The remaining specific names given by Say among the Carabidae are 147 in number. Leng (1920) gives the same figure, but two of his names (Calosoma in- distincturn, HarpaZus tricolor), listed as synonyms, are nomina nuda. On the other hand, Calathus gregarius, in Leng, was attributed to Dejean instead of to Say, and Harpalus similis (Anisotarsus s.) was omitted.
Of the two authors of the present paper, R. Freitag is responsible for the 5 species belonging to genus (or subgenus) Evarthrus among the Pterostichini, whereas C. H. Lindroth studied the remaining species with kind help, in some cases, from the following experts, who selected neotypes among their specialities: Prof. R. T. Allen, University of Arkansas, Fayetteville, Ark., Loxandrus rectus.
Mr. T. L. Erwin, University of Alberta, Edmonta, Alta., Brach- inus cyanipennis and B. stygicornis.
Mr. T. F. Hlavac, Harvard University, Cambridge, Massachu- setts, Clivina pallida.
Say's descriptions were usually good, sometimes excellent for his time. Nevertheless, many of them, notably of course in critical genera, cannot be reliably interpreted. Unfortunately, Say's private collection was entirely destroyed after his death (Leconte 185ga, p. VI; Ord 1859, p. XIX, footnote), and a comparison with authentic Say specimens is therefore excluded (but see below). For the correctness of the present applications of his names, we are indebted primarily to John L. Leconte who, in 1859, edited "The Complete Writings of Thomas Say," with comments as to the right interpretation and taxonomic position of most Say species of Coleoptera. The Leconte Collection, in the Museum of Comparative Zoology ( MCZ) , Cam- bridge, Mass., provides a complete picture of how the Say species were interpreted by him.
In order to stabilize future use of specific names given by Say, it is highly desirable to designate type specimens. The aim of the present paper is to do so, as far as ground-beetles are concerned. It might seem, considering the total destruction of the Say Collection (see above), that a selection of neotypes would be the only solution. However, it so happened that Say distributed specimens from his own collection to at least one of his contemporaries abroad, Count



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328 Psyche [September
P. F. M. A. Dejean in France. This is apparent from Dejean's famous work, "Species General des Coleopt&-es" (Vols. I-V, 1825- 31). When he wrote the first volume (1825), Dejean had not acquired contact with Say, but in the preface to Vol. 2 (1826, p.
VIII) he acknowledges, in a general way, having received many Say specimens. In this, as well as in the later volumes of his work, when re-describing certain Say species, Dejean mentions that Say had sent him specimens.
This is the case for the species listed in Table I. Under many other species names, here omitted, Dejean quotes Say as the author but without mentioning specimens received from him.
One of my assistants, Mr. Reinhold Cha,rpentier, when visiting the Museum National d'Histoire Naturelle in Paris (MNP) in 1967, kindly undertook to study the Oberthiir Collection, which includes the Dejean Collection, to search for genuine Say specimens. Dejean specimens are easily recognized by their bright green labels (see Lind- Table 1. Say species mentioned by Dejean in "Species General des Coliop- teres," Parts 11-V, as received from Say (modern generic names used). An * = with Say's name on the specimen.
Species Say description Dejean reference Agonum decentis
1823b: 53 111.1828 :lo7
Amara impuncticollis
1823b: 36 111.1828 :466
*A. musculis
1823b: 35 111.1828 :478
*Anisodactylus agricola
1823b: 33 IV.1829:151
A. baltimoriensis
1823b: 33 IV.1829 :I54
*A. caenus
1823b: 34 IV.1829 :I59
A. rusticus
1823b: 32
IV.1829 :I57
%sotarsus terminatus
1823b: 48 IV.1829 :356
*Bembidion contractum
1823b: 85 V.1831 :I24
*B. dorsale
1823b: 84 V.1831: 72
*B. levigatum
1823b: 84 V.1831 :I51
Calosoma luxatum
1823a :l49 11.1826 :I97
*Carabus sylvosus
1823b: 75 11.1826 :I52
*Chlaenius emarginatus 1823b: 63
11.1826 :367
Dyschirius globulosus
1823b: 23 11.1826 :480
D. pallipennis
1823b: 24 11.1826 :481
D. sphaericollis
1823b: 23 11.1826:480
Harfalus erraticus
1823b: 27 IV.1829 :258
*H. faunus
1823b: 28 IV.1829 :254
*Lebia tricolor
1823b: 11 11.1826 :454
Olisthopus parmatus
1823b: 49 111.1828 :I82
*Pasimachus subsulcatus
1823b: 19 11.1826 :471
Scaphinotus bilobus
1823b: 73 11.1826: 17
*Sten0 'ophus ochropezus
1823b: 54 IV.1829 :424
Synuchus impunctatus
1823b: 45 111.1828 :469
Tachys flavicauda
1823b: 87 V.1831: 54
T. inornatus
1823b: 87 V.1831: 53




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19691 Lindroth - Ground Beetles 329
roth 1955a), and in some cases the pin bears a small additional square of the same color with "Say" or "D. Say" in Dejean's hand, or this note is added on the first label, implying that Say's name was written there twice. These specimens, belonging to species marked with an asterisk (*) in Table I, must be accepted as authentic Say specimens and are available for selection as lectotypes. Mr. Char- pentier was able to find representatives of g such species. Afterwards, Mr. A. Descarpentries, of the Paris Museum, was kind enough to make a complementary search for Say specimens in the Oberthur collection, and the rediscovery of three of the species (Chlaenius emarginatus, Pasimachus subsulcatus, Stenolophus ochropezus) was due to his efforts. For reasons mentioned below under each species, lectotypes were not selected for the following Say species represented in the Dejean collection : Anisodacty Zus agricola, Bembidion con- tractum, B. dorsale, Harpalus faunus.
Article 75 of the International Code of Zoological Nomenclature sets the following limitations and conditions for designating neotypes. They are to be designated only if no holo-, lecto-, or syntypes exist: this condition is satisfied in the present case by information given in preceding paragraphs. They are to be designated only in connection with revisory work: present designations are connected with the senior author's revisory work on the ground-beetles of Canada and Alaska (Lindroth I 961 -I 968), which includes almost all species of northern United States too. They are to be designated only in "ex- ceptional circumstances," in the interests of stability of nomenclature : for reasons given in preceding paragraphs, neotypes are considered necessary to stabilize use of Say's names. Characters regarded as differentiating the taxa for which neotypes are designated are given in bibliographic references in square brackets; the reference is usually to Lth 1951-1968. Evidences are given for believing that the neo- types are consistent with what is known of the original type material (according to the descriptions and with reference to localities). And the neotypes have been marked as such for recognition, and are the property of the Museum of Comparative Zoology. The proposal to designate neotypes for Say's Carabidae is known to and approved by several of the most active specialists in North America. ENUMERATION OF SPECIES~
In the following pages, all Say names given to North American Carabidae other than Cicindelinae are treated under Say's original 'Aretkarea helluon'is Say (1834, p. 411) is evidently an artifact (see Lee. 1859b, p. 524; Chd. 1871, p. 287). It is omitted from the present list.



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330 Psyche [September
genus names but in the order of Leng's Catalogue (1920), the Leng number being given in each case.
Of 8 species, lectotypes (in the Paris Museum, MNP) are desig- nated; of the remaining species, neotypes are designated and deposited in the Museum of Comparative Zoology (MCZ). Exceptions are 2 nonzina nuda (names without descriptions) and 7 nomina dubia (names not interpretable).
In the majority of cases, a specimen in the Leconte Collection (MCZ) could have been chosen as neotype. We have, however, avoided doing this for two reasons : ( I ) it seemed more convenient to keep all the Say neotypes together, as a separate collection, and 2 ) it seemed important to designate a type locality if possible, or at least a restricted type area, and to select specimens with appropriate locality labels. Such specimens are seldom available in the Leconte Collection. This has allowed us to select specimens from as close as possible to the parts of the country from which Say's specimens came, when he gives this information.
For one species, Patrobus longicornis, a neotype has been selected by Darlington, 1938 ( MCZ) , and type localities or type areas have been designated by Lindroth ( 1961, 1963, 1966, 1968) for many other Say species.
134~ Cychrus bilobus ( 1823b: 73). Type area "Missouri" or "North-Western Territory." Say's description and the later colored plate (1828: XLV: 3) do not permit a reliable separation from allied species, cavicollis Lec. and fissicollis Lec. The ~rovenience "Missouri" (ace. to Schwarz, 1895, p. 270, probably = N Nebraska) suggests fissicollis, represented from Kansas and Missouri in the MCZ. "North-Western Territory," however (ace. to Schwarz, I.c.), is probably Minnesota. This is herewith designated as type area. As neotype I have selected a from a locality as close to Minnesota as possible : Nipigon, W. Ont. - Scaphinotus momaretus) bilo bus [Lth. 1961, p. 191.
170 Carabus sylvosus ( 1823b: 75). No type area given. Desig- nated type loc.:
Asheville, N. C. (L'th., 1961, p. 41). The inter- pretation of Say's name is clear from his description of the dilated palpi and the elytral sculpture. The species is geographically uni- form, except in the south (see Van Dyke, 1945). In MNP is a Dejean 8 with two of the usual green labels: (a) "sylvosus Say"; (b) "D. Say." It agrees with the present concept of the species and 'The number preceding each name is the number assigned in the Leng (1920) list.




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19691 Lindroth - Ground Beetles 331
I have designated it as iectotype. - Carabus syZvosus [Lth. 1 961, P- 411-
171 Cmabus serratus ( 1823b: 77). No type area given. Ashe- ville, N. C., designated as type loc. (Lth. 1961, p. 40). The inter- pretation of Say's name is clear from his description of the serrate elytral margin, from which his name was derived. A 8 from the type Ioc. designated as neotyfie. - Carabus serrat,us [Lth. 1961, P. 401.
172 Cambus Zimbatus ( 1823b : 77). Type area Maryland. The application of Say's name is clear from his description of the eIytra1 sculpture. The species has no tendency of geographical variation. A d from Charles Co., Md., designated as neotype and the place as type loc. - Carabus Zim batus [Lth. I 96 I ) p. 351. 173
(syn.) Cm-ah interruptus ( 1823b : 62). No type area given. The interpretation is clear from the description of the elytral sculp- ture, and the synonymization with vinctus Weber ( 1801) was ac- cepted by Say himself (1834, p. 416). A 8 from Germantown, Penn., designated as neotype and the place as type loc. - Carabus vinctus Web. [Lth. 1961, p. 341.
I 76 Carabus externus ( I 823a : I 50). Type area "Arkansa." Say's description of the form of prothorax and of the elytral sculpture seems sufficient for an identification. A Q from Little Rock, Ark., designated as neotype and the place as type loc. - Caiosoma externum [Lth. 1961, p. 491.
200 CaZosoma obsoleta (1823a: 149). Type area "Arkansa" and "near the Rocky Mountains"; apparently the Arkansas River is meant. Say's description of the sculpture and the bluish foveolae of the elytra seems to exclude other species. A ? from Fort Reynolds, Colo., designated as neotype and the place as type loc. - Calosoma obsoZetum [Lth. 1961, p. 491.
200 (syn.) Caiosoma indistincturn. This is a nornen nudurn, ap- parently never described by Say (in spite of the quotation "1825 : 151'' in Leng, 1920). It was mentioned by Lec. (1845, p. 208) being the same as Zuxatum Dej. (1826, p. 126; nec Say)) that is obsoietum Say.
219
Caiosoma /uxata ( 1823a :
149). Type area "Arkansa," no
doubt in the same sense as for C. obsoleta, described in the same paper. The iuxatunz group of CaZosonza (CdZisthenes) is extremely di~fficult and it is important to fix the ropert ties of the true iuxatum Say. The original patria, "Arkansa," no doubt aims at the upper parts of the Arkansas River, that is, in Colorado. According to the revision of Mrs. Gidas~ow ( 1959, map, fig. I I, p. 312)) only what



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332 Psyche [September
she regards as the true luxatu?n has been found in Colorado. A c? labeled ('Douglas Spring, Routt Co.? Colo." (coll. Fall) has been selected as neotype and the place as type loc. [Lth. 1961, p. 541. Description of d neotype. - Chaetotax~. Head: 2 fix-points (setae broken) each side inside hind-margin of eye. Prothorax: all setae broken but? judging from fix-point, there have been, on each side? only I at hind-angle and I at middle laterally. (On the high vari-
ability of this character, see Lth.? 1961, p. 55.) - Form of prothorax approximately as in Gidaspow's fig. 55 ; but greatest width clearly before middle and sides faintly sinuate in basal half. - Elytra with strong sculpture, striae regular in frontal half, tegulae subquadrate, alnlost flat near the suture anteriorly? becoming rounded and very convex (granulate)
laterally and apically. - 3 : also I. pro-tarsal segment with well-developed brush (as described by Gidaspow, p. 313, for a sample from Colo.). Penis apex as in a d' from Utah, figured by Gidaspow (fig. 166). - Calosoma Z,uxatum [Lth. 1961, P. 541.
228
Elaphrus fuliginosus ( I 834 : 41 7). Type area Pennsylvania. Say's description was based on a specimen with head and prothorax mutilated ; he therefore used only elytral characters when comparing it with the Europem uliginosus I?. Leconte first (1859b, p. 530) suggested identity with cic~tricosus Lec. but in his collection (MCZ) applied the name as n0~7 unanimously used (apparently following Crotch, in Horn, 1876). Though Say's description cannot be inter- preted? this practice should be retained. No specimen from Penn. could be found.
A d? Rumney? N. H.? designated as neotype.- Elaphrus fuZiginosus [Lth., 1961, p. I 141. 233
Eluphrus mscarius ( 1834 : 41 7). Type area Pennsylvania, named in the first place and designated by Lth. (1961, p. 119). The identity of Say's species seems clear from his comparison with Euro- pean specimens of riparius L. A d from Columbia, Penn,, designated as neotyfle and the place as type loc. - Ehphrus ruscarius [Lth. 1961? P. 1191.
246 (syn.) Notiophilus florrectus ( 1834: 418). No type area given. The description of the form of prothorax and the pale color of the legs confirms the view introduced by Lec. (1863) that this is a synonym of aeneus Hbst. (1806). A 3 from Pennsylvania desig- nated as neotyfie and this state as type area. - A~otiophilus aeneus Hbst* [Lth. 1961, p. 931.
247
Noiio@hilus se?nistriatm ( 1823b : 81 ) . No type area given. Llarquette, Slich., was designated by Lth. (1961, p. 94) but, since this specimen could not be rediscovered at the MCZ? a new type



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19691 Lindroth - Ground BeetZes 333
loc. is proposed below. - Say's description cannot be interpreted. His forma typica (nominate form) ) because it has "feet black," is more likely to be referred to, aquaticus L., whereas his "Var. a)'' with "tibiae piceous)" may be semistriatus auct. The 'Var. P" of 1823 was Iater (1834) by Say separated as distinct under the name of porrectus (= aeneus Hbst.). Say's concept of semistriatus was thus composite and, though Lec.
(e. g. 1848, p. 450) confused it with
his novemstriatus, it seems permissable to retain the name as generally used in this century) that is) for the species with a single preapical elytral puncture and pale tibiae. Fall (1906) pp. 79, 84) reports that F. Blanchard saw "undoubtedly authentic exponents of Say's sernistriatus" in the collections of Harris, L3elsheimer and Ziegler) and that these agreed with the present concept of the species' name. - A cf from Fairfax) Va., with genital slide, was designated as neotype and this place as new type loc. (see above). -Notiophilus semi- striatus [Lth. 1961) p, 941.
292
Nebria paZZipe.r ( 1823b : 78). No type area given ; Boston, Mass.) designated as type loc. by Lth. ( 1961, p. 76). Say mentions the two pale frontal spots and the interpretation of his name is there- fore certain. A d from Monterey) Mass.) designated as neotype. - Nebria paZZipes [Lth. I 961, p. 761.
305 Pasinzachus subsulcat,us ( 1823b : 19). Type areas "Georgia and Florida.''
Dej. (1826, p. 471) mentions that he has received this species from Say and in MNP is a specimen (sex not determined) with one of the characteristic green Dej. labels : "subsulcatus Say) in Amel-. bor. - D. Say." It agrees with Say's description, except that the "obsolete rudiments of punctures)' on the elytra are virtua~ly absent.
I have designated this specimen as lectotype. The species is musually variable, as described by Leng ( 1915, p. 566) and Ban- ninger (1950, pp. 495, 502). The lectotype belongs to what the latter author regards as the typical form: the inner el~tral intervals are clearly indicated, nrs. 3 & 5 broader and more convex, It agrees fairly well with ex. nr. 4 in coll. Lec., whereas his exx. nrs. 1-3 belong to what Banninger (LC., p. 495)) with some doubt, regards as sbsp. subnitens Csy. The lectotype also agrees with I ex., Sanford, Fla. (h4CZ)) except that, in this) the costae and punctures of elytra are better developed. Florida is herewith designated as type area. - Pasimachus xu bsuZcatus [Banninger I 950, pp. 495, 5021. 331 CZivina gZobuZosa (1823b: 23). No type area given. It is a Dyschirius belonging to a very difficult group and Say's description cannot be interpreted. The name is here applied to the commonest and most widespread member of the group) as described by Lth. (1961)



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334 Psyche [September
p. 154). A macropterous Q from Arlington, Mas., herewith desig- nated as neotype and the place as type loc. ("Boston area" already designated by Lth,, 1.c.). - Dyschirius globulosus [Lth. 1961, p. 154-1-
339 CZivina sphaericoZZis ( I 823b : 23). No type area given ; Rum- ney, N. H., designated as type loc. by Lth. (1961, p. 145). Say's description cannot be interpreted. His name is used here according to general practice, as expressed in coll. Lec. (MCZ). A Q from Rumney, N. H. designated as neotype. - Dyschirius sp haericollis [Lth. 1961, p. 1451.
346 CZivina paZZipennis ( 1823b: 24). Type loc. Egg Harbour, N. J., herewith designated ; also Virginia & Florida mentioned. Though Say's description does not exclude seZZatus Lec., it seems per- missible to use his name according to general practice, that is, for the species without dorsal puncture on the elytra. A from Anglesea, N. J., designated as neotj~pe. - Dyschirius pallipennis [Lth. 1961, P. 1421.
365
CZivina pallida ( I 823b : 22). Type loc. Chinquoteague Island, Va., "under yellow pine bark." Mr. T. F. Hlavac, who is revising this group of CZivina, has preserved Say's name for the species char- acterized by slender pro-tibiae with short lateral teeth, thus separated e. g. from rubicunda Lec. C. rufescens Dej. has been regarded as a probable synonym of paZZida (Lec., 1859b, p. 452) and specimen nr. I in coll. Lec. (MCZ), labeled "C* rufescens Dej. ? pallida Say," belongs to paZZida Say as defined by Hlavac. Since no authentic specimen of rufescens Dej. is present in the Paris Museum (Lth., I 955a, p. I 3), a correct application of the name may be difficult. AS neotype, Hlavac has designated a 3 from South Carolina (~011. Liebeck). - CZivina paZZida [as characterized above]. 383
CZivina ZineoZata ( 1823b: 22). No type area given; Alle- gheny, Penn., designated as type loc. by Lth. ( 1961, p. 166). Say's description could as well be applied to suZcifrons Putz. ( 1846) which, by Lec. ( 185gb, p. 45 I ) and also by Putzeys himself ( 1866, p. 223), was regarded a. synonym (see Lth., 1. c.). The interpretation here made is in accordance with that of the coll. Lec. (MCZ). A d (with genitalia dissected) from Allegheny, Penn., designated as neo- type. - Schizogenius heolatus [Lth. 1961, p. 661. 395 CZivina viridis ( I 823b : 21 ) . No type area given. It is almost certain that Sa,y described viridis auct. He would hardly have over- looked the coarse puncturec on the disc of prothorax in puncticolZis Dej.; and the remaining three N. American species of Ardistomis -
have striate, not seriately punctate elytra. A c? (genital slide made)



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19691 Lindroth - Ground Beetles 335
from Philadelphia Neck, Penn.) designated as neotype and the place as