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THE CARNEGIE MUSEUM OF NATURAL HISTORY
4400 FORBES AVENUE • PITTSBURGH, PENNSYLVANIA 15213
VOLUME 71
26 FEBRUARY 2002
NUMBER 1
CONTENTS
ARTICLES
New specimens of picromomyids (Plesiadapiformes, Primates) with descrip-
tion of a new species of Alveojunctus
Mary T. Silcox, Kenneth D. Rose, and Stephen L. Walsh 1
Review of the Neotropical genus Bredinia (Trichoptera: Hydroptilidae:
Stactobiini)
.... Steven C. Harris, Ralph W. Holzenthal, and Oliver S. Flint, Jr. 13
New species of microcaddisflies (Trichoptera: Hydroptilidae) from northern
Florida Steven C. Harris 47
FROM THE ARCHIVES AND COLLECTIONS
A Miocene bone bed from Agate Fossil Beds National Monument
W. Orr Goehring and Mary R. Dawson 59
^^^^hson/Tat
A1 1 3
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ANNALS OF CARNEGIE MUSEUM
VoL. 71, Number 1, Pp. 1-11
26 February 2002
NEW SPECIMENS OF PICROMOMYIDS
(PLESIADAPIFORMES, PRIMATES) WITH DESCRIPTION OF A NEW
SPECIES OF ALVEOJUNCTUS
Mary T Silcox^
Kenneth D. Rose^
Research Associate, Section of Vertebrate Paleontology
Stephen L. Walsh^
Abstract
Two new specimens attributable to the plesiadapiform family Picromomyidae are documented. One,
a lower molar, represents a new species here named Alveojimctus bowni. This species is the largest
and most derived picromomyid currently known, with an extremely simplified molar morphology. The
second new specimen is a fragmentary lower jaw attributed to Picromomys petersonorum; the speci-
men includes the first complete lower central incisor known for a picromomyid. This tooth shows
none of the derived features seen in microsyopid IjS, providing no support for a tie to this family.
Instead, it is most similar to I, in Tinimomys graybulliensis, supporting a close relationship between
Picromomyidae and Micromomyidae.
Key Words; Incisors, Picromomyidae, Micromomyidae, Plesiadapiformes, Primates, Teeth
Introduction
In 1996 Rose and Bown recognized a new family of plesiadapiform based on
the discovery of associated right and left dentaries representing a new species,
Picromomys petersonorum. This species is characterized by an enlarged P4 with
a wide, oddly flattened talonid and unusual, cusp-like, mesiobuccal expansions
on the trigonids of Mj.2. P. petersonorum is also exceptional as one of the smallest
primate-like animals ever described. The recent description of the “smallest pri-
mate” by Gebo et al. (2000) does not discuss Picromomys, presumably because
the authors do not consider plesiadapiforms to be primates. A comprehensive
phylogenetic analysis of dental, cranial, and postcranial features of a wide diver-
sity of plesiadapiforms, euprimates, and other archontans indicates that plesia-
dapiforms are most appropriately considered primates (Silcox, 2001). The body
mass estimates of P. petersonorum and the smallest Shanghuang primate dis-
cussed by Gebo et al. overlap, so it is unclear which species is actually the
smallest described primate.
Rose and Bown (1996) also included a previously described species of plesia-
dapiform, Alveojunctus minutus Bown, 1982, in the Picromomyidae on the basis
of similarities in the morphology of the enlarged P4 and the shared presence of
’ Department of Anthropology, Penn State University, 409 Carpenter Bldg., University Park, PA 16802.
^ Department of Cell Biology and Anatomy, Johns Hopkins School of Medicine, 725 N. Wolfe St.,
Baltimore, MD 21205.
^ Department of Paleontology, San Diego Natural History Museum, P.O. Box 121390, San Diego, CA
92112.
Submitted 13 April 2001.
1
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Annals of Carnegie Museum
VOL. 71
low-crowned lower molars with shallow or absent hypoflexids, strongly mesially
canted trigonids, twinned entoconid and hypoconulid separated by an entoconid
notch, and weak or absent mesiobuccal cingulids. Hooker et ah (1999) challenged
the link between Picromomys and Alveojunctus, suggesting that the relationships
of the latter “will remain obscure until more material is found (p. 393).” These
authors did not account for the close similarity in P4 morphology seen in Picro-
momys and Alveojunctus, or for the molar characteristics shared by these two
genera. Although further material of Alveojunctus may suggest a different reso-
lution, the current evidence strongly supports the inclusion of both species in a
taxonomic grouping to the exclusion of any other known plesiadapiform.
Rose and Bown (1996) presented a cladistic analysis to elucidate the wider
relationships of the Picromomyidae. In spite of some general resemblances to
Picrodontidae (e.g., low-crowned molars and strongly mesially canted molar tri-
gonids), the results of the analysis suggested that these similarities are due to
convergence and that picromomyids are most closely related to either uintasori-
cine microsyopids or micromomyids. Although the authors slightly favored the
micromomyid tie, they conceded that formalizing this relationship taxonomically
was premature given the paucity of the material available at that time. Subse-
quently, Stafford and Szalay (2000) opined that the Picromomyidae were a group
of microsyopids, although this view was not supported by new specimens, a
revised cladistic analysis, or a discussion of any character evidence.
This paper documents new specimens of the Picromomyidae, including the
description of a new species of Alveojunctus based on a single lower molar from
the Uintan of California. We recognize that naming a new species based on a
single tooth is ordinarily an undesirable practice. In this instance, however, we
deem it appropriate because of the extreme scarcity of this group, the diagnostic
differences between the new specimen and known picromomyids, and the tem-
poral and geographic separation between this specimen and the rest of the known
material of the Picromomyidae. The other species of Alveojunctus, A. minutus, is
known from only a few isolated teeth, and its holotype is also a lower molar,
allowing the most appropriate comparisons to be made (Bown, 1982). To date,
only nine specimens that can be confidently referred to the Picromomyidae (see
below) have been discovered, seven of which are isolated teeth of Alveojunctus.
This material is known from Wasatchian, Bridgerian, and Uintan deposits, indi-
cating that the family existed for at least six million years. In light of the derived
morphology exhibited by known picromomyids, compared to that of their likely
closest relatives, the family probably originated significantly earlier. Previous to
this report the geographic range of the Picromomyidae was limited to Wyoming —
with the description here of a specimen from California the geographic range of
this family is significantly increased. All of these indications show that the family
is so rare that there can be no guarantee that any better specimens of the new
species will ever be recovered.
Dental characteristics of known picromomyids are suggestive of a diet similar
to that of the tiny extant marsupial feathertail glider, Acrobates (Rose and Bown,
1996), which feeds on a diet of insects, nectar, pollen, and possibly tree exudates
(Fleay, 1947; Woodside, 1995; Nowak, 1999), although further study is required
to rule out other possibilities (e.g., a diet of soft fruits).
A new specimen of Picromomys petersonorum that includes the first complete
lower central incisor is also documented here. The morphology of this tooth pro-
vides new evidence on the wider relationships of the family.
2002
SiLCOX ET AL. — New Picromomyids
3
Abbreviations used in text are as follows: FMNH, Field Museum of Natural
History, Chicago, Illinois; SDSNH, San Diego Natural History Museum, San
Diego, California; USGS, United States Geological Survey, Denver, Colorado;
USNM, United States National Museum of Natural History, Smithsonian Insti-
tution, Washington D.C.
Systematic Paleontology
Order Primates Linnaeus, 1758
Suborder Plesiadapiformes Simons and Tattersall, in Simons, 1972
Family Picromomyidae Rose and Down, 1996
Genus Alveojunctus Bown, 1982
Alveojunctus bowni, new species
(Fig. lA)
Holotype. — SDSNH 31788, right Mj or M2. Only known specimen; collected
by M. A. Boeder and S. L. Walsh in 1986.
Locality. — San Diego Society of Natural History locality 3373, early Uintan,
upper tongue of the Friars Formation, Carmel Mountain Ranch housing devel-
opment, Unit 15, San Diego Co., California.
Diagnosis. — M, ^^2 approximately 25-30% larger in linear dimensions than Mj oj.2
in Alveojunctus minutus. Lower molar paracristid less well defined than in A.
minutus. Strong, straight valley present between the protoconid and paraconid/
metaconid, missing in A. minutus. Contrasts with Picromomys petersonorum (but
not A. minutus) in the lack of cingulids on the lower molar, the buccal cristid
obliqua resulting in the absence of a hypoflexid, the poor definition of the buccal
cusps, and the absence of a protocristid and postvallid.
Etymology. — For Thomas M. Bown, in recognition of his valuable contributions to knowledge of
Paleogene mammals generally, and particularly to the discovery and study of plesiadapiform primates
including Picromomyidae.
Description. — The single known isolated tooth cannot be definitively identified as M, or M2 (see
Discussion). The lack of a distal expansion associated with an enlarged hypoconulid, present on M3
of all plesiadapiforms except picrodontids, indicates that it is not likely to be M3. It is very low
crowned. The tooth has a straight mesial border, associated with a poorly demarcated paracristid and
a paraconid that is situated far lingually on the tooth, being positioned just mesial to the metaconid.
The paraconid and metaconid, although closely appressed, are separated by a notch, so that each cusp
is distinct. A narrow talonid notch separates the metaconid and entoconid. These cusps are closely
positioned, compressing the talonid notch, and effectively closing the talonid basin lingually (in spite
of the absence of an entocristid). The lingual cusps are better defined than the buccal cusps, although
all the cusps are low, broad, and bulbous swellings rather than sharp, pyramidal structures. The broadly
swollen bases of the lingual cusps create a scalloped border to the central basin of the tooth (see
below).
A chip of enamel is missing on the mesiobuccal margin of the protoconid, but the cusp appears to
have been broadly based and displaced mesially, so that it is almost directly buccal to the paraconid.
A faint postprotocristid extends down the distal face of the protoconid and merges with the cristid
obliqua, which is fully buccal in position. The buccal border of the tooth is convex (although this is
accentuated by the minor damage to the protoconid), so there is no sign of an indentation (hypoflexid)
distinguishing the trigonid from the talonid. The presence or absence, or precise position, of a hypo-
conulid is unclear due to slight chipping of the enamel. The lingual cant to the back of the tooth is
suggestive of a lingually positioned hypoconulid, but this cannot be confirmed. If a hypoconulid was
present it must have been small, because the damage is very limited and there is no evidence of such
a feature extending into the talonid basin.
There is no evidence of a protocristid or postvallid, making the trigonid and talonid basins contin-
uous with one another. There is a straight, distinct valley between the protoconid and paraconid/
metaconid that extends this composite basin even further mesially, to the front of the tooth. These
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Annals of Carnegie Museum
VOL. 71
Fig. 1. — A. Alveojiinctus bowni, new species, SDSNH 31788 (holotype), RM, 2^ stereophotograph.
B. Alveojunctus minutus, USGS 2005 (holotype), RM, 2^ stereophotograph. Mesial is to the top of
the figure and lingual to the left.
features make the crown of the tooth approximate a single, large, central basin surrounded on the
periphery by low cusps and crests.
Measurements (mm) of the Holotype. — M, 2 maximum width = 1.60 mm,
maximum length = 2.25 mm. Measured with an ocular micrometer to the nearest
0.05 mm.
Discussion. — SDSNH 31788 was tentatively assigned by Walsh (1996:85) to
the ?Palaeanodonta. Its revised identihcation here as pertaining to a new species
2002
SiLCOX ET AL. — New Picromomyids
5
of Alveojunctus thus eliminates any record of palaeanodonts from the Uintan of
southern California.
SDSNH Loc. 3373 has yielded a diverse assemblage of small mammals of
early Uintan age. It was collected by screen-washing approximately 5000 kg of
bulk matrix from a 1.2 m thick, caliche nodule-bearing, olive greenish gray silt-
stone paleosol. Other primate taxa represented at this locality include Omomys
sp. cf. O. carteri, Washakius woodringi, and Uintasorex montezumicus (see Lil-
legraven, 1980; Honey, 1990; Walsh, 1996). The stratigraphic and geographic
position of SDSNH Loc. 3373 was shown schematically by Walsh et al. (1996:
fig. 3).
The identification of SDSNH 31788 as Mj or M2 is complicated by uncertainty
surrounding the identification of the holotype of Alveojunctus minutus, USGS
2005, This tooth was originally identified as an Mj (Bown, 1982), presumably on
the basis of the triangular trigonid that is relatively narrow compared to the tal-
onid. M2S in plesiadapiforms tend to have trigonid and talonid regions that are
more nearly equal in width. Also, in M2 the trigonid is generally less triangular
than in Mj as a result of its mesiodistal compression, a flatter mesial border, and
a lingually positioned paraconid. These are also distinctions between USGS 2005
and the new specimen. In USGS 2005 there is a shallow hypoflexid, distinguishing
the trigonid from the talonid (see Fig. IB), that is missing in SDSNH 31788 (see
Fig. lA). Also, the flatter mesial border of SDSNH 31788 and more fully lingual
position of the paraconid relative to USGS 2005 are consistent with differences
between tooth positions (although there are some exceptions to these generalities,
as seen in the lower molars of Purgatorius janisae and Palenochtha minor). Based
on this reasoning it seems probable that USGS 2005 is an Mj and SDSNH 31788
an M2.
Rose and Bown (1996) suggested, however, that USGS 2005 is more likely
M2, based on its similarity to that tooth in Picromomys petersonorum. That taxon
has a trigonid and talonid of Mj that are subequal in width, whereas on M2 the
trigonid is clearly narrower than the talonid. Picromomys also has a relatively
longer trigonid on M, than on M2. In both of these features USGS 2005 is more
similar to M2 of Picromomys than to M,. The straighter mesial border and more
nearly equal width of the trigonid and talonid in SDSNH 31788 may make this
more plausibly an Mj, based on the comparison with Picromomys. It is not clear,
however, whether the unusual pattern of differences between M, and M2 in Pi-
cromomys stems solely from the different expression of the mesiobuccal accessory
cusp in this taxon. Particularly, the greater development of this cusp on Mj, rel-
ative to M2, makes the mesial border of the tooth appear more squared off, and
adds to the length and width of the trigonid. Since this cusp is missing in Alveo-
junctus it might be expected to show the more typical plesiadapiform pattern of
differences between Mj and M2. For these reasons, the attribution of the tooth is
left open until further material becomes available.
This discussion may prompt the criticism that the differences between USGS
2005 and SDSNH 31788 can be attributed to tooth position alone. The pronounced
size disparity between the specimens (we measured USGS 2005 as 1.80 mm long
and 1.20 mm wide; SDSNH 31788 is therefore 25% longer and 33% wider than
USGS 2005) argues against this. The size difference substantially exceeds that
generally seen between tooth positions in plesiadapiforms — in Picromomys pe-
tersonorum, for example, Mj is only 10% longer than M2, and M2 is slightly
wider (7%) than M,. This dental size disparity also translates into a significant
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Annals of Carnegie Museum
VOL. 71
difference in body size. If it is assumed that both molars are Mj, body mass
estimates for Alveojunctus minutus range from 50-111 g (Conroy, 1987: all pri-
mate or prosimian regressions = 50 g; Gingerich et aL, 1982: primate regression
= 1 1 1 g), whereas the range of estimates for Alveojunctus bowni suggests it may
have been twice the mass of A. minutus (Conroy, 1987: all primate regression ==
124 g, prosimian regression = 114 g; Gingerich et ah, 1982: primate regression
= 238 g). These estimates do not differ significantly if one of the specimens is
an M2 rather than an M^.
Additionally, the presence of the deep valley between the protoconid and para-
conid/metaconid in SDSNH 31788 (missing in USGS 2005), argues against the
view that both of these teeth belong to the same species. The presence or absence
of a valley between the protoconid and paraconid/metaconid is not a difference
that is seen between tooth positions, and this feature is missing in all related
forms. The differences between USGS 2005 and SDSNH 31788 seem to be com-
prehensible as a further expansion, in the latter, of the broad central basin that is
characteristic of Alveojunctus. In A. minutus this feature is formed because the
protocristid and postvallid are absent, allowing continuity between the trigonid
and talonid basins. In A. bowni, in addition to these features, there is a further
expansion forward created by the strong valley between the protoconid and para-
conid/metaconid. The reduction in the paracristid allows this valley to extend to
the front of the tooth. The mesial displacement of the protoconid in A. bowni
relative to A. minutus is probably also related, since this broadens the part of the
trigonid basin that is directly continuous with the talonid basin. In all, the differ-
ences between the two species can be explained as an accentuation of the basic
adaptive pattern that characterizes Alveojunctus, in which the molars became al-
most flat basins surrounded by weak bulbous cusps and low crests. The geolog-
ically younger age of A. bowni is also consistent with this view.
Alveojunctus bowni extends both the known temporal and geographic range of
the Picromomyidae. To date, the genus Alveojunctus is definitively known only
from middle Bridgerian and early Uintan deposits. Bown (1982) suggested that
FMNH PM28689 might belong to A. minutus. This tooth, from the Cathedral
Bluff Tongue (early Bridgerian), was attributed to Niptomomys sp. by West and
Dawson (1973). West and Dawson identified the tooth as a right M3, but the molar
illustrated is a left M3. Unfortunately, the specimen could not be located for study
(and may be lost). The larger size of this tooth, relative to A. minutus, might
suggest that the specimen could belong to A. bowni. FMNH PM28689 is unusual
in lacking a paraconid, a feature that is present in both SDSNH 31788 and USGS
2005. The paraconid is absent on M3 of paromomyid plesiadapiforms, and it is
indistinct in Niptomomys doreenae (see Rose et ak, 1993:fig. 1). In Picromomys
and micromomyids known from serially associated dentitions (Tinimomys gray-
bulliensis [Rose et aL, 1993:fig. 2] and Micromomys fremdi [Fox, 1984:fig. 1]),
however, a paraconid is clearly demarcated on M3. Since we consid^i Alveojunctus
to be more closely related to Picromomys and micromomyids than to Niptomomys
or paromomyids, an indistinct M3 paraconid would not be expected as a feature
of the genus. These indications suggest that this tooth may be attributable to a
taxon that is more closely related to Niptomomys (as originally indicated by West
and Dawson) than to Alveojunctus.
Stucky (1982, 1984) referred an additional specimen to Alveojunctus. This iso-
lated tooth, a right M], (UCM 44681) was not available for study. Stucky (1982)
notes that this tooth has more clearly delineated cusps and a higher trigonid than
2002
SiLCOX ET AL. — New Picromomyids
7
Alveojunctus minutus. In light of the fact that Alveojunctus bowni differs from A.
minutus in precisely the opposite ways (i.e., it has a less well defined cusps and
a lower trigonid), it seems unlikely that this specimen could be referred to the
new species.
Picromomys petersonorum Rose and Bown, 1996
(Fig. 2)
Discussion. — A fragmentary left dentary with a nearly complete Ij, P4 talonid,
Ml, and alveoli for M2.3 (USNM 494979; Fig. 2) was collected in 1996 by M. T.
Silcox from Dorsey Quarry (U.S.G.S. locality D-2035Q; 397m above the base of
the Willwood Formation; see Bown et ah, 1994, and Silcox and Rose, 2001) in
the Bighorn Basin of northwestern Wyoming. Although the Ii has a break between
the root and the rest of the tooth, and the crown is now displaced relative to the
dentary, it was found in continuity with the dentary and is clearly part of the
same specimen. The morphology of the P4 talonid and Mj matches that of the
holotype of Picromomys petersonorum.
The root of Ij is enlarged, laterally compressed, and is in a nearly horizontal
position in the mandible, as in the holotype (Rose and Bown, 1996). It bears a
distinct crest close to the lateral margin of the dorsal surface. Near the base of
the crown this dorsal crest crosses from its lateral position to the medial side of
the crown, where it descends onto the medial surface and runs distally to a point
approximately halfway from the base of the tooth, before fading away. Near the
point of this cross-over, another crest begins at the dorso-lateral extent of the
crown. The two crests present on the crown of the tooth define a surface that
exists in two planes, facing dorsally and medially.
Rose and Bown (1996) noted some basic differences between the morphology
of the lower central incisor preserved in the holotype of Picromomys petersono-
rum and that seen in microsyopids. Although the holotype preserves only a part
of the root of Ij, it is clear that Ij lacks the sharp dorsal border and basal expansion
characteristic of microsyopid incisors. Rose and Bown pointed out that the dis-
tinctive morphology of microsyopid incisors has resulted from a medial rotation
around the long axis of the tooth, so that the homolog of the lateral border of Ij
in most plesiadapiforms forms a sharp dorsal crest in microsyopids. Rose and
Bown (1996) argued that such a rotation had not occurred in Ii of picromomyids.
It remained possible, however, that when the rest of the crown became known it
would be more microsyopid-like, or else represent a recognizable intermediate
stage between the morphology observed in other plesiadapiforms and microsy-
opids.
The morphology of USNM 494979 does not support this view. The dorsal crest
noted by Rose and Bown (1996) on the root is continuous with the medial crest
on the crown of the Ij in USNM 494979. In microsyopids, however, the dorsal
crest is homologous with the lateral crest on the crowns of more typical plesia-
dapiforms (e.g., Plesiolestes problematicus; see Bown and Gingerich, 1973). This
indicates that the Ii morphology in Picromomys is not an intermediate on the path
to the microsyopid condition. Furthermore, the shape of this incisor differs mark-
edly from that observed in microsyopids in lacking a basal expansion of the
crown. Also, the portion of the crown that is delimited by crests is not flat, as in
microsyopids, but incorporates two distinct planes. In all, the picromomyid I,
morphology is less consistent with a relationship to microsyopids than that ob-
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Annals of Carnegie Museum
VOL. 71
Fig. 2. — Picromomys petersonorum, USNM 494979. Incomplete left dentary containing I,, talonid of P4, M,, and alveoli for M2.3 in oblique buccal view
(below), with occlusal view of Ij (above). Scale bar = 1 mm.
2002
SiLCOX ET AL. — New Picromomyids
9
served in some palaechthonids. Even some primitive plesiadapoids have a more
closely comparable incisor morphology (e.g., Elphidotarsius sp. cf. florencae, in
which I, is broad at the base but not rotated medially; see Rose, 1975), in spite
of the fact that plesiadapoids are not very closely related to microsyopids (Silcox,
2001).
The Ij of Picromomys is similar to this tooth in a number of plesiadapiforms,
including Saxonella, Picrodus, paromomyids, micromomyids, and possibly PaP
enochtha minor (see Gunnell, 1989:fig. 5), in being slender, laterally compressed,
and elongate. The evidence available for Purgatorius is also consistent with the
presence of this morphology, although the tooth itself is still unknown for this
basal plesiadapiform (Silcox, 2001). The widespread distribution of this basic
morphology suggests that it may be primitive for plesiadapiforms (Silcox, 2001).
The lower incisor of Picromomys is particularly similar to that of the only mi-
cromomyid for which this tooth has been published, Tinimomys graybulliensis
(i.e., USNM 425583, Beard and Houde, 1989). As in Picromomys, Ij in Tini-
momys shows a similar reorientation of a dorsolateral crest in the root region to
a medial position on the crown. This change in direction in the dorsal crest is not
present in any of the other plesiadapiforms examined. When added as a character
to the dataset published by Rose and Bown (1996) (see Appendix), an exhaustive
search using PAUP* 4.0p6 (Swofford, 2001) resulted in a single most parsimo-
nious tree corresponding to their Tree B (length = 73 steps. Cl = .49, RI = .537,
statistics from PAUP*; see Rose and Bown, 1996:fig. 5). This tree supports a
clade including Picromomys, Alveojunctus, and Tinimomys, to the exclusion of
Niptomomys, and portrays picromomyids as most closely related to micromo-
myids.
Measurements (mm) of USNM 494979. — P4 width = 0.80 mm; Mj trigonid
width = 0.70 mm, Mj talonid width = 0.80 mm, M, length = 1.20 mm. Measured
with an ocular micrometer to the nearest 0.05 mm.
Conclusions
This paper documents the eighth and ninth confidently attributed specimens of
the Picromomyidae. The type specimen of the new species Alveojunctus bowni
from the early Uintan of California extends the temporal and geographic range
of the family. This specimen demonstrates an even more extreme version of the
basic pattern of picromomyid dental structure, leading to an extraordinarily flat
and simplified lower molar morphology. This is consistent with an increasingly
specialized diet. The second recognized specimen of Picromomys petersonorum
provides new information on the incisor morphology and suggests a stronger tie
to the Micromomyidae than to the Microsyopidae. This may imply that Micro-
momyidae is a paraphyletic group (Hooker et ah, 1999). Specifically, picromo-
myids and Tinimomys may be sister taxa to the exclusion of Micromomys (as
indicated in Tree B of Rose and Bown, 1996:fig. 5). In light of this, it may
eventually be appropriate to make Picromomyidae a subfamily of the Micromo-
myidae. Making definitive conclusions about the relationships of micromomyids
and picromomyids is hampered, however, by the very limited information avail-
able for most species of Micromomys (Beard and Houde, 1989), and the lack of
specimens from the upper dentition of Picromomyidae. Consequently, we prefer
to maintain Picromomyidae and Micromomyidae as distinct families, pending
further discoveries or additional analyses.
10
Annals of Carnegie Museum
VOL. 71
Acknowledgments
Figure 1 was taken at the Johns Hopkins Microscope Facility with supplies generously provided by
M. F. Teaford. Figure 2 was expertly prepared by Elaine Kasmer. Fieldwork in the Bighorn Basin of
Wyoming was carried out under BLM permits and supported by NSF grant IBN-94 19776 to KDR.
Comparative studies of plesiadapiforms by MTS were supported by the Paleobiological Fund, the
Wenner-Gren Foundation for Anthropological Research, Sigma Xi, and NSF Doctoral Dissertation
Improvement Grant 9815884. Collection of SDSNH 31788 was made possible by Carmel Mountain
Ranch. This paper was significantly improved by the comments of three anonymous reviewers.
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Rose, K. D., and T. M. Bown. 1996. A new plesiadapiform (Mammalia: Plesiadapiformes) from the
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Plesiadapiformes. Annals of Carnegie Museum, 62:351-361.
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SiLCOX ET AL. — New Picromomyids
1 1
Early Eocene of the Bighorn Basin, Wyoming. Pp. 131-164, in Eocene Biodiversity: Unusual
Occurrences and Rarely Sampled Habitats (G. E Gunnell, ed.). Plenum Press, New York, New
York.
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and biostratigraphy of the upper part of the Wind River Formation, Northeastern Wind River
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Sinauer Associates, Inc, Sunderland, Massachusetts.
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eds.). Cambridge University Press, Cambridge, United Kingdom.
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Smithsonian Institution Press, Washington D. C.
Appendix
New character added to cladistic analysis from Rose and Bown, 1996. 0 = lateral crest (or its
homologue) in the root region of I, runs straight to the tip of the tooth, without crossing medially;
1 = A dorsolateral crest present in the root region of I; becomes re-oriented to lie on the medial side
of the crown.
Purgatorius 1
Palaechthon ?
Palenochtha ?
Picromomys 1
Alveojunctus ?
Paromomys ?
Ignacius 0
Navajovius 0
Micromomys ?
Tinimomys 1
Niptomomys 0
Picrodus 0
TT— 7v’
■’M
x'r W
'"fi!
V,
a ■ ''wM'’-"' /V. "•JiSL '.' 'sk-v' - "^i . '■ ' A
■ '■*">'.• t:' #>.'4;* '-Vi; ■■^■' •,.>,, . ■
t##.. v-i^3sw«ji' *' ;' .•jj.'.dtL %■•;♦’'*' ”*
'— * '**.. .-■■.fjif •/ t# . -j
ANNALS OF CARNEGIE MUSEUM
VoL. 71, Number 1, Pp. 13-45
26 February 2002
REVIEW OE THE NEOTROPICAL GENUS BREDINIA (TRICHOPTERA:
HYDROPTILIDAE: STACTOBIINI)
Steven C. Harris’
Research Associate, Section of Invertebrate Zoology
Ralph W Holzenthal^
Oliver S. Elint, Jr.^
Abstract
The Neotropical genus Bredinia is reviewed, with descriptions and figures provided for the 16
species presently known, including 13 new species: B. guanacasteca n.sp., B. emarginata n.sp., and
B. selva n.sp. from Costa Rica; B. davenporti n.sp. and B. pilcopata n.sp. from Peru; B. manabiensis
n.sp. and B. spangleri n.sp. from Ecuador; B. alza n.sp. from Paraguay; B. mexicana n.sp. from
Mexico; B. sucrensis n.sp. and B. zulia n.sp. from Venezuela; B. espinosa n.sp. from Brazil, Ecuador,
and Venezuela, and B. venezuelensis n.sp. from Venezuela, Peru, and Ecuador. New records and
illustrations are provided for B. costaricensis (Elint), B. dominicensis Elint, and B. appendiculata Elint
and Sykora. A key is provided to separate the males and the associated females.
BCey Words: Trichoptera, Hydroptilidae, Neotropics, Bredinia, new species, key, systematics
Introduction
This paper on the genus Bredinia represents an addition to a series initiated to
describe the microcaddisflies of Costa Rica. However, to place the systematics of
the group under review in context, we have expanded the scope of this study to
encompass the entire Neotropical region. The genus Bredinia was previously
thought to be small in terms of species and restricted in distribution (Elint et al.,
1999fl), but these ideas are revised with this paper. Herein we describe 13 new
species from the Neotropical region and provide illustrations and redescriptions
of the three species previously described, and a key to the species of males and
known females.
The genus Bredinia was erected by Flint (1968) for B. dominicensis, which
was originally collected near the mouth of the Hodges River in Dominica. A
second species, B. appendiculata Flint and Sykora from Grenada, was added in
1993. More recently, a third species, B. costaricensis (Flint), was transferred from
Neotrichia into Bredinia (Flint et ak, \999b). Material examined in this work
comes largely from collections at the University of Minnesota and the National
Museum of Natural History.
Bredinia is assigned to the tribe Stactobiini, which as currently defined contains
ten genera, most of which occur in the Old World. In the New World Stactobiini,
there are three genera presently known: Stactobiella Martynov in the Nearctic
‘Department of Biology, Clarion University, Clarion, PA 16214.
^Department of Entomology, University of Minnesota, St. Paul, MN 55108.
^Department of Entomology, National Museum of Natural History, Smithsonian Institution, Washing-
ton, D.C. 20560.
Submitted 17 December 2001.
13
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Annals of Carnegie Museum
VOL. 71
region, and Bredinia and Flintiella Angrisano in the Neotropics (Flint et ah,
1999a). Two additional genera from the Neotropics are described and a key pro-
vided for the New World Stactobiini in Harris et al. (In Press).
Types of species described in this paper, as indicated by acronyms in the text,
are deposited in the collections of the National Museum of Natural History,
Smithsonian Institution, Washington (NMNH), University of Minnesota Insect
Collection, St. Paul (UMSP), Cornell University, Ithaca (CU), Universidad Na-
cional Autonoma de Mexico, Mexico City (UNAM), Florida State Collection of
Arthropods, Gainesville (FSCA), Academy of Natural Sciences, Philadelphia
(ANSP), Carnegie Museum of Natural History, Pittsburgh (CMNH), and Instituto
Nacional de Biodiversidad, Heredia, Costa Rica (INBIO). Terminology follows
that of Marshall (1979). Length is measured from the tip of the head to the end
of the wings and is given as a range within a series of specimens.
Systematic Entomology
Bredinia Flint
(Fig. 1)
Bredinia Flint, 1968:50. Type species. Bredinia doniinicensis Flint, 1968, original designation.
Diagnosis. — Adults of Bredinia are minute (1—2 mm in length), and typically
gray in color, with silvery bands of hairs on the forewings. There are 3 ocelli on
the head (Fig. lA) with simple antenna having 17-19 segments. The mesoscu-
tellum is divided by a transverse suture; the metascutellum is narrow and rectan-
guloid, and nearly as wide as scutum. Legs have a tibial spur formula of 0, 2, 4
(Fig. IB). Wings are narrow, with reduced venation (Fig. 1C). The tibial spur
formula and the presence of ocelli serve to separate Bredinia from other New
World genera. Unique to males in this genus is the presence of an elongate,
thickened seta from the posterior of the inferior appendages and elongate apo-
demes from the anterior of abdominal segment IX. Females of most of the Neo-
tropical stactobiines are unknown, so it is premature at this point to speculate on
distinguishing characters. Larvae have been described by Angrisano (In Press),
but as with female stactobiines, too few species are known in the larval stage to
permit diagnoses.
Male. — Abdominal segment VII annular, without a ventromesal process. Seg-
ment VIII annular and complete dorsally, often with anterolateral margins devel-
oped as thin apodemes. Segment IX usually reduced ventrally with anterolateral
margins developed as apodemes which vary in length. Inferior appendages each
with heavy, elongate seta originating on posteroventral margin. Subgenital plate
shelflike. Phallus tubular with apex distinctive for the species.
Female. — Abdominal segment VII annular. Segment VIII rectangular, ring of
setae on posterior margin, pair of mesal and lateral apodemes. Segment IX narrow
and rectangular, pair of lateral apodemes connecting anteriorly with apodemes of
VIII. Segment X short and conical, bearing pair of apical papillae. Bursa copu-
latrix round, rectangular or triangular in ventral aspect, with or without posterior
extensions; anteriorly with pair of elongate lateral processes attached to and par-
alleling narrow mesal process.
2002
Harris et al. — Review of the Neotropical Genus Bredinia
15
Fig. 1. — Bredinia costaricensis (Flint), adult: A. Head and thorax, dorsal. B. Fore, mid and hindleg.
C. Fore and hindwing.
Bredinia dominicensis Flint
(Figs. 2, 3)
Bredinia dominicensis Flint, 1968:51.
Diagnosis. — -Bredinia dominicensis, along v^ith B. selva n. sp., and B. guana-
casteca n. sp., is recognized by the elongate rods extending from the margins of
segment IX. Bredinia dominicensis is most similar to B. selva, but is separated
by the acute ventral margin of the inferior appendages in lateral aspect, and the
structure of the phallus apex, which has contiguous inner processes. We have
records of B. dominicensis from Costa Rica, Dominica, Ecuador, Panama, and
Trinidad.
Male. — Length 1.5-1. 6 mm. Antenna with 18 segments. Silvery gray in color,
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Annals of Carnegie Museum
VOL. 71
2
Fig. 2. — Bredinia dominicensis Flint, male genitalia, redrawn from paratype: A. Lateral. B. Ventral C.
Dorsal. D. Phallus, dorsal. E. Phallus, lateral.
Fig. 3. — Bredinia dominicensis Flint, female genitalia. A. Terminal abdominal segments, ventral. B.
Bursa copulatrix, ventral.
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Harris et al. — Review of the Neotropical Genus Bredinia
17
forewing with bands of white hair apically and basally. Genitalia as in Fig. 2.
Segment VIII annular. Segment IX reduced to narrow bridge ventrally; dorsally
round and fused with segment X, anterior margins extended anteriorly as elongate
rods which pass through segment VIIL Segment X truncate and membranous.
Inferior appendages in lateral view each narrowing to acute apex postero ventrally;
in ventral view square. Subgenital plate narrow, shelflike in lateral view; triangular
in ventral view. Phallus tubular, apex flattened and tridentate, ejaculatory duct
short and triangular.
Female. — Length 1.7 mm. Antenna with 18 segments. Color and features of
the head, thorax and wings as in male. Genitalia as in Fig. 3. Segment VII annular.
Segment VIII rectangular, ring of setae on posterior margin, pair of mesal apo-
demes extending into segment VII, second pair of elongate lateral apodemes orig-
inating at anterolateral margins and extending into segment VII. Segment IX
narrow and rectanguloid, with pair of lateral apodemes connected anteriorly with
mesal apodemes of VIIL Segment X short and rounded apically, bearing pair of
apical papillae. Bursa copulatrix rectanguloid with short posterior extension; an-
terior with wide lateral processes turned outward at apices, mesal process divided
apically.
Material Examined. — COSTA RICA. Puntarenas: Quebrada Potrero near Potrero Grande, 5 July
1992, T. Shepard, 1 3 (CMNH).
DOMINICA. Hodges River, at mouth, swamp forest, 27 February 1965, W. Wirth, 2 d, 2 $
paratypes (NMNH).
ECUADOR. Esmeraldas: La Union, 3 February 1979, J. Anderson, 16 d (4 d NMNH, 4dUMSP,
4dCMNH). Pichincha: Santo Domingo de los Colorados (29 km W), 6 May 1975, R Spangler, 1 d
(NMNH).
PANAMA. Darien: Rio Tuira at Rio Pucura, 16-17 February 1983, J. Louton, 1 d (NMNH).
TRINIDAD. Tacariqua River, Caura Recreation Area, 10°43' N, 61°17' W, 22 June 1993, O. Flint,
N. Adams, 1 d, 4 $ (NMNH).
Bredinia guanacasteca, new species
(Fig. 4)
Diagnosis. — Bredinia guanacasteca is most similar to B. dominicensis and B.
selva n.sp., each having elongate anterior rods from segment IX. The new species
is distinguished by the truncate apical margin of the inferior appendages, seen in
lateral aspect, and the structure of phallus apex, which terminates in a pair of
elongate inner processes. Bredinia guanacasteca is known only from the type
locality in Costa Rica.
Male. — Length 1.8 mm. Antenna with 18 segments. Brown in alcohol. Geni-
talia as in Fig. 4. Segment VIII annular, venter emarginate posteriorly. Segment
IX reduced to narrow bridge ventrally; dorsally round; anterior margins extended
anteriorly as elongate rods passing through segment VIIL Segment X elongate,
shelflike in lateral view; dorsum rectangular, slightly emarginate on posterior mar-
gin. Inferior appendages nearly square in lateral view, sinuate on posterior margin;
square in ventral view, diverging on mesal margin. Subgenital plate narrow, shelf-
like. Phallus tubular, apex with pair of narrow, elongate inner processes, lateral
processes short and contiguous, ejaculatory duct short and wishbone-shaped.
Female. — Unknown.
Type Material. — Holotype d. COSTA RICA. Guanacaste: Rio Tempisquito, ca 3 km S route 1,
10.790°N, 85.552°W, el. 75 m, 6 March 1986, Holzenthal and Fasth (NMNH).
Etymology. — Named for the Costa Rican province of Guanacaste.
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Annals of Carnegie Museum
VOL. 71
Fig. 4, — Bredinia guanacasteca new species, male genitalia: A. Lateral. B. Ventral. C. Dorsal. D.
Phallus, dorsal. E. Phallus, lateral.
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Harris et al. — Review of the Neotropical Genus Bredinia
19
D
Fig. 5. — Bredinia selva new species, male genitalia: A. Lateral. B. Ventral. C. Dorsal. D. Phallus,
dorsal. E. Phallus, lateral.
Bredinia selva, new species
(Fig. 5)
Diagnosis. — Bredinia selva is separated from the similar B. dominicensis by
the shape of the inferior appendages and phallus apex. The inferior appendage of
B. selva ends in a rounded upturned projection from the ventrololateral margin,
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Annals of Carnegie Museum
VOL. 71
whereas in B. dominicensis the ventrolateral margin is acute. The apex of the
phallus of both species is similar, but in B. selva the inner processes are strongly
divergent. Bredinia selva is known only from the type locality in Costa Rica.
Male, — Length 1.7 mm. Antenna with 18 segments. Brown in alcohol. Geni-
talia as in Fig. 5. Segment VIII bearing heavy setae ventrolaterally; venter slightly
emarginate posteriorly. Segment IX reduced to narrow bridge ventrally, dorsally
rounded and fused with segment X; anterior margins extended anteriad as elongate
rods passing through segment VIII. Segment X shelflike, narrowing distally; in
dorsal view rectangular, slightly emarginate posteriorly. Inferior appendages in
lateral view each narrowing poster© ventrally to rounded, upturned knob; in ventral
view triangular in shape with thumb-like process posteriorly, diverging along
mesal margin. Subgenital plate in lateral view rounded posteriorly, with down-
turned distal hook; in ventral view a rectangular plate. Phallus tubular, apex with
bifid lateral processes, inner processes strongly diverging, outer processes curved
and nearly as long as inner processes; ejaculatory duct narrowly rectangular.
Female. — Unknown.
Type Material — Holotype S. COSTA RICA. Heredia: Estacion Biologica La Selva, 28 April
1989, Agriculture Canada (NMNH). Paratype: COSTA RICA. Same locality as holotype, but 21-24
August 1999, D. and M. Davis, 1 S (NMNH).
Etymology. — Named for the type locality, La Selva Biological Station of the Organization for Trop-
ical Studies.
Bredinia espinosa, new species
(Figs. 6, 7)
Diagnosis, — The prominent, spinelike setae from the sides of segment VIII are
distinctive and unique for this species. Bredina espinosa is known from Brazil,
Ecuador, and Venezuela.
Male. — Length 1,4- 1.8 mm. Antenna with 18 segments. Brown in alcohol.
Genitalia as in Fig. 6. Segment VIII bearing thickened, spinelike setae laterally;
venter emarginate on posterior margin, anterior margins produced into lateral rods
extending midway into segment VIL Segment IX reduced to narrow bridge ven-
trally; dorsally rectangular, rounded apically and emarginate laterally; anterior
margins with short lateral rods. Segment X shelflike laterally; rounded lobe in
dorsal and ventral views. Inferior appendages round in lateral view; in ventral
view spatulate, diverging on mesal margin, emarginate on lateral margins. Sub-
genital plate in lateral view curving downward and narrowing posteriorly; in ven-
tral view narrowing to mesal beak. Phallus tubular, apex flattened and widened,
large lateral flaps with apex of ejaculatory duct mesal and prominent.
Female. — Length 1.8-2. 0 mm. Antenna with 18 segments. Color and features
of head, thorax and wings as in male. Genitalia as in Fig. 7. Segment VII annular.
Segment VIII elongate, ring of setae on posterior margin, pair of elongate mesal
apodemes extending through segment VII, second pair of elongate apodemes orig-
inating at anterolateral margins and extending into segment VIL Segment IX short,
emarginate on posterior margin; pair of lateral apodemes which connect anteriorly
with mesal apodemes of VIII. Segment X short and conical, bearing pair of apical
papillae. Bursa copulatrix triangular, narrowing posteriorly, pair of sclerotized
internal plates posteriorly; lateral anterior processes narrow, sinuate, and mem-
branous apically, inner process originating as oval plate, tapering anteriorly.
Type Material. — Holotype 8. ECUADOR. Los Rios: Quevedo (56 km N), Rio Paleeque Biological
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Harris et al. — Review of the Neotropical Genus Bredinia
21
Fig. 6. — Bredinia espinosa new species, male genitalia: A. Lateral. B. Ventral. C. Dorsal. D. Phallus,
dorsal. E. Phallus, lateral.
Fig. 7. — Bredinia espinosa new species, female genitalia: A. Terminal abdominal segments, ventral.
B. Bursa copulatrix, ventral.
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Annals of Carnegie Museum
VOL. 71
Station, blacklight at riverbed, el. 250 m, 28-29 July 1976, J. Cohen (NMNH). Paratypes; ECUADOR.
Same data as holotype, 13 d, 1 $ (8(3,1$ NMNH, 5c3CMNH). Pichincha; Santo Domingo, (47 km
S), 29 July 1976, J. Cohen, 2 d, 2 $ (UMSP). Cotopaxi: Quevedo (36 km NE), el. 1100 m, 21 July
1976, J. Cohen, 8 <3, 12 $ (4c3, 8$ NMNH, 4c3, 4$CMNH). Manabi: Santo Domingo (29 km SW),
Rancho Ronald, 6 September 1978, J. Anderson, 4 c3 (NMNH), same locality, but 20 July 1978, J.
Anderson, 1 $ (NMNH). Guayas: Daule, 23 December 1977, J. Anderson, 9 c3 (5c3NMNH,
4(3 UMSP).
VENEZUELA. Amazonas: San Carlos de Rfo Negro, 1°56' N, 67°03' W, 13-17 December 1984,
R. Brown, 1 (3, 1 $ (NMNH), Cerra de la Neblina, basecamp, el. 140 m, 0°50' N, 66° 10' W, 10
February 1985, blacklight in rainforest clearing near Rio Baria, W. Steiner, 1 6 (NMNH).
BRAZIL. Mato Grosso: bridge on Rio Arica, on Cuiaba to Rondonoplis road near km 391 marker,
22 April 1981, D. Wojcik, 1(3 (NMNH). Rondonia: creek 8 km S Cacaulandia, 21 November 1991,
D. Petr, 13,1$ (NMNH).
Etymology. — Derived from the Spanish, ''espina — bearing spines,” referring to the lateral setae of
abdominal segment VIII.
Bredinia appendiculata Flint and Sykora
(Figs. 8, 9)
Bredinia appendiculata Flint and Sykora, 1993:56.
Diagnosis. — Bredinia appendiculata is most similar to B. davenporti n.sp.
Unique to both are the sclerotized dorsal margins of the inferior appendages,
which form an acute process extending posteriad, and both have the apex of the
subgenital plate downturned. Bredinia appendiculata is separated from B. dav-
enporti on the basis of having the dorsal process of the inferior appendage elon-
gate, and the more hooklike subgenital plate. Originally described from Grenada,
we have also seen specimens of this species from Peru and Venezuela.
Male. — Length 1.5-1. 7 mm. Antenna with 18 segments. Brown in alcohol.
Genitalia as in Fig. 8. Segment VIII annular; venter deeply emarginate posteriorly,
anterior margins produced into short lateral rods. Segment IX reduced to narrow
bridge ventrally; dorsally truncate and fused with X; anterior margins with short
lateral rods. Segment X shelflike in lateral view; dorsum truncate, with mesal
incision posteriorly. Inferior appendages in lateral aspect each with ventral margin
crenate, dorsal margin modified into sicklelike process which narrows posteriorly
and curves ventrad; in ventral view acute posterior process curving mesad, pos-
teriorly strongly diverging mesally. Subgenital plate in lateral view narrowed pos-
teroventrally into hooklike process. Phallus tubular, apex divided into pair of
narrow lobes, with acute apices and inner margins which vary from being crenate
to spinose; ejaculatory duct short and triangular.
Female. — Length 1.5-1. 8 mm. Antenna with 18 segments. Color and features
of the head, thorax and wings as in male. Genitalia as in Fig. 9. Segment VII
annular. Segment VIII square, ring of setae on posterior margin, pair of elongate
mesal apodemes extending into segment VI, second pair of elongate apodemes
originating at anterolateral margins and extending into segment VI. Segment IX
short; pair of lateral apodemes connecting anteriorly with mesal apodemes of VIII.
Segment X short and conical, bearing pair of apical papillae. Bursa copulatrix
triangular, narrowing posteriorly, pair of membranous lobes posteriorly, oval plate
anteriorly to which pair of flipperlike processes attach; mesal process elongate
and divided apically.
Material Examined. — Holotype 3, GRENADA. Parish St. Andrews: Balthazar Estate, 1 June
1990, J. Frank, A. Thomas (FSCA).
PERU. Madre de Dios: Manu, Pakitza, 12° 7'S, 70° 58'W, el. 250 m., trail 2, first stream, malaise
trap, day collection, 14-23 September 1988, O. Flint, N. Adams, 6 3, 3 $ (43, 2$ NMNH, 23,
2002
Harris et al. — Review of the Neotropical Genus Bredinia
23
Fig. 8. — Bredinia appendiculata Flint and Sykora, male genitalia, redrawn from holotype: A. Lateral.
B. Ventral. C. Dorsal. D. Phallus, dorsal. E. Phallus, dorsal (specimen from Peru). F. Phallus, lateral.
Fig. 9. — Bredinia appendiculata Flint and Sykora, female genitalia: A. Terminal abdominal segments,
ventral. B. Bursa copulatrix, ventral.
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Annals of Carnegie Museum
VOL. 71
1 5CMNH), same locality, but night collection, 2 <5, 1 $ (NMNH), Hostel Erica, near Salvacion, 12°
53'S, 71° 14'W, 3 and 5 September 1989, J. Collins, 1 9 (NMNH).
VENEZUELA. Sucre: Rio Cocollar, 1.5 km SE Las Piedras de Cocollar, 10° 09.67 1'N, 63°
47.605 'W, el. 810 m., 7-8 April 1995, R. Holzenthal, O. Elint, 2 d, 5 9 (UMSP). Zulia: Cano
Carichuano, 3.4 km SE Carbones del Guasare, 11.002°N, 72.285°W, el 70 m, 12-13 January 1994,
Holzenthal, Cressa, Rincon, 2 1ff(', : ■’ - '/m ..!?• “•«
■H'. ■ .'„ ;-5 ;■ Vf;. U , ■■' <7f r, ait,J| ''• ■ ' ••\ ,'j f , '• •, ‘*1^
''‘^' EMrt ' ' "' ' ' "‘■‘'^ i.V|^j^iJ •■,►.•/■ /.y,, . f
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ANNALS OF CARNEGIE MUSEUM
VoL. 71, Number 1, Pp. 47-57
26 February 2002
NEW SPECIES OF MICROCADDISFLIES (TRICHOPTERA:
HYDROPTILIDAE) FROM NORTHERN FLORIDA
Steven C. Harris*
Research Associate, Section of Invertebrate Zoology
Abstract
Seven new species of microcaddisflies, Oxyethira chrysocara, Hydroptila eglinensis, H. hribriae,
H. sarahae, H. okaloosa, H. hamiltoni and H. sykorai from northern Florida are described and illus-
trated. New figures for previously described species, H. carolae, H. circangula and H. roberti, are
provided to facilitate identifications of the new species.
Key Words: Trichoptera, Hydroptilidae, microcaddisflies, new species, Florida
Introduction
Ongoing collecting in spring-fed streams and other aquatic habitats in northern
Florida by Manuel Pescador and Andrew Rasmussen of Florida A&M University
have yielded several new species of caddisfiies. In this paper, seven new micro-
caddisflies are described, one in the genus Oxyethira and six in the genus Hy-
droptila. With these new species, microcaddisflies in Florida now total 60 species
(Pescador et ah, 1995; Harris et ah, 1998). The distribution of most of the new
species is apparently restricted to small streams and springs on Eglin Air Force
Base, an area of northern Florida noted for several endemic caddisflies (Harris et
ah, 1982; Gordon, 1984), three of which are microcaddisflies. Several of the new
species are very similar to more widely occurring southeastern species, suggesting
that this region of northern Florida may have contained isolated relict populations
at some point in time. Terminology used in the descriptions follows that of Mar-
shall (1979), with length measured from the tip of head to the wing end. Type
material will be deposited at the National Museum of Natural History, Smithson-
ian Institution (NMNH), Carnegie Museum of Natural History (CMNH), Florida
State Collection of Arthropods (FSCA), Illinois Natural History Survey (INHS),
Clemson University Arthropod Collection (CUAC), Florida A&M University
(FAMU), and Clarion University (CU).
Systematic Entomology
Oxyethira chrysocara, new species
(Fig. 1)
Diagnosis. — In many respects, Oxyethira chrysocara is very similar to O. dun-
bartonensis Kelley from South Carolina and Georgia. Both species have reduced
inferior appendages and elongate processes from the phallus apex, but the two
species differ in the details of these structures. In O. dunbartonensis the phallic
processes are membranous and linear, whereas in O. chrysocara they are sclero-
‘Department of Biology, Clarion University, Clarion, PA 16214.
Submitted 3 October 2001.
47
48
Annals of Carnegie Museum
VOL. 71
Phallus, lateral.
tized and curving; and the inferior appendages differ in the number of setiferous
lobes, two in O. dunbartonensis, six in O. chrysocara. As well, the new species
lacks the ventromesal process from abdominal segment VII, which is present in
O. dunbartonensis. The species is known only from the type locality, a small
headwater stream in Clay County.
Description. — Male. Length 2.4 mm. 25 antennal segments. Brown in alcohol. Abdominal segment
VII annular in lateral view; emarginate posteriorly in ventral view, lacking ventromesal process. Seg-
ment VIII in lateral view narrowing posteromesally to truncate knob; deeply incised ventrally, shal-
lowly emarginate dorsally. Segment IX in lateral aspect narrowing anteriorly, posterodorsally a thin
bridge, posteroventrally narrowing to acute process; in ventral view tapering anteriorly, truncate pos-
teriorly; dorsally fused with X. Segment X rectanguloid in lateral aspect; dorsally a membranous lobe,
gradually tapering distally to truncate apex. Inferior appendages reduced to triangular knob on the
inner surface of segment IX, thin in ventral view with numerous setiferous lobes along posterior
margin. Subgenital plate tonguelike in lateral view and strongly downturned; in ventral view square,
fused along mesal margin, narrow bilobed process ventrad. Phallus long and narrow with broad medial
constriction; apical portion bulbous with pair of crossing sclerous fingerlike processes, elongate lip
ventrally; long paramere at base of medial constriction and encircling shaft.
Type Specimens. — Holotype, 6 (NMNH): FLORIDA. Clay County: Gold Head Branch near old
mill crossing, 29°49'56"N, 81°56'45"W, 1 May 1998, A. Rasmussen.
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Harris — New Microcaddisflies from Florida
49
Fig. l.—Hydroptila eglinensis new species, male genitalia: A. Lateral. B. Ventral. C. Dorsal. D.
Phallus, ventral.
Etymology. — From the Greek “gold head,” referring to the type locality of the new species.
Hydroptila eglinensis, new species
(Figs. 2, 3A)
Diagnosis. — In many features, H. eglinensis is similar to H. disgalera Holzenthal
and Kelley. Both species lack the eversible scent glands on the heads of the males,
and genitalic features are similar. Hydroptila eglinensis is readily distinguished by
the saberlike inferior appendages, which are clublike in H. disgalera. The new
species is widely distributed in the cold water streams on Eglin Air Force Base.
Description. — Male. Length 2. 2-2. 7 mm. 28 antennal segments. Head lacking eversible glands
under the dorsal scent caps. Brown in alcohol. Abdominal segment VII annular with short posteromesal
process from venter. Segment VIII annular; ventrally with slight emargination on posterior margin.
Segment IX roughly rectangular in lateral aspect, rounded anteriorly, posteriorly with mesal emargi-
nation; square in ventral view, truncate anteriorly, posteriorly with wide mesal incision, elongate mesal
sclerite internally; dorsum with wide incision on posterior margin. Segment X elongate and tapering
distally in lateral view, fingerlike process from dorsum; dorsally with deep mesal excision, lateral
lobes abruptly narrowing at midlength. Inferior appendages thin and tapering to slightly upturned apex
in lateral view; triangular in ventral view, diverging apically. Subgenital plate triangular in lateral and
ventral views, sclerous along posterior margin. Phallus elongate and tubular, composed of three sec-
tions; anterior section wide basally, bearing short paramere below juncture with short, tubular median
section; distal portion about same length as basal portion, wide at juncture, slender apically.
50
Annals of Carnegie Museum
VOL. 71
Fig. 3. — Eversible scent glands on heads of male Hydroptila: A. Hydroptila eglinensis new species.
B. Hydroptila brihriae new species. C. Hydroptila carolae Holzenthal and Kelley. D. Hydroptila
sarahae new species. E. Hydroptila circangula Harris.
Type Specimens. — Holotype S (NMNH): FLORIDA. Okaloosa County: Rogue Creek, 0.6 km S
Base Rd. 232, Eglin Air Force Base, 30°33'19"N, 86°34'51"W, 21 May 1998, M. Pescador, A. Ras-
mussen. Paratypes: FLORIDA. Same locality and data as holotype, 28 6 (lOd NMNH, lOd CMNH,
4d INHS, 46 FSCA), same locality, but 28 October 1998, 3 6 (CU), Juniper Creek, at Base Rd.
221, Eglin Air Force Base, 1.3 km E State Rd. 85, 30°36'29"N, 86°31'24"W, 21 May 1998, M.
Pescador, A. Rasmussen, Id (CU), Juniper Creek at head, west side of Base Rd. 231, Eglin Air Eorce
Base, SE Duke Field, 30°36'21" N, 86°30'05" W, 21 May 1998, M. Pescador, A. Rasmussen, 3 6
(CUAC), East Turkey Hen Creek at Base Rd. 601, Eglin Air Eorce Base, 30°39'27" N, 86°34'05" W,
28 October 1998, M. Pescador, A. Rasmussen, 2 6 (FAMU), East Turkey Hen Creek at head, 0.3 km
W Okaloosa Lookout Tower, 30°38'48" N, 86°33'23" W, 25 May 1998, M. Pescador, A. Rasmussen,
13 d (5d NMNH, 5d CMNH, 3d INHS), unnamed tributary to Turkey Creek at Base Rd. 634, Eglin
Air Force Base, 30°35'29" N, 86°35'58" W, 27 October 1998, M. Pescador, A. Rasmussen, 4 d (2d
CUAC, 2d FAMU).
Etymology. — From Eglin, in reference to this species occurring on Eglin Air Force Base.
Hydroptila bribriae, new species
(Figs. 3B, 4A-D)
Diagnosis. — This species and the following can be grouped with H. carolae
Holzenthal and Kelley and H. circangula Harris on the basis of genitalic features.
2002
Harris — New Microcaddisflies from Florida
51
Fig. 4. — Hydroptila bribriae new species, male genitalia: A. Lateral. B. Ventral. C. Dorsal. D. Phallus,
ventral. E. Hydroptila carolae Holzenthal and Kelley, phallus, ventral.
All four have the subgenital plate ending in a nipplelike projection; all four have
club-shaped inferior appendages; and all have a similar phallic structure. The
species are separated by a combination of genitalic structures, and the structure
of scent glands on the head. Hydroptila bribriae is separated from H. carolae by
phallic structure, which in the latter is much more elongate basally (Fig. 4E), and
from both H. carolae and H. circangula by the scent glands of the head. The
filaments of these glands are pipelike in both H. bribriae (Fig. 3B) and H. carolae
(Fig. 3C), and linear in H. circangula (Fig. 3E). However, in H. bribriae the
filaments at the end of the eversible lobes are replaced by elongate hairs. Hy-
droptila bribriae was widely distributed across Eglin Air Force Base.
Description. — Male. Length 2.2-2. 7 mm. 30 antennal segments. Eversible scent glands of head with
elongate pipe-shaped filaments, except at tip where replaced with long hairs. Brown in alcohol. Abdom-
inal segment VII annular with short posteromesal process from venter. Segment VIII annular; ventrally
with slight emargination on posterior margin. Segment IX roughly rectangular in lateral aspect, rounded
anteriorly, posteriorly with mesal emargination; in ventral view, deeply incised anteriorly, posteriorly
with wide, shallow incision mesally, narrow internal mesal sclerite; dorsum with thin lateral lobes pos-
teriorly. Segment X elongate and tapering distally in lateral view, central lobe appearing as short fin-
gerlike process from dorsum; dorsally trifid, with elongate lateral lobes narrowing abruptly at midlength.
52
Annals of Carnegie Museum
VOL. 71
mesal lobe membranous and round. Inferior appendages club-shaped in lateral view, widening towards
apex; roughly rectanguloid in ventral view, diverging apically, sclerotized point on outer apical margin,
elongate seta from basal margin. Subgenital plate triangular in lateral view; in ventral view rounded
basally, apex narrowing mesally to nipplelike projection bearing pair of stout setae. Phallus tubular,
composed of two sections; anterior section wide basally, bearing short paramere below juncture with
posterior section, which is tubular at base, then narrowing abruptly to slender curved apex.
Type Specimens. — Holotype 6 (NMNH): FLORIDA. Santa Rosa County: Indigo Creek, at Base
Rd. 213, Eglin Air Force Base, 19 March 1998, M. Pescador, A. Rasmussen. Paratypes: FLORIDA.
Same locality and data as holotype, 5 c? (3(3 NMNH, 2S CMNH). Okaloosa County: Juniper Creek,
at Base Rd. 221, Eglin Air Force Base, 1.3 km E State Rd. 85, 30°36'29"N, 86°31'24"W, 21 May
1998, M. Pescador, A. Rasmussen, 17 S (56 CMNH, 3 <3 FSCA, 36 CUAC, 3c3 CU, 3c3 INHS),
same, but 19 March 1998, 2 6 (FAMU), Juniper Creek at head, west side of Base Rd. 231, Eglin Air
Force Base, SE Duke Field, 30°36'21" N, 86°30'05" W, 21 May 1998, M. Pescador, A. Rasmussen, 1
6 (CU), East Turkey Hen Creek at head, 0.3 km W Okaloosa Lookout Tower, 30°38'48" N, 86°33'23"
W, 25 May 1998, M. Pescador, A. Rasmussen, 1 6 (FAMU). Walton County: Rocky Creek, at
headwaters, Eglin Air Force Base, 25 April 1979, J. Scheiring, 3 6 (26 CMNH, 1 c3 NMNH).
Etymology. — Named for Briana Kriebel, affectionately nicknamed “bribri” and “bug” by her fam-
ily, in recognition of the Kriebel family’s support of Clarion University.
Hydroptila sarahae, new species
(Figs. 3D, 5)
Diagnosis. — This species appears to be closest to Hydroptila circangula Harris,
on the basis of genitalic features. The appearance of the inferior appendages and
the phallus are nearly identical, but subtle differences can be found. Both have
the subgenital plate ending in a nipplelike projection, but this projection is much
longer in the new species, and although the inferior appendages are similar in
ventral aspect, an elongate seta at midlength is present in H. sarahae, but absent
in H. circangula. The new species can also be separated by the arrangement of
the scent glands on the head, which have linear filaments in H. circangula (Fig.
3E) as compared to the pipe-shaped filaments of H. sarahae (Fig. 3D). As with
the previous two species, H. sarahae is widely distributed in the streams on Eglin
Air Eorce Base.
Description. — Male. Length 2. 0-2. 5 mm. 26 antennal segments. Eversible scent glands of head with
elongate pipe-shaped filaments. Brown in alcohol. Abdominal segment VII annular with short poster-
omesal process from venter. Segment VIII annular. Segment IX roughly rectangular in lateral aspect,
rounded anteriorly, posteriorly with slight ventral emargination; in ventral view, deeply incised ante-
riorly, posteriorly with pair of lateral incisions, narrow mesal sclerite; dorsum truncate posteriorly.
Segment X elongate and hoodlike in lateral view; dorsally with deep mesal incision posteriorly, lateral
margins sclerous. Inferior appendages boat-shaped in lateral view, widening to rounded apex; triangular
in ventral view, diverging distally with apices rounded, elongate seta laterally at midlength. Subgenital
plate elongate and slender in lateral view; in ventral view triangular, apex narrowing mesally to long
nipplelike projection bearing pair of stout setae. Phallus tubular, evenly divided into two sections;
anterior section wide basally, bearing short paramere below juncture with posterior section, which is
wide and tubular basally, with slender ejaculatory duct protruding apically.
Type Specimens. — Holotype 6 (NMNH): FLORIDA. Okaloosa County: Rogue Creek, 0.6 km S
Base Rd. 232, Eglin Air Force Base, 30°33'19"N, 86°34'51"W, 21 May 1998, M. Pescador, A. Ras-
mussen. Paratypes: FLORIDA. Same locality and data as holotype, 15 6 (66 NMNH, 6c3 CMNH,
36 INHS), same locality, but 28 October 1998, 4(3 (CU), Rogue Creek at Base Rd. 233, Eglin Air
Force Base, 14 August 1985, B. Armitage, 2 6 (CU), Juniper Creek, at Base Rd. 221, Eglin Air Force
Base, 1.3 km E State Rd. 85, 30°36'29"N, 86°31'24"W, 21 May 1998, M. Pescador, A. Rasmussen, 1
(3 (CUAC), East Turkey Hen Creek at head, 0.3 km W Okaloosa Lookout Tower, 30°38'48" N,
86°33'23", 21 May 1998, M. Pescador, A. Rasmussen, 1 6 (FSCA), East Turkey Hen Creek at Base
Rd. 601, Eglin Air Force Base, 30°39'27" N, 86°34'05" W, 26 October 1998, M. Pescador, A. Ras-
mussen, 1 6 (FAMU), Turkey Creek at Base Rd. 232, Eglin Air Force Base, 30°33'42" N, 86°32'10"
W, 27 October 1998, M. Pescador, A. Rasmussen, 2 6 (FSCA), Turkey Creek at Base Rd. 232, Eglin
Air Force Base, 30°33'42" N, 86°32'10" W, 21 May 1998, M. Pescador, A. Rasmussen, 2 6 (CUAC),
Turkey Creek at Base Rd. 233, Eglin Air Force Base, 19 August 1983, B. Armitage, 41 6 (14(3 NMNH,
2002
Harris — New Microcaddisflies from Florida
53
Fig. 5. — Hydroptila sarahae new species, male genitalia: A. Lateral. B. Ventral. C. Dorsal. D. Phallus,
ventral.
13(3 CMNH, 103 INHS, 103 FAMU), unnamed tributary to Turkey Creek at Base Rd. 619, Eglin
Air Force Base, 14 August 1985, B. Armitage, 11 3 (43 FSCA, 43 CUAC, 33 CU).
Etymology. — Named for my daughter Sarah on the occasion of her 21st birthday.
Hydroptila okaloosa, new species
(Fig. 6)
Diagnosis. — Hydroptila okaloosa bears some resemblance to H. pecos Ross
and H. ajax Ross in the shape of the downturned inferior appendages, but the
large, bulbous segment X is more similar to that of H. protera Ross. The new
species is quite distinctive, particularly in the appearance of the phallic apex, but
the combination of the ventrad curving inferior appendages and deeply incised
tenth tergum will also separate H. okaloosa. Despite widespread collecting in the
streams of Eglin Air Force Base, H. okaloosa was only found in Rogue Creek.
Description.- — Male. Length 2.9-3. 1 mm. 30 antennal segments. Eversible scent glands of head with
long pipe-shaped filaments, except at tip where replaced with long hairs (as in Fig. 3B). Brown in
alcohol. Abdominal segment VII annular with short posteromesal process from venter. Segment VIII
tapering posteroventrally in lateral aspect, setose lobe posterodorsally; ventrally with mesal incision
on posterior margin; rectanguloid dorsally. Segment IX short, largely contained within VIII, rounded
54
Annals of Carnegie Museum
VOL. 71
Fig. 6. — Hydroptila okaloosa new species, male genitalia; A. Lateral. B. Ventral. C. Dorsal. D. Phallus,
ventral.
anteriorly; in ventral view deeply incised anteriorly, posteriorly with wide emargination; dorsum nar-
rowing to rounded lobe which is apparently fused with X. Segment X elongate and bulbous in lateral
view, sclerotized band dorsolaterally; in dorsal view deeply emarginate, lateral extensions tapering
distally to acute out-turned apices, outer margins sclerous. Inferior appendages enlarged distally and
downturned in lateral view; ventrally long and fingerlike, contiguous basomesally, inner margins slight-
ly concave. Subgenital plate long and thin in lateral aspect, bearing elongate seta apically; ventrally
a thin, narrow shelf, emarginate and sclerous on posterior margin. Phallus tubular, apex divided into
two elongate processes, outer process curved at apex, inner process rectanguloid and bearing ejacu-
latory duct which protrudes apically; paramere absent.
Type Specimens. — Holotype 6 (NMNH): FLORIDA. Okaloosa County: Rogue Creek, 0.6 km S
Base Rd. 232, Eglin Air Force Base, 30°33'19"N, 86°34'51"W, 21 May 1998, M. Pescador, A. Ras-
mussen. Paratype: FLORIDA. Same locality as holotype, but 7 April 1999, 1 S (CMNH).
Etymology. — Named for the type locality in Okaloosa County.
Hydroptila hamiltoni, new species
(Fig. 7A-D)
Diagnosis. — In overall genitalic features, this new species is nearly identical to
H. roberta Hamilton and Holzenthal from Georgia. Only the structure of the
phallus will serve to separate the two species; in H. roberta (Fig. 3E) the basal
and apical sections of the phallus are the same length, with the phallic apex
2002
Harris — New Microcaddisflies from Florida
55
Fig. 7. — Hydroptila hamiltoni new species, male genitalia: A. Lateral. B, Ventral. C. Dorsal. D. Phal-
lus, ventral. E. Hydroptila roberta Hamilton and Holzenthal, phallus, ventral.
elongate and thin, while in H. hamiltoni, the basal section of the phallus is about
four times the length of the apical section, with the phallic apex short. The new
species was widely distributed in the streams on Eglin Air Force Base.
Description. — Male. Length 2.5-2. 8 mm. 27 antennal segments. Eversible scent glands of head with
elongate pipelike filaments, except at tips (as in Fig. 3B). Brown in alcohol. Abdominal segment VII
annular with short posteromesal process from venter. Segment VIII annular. Segment IX in lateral
aspect rounded anteriorly, posteriorly with dorsal protuberance; ventrally with deep mesal incision
anteriorly, truncate posteriorly; dorsum lobate posteromesally with narrow incision. Segment X elon-
gate and bulbous in lateral view; narrow dorsally with deep mesal excision posteriorly. Inferior ap-
pendages thin and nearly parallel-sided in lateral view, strongly curved at base which bears thin
setiferous lobe; in ventral view elongate and narrow, tapering distally with sclerotized lateral point at
apex, widely separated basally. Subgenital plate long and slender in lateral view; ventrally a thin,
narrow shelf, emarginate and sclerous on posterior margin. Phallus elongate and tubular, distal portion
about 14 length of basal portion, widening at juncture, narrowing apically to recurved tip; short par-
amere encircling shaft below juncture of sections.
Type Specimens. — Holotype 6 (NMNH): FLORIDA. Okaloosa County: Rogue Creek, 0.6 km S
Base Rd. 232, Eglin Air Force Base, 30°33'19"N, 86°34'51"W, 21 May 1998, M. Pescador, A. Ras-
mussen. Paratypes: FLORIDA. Same locality and data as holotype, 13 S (56' NMNH, 46 CMNH,
56
Annals of Carnegie Museum
VOL. 71
Fig. 8. — Hydroptila sykorai new species, male genitalia: A. Lateral. B. Ventral. C. Dorsal. D. Phallus,
ventral.
A 6 INHS), Juniper Creek, at Base Rd. 221, Eglin Air Force Base, 1.3 km E State Rd. 85, 30°36'29"N,
86°3L24"W, 21 May 1998, M. Pescador, A. Rasmussen, 6 6 {26 FAMU, 2c? CUAC, 2(? CU), East
Turkey Hen Creek at head, 0.3 km W Okaloosa Lookout Tower, 30°38'48" N, 86°33'23" W, 25 May
1998, M. Pescador, A. Rasmussen, 3 (2 c? FAMU, 1 c? FSCA).
Etymology. — Named for Steven W. Hamilton, co-describer of H. roberta, in recognition of his many
contributions to the study of caddisflies.
Hydroptila sykorai, new species
(Fig. 8)
Diagnosis. — This new species is very similar in overall appearance to Hydrop-
tila Ouachita Holzenthal and Kelley, which is apparently endemic to Schoolhouse
Spring, a small artesian spring, in Louisiana (Holzenthal and Kelley, 1983). Both
species have heavy spines from the margins of abdominal segment VIII, but while
these spines are short and curved inward in H. ouachita, they are much longer
and straight in H. sykorai. Both species also have abdominal segment X divided
distally, but in H. ouachita this division is shallow, whereas in H. sykorai it is
very deep, creating a pair of narrow rods. The locality of this new species is a
small, cold, sand-bottom springrun.
2002
Harris — New Microcaddisflies from Florida
57
Description. — Male. Length 1.9-2. 3 mm. 28 antennal segments. Eversible scent glands of head with
elongate pipe-shaped filaments (as in Fig. 3B). Brown in alcohol. Abdominal segment VII annular
with short posteromesal process from venter. Segment VIII in lateral view elongate and tapering
distally; deeply emarginate dorsally, with series of stout elongate spines from posterolateral margins;
posterior margin with shallow mesal incision ventrally. Segment IX short, retracted within segments
VII and VIII; narrowing dorsally in lateral view; in dorsal and ventral views square, with deep, rounded
mesal incision posteriorly and shallow incision anteriorly. Segment X elongate and thin in lateral view;
dorsally with deep mesal incision distally, creating acute, sclerous lateral rods. Inferior appendages
long and thin in lateral view, widening to rounded apex distally; thin in ventral view, diverging distally
with apices rounded and bearing darkened point laterally. Subgenital plate not evident. Phallus very
narrow and tubular, with thin paramere encircling shaft near midlength.
Type Specimens.- — ^Holotype male. FLORIDA. Gadsden County: headwaters of Quincy Creek, 7
km N Quincy at Florida A&M Research and Extension Center, 30°39'27" N, 84°36'50" W, 7 June
1999, A. Rasmussen, emergence trap (NMNH). Paratypes: FLORIDA. Same locality as holotype, but
17 January 1998, 1 S (CMNH), same, but 20 March 1999, 1 S (FAMU), same, but 4 April 1999, 1
c? (INHS), same, but 21 June 1999, 1 c? (CUAC).
Etymology. — Named for Jan Sykora, friend and colleague, in recognition of his contributions to the
study of caddisflies.
Acknowledgments
Appreciation is expressed to Manny Pescador and Andy Rasmussen of Florida A&M University
for allowing me to examine microcaddisflies in their collections from northern Florida and for re-
viewing the resultant manuscript. John Morse of Clemson University kindly lent paratypes of Hy-
droptila roberti, H. carolae, and H. ouachita for examination. Brian Armitage of the Ohio Biological
Survey and Joseph Scheiring provided some material from their collections on the Eglin Air Force
Base. The comments of the anonymous manuscript reviewers and editors at the Carnegie Museum of
Natural History are also greatly appreciated.
Literature Cited
Gordon, A. E. 1984. The Trichoptera of Florida: A preliminary survey. Pp. 161-166, in Proceedings
of the Fourth International Symposium on Trichoptera, 1983 (J. C. Morse, ed.). Dr. W. Junk
Publishers, The Hague.
Harris, S. C., P K. Lago, and J. F. Scheiring. 1982. An annotated list of Trichoptera of several
streams on Eglin Air Force Base, Florida. Entomological News, 93:79-84.
Harris, S. C., M. L. Pescador, and A. K. Rasmussen. 1998. Two new species of microcaddisflies
(Trichoptera: Hydroptilidae) from northern Florida. Florida Entomologist, 81:221-224.
Holzenthal, R. W., and R. W. Kelley. 1983. New micro-caddisflies from the southeastern United
States (Trichoptera: Hydroptilidae). Florida Entomologist, 66:464-472.
Marshall, J. E. 1979. A review of the genera of the Hydroptilidae (Trichoptera). Bulletin of the
British Museum of Natural History (Entomology), 39:135-239.
Pecador, M. L., a. K. Rasmussen, and S. C. Harris. 1996. Identification manual for the caddisfly
(Trichoptera) larvae of Florida. Florida Department of Environmental Protection, Tallahassee,
Florida.
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FROM THE ARCHIVES AND COLLECTIONS
A MIOCENE BONE BED FROM AGATE FOSSIL BEDS
NATIONAL MONUMENT
W. Orr Goehring^ and Mary R. Dawson^
“In 1904 a field party of the Carnegie Museum, the writer in charge, was
extremely fortunate in discovering what will undoubtedly prove to be one of the
most important quarries of fossils as yet discovered in the Miocene of North
America” (Peterson, 1906). So begins former field collector for Carnegie Museum
O. A. Peterson’s account of the discovery of Agate Spring Fossil Quarry, located
on the Upper Niobrara River in Sioux County, Nebraska.
Peterson’s claim that the quarry’s discovery was “extremely fortunate” is no
exaggeration. The quarry may never have been discovered at all had it not been
for the curious eye of James H. Cook, owner of the Agate Springs Ranch. In
1885, two years before he purchased the ranch from his father-in-law. Cook spot-
ted some bone fragments scattered under a rock shelf near the ranch house (United
States National Park Service, 1980). Not certain of the origin of the bones or
their significance, Cook did not report the find until after he and his wife Kate
bought the ranch from Dr. Elisha B. Graham in 1887 (United States National Park
Service, 1980).
Though he provides a different date for the discovery of the bone fragments,
Peterson (1906) gives a similar account of Cook’s find, and he goes on to describe
the enthusiasm with which he and other members of the Carnegie Museum field
party began their initial excavation:
In 1890 Mr. James H. Cook, on whose property the Agate Spring
Fossil Quarry is located, discovered many small bones and fragments in
the talus from the fossil-bearing stratum from the hills, in which the
quarry is located. Very naturally he thought that the bones were those
of Indians interred together with their horses. Mr. Cook accompanied me
to this place in August, 1904. Realizing that this was a discovery of
much paleontological promise we immediately began work on the de-
posits and resumed work early in the season of 1905.
According to Peterson (1906), Cook began his own excavation of the site in
the fall of 1904. Anxious to preserve the specimens using the most modern meth-
ods, Peterson wrote Cook a letter asking him to refrain from his work until a
party from the Carnegie Museum could return to the site in the spring of 1905.
Cook complied with the request, and excavation resumed in April of 1905 (Fig.
1). Peterson (1906) writes:
' Managing Editor, Scientific Publications.
^ Curator, Section of Vertebrate Paleontology.
59
60
Annals of Carnegie Museum
VOL. 71
Fig. 1. — O. A. Peterson excavating at Carnegie Hill in Agate Fossil Beds, Sioux County, Nebraska,
circa 1909. Section of Vertebrate Paleontology photo archives, negative no. 6.
During the season of 1905 the party uncovered an area of 45 X 20
feet in the quarry. This area was plotted out in squares five feet in
dimension, and a diagram was made representing this arrangement. The
bones are found mostly disarticulated, much mixed, and thickly distrib-
uted through this layer of sandstone. It was soon discovered that the
most intelligent way in which to secure this tangled mass of material
was to take out blocks of sandstone which contain the fossils.
Fossils of the rhinoceros Menoceros were the most abundant in the excavation
area that became known as Carnegie Hill, followed by those of the large ente-
lodont Dinohyus and the chalicothere Moropus. One of the slabs containing this
jumble of bones has been kept together as a bone bed, preserving the fossils just
as they were found in the sandstone block collected by Peterson and his team.
2002
Goehring and Dawson — From the Archives and Collections
61
Fig. 2. — A small section of the Early Miocene Bone Bed from Agate Fossil Beds, Sioux County,
Nebraska. The bone bed, newly renovated for exhibit in 2001, measures approximately 4 X 8.5 feet.
Photograph by Norman Wuerthele, CMNH.
The newly renovated block (Fig. 2), one of the most impressive specimens from
the quarry, contains hundreds of dissociated bones, probably carried to their place
of deposition by moving water.
The locality that Peterson (1906) described as “one of the most important
quarries of fossils as yet discovered” has proven to be just that: in 1965 the
United States Congress designated the quarry and the area surrounding it along
the Niobrara River Agate Fossil Beds National Monument.
Acknowledgments: Elizabeth A. Hill and Norman Wuerthele, CMNH, contrib-
uted to the preparation of figures 1 and 2, respectively.
Literature Cited
Peterson, O. A. 1906. The Agate Spring Fossil Quarry. Annals of Carnegie Museum, 3:487-494.
United States National Park Service. 1980. Agate Fossil Beds; Agate Fossil Beds National Mon-
ument, Nebraska. U. S. Department of the Interior, Washington, D. C.
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ANNALS
of CARNEGIE MUSEUM
THE CARNEGIE MUSEUM OF NATURAL HISTORY
4400 FORBES AVENUE • PITTSBURGH, PENNSYLVANIA 15213
VOLUME 71
28 MAY 2002
NUMBER 2
CONTENTS
ARTICLES
Comparing pottery from the Proto-historie MeKees Rocks Village and
Eisiminger sites of southwestern Pennsylvania
. Richard L. George 63
Fleas (Siphonaptera) from Ancash Department, Peru with the description of a
new species, Ectinorus alejoi (Rhopalopsyllidae), and the description of the
male of Plocopsylla pallas (Rothschild, 1914) (Stephanocircidae)
Michael W. Hastriter, Michael D. Zyzak, Ruler Soto,
.... Roberto Fernandez, Nelson Solorzano, and Michael F. Whiting 87
Review of Leptomeryx (Artiodactyla, Leptomeryicidae) from the Orellan
(Oligocene) of Nebraska
William W. Korth and Margaret E. Diamond 107
FROM THE ARCHIVES AND COLLECTIONS
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ANNALS OF CARNEGIE MUSEUM
VoL. 71, Number 2, Pp. 63-86
28 May 2002
COMPARING POTTERY FROM THE PROTO-HISTORIC McKEES ROCKS
VILLAGE AND EISIMINGER SITES OF
SOUTHWESTERN PENNSYLVANIA
Richard L. George
Staff Archaeologist/Archaeology Collection Manager, Section of Anthropology
Abstract
Comparison of pottery from two Proto-historic sites in southwestern Pennsylvania, McKees Rocks
Village and Eisiminger, disclosed important differences. McKees Rocks Village, occupying a promi-
nent Ohio River escarpment in Allegheny County, produced distinctive pottery featuring decorated
added rim-strips and simple stamping, while cordmarked Eisiminger pottery had lips that were scal-
loped or impressed in their interior by cord-wrapped paddle impressions applied perpendicularly to
the lip. It is hypothesized that Eisiminger, located on a hill in Greene Country, represented remnant
Moeongahela populations, whereas McKees Rocks Village was established by an intrusive people
with affinities to Wellsburg Phase sites in Ohio and West Virginia. It is also suggested that the
Wellsburg Phase people may have been of Iroquoian origin, based on discussed artifacts.
Key Words: Proto-historic, rim-strips, Foley Farm Phase, Wellsburg Phase
Introduction
The McKees Rocks Village site, 36 AL16, and the George Eisiminger site,
36GR2, are Proto-historic sites located 67 km apart in southwestern Pennsylvania
(Figs, 1:1&2). Pottery indicates the sites had dissimilar antecedents in spite of
their proximity and apparent contemporaneous occupation. The purposes of this
paper are to review the archaeological contexts of the two sites, describe and
compare their pottery forms in order to illustrate the differences, and discuss the
cultural affiliations of the inhabitants of the sites. The McKees Rocks Village site
and the George Eisiminger site were occupied at a critical span of time for the
Upper Ohio Valley region of North America, which in succeeding years came to
be abandoned by Native Americans for almost a century.
The Sites
The McKees Rocks Village Site
The McKees Rocks Village site was located on the south side of the Ohio River
about 4.8 km downstream (northwest) of its origin at the confluence of the Al-
legheny and Monongahela rivers (Fig. 2). It was situated a short distance from
the mouth of Chartiers Creek, a mainly northward flowing stream that drains
approximately 834 square km. Anyone traveling down Chartiers Creek and en-
tering the Ohio River would immediately confront Brueot Island, a locality that
would have provided the site inhabitants with access to riverine resources since
the channel between the island and the mainland was relatively shallow except
during early spring flooding. The McKees Rocks Village site was elevated 24 m
above the Ohio River, on a high terrace providing a commanding view of the
surrounding area.
Submitted 3 September 2001.
63
64
Annals of Carnegie Museum
VOL. 71
Fig. 1. — Map of U.O.V. with archaeological sites referred to in text. 1. McKees Rocks Village,
36AL16. 2. Eisiminger, 36GR2. 3. Foley Farm, 36GR52. 4. Household, 36WM61. 5. Johnston, 36IN2.
6. Georgetown, 36BV29. 7. Lang, 36WH48. 8. Portman, 36AL40. 9. Sony, 36WM151. 10. Throck-
morton, 36GR160. 1 1. Wellsburg, 46BR2. 12. Godwin-Portman, 36AL9.
2002
George — McKees Rocks Village, Eisiminger Sites
65
Fig. 2. — Portion of U.S.G.S. Pittsburgh West Quadrangle (1948) showing location of the McKees
Rock Village site, 36AL16.
The area occupied by the McKees Rocks Village site was referred to as “Fort
Hill” on a map prepared by George Mercer for the Ohio Company in 1753
(Brown 1959:P1. 17). Mercer also noted that:
There had been an Indian Fort there some years ago. The ditch is now
to be seen. There the Indians always fled upon an Alarm as it was
reckoned the strongest Fort they had. Several thousands have lost their
lives in the Attack of it but was never taken (Brown, 1959:PL 17).
Also located on the same terrace was the McKees Rocks Mound, 36AL6, exca-
66
Annals of Carnegie Museum
VOL. 71
vated by Carnegie Museum of Natural History (CMNH) in 1896 (Mayer-Oakes,
1955:145-153).
The ceramic artifacts (and some triangular points) used in this analysis were
recovered by William Buker during a 1958-64 salvage excavation of the McKees
Rocks Village site. Buker’s work was hampered by factors such as vandalism,
past disturbance of the general area containing the dwellings and a cemetery, and
the lack of excavation continuity stemming from seasonal shutting down of the
fieldwork (Buker, 1968:3-4). The important assemblage of artifacts Buker man-
aged to salvage, despite these deterrents, provided the basis for the recent attri-
bution of the McKees Rock Village site to the Wellsburg Phase (Carskadden and
Morton, 2000:183). Buker (1968:46-49) recognized that McKees Rocks had af-
finities to the Wellsburg and Riker sites of Fort Ancient. Buker’s (1968:3-49)
report in Pennsylvania Archaeologist was followed by reports by Lang (1968:50-
80) on “The Natural Environment and Subsistence Economy of the McKees
Rocks Village Site,” and Jones (1968:81-86) on “Corn from the McKees Rocks
Village site.” Thus, Buker and his colleagues provided important contributions to
the prehistory of the Upper Ohio Valley even though much of the site they ex-
cavated had been destroyed before their work began.
The sole radiocarbon date for the McKees Rocks Village site was obtained
from wood charcoal submitted in 1968 by Don Dragoo to the University of Mich-
igan Radiocarbon Laboratory. A date of 620±100 B.R, Cal. A.D. 1354 (M-2201)
was obtained (James B. Griffin, letter to Dragoo, 5/21/69). This assay, from an
unknown on-site provenience, is too early for the Wellsburg Phase artifacts herein
discussed. The date might relate to an earlier Late Prehistoric occupation since
most southwestern Pennsylvania sites were utilized more than once during that
period (George, 1983; Buker, 1993; Eisert, 1993).
The genesis of the theory for the Proto-Historic age of the Wellsburg compo-
nent of the McKees Rocks Village site came from James Morton (personal com-
munication, 1995), who developed his idea while examining comparable pottery
from known A.D. 17th century sites in Ohio and West Virginia. After subsequent
conversations with Buker, the author re-examined CMNH catalog listings of metal
scraps found during the 1958-64 excavation. The results are that one brass bead
(Pig. 3A) and four brass scraps (Pig. 3B) were recorded from three units — Pea-
tures 32, 36 and 41. Further supporting a Proto-historic locus at the McKees
Rocks Village site are another piece of scrap brass (Fig. 3C) and a brass arrow
point (Fig, 3D) recovered by Donald Tanner from old spoil piles.
CMNH eurates a small collection of artifacts from the Wellsburg site, 46Br2,
donated by W. Singer in 1954. Among the 51 artifacts are 23 shell tempered rim
sherds that feature sub-lip decoration like those found at the McKees Rocks Vil-
lage site. When viewed by the author in 1993, the Wellsburg site (Fig. 1:11)
appeared to be severely impacted by commercial development.
The George Eisiminger Site
The George Eisiminger site, 36GR2, is located at an elevation of 366 m on a
hill in Greene County (Pig. 1 :2). Originally it was recorded as the Jesse Lapping
site. The Eisiminger site occupies the southwestern slope of the hill (Fig, 4), but
its extent and exact boundaries are difficult to discern because the field has not
been plowed for many years. A spring-head and an unnamed run are situated 25
m to the southeast; Smith Creek, a north-flowing, fourth order stream, lies 1.2
2002
George — McKees Rocks Village, Eisiminger Sites
67
Fig. 3. — Proto-historic artifacts from the McKees Rocks Village site excavated by Buker. A. Brass
bead. B. Brass scraps. C. Brass scrap, donated by Tanner. D. Brass arrowhead, donated by Tanner.
68
Annals of Carnegie Museum
VOL. 71
Fig. 4. — Portion of U.S.G.S. Oak Forest Quadrangle (1973) showing location of the Eisiminger site,
36GR2.
km to the west (Fig. 4). Mayer=Oakes (1955:122) first called attention to the
Eisiminger site as one of . . at least five sites that are clustered in an area just
south of Waynesburg, PA. ...” that produced “ . . . [a] fair amount of European
trade material . . . .” CMNH possesses a small amount of surface collected arti-
facts from the Eisiminger site; a much larger assemblage is curated at the Paul
R. Stewart Museum, Waynesburg College.
The Eisiminger site artifacts at Waynesburg College were excavated by Paul
2002
George — McKees Rocks Village, Eisiminger Sites
69
Stewart ie the early 1950s. In an unpublished manuscript entitled “Excavation of
Pottery House” (see Appendix), Stewart (n.d.:2) described a structure that .
was approximately 18 feet long by 12 feet wide, almost rectangular, and the main
floor almost completely paved with small stones ranging from. 2 to 3 square inches
to 50 or 50 square inches, the average being approximately 10 square inches.”
Stewart (n.d.:2-3) also recorded the existence of a . number of lobes from 2
to 3 inches to as much as 8 or 10 inches below the paved surface,” Apparently,
there were six of these “lobes” that were . from two to three feet across and
were found ... all around the structure except at the southwest, where presumably
the entrance had been established.” Stewart (n.d.:2) considered these features to
be firepits, but their numbers and positions around the structure suggest instead
that this was a “petal house,” a structure type originally described as a “Food
Warehouse or Big House” by Herbstritt (1983: 1 14-1 17) at the Throckmorton site,
36GR160 (Figure 1:10). Similar features have been recorded at the Foley Farm
site, 36GR52 (Fig. 1:3) (Herbstritt, 1984), and the Sony site, 36WM151 (Fig. 1:
9) (Davis and Wilkes, 1997:33-36), but the appendages in these cases were at=
tached to round houses (detected from their circular postmold patterns). Thus, the
“Pottery House” at the Eisiminger site, being rectangular, is apparently anoma-
lous ie form compared to those of the other sites. Stewart (n.d.:5) states that “. . .
no well-defined post holes were found ...” during his excavation.
Stewart (n.d.:3) noted the . . amazing wealth of potsherds . . from the
structure that, otherwise, had an . . almost complete absence of artifacts except
pottery . . . For this study, 61 rim sherds provenieeced to the Eisiminger site
were borrowed from the Paul R. Stewart Museum, and many of them presumably
were recovered during Stewart’s excavation of the Pottery House. Mayer-Oakes
(1955:127-128) illustrated Waynesburg College pottery from the Eisiminger site
that included rim sherds of three bowls, two reconstructed “jars,” and a section
of a compound vessel (Fig. 5). The larger of the two jars (Mayer-Oakes, 1955:
128, PL 68, A), has a scalloped rim.
Among the few non-ceramic items found was “the small roughly-rolled bit of
copper kettle of settler origin” (Stewart, n.d.:3). Mayer-Oakes (1955:123) listed
Proto-historic artifacts as “. . . blue glass beads, two red beads with a blue core,
and several perforated fragments of small copper spheres, perhaps crudely hand-
made bell fragments . . . He also alluded to . a small scrap of silver ...”
that was considered, at that time, as providing the best evidence of a Proto-historic
artifact (Mayer-Oakes, 1955:122"-! 23). The blue glass beads are a type Lapham
(1995:24) designates “Washington Boro Blue.” The type is the most commonly
found bead on three Pennsylvania and one New York sites dating between A.D.
1600 and 1625 (Lapham, 1995:Fig. 4). Two of these sites, Spragg, 36GR12, and
Foley Farm, 36GR52, are Proto-historic sites located in Greene County, Penn-
sylvania.
Comparisons
The McKees Rocks Village and Eisiminger sites are dissimilar in many ways
and the data gathering process for each site also differed. The author has analyzed
1,221 rim sherds from 14 Late Prehistoric and Proto-historic Monongahela sites
as part of a larger study, and the data from the Eisiminger site reproduced in
Table 1 are extracted from that analysis. The McKees Rocks Village site data are
derived mostly from Buker’s site report, with special attention given his Tables 3
70
Annals of Carnegie Museum
VOL. 71
0 cm 5
Fig. 5. — Portion of a double pottery vessel from the Eisiminger site.
Table 1. — Comparison of pottery attributes for the McKees Rocks Village and Eisiminger sites.
Trait
McKees Rocks Village’
Eisiminger
Shell Tempering
Exterior Treatment, rim sherds:
98.8%
98.0%
Cordmarked
86.0%
96.0%
Plain
10.0%
2.0%
Simple Stamped
4.0%
0.0%
Striated
0.0%
2.0%
Incidence of Rim Strips
Final Cordage Twist:
85.0%
3.0%
“Z”
21.0%
70.0%
“S”
79.0%
30.0%
(N = 34)
(N = 46)
Bowls
Present (not common)
Present (not common)
Loop Handles
Present (not common)
0.0%
Nodes
Present
0.0%
Lugs
Present
Vertical Compound Vessels
0.0%
2.0%2
' Only shell tempered rim sherds were used for more viable comparisons.
^ There are two examples of vertical compound vessel sherds in the Eisiminger sample, but only one
was available for study.
2002
George — McKees Rocks Village, Eisiminger Sites
71
and 4 (Buker, 1968:21, 24)„ The author performed the final twist analysis on 34
rim sherds since that analysis technique was not yet utilized in the 1960s. Buker
kindly donated most of the rim sherds used in the analysis.
The McKees Rocks Village site and Eisiminger site samples are comparable in
having high percentages of shell tempering and cordmarking, a not unexpected
occurrence for Upper Ohio Valley Proto-historic sites, but the samples differ
markedly in all other attributes. The Eisiminger vessel sample discloses a ceramic
'‘profile” consisting of a shell tempered, cordmarked vessel with a high percent-
age (70%) of Final “Z” twist cordage impressions. In addition, the vessel would
have an elongated jar shape with a slightly flaring rim (Mayer-Oakes, 1955:128,
Fig. 68 A). Punctations are present on 16% and one rim sherd exhibits fingernail
incising. The McKees Rocks Village vessel sample indicates a vessel that was
most often cordmarked, but with a small percentage (4%) being simple stamped.
The typical McKees Rocks Village vessel also had a decorated, added rim strip,
or filet, and a globular form (Buker, 1968:14-16). The final cordage twist type is
usually (79%) Final “S” twist in the McKees Rocks Village sample, a finding
that sharply contrasts with the Eisiminger data. The McKees Rocks Village site
ceramic inventory includes lugs, nodes, and a few loop handles, although “Loop
handles from the site are usually small and poorly made” (Buker, 1968:24).
There is no question that the pottery vessels from the two sites represent two
distinct potter y-making traditions. In the following section, these differences will
be further discussed.
Not All Rim Strips Are Created Equal
Differences in Added Rim Strips
There is a significant difference between the frequency of added rim strips for
the Eisiminger site (3%) and McKees Rocks Village site (85%) ceramic samples
(Table 1). There are qualitative as well as quantitative differences between the
rim strips.
Eisiminger
Only four rim sherds incorporate rim strips. Two rim strips exhibit oblique
cordmarking. One of the cordmarked rims (Fig. 6A) is decorated with an uneven
row of ovate punctations that exist above as well as at the bottom of the rim strip.
The second cordmarked rim (Fig. 6B) has a row of crude punctations more or
less in the mid-section of the strip. The punctations appear to have been applied
with a bifurcated tool. A third rim sherd (Fig. 6C) has punctations applied at the
bottom of its wider rim strip. These punctations were done with a circular tool
that had a central blade-like middle.
The fourth Eisiminger rim strip (Fig. 7) is represented by two, cross-mended
sections of a rim sherd with a single, incipient castellation. The rim strip is thicker,
decorated more prominently with a row of punctations, and horizontally cork-
marked; the sub-rim strip area is vertically cordmarked. This fourth Eisiminger
rim sherd is the most similar in overall appearance and configuration to the
McKees Rocks Village site forms,
McKees Rocks Village Site
Buker (1968:13-25) provides detailed descriptions of four varieties of rim strips
from the McKees Rocks Village site. Buker's illustrations and descriptions and
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Fig. 6. — Eisiminger site rim sherds. A. Uneven row of punctations at bottom of rim strip. B. Crude
punctations in central area of narrow rim strip. C. Wide rim strip with crude punctations at bottom.
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George — McKees Rocks Village, Eisiminger Sites
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c
112 3^
1 5 cm
Fig. 7. — Eisiminger site rim sherds from same vessel with prominent punctations on a narrow strip
and a low, rounded castellation.
the author’s study of McKees Rocks Village site specimens (Fig. 8) leave no
doubt that the McKees Rocks Village potters were crafting these decorative and
practical additions to their pottery in a much more refined and careful manner,
quite possibly for a long time. The contrast between the rim strips from the two
sites is striking. Whereas the Eisiminger rim strips are generally unevenly deco-
rated, the McKees Rocks Village samples are the opposite, indicating a well-
developed competence. Punctations are prominent and pleasing (Fig. 8), as if the
individual potters were “signing” their products with their “maker’s marks,” in
the author’s view.
The Pots, The People, and the Questions of Origins
Eisiminger Site
The five Proto-historic sites identified by Mayer-Oakes (1955:122) for Greene
County have risen to twelve sites, including two reported from West Virginia, one
in bordering Monongalia County, and another in Ohio County (Babich et ak,
1996). The Eisiminger site is included in these twelve sites. Johnson (1990:9)
assigned these sites to the Foley Farm Phase of Monongahela following Herbs-
tritt’s (1984) popular report descriptions based on the partial excavation of the
Foley Farm site, 36GR52 (Fig. 1:3). However, none of these authors have pro-
vided a definition of the Foley Farm Phase that entails artifact and settlement
pattern trait frequencies for a number of sites. Thus, we are left with an extremely
spotty database for, at least, one of the terminal phases of Monongahela.
The data from the Eisiminger site does, however, invite contemplation about
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t # #
Fig. 8. — Three rim sherds from the McKees Rocks Village site.
the possible cultural make-up of the Foley Farm Phase people. In discussing the
results of the Foley Farm site excavation, Herbstritt states that the fieldwork:
. . . uncovered samplings of the Monongahela’s material culture which
have, among other things, proven a long tradition of native technology.
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George — McKees Rocks Village, Eisiminger Sites
75
There seems to have been very little change in the culture’s ceramic,
stone, bone and shell industries. (Herbstritt, 1984:30).
Certainly, the validity of Herbstritt’s proposition holds true for the ceramic arti-
facts representing Eisiminger. Although shell tempering and cordmarking are traits
that Monongahela shares with Fort Ancient, these attributes are nonetheless typ-
ically Monongahela, especially of later components. A general lack of append-
ages, such as lugs or loop handles, is also characteristic of non-Drew Phase Mo-
nongahela ceramic assemblages. A minor incidence of non-decorated castellations
occurs in many Monongahela samples, and Eisiminger is no exception.
During the aforementioned study of 1,221 rim sherds from 14 sites, it was
determined that 20% of the Eisiminger site’s rim sherds were scalloped. What can
be considered significant about this figure is the fact that the Late Monongahela
Household site, 36WM61 (Fig. 1:4), had a 94% incidence of scalloped, or deep,
tool-impressed lips (George et ak, 1990:51). These occurred on cordmarked, shell
tempered vessels that had slightly flared rims (George et ak, 1990:50). The House-
hold site, with a date of 325±80 B.P., Cal. A.D. 1562 (UGA-3453), was
occupied just a few years before European trade goods started to appear in the
Upper Ohio Valley. Why, then, is there only a small percentage of scalloped lips
at Eisiminger? Why, also, are there punctations, added rim strips and castellations
in the Eisiminger site sample, whereas none of these attributes were at Household
(George et ak, 1990:50-51)?
It is here proposed that Eisiminger represents a gathering of remnant Monon-
gahela populations who brought their pottery traditions with them and that the
various attributes constitute continuations of these. This would be in agreement
with Herbstritt’s (1984:30) suggestion that there was “little change in the culture’s
ceramic ...” industry. Thus, the tradition of tool-impressed or scalloped lips,
present at Eisiminger (Fig. 9A), if not in great numbers, seems to imply that some
potters carrying the Household site ceramic mental template were a part of the
Eisiminger site population.
Another lip decoration, that of perpendicularly applied, cord-wrapped paddle
edge impressions on the inner lip (Fig. 9B), occurs on 15% of Eisiminger rim
sherds. This attribute is also present on rim sherds from Foley Farm (32%) and
Throckmorton (14%) (Figure 1:10) (Herbstritt, 1983:153-154). On one Eisimin-
ger rim sherd the interior lip notching was deep enough to result in a scalloped
lip (Fig. 9C), suggesting, perhaps, that scalloping gave rise to the inner lip, cord-
wrapped paddle impressions, that is considered to be a Proto-historic Foley Phase
attribute.
The most unusual rim sherds in the Eisiminger sample are portions of vertical
compound vessels in which only the lower of two rim sherds are intact (Fig. 5).
The larger of these consists of three joined sections, two of which exhibit a flared
rim with a perpendicularly notched lip. The lower section of the vessel has almost
horizontal cordmarking, whereas the less intact upper vessel has convoluted cord-
marking that tends toward vertical. The plain interior, while carinated at the joined
area, exhibits no evidence of an intersecting joint. Mayer-Oakes (1955:128, Pk
68B) illustrated the same compound vessel portion that was attributed to Pottery
House at Eisiminger. The second example (Fig. 9D) is a single sherd, with a
protruding lip, that exhibits shallow notching. The upper portion of the vessel is
not large enough to determine surface treatment, whereas the lower section has
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Fig. 9. — Eisiminger site rim sherds. A. Rim sherd with scalloped lip. B. Perpendicular cord-wrapped
paddle edge applied to inner lip. C. Same as “B” except that application resulted in scalloped lip. D.
Rim sherd of a second double pot.
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George — McKees Rocks Village, Eisiminger Sites
77
fine, horizontal cordmarking (Fig. 9D). This rim sherd differs from the one de-
scribed above (Fig. 5) because the interior is not carinated.
The only other vertical compound vessel that is known for the Upper Ohio
Valley is a McFate Incised vessel found associated with an adult female burial at
the Johnston site, 36IN2 (Fig. 1:5) (Dragoo, 1955:95, 136). Griffin (1966:137 &
PL LXIX) discussed two double vessels recovered from the Madisonville site,
one of which was associated with a child burial. With the paucity of data available,
one can only conclude that compound or double vessels are so rare that there are
too few of them known for anyone to undertake a study of this enigmatic yet
intriguing artifact type that appears on Proto-historic, and slightly earlier, sites.
McKees Rocks Village Site
In the “Interpretations” section of his site report, Buker (1968:46) states, “If
McKees Rocks cannot be placed satisfactorily within the framework of known
Monongahela culture, even when all the variations in Monongahela are consid-
ered, some of the answers probably lie elsewhere.” The question we still ask is
where is elsewhere? The author herein makes a few suggestions about the McKees
Rocks Village site and its possible relationships, although that question cannot be
answered with certainty.
In its present context, McKees Rocks Village site appears to have represented
an isolated village far removed from its nearest known affiliate, the Wellsburg
site (Fig. 1:11), which is 1 1 1 river km downstream on the Ohio River in West
Virginia. This might not have been so. A few artifact assemblages are of interest,
although the data are limited because the upper Ohio River and Chartiers Creek
valleys have been devastated by urbanization and industrialization.
The most significant of these is from the Georgetown site, 36BV29, located on
an Ohio River terrace in Beaver County, Pennsylvania, just east of the West
Virginia State line (Fig. 1:6). Mayer-Oakes (1955:178) excavated a series of five-
foot squares to various levels at Georgetown to help formulate pottery sequences
for the Upper Ohio Valley. Among artifacts from the site, donated to CMNH by
Emil Alam, are six rim sherds (in two restored sections) of a shell tempered vessel
with sub-lip fillet, and a notched lip (Fig. lOA); the extant neck area exhibits
faint, vertical cordmarking. The sherds’ close resemblance to a McKees Rocks
Village site rim sherd (Fig. IOC) implies its Wellsburg Phase affinities.
Another rim sherd from the Georgetown site consists of a shell tempered cas-
tellation with three sub-lip vertical punctations (Fig. lOB) on a plain surface. Both
the prominent and uniform punctations and the almost pointed castellation, similar
to those illustrated by Buker (1968:Fig. lOA, Fig. 12A, and 12D), indicate a
Wellsburg Phase origin. Unfortunately, the Georgetown site has been recently
devastated by severe flood-induced bank cave-ins as well as uncontrolled looting,
so determining the extent of any Wellsburg Phase component is highly unlikely.
The incidence of simple stamped pottery in the Upper Ohio Valley is very rare,
except for the McKees Rocks Village site. In the Chartiers Creek valley, the multi-
Late Prehistoric component Lang site, 36WH48 (Fig. 1:7), salvage excavated in
1986-87, produced nine simple stamped body sherds from three features. A single
radiocarbon date of 490±110 B.P, Cal. A.D. 1430 (Beta-19,162) is considered to
be a valid temporal marker for the Late Monongahela occupation. Shell tempered
rim sherds from the Lang site, relating to the Late Monongahela occupation, have
sub-lip punctations that are usually oblique, pseudo-collars and protruding cas-
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George — McKees Rocks Village, Eisiminger Sites
79
tellations. The Lang site simple stamped body sherds have some resemblance to
Wellsburg Phase sherds, but the decorative techniques of rim sherds are not iden=
ticaL
Another site with similar rim sherds is the Portman site, 36AL40 (Fig. 1:8),
partially excavated by Allegheny Chapter, Society for Pennsylvania Archaeology
(SPA) members and directed by William Buker in 1968-69. Like the Lang site,
Portman had multiple Late Prehistoric occupations (Buker, 1993:13). Two radio-
carbon dates of 530±90 B.R, Cal. A.D. 1412 (Uga-1643) and 560±60 B.P., Cal.
A.D. 1403 (Beta-57298) document contemporaneity with the Lang site. One sim-
ple stamped body sherd is recorded for Portman (Buker, 1993:23). It is believed
that the late components of the Lang and Portman sites, separated by only 5.5
km, represent a yet unnamed Late Monongahela phase that, in some ways, is
related to the Wellsburg Phase. Its spatial extent is presently unknown.
There are two simple stamped, shell tempered rim sherds recorded from the
Godwin-Portman site, 36AL39 (Fig. 1:12), another Chartiers Creek valley locus.
A recently obtained radiocarbon assay of 520±60 B.R, with a calibrated intercept
date of A.D. 1420 (B- 153248), may relate to the simple stamped rim sherds. The
site, which is 0.25 km north of the Portman site, was partially excavated by
CMNH in 1968 (Dragoo et ak, 1969) and salvage excavated in 1978-79 by mem-
bers of the SPA just prior to its destruction. One of the small rim sherds has
shallow punctations on the lip whereas the lip of the other is plain.
Further afield, there are four simple stamped, shell tempered body sherds from
the Johnston site, 36IN2 (Fig. 1:5), that are listed in the CMNH Field Catalog
(#760) as “simple stamped?”, but are not mentioned in the site report by Dragoo
(1955:101-105). Re-examination of the sherds by the author verifies that they are
simple stamped. The Johnston site is obviously late, based on its inclusion in the
Johnston Phase of Monongahela that, characteristically, produces both Monon-
gahela and McFate pottery (George, 1997).
Discussion
The coming of the Wellsburg Phase people to McKees Rocks Village, probably
during the late sixteenth century A.D., to one of the most prominent and defensive
locations along the upper Ohio River valley, must have created consternation
among indigenous populations. Buker (n.d.) noted that the abundant subsistence
remains at McKees Rocks suggest that the village population was large enough
to control an adequate sized catchment area. The resource-rich and accessible
Chartiers Creek Valley may not have been occupied by indigenous populations
when the McKees Rocks Village was settled. It is also especially interesting that
the generally upland area north of the site, and away from the river, is devoid of
Late Prehistoric village sites, a fact emphasized by Buker (personal communica-
tion). This area, now known to Pittsburghers as the “North Hills,” would have
provided extensive and accessible hunting grounds necessary for sustaining a
large village population.
Fig. 10. — Georgetown and McKees Rocks Village sites rim sherds. A. Shell tempered rim sherd with
a sub-lip fillet and notched lip from Georgetown. B. Shell tempered castellation with sub-lip vertical
punctations from Georgetown. C. McKees Rocks Village site shell tempered rim sherd similar to “A”
from the Georgetown site.
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Ceramic influences, such as the McKees Rocks-like decorated rims strip at
Eisiminger and the curious, but somewhat similar, pottery from Lang and Port-
man, suggest that there were degrees of cultural integration in force in south-
western Pennsylvania, and elsewhere, slightly before contact, as had been sug-
gested by other archaeologists (Pollack and Henderson, 1992:289). It appears that
the McKees Rocks Village potters had some influence on indigenous peoples
based on the seemingly random spread of simple stamped wares.
The author also suggests that there is some evidence that the Wellsburg Phase
people were Iroquoian. Buker (1968:Fig. 15K) illustrated a portion of an Iroquois
acorn-ring pipe from McKees Rocks. An Iroquoian affiliation for the Wellsburg
Phase might also be inferred by the lack of biconcave discoidals in the McKees
Rocks Village site sample since this artifact type is not present on northern Iro-
quoian sites (George, 2001:4). The McKees Rocks Village site contained trian-
gular points that are the small Madison forms one would expect from a late Late
Prehistoric/Proto-historic site in the Upper Ohio Valley. However, there is also a
sample of parallel-sided triangular points (Fig. 11 A), the type that Railey (1992:
161-163) referred to as “Type 5, Fine Triangular; Straight-Sided” in “Fort An-
cient Cultural Dynamics in the Middle Ohio Valley” (Henderson, 1992). The
latter is not the triangular point type found at the South Park Village site of
Whittlesey origin (Brose, 1994:94-98). A second point sample from McKees
Rocks Village (Fig. IIB) is what Railey (1992:156-158) designated “Fine Tri-
angular; Flared Base” and may represent an earlier Fort Ancient involvement
with the McKees Rocks Village site, possibly dated by the aforementioned A.D.
14th century radiocarbon date.
Conclusions
The separate cultural entities that are recognized by two distinct ceramic tra-
ditions, herein described, are symptomatic of the upheaval in the Upper Ohio
Valley at the beginning of the 17th century. Diminished Monongahela populations
occupied upland loci and the floodplains of minor streams well away from major
rivers. It is suspected that all of these were relatively small villages with limited
population, as exemplified by the Throckmorton site (Herbstritt, 1983). Paradox-
ically, a much stronger alien population was able to establish and maintain a large
defensive-positioned village, on a prominent escarpment, and able to control an
extensive catchment area. There is now some evidence that there were Wellsburg
Phase settlements between Wellsburg and McKees Rocks. This belies the sup-
posed spatial isolation of the latter. Unfortunately, the heavy industrialization of
the upper Ohio River valley may have eliminated other possible Wellsburg Phase
sites. The appearance of simple stamped sherds on a few sites, plus the existence
of at least one Eisiminger rim sherd with an added rim strip, implies social in-
terchange and, perhaps, a non-belligerent relationship between McKees Rocks
villagers and some indigenous people. As always, we have much to learn.
Acknowledgments
This paper would not have been possible without the substantial efforts of Bill Buker, who, with
other volunteers, excavated portions of the McKees Rocks Village site while enduring the most trying
of conditions. Such dedication is seldom properly recognized. I also thank Jim Morton of Columbus,
Ohio, who prodded us into the recognition of the Proto-historic age of the McKees Rocks Village.
James D. (Fuzzy) Randolph, Curator of the Paul R. Stewart Museum at Waynesburg College, loaned
me the artifacts and the unpublished report on the “Pottery House” at Eisiminger. Stewart’s unpub-
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George — McKees Rocks Village, Eisiminger Sites
81
0 12'
1 ^
1 5 cm
Fig. 11. — McKees Rocks Village site triangular points. A. Straight sided. B. Elongated, flared base
type.
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lished report deserves wider recognition; it is appended to this article to increase its availability.
Randolph’s interest and cooperation are greatly appreciated, as is his upbeat attitude. The author is
also appreciative of the help in the form of donated collections, provided by long-time Society for
Pennsylvania Archaeology members Emil Alam and Donald P. Tanner. I thank Carnegie Museum of
Natural History people as follows: Curator-in-Charge David R. Watters for darkroom help and editorial
assistance; retired Associate Curator Verna L. Cowin, who made editorial suggestions; and Section
Secretary Charmaine C. Steinberg for typing the various drafts of this paper. I express my sincere
gratitude to Fuzzy Randolph for permission to include Stewart’s “Pottery House” manuscript.
Literature Cited
Babich, J., N. M. Kirshner, W. C. Johnson, and T. S. Kirshner. 1996. Archaeological investigations
at the Kirshner site (36WM213), a Middle Monongahela Period village in the Youghiogheny
River Valley: An interim report. Paper presented at the Sixty-Third Annual Meeting of the Eastern
States Archaeological Federation, Huntington, West Virginia, October 24-27.
Brose, D. S. 1994. The South Park Village site and the Late Prehistoric Whittlesey tradition of
northeast Ohio. Prehistoric Press Monographs in World Prehistory, 20.
Brown, A. L. 1959. Early Maps of The Ohio Valley. University of Pittsburgh Press, Pittsburgh,
Pennsylvania.
Buker, W. E. 1968. The archaeology of McKees Rocks Late Prehistoric village site. Pennsylvania
Archaeologist, 38(1-4): 1-49.
. 1993. The Portman site (36AL40). Pennsylvania Archaeologist, 63(2):7-52.
. n.d. The Kelso site: A Drew Phase Monongahela settlement in southwestern Pennsylvania.
Manuscript on file at Carnegie Museum of Natural History, Pittsburgh, Pennsylvania.
Carskadden, J., and J. Morton. 2000. Fort Ancient in the central Muskingum valley of eastern
Ohio: A view from the Philo II site. Pp. 158-193, in The Late Prehistory of Ohio and Surrounding
Regions (R. A. Genheimer, ed.). The Ohio Archaeological Council.
Davis, C. E., and A. K. Wilkes. 1997. Phase III data recovery {at the} Sony site, 36Wml51,
Westmoreland County, Pennsylvania ER#91-4617-129. Report submitted to the Bureau for His-
toric Preservation, Pennsylvania Historical and Museum Commission, Harrisburg, Pennsylvania.
Dragoo, D. W. 1955. Excavations of the Johnston site, Indiana County, Pennsylvania. Pennsylvania
Archaeologist, 24(2):85-141 .
Dragoo, D. W., R. L. George, and D. P. Tanner. 1969. Excavations at selected archaeological sites
in the Chartiers Valley in western Pennsylvania. Report submitted to the National Park Service
on Project No. 14-10-5-950-11, Chartiers Creek Flood Control Project.
Eisert, R. W. 1993. An unusual feature type from the Chartiers Valley. Pennsylvania Archaeologist,
63(l):l-9.
George, R. L. 1983. The Gnagey site and the Monongahela occupation of the Somerset Plateau.
Pennsylvania Archaeologist, 53(4): 1-97.
. 1997. McFate artifacts in a Monongahela context; McJunkin, Johnston, and Squirrel Hill.
Pennsylvania Archaeologist, 67(l):35-44.
. 2001. Of discoidals and Monongahela: A league of their own? Archaeology of Eastern North
America, 29:1-8.
George, R. L., J. Babich, and C. E. Davis. 1990. The Household site: Results of a partial excavation
of a Late Monongahela village in Westmoreland County, Pennsylvania. Pennsylvania Archaeol-
ogist, 60(2):40-70.
Griffin, J. B. 1966. The Fort Ancient aspect: Its cultural and chronological position in Mississippi
Valley Archaeology. University of Michigan Museum of Anthropology Anthropological Papers, 28.
Henderson, A. G. (ed.). 1992. Fort Ancient cultural dynamics in the Middle Ohio Valley. Prehistory
Press Monographs in World Archaeology, 8.
Herbstritt, j. T. 1983. Excavation of Two Monongahela Sites: Late Woodland Gensler (36GR63)
and Proto-Historic Throckmorton (36WM160). Report submitted to Consolidated Coal Company,
Pittsburgh, Pennsylvania.
. 1984. The mystery of the Monongahela culture. Pennsylvania Heritage, 5(3):26-3L
Johnson, W. C. 1990. The Protohistoric Monongahela and the case for an Iroquois connection. Paper
presented at the Fifty-Seventh Annual Meeting of the Eastern States Archaeological Federation,
Columbus, Ohio, November 8-1 1.
Jones, V. H. 1968. Corn from the McKees Rocks Village site. Pennsylvania Archaeologist, 38(1-4):
81-86.
Lang, R. W. 1968. The Natural Environment and Subsistence Economy of the McKees Rocks Village
site. Pennsylvania Archaeologist, 38(l-4):50-80.
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Lapham, H. a. 1995. The analysis of European glass trade beads recovered from Monongahela sites
in Greene County, Pennsylvania. Report submitted to The Bead Society of Greater Washington,
Washington, D. C.
Mayer-Oakes, W. J. 1955. Prehistory of the Upper Ohio Valley. Annals of Carnegie Museum, 34.
Pollack, D., and A. G. Henderson. 1992. Toward a model of Fort Ancient society. Pp. 281-294,
in Fort Ancient Cultural Dynamics in The Middle Ohio Valley (A. G. Henderson, ed.). Prehistoric
Press Monographs in World Archaeology, 8.
Railey, J. a. 1992. Chipped stone artifacts. Pp. 137-169, in Fort Ancient cultural dynamics in the Middle
Ohio Valley (A. G. Henderson, ed.). Prehistoric Press Monographs in World Archaeology, 8.
Stewart, P. R. n.d. Excavation of Pottery House. Portion of the “Indian Archaeology” manuscript
on file at Paul R. Stewart Museum, Waynesburg College, Waynesburg, Pennsylvania.
Appendix
While reviewing “Comparing pottery from the Proto-historic McKees Rocks
Village and Eisiminger sites of southwestern Pennsylvania,’’ David R. Watters,
Curator of Anthropology and the Anthropology Editor for Scientific Publications,
suggested incorporating the following unpublished manuscript, entitled “Exca-
vation of Pottery House” by Dr. Paul R. Stewart, since the information therein
contributes significantly to an understanding of the Eisiminger site. Stewart’s early
research at the Pottery House assumes even more importance because of James
Herbstritt’s excavation of the Proto-Historic site (36Grl60) in 1982, where settle-
ment pattern details seemingly duplicate those at Eisiminger,
The “Excavation of Pottery House” is a part of Stewart’s undated, sixteen-
page manuscript entitled “Indian Archaeology,” an important document, com-
pleted in the middle 1950s, that is on file at the Paul R, Stewart Museum of
Waynesburg College. Permission to publish the “Excavation of Pottery House”
portion of the manuscript was granted by James D. Randolph, Director of the
Stewart Museum. On the last page of the “Indian Archaeology” manuscript is
this statement:
“Unfinished manuscript by Dr. Paul R. Stewart who, with James D.
Randolph, made this study. The illustrations referred to in the article
were never made. The pottery pieces are in the Waynesburg College
Museum.”
The content of the manuscript is reproduced exactly as it appears in the original,
including blank spaces ( ) where data or other information were not in-
cluded in the original.
EXCAVATION OF POTTERY HOUSE
[by Paul R, Stewart]
THE PROBLEM
A village site, which for the present we will call the Eisiminger focus of the
Monongahela Woodland culture in harmony with the suggestions made by Mary
Butler in Bulletin No. 753 of the Pennsylvania Historical Commission, lies ap-
proximately three and one-half miles south of Waynesburg, Pennsylvania, three-
fourths of a mile east of the Waynesbug-Blacksville road. The site occupies por-
tions of two fields on what is now known as the George Eisiminger property.
Figure 1 shows the contoured map of the village site.
The site was partially plowed in the summer of 1950 and completely plowed
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VOL. 71
in the summer of 1951. Previous to the year mentioned, a large number of des-
ultory excavations had been made on the south side of the spot marked “Lapping
Cemetery.” A fence had previously divided the field into a north and south portion
just to the south of this cemetery. When the fence was taken away and the entire
village site plowed, the author with a number of his students undertook to explore
a few pits disclosed by probing just to the northwest of the former Lapping
Cemetery.
In the summer of 1950, Mr. William Mayer-Oakes and James Swauger of the
Carnegie Museum, had been taken to the site by the author, Mr. Mayer-Oakes
continued a superficial examination through the summer of 1950. Our conclusions
were that this site represented an occupation of the Monongahela Woodland cul-
ture mentioned above. Certain artifacts of brass and copper indicated European
settler contact. Fort Ancient culture never seemed definitely to be established. The
excavations at the locality mentioned above were for the purpose of further de-
termining the type of culture, [end of page 1 in original]
THE EXCAVATION
Upon the excavation of the first pit probed, it was discovered that we were not
exploring a simple firepit, but a complex situation disclosed in Figures 2 and 3,
where it will be observed that instead of separate firepits, we were in fact ex-
ploring an ancient house which we have been pleased to call “Pottery House.”
As will be seen by the drawing, the structure was approximately 18 feet long
by 12 feet wide, almost rectangular, and the main floor almost completely paved
with small stones ranging from 2 or 3 square inches to 50 or 60 square inches,
the average being approximately 10 square inches. These stones for the most part
showed no indication of firing, but had evidently been taken from a sandy shale
which underlies most of the village site. Here and there among the pavement
stones were other stones showing the hematitic red caused by firing. The stones
had probably been tramped in between the flatter stones of the main pavement.
In a very few places the pavement was double as if certain of the stones had
sunk and the inhabitants had tramped a few extras in above them. The entire
pavement was laid in wood ashes.
This pavement was approximately level and had obviously been graded back
into the slight slope, giving approximately a depth of 2 1/2 [two and one half]
feet at the upper end and 1 foot at the lower side.
In addition to what we call the main structure (20 feet by 12 feet) there were
a number of lobes of from 2 or 3 inches to as much as 8 or 10 inches below the
paved surface. Into these pits the pavement extended downward from the main
pavement into the lobes. There were six of these lobes, which were doubtless
firepits, the ashes being much deeper in them than over the main structure. These
pits were from two to three feet across and were found, as Figure 4 shows, all
around the [end of page 2 in original] structure except at the southwest, where
presumably the entrance had been established.
Smaller fire pits were found near the structure and were assumed to be related
to it. On the upper side of the pavement (See Figure 3) an embankment of ex-
cavated fragments of shaly sandstone had been piled and later, presumably after
the structure was deserted, had caved in.
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George — McKees Rocks Village, Eisiminger Sites
85
ARTIFACTS
One of the noteworthy observations was the almost complete absence of arti-
facts except pottery. Two arrowheads, one notched and the other triangular, were
found (See Figure 5). One small bone awl (animal species undetermined) and
another awl made from the bone of an were excavated. One slightly
decorated bit of mussel shell was found and one very well-formed flaking imple-
ment. Added to these artifacts was the small roughly-rolled bit of copper kettle
of settler origin. This was found in such a position that it could possibly have
been intrusive.
DESCRIPTION OF THE POTTERY
The outstanding feature of this dwelling was the amazing wealth of potsherds.
This led to our naming it the “Pottery House.” rimsherds were found in
the various pits and scattered over the floor. Usually more or less wood ash was
found over the potsherds whether they happened to be in the firepit lobes or on
the main floor. Approximately sherds other than rimsherds were found in
the dwelling, not counting the sherds which were found in sufficient number to
reconstruct vessels. Figure 6 shows a group of the most interesting rimsherds.
Attention is called to rimsherds a, b, c, d, e, which were definitely crenate, as were
rimsherds used in reconstructing vessel No. shown in Figure In
surface discoveries, no crenate rimsherds had been found previously by the author
or by Mayer-Oakes.
Most noteworthy of all the sherds were Nos. f, g, h, which neither the author
or [sic] Mayer-Oakes had ever seen before in any of the village sites [end of page
3 in original] examined in this area. Evidently a secondary flange was modeled
part of the way down from the rim of the pot. Unfortunately, the primary rim
could not be discovered in any of these three unusual sherds. From the rapid
thinning of the pottery above the secondary ledge, it was estimated that this vessel
probably held in the neighborhood of one and a half pints or a quart.
A noteworthy feature of the pottery in Pottery House is the fact that not one
of the rimsherds had any notch crease or lip designed for facilitating
pouring, although several such devices had been found in the surface finds of the
site.
There were sufficient sherds to reconstruct with confidence two complete pot-
tery vessels shown in Figure 7. The left shows interrupted cord shaping with
either cylindrical or flat corded paddle. To the right was the most important find
of all. A little more than three-fourths of the entire rim was found and was
beautifully and rather regularly crenate with a slight twist such as is used by
modern housewives in edging a pie crust. The sides of the vessel had been fash-
ioned with rather continuous diagonal strokes of a corded paddle, averaging about
— degrees departure from the edge of the pot. Toward the bottom of the
bulge of this pot, the cording had been changed to cross as well as oblique strokes.
This pot had been extremely well-fired because the potsherds were practically
unweathered and when dried made a clinking sound when stroked. This was in
contrast with the pot represented by rimsherd No. j, which had almost exactly the
same design, but in this case the pot had been very poorly fired so that most of
its sherds have weathered beyond the possibility of much reconstruction, [end of
page 4 in original]
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VOL. 71
Note: (Excavation)
No well-defined post holes were found although along the upper side where a
trench, possibly for drainage, had been dug, there were lower spots in two of
which were there were found fairly large stones which might have been used in
wedging saplings in place. The question, of course, arises whether this was a
wigwam or hogan type made by bending poles over the structure, and roofing
and siding it with bark and latticed branches, or whether advantage had been
taken of trees growing in the neighborhood, [end of page 5 in original, except
for the “unfinished manuscript by Dr. Paul R. Stewart ...” quotation that appears
before the text in the Appendix of the present paper]
ANNALS OF CARNEGIE MUSEUM
VoL. 71, Number 2, Pp. 87-106
28 May 2002
FLEAS (SIPHONAPTERA) FROM ANCASH DEPARTMENT, PERU WITH
THE DESCRIPTION OF A NEW SPECIES, ECTINORUS ALEJOI
(RHOPALOPSYLLIDAE), AND THE DESCRIPTION OF THE MALE OF
PLOCOPSYLLA PALLAS (ROTHSCHILD, 1914) (STEPHANOCIRCIDAE)
Michael W. Hastriter*
Research Associate, Section of Invertebrate Zoology
Michael D. Zyzak^
Ruler Soto^
Roberto Fernandez^
Nelson Solorzano^
Michael E Whiting'’
Abstract
A collection of 358 fleas representing 6 families, 19 genera, and 21 species was made from eight
species of mammals in seven provinces within Ancash Department, Peru. Specimens were also ob-
tained from bank cavities and water flea traps. A new species of flea, Ectinonis alejoi (Rhopalopsyl-
lidae) collected from Lagidium peruana Meyen, 1833 (vizcacha) is described from Recuay Province,
bringing the total number of species belonging to the genus Ectinorus to 32. The male of Plocopsylla
pallas (Rothschild, 1914) (Stephanocircidae) is also described for the first time. Plocopsylla pallas
(16%), Neotyphloceras crassispina hemisus Rothschild, 1914 (14%), and Cleopsylla townsendi Jordan,
1936 (9%) comprised 39% of the total number of fleas collected. Three fleas are reported from Peru
for the first time (D. stejnegeri, P. achilles, and E. alejoi). Eleven additional species represent new
records for Ancash Department (A. thurmani, C. townsendi, C. m. minerva, E. gallinacea, E. claviger,
N. c. hemisus, P. pallas, P. d. quitanus, S. inca, T. bleptus, and T. titschacki).
Key Words; Ectinorus alejoi, Ectinorus claviger, Plocopsylla pallas, flea, Siphonaptera
Introduction
The flea fauna of South America is drastically understudied and Peru is no
exception. Johnson’s (1957) monographic work of the South American flea fauna
comprises most of the known Peruvian records, which include 59 recognized taxa.
Four additional Peruvian taxa were recorded by Hopkins and Rothschild (1966),
ten by Smit (1970, 1976, 1978, and 1987), and three by Schramm and Lewis
(1987, 1988), providing a total of 76 taxa. Our report provides a record of flea
' Monte L. Bean Life Science Museum, Brigham Young University, 290 MLBM, RO. Box 20200,
Provo, Utah 84602-0200, e-mail: hastritermw@sprintmail.com.
2 Uniformed Services University of the Health Sciences, Department of Preventive Medicine and
Biometrics, 4301 Jones Bridge Road, Bethesda, MD 20814.
3 Naval Medical Research Center Detachment, Lima, Peru, APO AA 34031.
Av. 9 de Octubre, S/N Hospital de Caraz, Caraz-Ancash-Peru.
^ Department of Zoology, 574 Widtsoe Building, Brigham Young University, Provo, Utah 84602.
Submitted 9 November 2001.
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taxa recently collected in Ancash Department. Prior to this study, only 10 species
(including cosmopolitan species) were recorded in Ancash Department. Miscel-
laneous fleas were opportunistically collected in September 1998 by Dr. Pat Car-
ney, Uniformed Services University of the Health Sciences (USUHA) during bio-
medical research studies. They were also collected during a medical entomology
training program conducted 2-6 September 1999 by personnel of USUHS, Meth-
esda, MD in cooperation with the Ministerio de Salud, Hospital de Caraz (USUHS
Team). Following the 1999 training program, medical technologists from several
provincial hospitals also submitted fleas. During 23-30 March 2000 hantavirus
studies were conducted (Hastriter Team) and fleas and tissue samples were col-
lected (results of the hantavirus investigations are published elsewhere). Among
our material are a new species of Ectinorus Jordan, 1942 (Rhopalopsyllidae) and
the undescribed male of Plocopsylla pallas (Rothschild, 1914) (Stephanocircidae).
A review of the genus Ectinorus was provided in Hastriter (2001). The description
of one additional species of Ectinorus brings the total number of species in this
genus to 32. This paper presents and discusses several new records for Ancash
Department and examines the current species known to occur in Peru to include
three new records for the country.
Materials and Methods
During the September 1998 collections and those made during the September
1999 training program, collection activities were confined to agricultural areas
along the Santa River near Choquechaca (Hauzlas Province). Vegetation included
bamboo thickets in wet areas along the river, and Agave sp. and numerous vari-
eties of cacti along the valley floor away from the river. Special emphasis was
placed on trapping in and around human dwellings and along rock fences ex-
tending through fields (elevation: 2100-2200 m). Habitats selected during the flea
and hantavirus research study (March 2000) included the following: 1) Chiquian
Province (3600-4000 m) — High elevation grassland known as “puna” consisting
of large fescue (Festuca orthophylla Pilger, 1898) and needle grass {Stipa ichu
Kunth, 1829). 2) Huaraz Province (1800-3700 m) — steep rocky slopes with low
growing vegetation. 3) Huaylas Province (3300-3700 m) — Parque Nacional
Huascaran was characterized by steep rocky slopes, heavily vegetated with low
growing shrubs. Region was small plot agriculture and sheep grazing. 4) Recuay
Province (3600-4400 m) — the lower elevations were over-grazed by sheep and
the upper regions were abundantly covered with F. orthophylla and S. ichu and
small scattered boulders. Habitats in Pomabamba and Huari Provinces were not
observed by the authors.
Standard aluminum collapsible Sherman traps baited with oatmeal or oatmeal
and peanut butter were used to trap small mammals. Trapped mammals were
shaken into cloth or plastic bags at each capture site and transported to a field
processing location. Each animal was brushed briskly with a toothbrush over a
white bucket to capture the fleas. Fleas were placed in 80% ethanol, returned to
the laboratory and prepared by soaking in 10% KOH for a period (—24 hours),
dehydrated in serial alcohols (70, 80, 95 and 100%, a minimum of 30 minutes
each), cleared in methyl salicylate (maximum of 20 minutes), and xylene (mini-
mum of one hour). Fleas were mounted on glass microscope slides in Canada
balsam using 12 mm cover-slips (#1 thickness). Mammals were euthanized by
cervical dislocation. Each was weighed, measured, preserved in 10% formalin.
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and later identified. Miscellaneous fleas were collected by hand-picking nesting
materials or substrates from earthen bank nest cavities. Water flea traps were used
to capture fleas in human dwellings. Traps were comprised of a shallow plastic
pan (2.5 cm deep X 45 cm wide) with a kerosene-wicked can set in the middle.
Water was added to the pan and the wick lighted. Fleas attracted to the burning
lantern were captured in the mote of water. Unless otherwise noted, all specimens
were collected in Ancash Department, Peru.
Total body lengths of males and females were measured from the foremost
portion of the frons to the apex of the st. VIII in males and to the posterior border
of the sensilial plate in females. The measurement of each sex is expressed as the
means of the composite number of specimens measured. Illustrations were pre-
pared with the aid of a camera lucida mounted on a Zeiss compound microscope.
With exception of the term processus basimeris ventralis adopted from Smit
(1987), the terminology of flea structures follows those of Rothschild and Traub
(1971).
Results
Ceratophyllidae
Dasypsyllus stejnegeri (Jordan, 1937)
Specimens Examined. — Recuay Province: 19 km W Recuay (9°44'6"S, 77°30'3"W), 4420 m, ex
“soil from nest cavity” of Colaptes rupicola Orbigny, 1840 (= Andean flicker) in earth bank, 29
March 2000, Hastriter Team, 3$; and road cut on highway 109 between Chiquian junction and Pav-
ilotica (10°9'36"S, 77°19'43"W), 3980 m, ex C. rupicola nest cavity in vertical road cut, 1 April 2000,
M. W. Hastriter and R. Soto, 1 S .
Remarks. — This flea is distributed from the Bering Straits of Alaska to the
southern tip of South America on a wide variety of avian hosts, particularly
Passerine land-birds (Smit, 1961). This is the first record of its occurrence in Peru
and on a species of Colaptes. Since little is known about the bionomics of this
flea, it seems prudent to record our observations during its collection. The adult
host bird was seen exiting the earth nest cavity (opening: —18 cm diameter, depth:
—60 cm) on several occasions. A ladder was acquired to access the nest 3.25 m
above the ground. Five hatchlings were present and fleas were not noted on them.
Approximately one liter of dry, course sand was removed from the earthen nest
cavity and was protected from the sun until examined. Each of the three female
fleas was rather inactive, failing to crawl or jump when probed; however, each
became very agitated when placed in alcohol. In comparison, the single male
removed from the nest cavity in the road cut along highway 109 was collected
from very fine moist soil. There were multiple holes in the road cut that were
made by C. rupicola, but few were accessible. It should be noted that the localities
of the two collection sites (within the same province) were separated by more
than 70 km. This would suggest that this flea is established and that C. rupicola
may be a significant host species throughout high elevations in the South Amer-
ican Andes. Smit (1961) suggested that long-range flying sea birds are responsible
for the wide distribution of this flea. This does not seem plausible as there is little
association between Passerine birds and sea birds and the species has never been
collected from sea birds. Our finding of this species in Peru has great significance
in demonstrating an established population between previously disjunct popula-
tions in northwestern South America and the Falkland Islands. Its presence in
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VOL. 71
central Peru supports the theory of distribution by “leap-frogging” by migrating
Passerine land birds as proposed by Traub, Rothschild, and Haddow (1983).
Plusaetis dolens quitanus (Jordan, 1931)
Specimens Examined. — Huaraz Province: Pariacota (9°33'10"S, 77°35'69"W), ex Phyllotis andium
Thomas, 1912, 24 March 2000, Hastriter Team, 23, 19. Huaylas Province: Parque Nacional Huas-
caran (9°30'40"S, 77°28'32"W), 3385 m, ex Phyllotis amicus Thomas, 1900, 25 March 2000, Hastriter
Team, 23, 19; Parque Nacional Huascaran (9°30'24"S, 77°27'69"W), 3475 m, ex P. andium, 25 March
2000, Hastriter Team, 29; Parque Nacional Huascaran (9°30'40"S, 77°28'32"W), 3385 m, ex Akodon
sp2 cf. mollis, 25 March 2000, Hastriter Team, 23. Pomabamba Province: Huayllan, ex Mus mus-
culiis Linnaeus, 1758, 17 February 2000, A Lopez, 13. Recuay Province: 4.6 km W Recuay
(9°43'36"S, 77°27'917"W), "-3660 m, ex Akodon spl cf. mollis, 29 March 2000, Hastriter Team, 39;
4.6 km W Recuay (9°43'36"S, 77°27'55"W), —3660 m, ex P. andium, 29 March 2000, Hastriter Team,
29.
Remarks. — The lateral aspect of t. VIII of males bears an anterior vertical row
of two setae and a vertical row of three just posterior to these. The telomere is
distinctly rounded along caudal margin opposed to that of P. equaotris (Jordan,
1933).
These records represent the most southern distribution of P. d. quitanus. John-
son (1957) elaborated on the distribution of earlier records approximately three
degrees north in Huancabamba Province, Department of Piura, Peru and material
from Ecuador (the type locality for the subspecies). It should be noted that Mac-
chiavello (1948) alluded to P. equatoris as an important vector of plague in rural
areas of northern Peru, but his specimens probably were P. d. quitanus.
Ctenophthalmidae
Neotyphloceras crassispina hernisus Jordan, 1936
Specimens Examined. — Chiquian Province: E of Chiquian, (10°9'54"S, 77°19'42"W), 3900 m, ex
Akodon spl cf. mollis, 27 March 2000, Hastriter Team, 1 9 ; E of Chiquian, (10°06'07"S, 77° 1 L07"W),
4190 m, 27 March 2000, Hastriter Team, 19; E of Chiquian, (10°08'06"S, 77°10'22"W), -3660 m,
27 March 2000, Hastriter Team, 1 9 . Hauzlas Province: Santa River Valley, Choquechaca (N of
Caraz), 2195 m, ex Oryzomys xantheolus Thomas, 1894, 3-4 September 1999, USUHS Team, 43,
5 9 ; ex Thomasomys sp., 4 September 1999, USUHS Team, 1 3, 29 ; ex P. andium, 2 September 1999,
USUHS Team, 23, 79; ex Akodon mollis Thomas, 1894, 3-4 September 1999, USUHS Team, 13,
1 9 ; ex O. xantheolus, 21 September 1998, R Carney, 1 9 ; ex A. mollis, 25 September 1998, P. Carney,
1 9; 9 km N Caraz (9°00'12"S, 77°49'36"W), 2440 m, ex P. andium, 30 March 2000, Hastriter Team,
53, 39. Huaraz Province: Pariacota (9°33'27"S, 77°38'82"W), 3630 m, ex P. andium, 24 March
2000, Hastriter Team, 13, 39; Pariacota (9°33'10"S, 77°35'69"W), 3715 m, ex P. andium, 24 March
2000, Hastriter Team, 1 9. Huaylas Province: Parque Nacional Huascaran (9°30'04"S, 77°25'3T'W),
3630 m, ex Akodon spl cf. mollis, 25 March 2000, Hastriter Team, 13. Recuay Province: 10 km W
Recuay (9°43'25"S, 77°29'13"W), 4010 m, ex P. andium, 29 March 2000, Hastriter Team, 13, 19;
19 km W Recuay (9°44'06"S, 77°30'03"W), 4420 m, ex Akodon spl cf. mollis, 29 March 2000,
Hastriter Team, 29; N of highway 109 between Chiquian junction and Pavilotica (10°9'14"S,
77°19'17"W), 3990 m, ex P. andium, 28 March 2000, Hastriter Team, 39.
Remarks. — This ubiquitous species is found on a variety of hosts throughout
its range and occurred at low (2195 m) and high (4420 m) elevations in all areas
sampled in Ancash Department. The apex of the basimere has been used as the
major feature to distinguish between the various subspecies of Neotyphloceras.
This character is variable and unreliable. Males of this subspecies may readily be
distinguished from N. c. chilensis Jordan, 1936, and N. rosenbergi (Rothschild,
1904) by the presence of spicules adorning the surface of the anterior sclerite
connecting the left and right basimeres. These structures become visible when
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focusing beyond the outer surface of the base of the basimere. They are also
present in specimens of N. c. crassispina Rothschild, 1914 from Concepcion,
Chile. Neotyphloceras c. hemisus may be a synonym of the nominate subspecies,
but the genus requires a comprehensive study for a definitive conclusion.
Leptopsyllidae
Leptopsylla segnis (Schonherr, 1811)
Specimens Examined. — Hauzlas Province: Santa River Valley, Choquechaca (N of Caraz), 2195
m: ex M. musciilus, 2 September 1999, USUHS Team, 1$; ex M. musculiis, 4 September 1999,
USUHS Team, AS, 69. Pomabamba Province: Huayllan, ex M. musculus, 17 February 2000, A.
Lopez, 5(5, 59.
Pulicidae
Ctenocephalides felis felis (Bouche, 1835)
Specimens Examined. — Hauzlas Province: Santa River Valley, Choquechaca (N of Caraz), 2195
m, ex domestic dog, 5 September 1999, USUHS Team.
Remarks. — Hundreds of C. f. felis were hand-picked from domestic dogs by
the students during the training program and countless others were trapped with
water flea traps. An account of the number of specimens was not maintained.
Echidnophaga gallinacea (Westwood, 1875)
Specimens Examined. — Hauzlas Province: Santa River Valley, Choquechaca (N of Caraz), 2195
m, water flea trap, 3 September 1999, USUHS Team, 35, 29.
Remarks. — It is interesting that these stick-tight fleas were actively ambulatory
and possibly attracted to the light of the water flea traps.
Hectopsylla suarezi C. Fox, 1929
Specimens Examined. — Huari Province: Progreso, ex Cavia porcellus (Linnaeus, 1758), 14 April
1999, D. Jodulhe, 19.
Remarks. — Most common hosts include caviid rodents, C. porcellus and Cavia
aperea Erxleben, 1777. Hiickinghaus (1961) considered C. porcellus as a syno-
nym of C aperea. This flea may support that conclusion, since it parasitizes both
taxa and peridomestic rodents associated with these caviids. Although this flea is
not often collected, its broad distribution is probably attributed to the common
transport and trade of C. porcellus throughout the Andes.
Pulex irritans Linnaeus, 1758
Specimens Examined. — Hauzlas Province: Santa River Valley, Choquechaca (N of Caraz), 2195
m, water flea trap, 3 September 1999, USUHS Team, 35, 19.
Remarks. — Johnson (1957) recognized only P. irritans in South America,
whereas Smit (1958) provided morphological justification that both P. irritans
and P. simulans Baker, 1895 occur in North America. The senior author has
examined material from localities adjacent to Lima (not reported in this study)
which includes specimens comparable to North American P. simulans. Species
belonging to Pulex are common around human habitations throughout Peru and
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VOL. 71
have been implicated in plague transmission in the highlands of northcentral Peru,
particularly in the high Andean valleys in the Department of Cajamarca. Detailed
studies of large series of this genus from diverse areas in South America are
needed to determine if the complex is comprised of P. irritans, P. simulans, and
possibly undescribed species, as alluded to by Smit (1958).
Tunga penetrans Linnaeus, 1758
Specimens Examined. — Hauzlas Province: Santa River Valley, Yuracota, (near brick making bat
cave, N of Caraz), 2225 m, ex domestic dog, 20 March 2000, R Lima, Id, 19. Pomabamba Province:
Pomabamba, “in abandoned house,” 9 January 1999, J. Valverde, 4d, 69.
Rhopalopsyllidae
Ayeshaepsylla thurmani (Johnson, 1957)
Specimens Examined. — Huaraz Province: (9°32'32"S, 77°47'50"W), 1860 m, ex P. andium, 23
March 2000, Hastriter Team, 1 9 .
Remarks. — Ayeshaepsylla was erected as a subgenus of Polygenis by Smit
(1987) and later elevated to genus by Linardi and Guimaraes (1993). Accordingly,
the genus Ayeshaepsylla is distinct from all other genera within the subfamily
Rhopallopsyllinae by the vestigial nature of the posterodorsal extension of the
metasternal furca that does not extend more than one-half the length of the me-
tapleural ridge. Smit (1987) incorrectly changed the spelling to “thurmanni” in
all reference to this species. To avoid future perpetuation of this error, the correct
species designation is A. thurmani, after Deed C. Thurman, Jr. (Johnson, 1957).
Several features of our female differ from those of A. thurmani illustrated by
Johnson (1957). The dorsal lobe along the caudal margin of st. VII is longer and
more acute and the subtending sinus much deeper. Sternum VII has 5 setae per
side instead of 4 and the vental margin of the spermatheca is angular rather than
smoothly concave. Johnson (1957) reports the length of the allotype as 3.0 mm,
whereas our specimen is about half that length (1.6 mm). Based on these varia-
tions, our specimen may prove to be a new species; however, a series to include
accompanying males is needed to evaluate this taxon.
This rarely-collected flea has been reported only from two disjunct localities in
southwestern Colombia (8c5, 7 9) and southern Peru (4d, 69X Since our new
record lies about half way between these localities, additional collecting will prob-
ably yield additional specimens throughout intermediate elevations (1465-1920
m) in Peru and Ecuador. Data are insufficient to establish host preferences for
this species; however, Linardi and Guimaraes (1993) consider Cricetine rodents
to be the primary hosts.
Ectinoriis (Ectinorus) alejoi Hastriter, new species
(Fig. lA-H, 2A)
Type Material. — Recuay Province: W of highway junction 3 and 109 (~10°03'S, 77°22'W), ~41 15
m, ex Lagidium peniamim Meyen,1833, 29 March 2000, T. Alejo, 126, 29. The holotype, allotype,
and 5 paratypes (56) are deposited in the Carnegie Museum of Natural History (CMNH), Pittsburgh,
Pennsylvania, and 7 paratypes (66, 19) are deposited in the senior author’s collection.
Diagnosis. — For those species that have no marginal spinelets on t. I, it most
closely resembles E. viscachae (Wagner, 1937) in the male by details of the clasp-
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Fig. 1. — Ectinorus alejoi n. sp. A. Male clasper (basimere and telomere), holotype. B. Male ninth
sternites, (L=left side, R=right side). C. Male eighth sternum, paratype. D. Apex of aedeagus, retract-
ed, paratype. E. Apex of aedeagus, protracted, holotype. F. Female seventh sternum, allotype. G. Bursa
copulatrix, allotype. H. Spermatheca, allotype. Ectinorus claviger. I. Bursa copulatrix. J. Female sev-
enth sternum. Scale = lOOp,.
er, St. VIII, and st. IX and in the female by the modified abdominal segments and
the shape of the spermatheca. It may be distinguished in the male by the presence
of a processus basimeris ventralis (Fig. lA), the caudal margin of the telomere is
convex throughout and the apex is only slightly broader than body of telomere,
an extended lobe (heel) is absent at the juncture of the proximal and distal arms
of St. IX, and the posterior apical margin of st. IX has only 3-4 stout setae (Fig.
IB). Females may be separated from E. viscachae by the presence of an isolated
lateral patch of 5-6 small setae on st. II {E. viscachae has a patch of many setae
merging with the ventral setae) and the perula of bursa copulatrix is strongly
reflexed and globular (Fig. IG). Females also differ from those of Ectinorus
claviger (Rothschild, 1914) by the presence of more than 20 trichobothria on the
sensilium.
Description. — Head, male holotype (Fig. 2A): Frontal tubercle close to oral angle by less than its
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VOL. 71
Fig. 2. — Ectinonis alejoi n. sp. A. Head and thorax, holotype. Plocopsylla pallas. B. Male seventh
and eighth tergites. C. Aedeagus. D. Male ninth sternum. E. Male clasper (basimere and telomere). F.
Enlargement of telomere, mesal view. G. Spermatheca. H. Female seventh sternum. Scale = 100|jl.
vertical width. Two placoid pits along frontal margin. Ocular setae 3, with single seta anterior to eye
(single seta lacking in female). Eye large and darkly pigmented with sclerotized ridge extending to
margin of gena associated with visible arch of tentorium. Three setae along lower margin of genal
lobe and 2 minute setae at apex. Postantennal area with 3 rows of setae per side (1, 1,5), intercalaries
in posterior row. Occipital groove as deep as width of first maxillary palpus. Third segment of max-
illary palpus only half length of other segments. Labial palpus of 5 segments extending to apex of
fore coxae, each of the 3 middle segments less than half the length of either segment 1 or 5; maxillae
sharply pointed apically. Antennal fossa bordered above by row of 21-22 minute setae. Scape with
apical fringe of 7 setae, pedicel with 2 very short setae, and clavus long, extending well onto prosterno-
some. Thorax (Fig. 2A): notal segments each with 2 rows of setae (anterior row of pronotum of only
2 dorsal setae per side). Mesonotal collar with 6-7 pseudosetae per side (variable in number in some
paratypes). Mesopleuron with 5 setae; mesosternum vestigial. Lateral metanotal area with 2 setae (1
large, 1 small). Pleural arch well developed. Metepisternum with single seta. Dorsal margin of me-
tasternum inconspicuous, ventral lobe not extending downward. Metepimeron with anterior vertical
row of 2 setae and posterior row of 3 setae, all below level of the foot-shaped spiracular atrium. Legs:
Fore coxa with 18-20 lateral setae, mid and hind coxae with numerous setae on mesal surface of
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anterior margins and lateral sulcus of mid coxa at most indicated by notch at posterior apical margin.
Pair of setae guarding femoral-tibial joint of fore femur about equal in length, lateral setae shorter of
pair on mid and hind femora. A single seta present over femoral pit. Surface sculpturing absent on
all femora. Mid and hind femora each with single lateral row of 5-6 setae. Tibiae each with 6 dorsal
notches and each with corresponding vertical row of 5-6 setae (female with 7 dorsal notches on hind
femur). Second segment of fore tarsus longer than each of segments 1, 3, and 4. Two apical bristles
of second segment of hind tarsus extending well beyond base of distitarsomere. Four lateral plantar
bristles on each segment, 2 preapical plantar bristles of fore and mid distitarsomeres spiniform but
less so on hind distitarsomere (1 spiniform and 1 hair-like seta on all distitarsomeres of female).
Unmodified Abdominal Segments; Tergum 1 with 2 rows of setae, anterior comprised of 1-2 setae
per side, t. II-VII each with single row of setae with intercalaries (female with 2 rows on each tergite).
Each row with one seta below level of very small spiracles (female has 1 or 3 below level of each
spiracle). Single marginal antesensilial bristle borne on tubercle. Lateral patch of 2-4 setae on st. II
and single seta per side on main row (female with lateral patch of 5-6 setae and 3 per side in main
row); St. III-VII each with 2-3 setae per side (female with two rows of 4-5 setae per side on st. III-
IV, two rows of 1 and 5 setae per side on st. V-VII). Modified Abdominal Segments, male (Fig. lA-
C): Tergum VIII reduced, without setae. Sensilium with 24-26 trichobothria per side. Manubrium of
t. IX narrow, parallel-sided, and turned upward at apex. Basimere nearly square with processus basi-
meris ventralis (see remarks) at ventral-caudal margin bearing 2 apical setae. Acetabular bristle single;
numerous long setae along periphery of basimere. Telomere slightly extending beyond basimere,
parallel-sided, and slightly convex along posterior margin (Fig. 1 A). Sternum VIII bilobed, with lateral
lobe bluntly rounded and medial lobe acutely pointed; lateral portion with 5-6 setae (Fig, IC). Distal
lobe of st. IX club-shaped; lobe of each side asymmetrical. The right distal arm bears a sclerotized
tooth on mesal side of apex not present on left side (Fig. IB). Caudal margin with 4 slender setae
and lateral area with scattered setae. Sternum IX lacks tendon. Aedeagus (Fig. ID-E): Aedeagal
apodeme broadly spatulate, bluntly rounded at apex, and with narrow neck preceding fulcral medial
and lateral lobes. Penis rods extend around apex of aedeagal apodeme. Orifice of sclerotized inner
tube oblique in lateral view; ringed at basal 14. Median dorsal lobe hooked apically with short blunt
disto-lateral lobes. Alpha portion of Ford’s sclerite “C”-shaped, heavily sclerotized. Securifer of Ford’s
sclerite claw-like with small tooth on anterior margin. Crochet sclerotized and rod-like. Paired ventral
lobes narrow and hook-like. Modified Abdominal Segments, female (Fig. IF— H): Tergum VIII with 4
setae above spiracle, lateral curved row of 5 long setae and 12 scattered lateral setae. Dorsal caudal
lobe of t. VIII conical with group of many setae both mesally and laterally. Conical lobe subtended
by broad sinus with 4 evenly spaced marginal setae. Caudal margin of st. VII with rounded lobe; 5-
6 lateral setae per side and anterior group of 5-6 smaller setae (Fig. IF). Sternum VIII broad, apically
rounded lobe without setae. Bursa copulatrix sclerotized anteriorly, strongly reflexed caudally; perula
globular (Fig. IG). Bulga of spermatheca nearly spherical; duct emerging at apex. Hilla nearly twice
length of bulga; slightly wider distally than proximally. Cribriform area expanded over half of bulga
(Fig. IH).
Length (slide mounted specimens). — Male: 2.0 mm (n=7, range= 1.9-2.2 mm);
Female: 2.9 mm (n=2, range=2. 7-3.0 mm).
Etymology. — This species is named after the collector, Teophilo Alejo, a local hunter and respected
conservationist in Recuay Province.
Remarks. — Specimens of E. alejoi collected from a single host animal were
found in association with a series of T. titschacki. The rarity of T. titschacki in
collections may be a reflection of the infrequent examination of its preferred host,
L. peruanum. It is predictable that E. alejoi occurs throughout the narrow high
elevation range of this chinchillid rodent, just as that of T. titschacki.
The use of the term processus basimeris ventralis in the key provided by Smit
(1987) describing some taxa within the genus Ectinorus may be misleading and
requires some explanation. As the term would imply, it includes a process that
arises from the basimere on its ventral margin. Among those species considered
to have a processus basimeris ventralis, the position of the process on the basimere
varies considerably from an extreme ventral position forming a deep cleft, or
sinus between the basimere and the process [E. claviger, E. budini (Jordan and
Rothschild, 1923), etc.] to a more caudal position that includes a process without
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the presence of a cleft, or sinus (E. ineptus Johnson, 1957, E. sentus, and E.
uncinatus Beaucournu and Gallardo, 1991). Nonetheless, any process along the
ventral or caudal margin of the basimere should be considered the processus
basimeris ventralis.
Ectinorus (Ectinorus) claviger (Rothschild, 1914)
(Fig. II-J)
Specimens Examined. — Recuay Province: W of highway junction 3 and 109 (~10°03'S, 77°22'W),
—4115 m, L. penianum, 29 March 2000, T. Alejo, 19. This specimen is deposited in the Carnegie
Museum of Natural History.
Remarks. — This female was collected from the same host species as E. alejoi
(described above). A very closely related species (if not conspecific), Ectinorus
sentus (Rothschild, 1914) was also reported from vizcacha (Lagidium Meyen,
1833 or Lagostomus Brookes, 1828) from the adjacent province of Junin. Wilson
and Reeder (1993) indicates that the monotypic genus Lagostomus does not occur
in Peru. Consequently, the Junin record was assuredly also from L. peruanum.
Neither E. claviger nor E. sentus possess marginal spinelets on t. I, and the sen-
silium of each has fewer than 20 trichobothria per side. The latter character sep™
arates them from the closely allied taxon, E. viscachae (Wagner, 1937). Females
of E. sentus are unknown. Our female was compared with the female description
of E. claviger provided by Wagner (1937) and found to be conspecific. The female
is unknown for three additional species within this group {E. budini (Jordan and
Rothschild, 1923), E. nomisis Smit, 1987, and E. pearsoni (Johnson, 1957)], but
their distributions are extralimital. Smit (1987), in couplet 19 of his key to the
species of the subgenus Ectinorus, overlooked the description of E. claviger fe-
males by Wagner (1937). A brief description of the female of E. claviger is
provided to supplement that of Wagner (1937).
Description. — Head: Frontal tubercle well developed. Ocular row of 3 setae, ventral seta the larger.
Labial palpus of 5 segments, extending to middle of trochanter. Genal lobe with marginal ventral row
of 5 small setae. Eye with deep ventral sinus. Occipital area with single minute seta dorsal to antennal
fossa and 3 setae per side along posterior main row. Scape with 5 marginal setae, pedicel with dorsal
lobe bearing several small setae and a single seta extending about Vs length of antenna. Thorax:
PronotLim with single row of setae, mesonotum with two rows of setae and 17-18 pseudosetae under
mesonotal collar, metanotum with two rows of setae. Dorsal portion of prosternum arched upward
forming depression and shield for terminal segments of antenna. Meso- and metasterna not extending
downward. Furca hair-like and pleural arch well developed. Metepimeron with single vertical row of
4 setae and single seta anterior to row. Legs: Mid coxa with lateral sulcus incomplete. Hind coxa with
fringe of setae mesally from base to apex. Lateral setae guarding femoral-tibial joint shortest on fore
femur and longest on mid and hind femora. A single seta extending over anterior pit on each femur.
Six dorsal notches on all tibiae with two minute setae between the fourth and fifth notches. Four
lateral plantar bristles on each of the distitarsomeres. Unmodified abdominal segments: Tergites lacking
marginal spinelets. Two rows of setae per tergite. One antesensilial seta per side set on small tubercle.
Patch of 12-13 small setae on st. II isolated from single basal seta per side. Sternum III with 6-7
setae per side, preceded by 5-6 scattered setae. Sternites IV-VI with 4-5, 3, and 4 per side, respec-
tively. Modified abdominal segments: Tergum VIII with 5-7 setae per side dorsal to atrium of spiracle.
Atrium with obvious expansion at base. Caudal margin of t. VIII rounded with 8 marginal setae, patch
of 6 setae anterior to these, and 10-12 short, stout setae mesally at apex. Caudal margin of st. VII
entire (Fig. IJ). Sternum VIII broad and obliquely rounded dorsally. Sides of anal stylet parallel, four
times as long as wide, bearing one long seta and 2 minute setae ventral to base of long seta. Ventral
anal lobe convexly arched ventrally with numerous stout marginal setae. Spermatheca slightly askew
but cribriform area appears to extend from surface to deep within bulga. Perula of bursa copulatrix
spherical, not ovoid (Fig. 11).
Length (single slide mounted specimen). — 2.5 mm
2002
97
Polygenis (Polygenis) Utargus (Jordan and Rothschild, 1923)
Specimens Examined. — Hauzlas Province: Santa River Valley, Choquechaca (N of Caraz), 2195
m, ex O. xantheolus, 2-3 September 1999, USUHS Team, 33, 4$; ex A. mollis, 3-4 September 1999,
USUHS Team, 29; ex P. andium, 2 September 1999, USUHS Team, 1 3 ; ex Thomasomys sp., 4
September 1999, USUHS Team, 13.
Remarks. — This flea was collected from inside a human dwelling from a water
trap and from sylvatic rodents. Macchiavello (1957) reported that P. Utargus was
the principal vector of plague among Sciurus stramineus Eydoux and Souleyet,
1841 in Lancones and Cazaderos, Piura Department, Peru. This flea should be
considered a potential link in transferring plague from wild rodent populations to
commensal, or peridomestic rodents such as O. xantheolus.
Tetrapsyllus (Phylliver) bleptus (Jordan and Rothschild, 1923)
Specimens Examined. — Huaraz Province: Pariacota (9°33'27"S, 77°38'82"W), 3630 m, ex P. an-
dium, 24 March 2000, Hastriter Team, 39; Pariacota (9°33'10"S, 77°35'69"W), 3715 m, ex P. andium,
24 March 2000, Hastriter Team, 1 9 .
Remarks. — Females of this species were collected from two adjacent sites from
three specimens of the same host species, P. andium. Known only from high
elevations (2745-4880 m), it has previously been reported from Junin Depart-
ment, Peru, south to Parincota Province, Chile and Catamarca (type locality) and
Jujuy Provinces, Argentina. Records from Ancash Department establish the most
northern account for T. bleptus.
Tiasmastus cavicola (Weyenbergh, 1881)
Specimens Examined. — Hauzlas Province: Santa River Valley, Choquechaca (N of Caraz), ex C.
porcellus, January 2000, P. Lima, 53, 209.
Remarks. — This flea is common on the domestic guinea pig, which is a staple
food source for the indigenous people of Peru. Little effort was expended in
collecting fleas from these hosts or in human habitations, although they are known
to occur in large numbers on C. porcellus.
Stephanocircidae
Cleopsylla townsendi Rothschild, 1914
Specimens Examined. — Chiquian Province: E of Chiquian (10°03'48"S, 77°19'27"W), 4180 m, ex
P. amicus, 27 March 2000, Hastriter Team, 13 29; E of Chiquian (10°06'07"S, 77°11'07"W), 4180
m, ex Akodon sp2 cf. mollis, 27 March 2000, Hastriter Team, 43 19; E of Chiquian (10°08'06"S,
77°10'22"W), —3995 m, ex P. andium, 27 March 2000, Hastriter Team, 19; E of Chiquian
(10°09'14"S, 77°19'17"W), 3990 m, ex P. andium, 27 March 2000, Hastriter Team, 13, 29. Huaraz
Province: Pariacota (9°33T0"S, 77°35'69"W), 3715 m, ex P. andium, 24 March 2000, Hastriter Team,
43. Huaylas Province: Parque Nacional Huascaran (9°30'04"S, 77°25'31"W), 3630 m, ex Akodon
spl cf. mollis, 25 March 2000, Hastriter Team, 13; Parque Nacional Huascaran (9°30'27"S,
77°26'13"W), 3935 m, ex Akodon spl cf. mollis, 25 March 2000, Hastriter Team, 23, 49; Parque
Nacional Huascaran (9°30'39"S, 77°27'69"W), 3475 m, ex P. andium, 25 March 2000, Hastriter Team,
13,29; and Parque Nacional Huascaran (9°30'04"S, 77°25'31"W), 3630 m, ex Akodon sp2 cf. mollis,
25 March 2000, Hastriter Team, 1 9. Recuay Province: 4.6 km W Recuay (9°43'36"S, 77°27'55"W),
—3660 m, ex P. andium, 29 March 2000, Hastriter Team, 29; 19 km W Recuay (9°44'6"S,
77°30'03"W), 4420 m, ex Akodon spl cf. mollis, 29 March 2000, Hastriter Team, 1 9 ; N of Santos
River (10°03'48"S, 77°19'27"W), 4180 m, ex P. amicus, 21 March 2000, Hastriter Team, 13, 29
Remarks. — The bulga of the spermatheca of C. townsendi extends well onto
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VOL. 71
the hilla, giving the false impression that the hilla is extended into the bulga. This
is not apparent in figures of the spermatheca illustrated by Hopkins and Rothschild
(1956) and Johnson (1957). The species has been reported from Chimborazo,
Equador (type locality) and Parincota, Chile, and in four Peruvian Departments
(Table 1). It was commonly collected on species of Phyllotis and Akodon in
Ancash Department, although it has been reported from a variety of hosts in other
high elevation regions (3660-4880 m).
Craneopsylla minerva minerva (Rothschild, 1903)
Specimens Examined. — Hauzlas Province: Santa River Valley, Choquechaca (N of Caraz), ex P.
andiiim, 2 September 1999, USUHS Team, \ 6 . Huaraz Province: Pariacota (9°32'32"S, 77°47'50"W),
1 860 m, ex P. andium, 23 March 2000, Hastriter Team, 1 9 . Huari Province: Yungilla, ex Rattus
“silvestre”, 15 April 1999, J. Cisamaro, 19. Huaylas Province: Parque Nacional Huascaran
(9°30'40"S, 77°28'32"W), 3385 m, ex P. amicus, 25 March 2000, Hastriter Team, 1 9 ; Parque Nacional
Huascaran (9°30'40"S, 77°28'32"W), 3380 m, ex Akodon sp2 cf. mollis, 25 March 2000, Hastriter
Team, IcJ, 29; Parque Nacional Huascaran (9°30'39"S, 77°28'69"W), 3475 m, ex Akodon spl cf.
mollis, 25 March 2000, Hastriter Team, 33, 19. Recuay Province: 10 km W Recuay (9°43'25"S,
77°29'13"W), 4010 m, ex P. andium, 29 March 2000, Hastriter Team, 13, 29; 10 km W Recuay
(9°43'25"S, 77°29H3"W), 4010 m, ex P. amicus, 29 March 2000, Hastriter Team, 23, 19.
Remarks. — Specimens of Craneopsylla minerva wolffhuegeli (Rothschild,
1909) referenced by Macchiavello (1948) from Ecuador, Peru and northern Chile,
undoubtedly belong to the nominate subspecies. Although the two subspecies are
sympatric in northern Argentina, C. m. wolffhuegeli probably does not occur in
Peru. Neither species have been reported in Bolivia.
Plocopsylla achilles (Rothschild, 1911)
Specimens Examined. — Chiquian Province: E of Chiquian (10°9'54"S, 77°19'42"W), 3900 m, ex
Akodon spl cf. mollis, 27 March 2000, Hastriter Team, 13, 19. Huaylas Province: Parque Nacional
Huascaran (9°30'27"S, 77°26'13"W), 3935 m, ex Akodon spl cf. mollis, 25 March 2000, Hastriter
Team, 23, 19; Parque Nacional Huascaran (9°30'04"S, 77°25'31"W), 3630 m, ex Akodon spl cf.
mollis, 25 March 2000, Hastriter Team, 19. Recuay Province: 10 km W Recuay (9°43'25"S,
77°29'13"W), 4010 m, ex P. andium, 29 March 2000, Hastriter Team, 13; 19 km W Recuay
(9°44'06"S, 77°30'03"W), 4420 m, ex Akodon spl cf. mollis, 29 March 2000, Hastriter Team, 23, 49.
Plocopsylla pallas (Rothschild, 1914), male description
(Fig. 2B-H)
Material Examined. — Huaraz Province: Pariacota (9°33'10"S, 77°35'69"W), 3715 m, ex P. andium,
24 March 2000, Hastriter Team, 13; Pariacota (9°33'27"S, 77°38'82"W), 3630 m, ex P. andium, 24
March 2000, Hastriter Team, 53,79. Recuay Province: N of Santos River (10°03'48"S, 77°19'27"W),
4180 m, ex P. amicus, 27 March 2000, Hastriter Team, 29 ; road cut on highway 109 between Chiquian
junction and Pavilotica (10°9'36"S, 77°19'46"W), 3980 m, ex “Berlese nest from cavity in earth bank,”
1 April 2000, M.W. Hastriter and R. Soto, 13; N of highway 109 between Chiquian junction and
Pavilotica (10°09'14"S, 77°19'17"W), 3990 m, ex Akodon sp2 cf. mollis, 28 March 2000, Hastriter
Team, 13; N of highway 109 between Chiquian junction and Pavilotica (10°9'14"S, 77°19'17"W),
3990 m, ex P. andium, 28 March 2000, Hastriter Team, 13, 89; 10 km W Recuay (9°43'25"S,
77°29'13"W), 4010 m, ex P. andium, 29 March 2000, Hastriter Team, 103, 12 9; 10 km W Recuay
(9°43'25"S, 77°29'13"W), 4010 m, ex P. amicus, 29 March 2000, Hastriter Team, 33, 59. Specimens
of this series are deposited in the CMNH (23, 29), the National Museum of Natural History, Wash-
ington, D.C. (23, 29), and the Robert E. Lewis collection (23, 29). The remaining specimens remain
in the senior author’s collection.
Remarks. — In his original description of P. pallas, Rothschild (1914) provided
only a brief description based on two females collected from a “rat” in Junin
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99
Department. His description of the female is herein supplemented (based on 34 9 )
and a description of the male is provided for the first time based on a large series
of 22d. Our records would indicate that P. andium is the preferred host. This flea
was not collected in other provinces (also high altitude sites), although P. andium
was prevalent.
Diagnosis. — Both sexes resemble P. viracocha Schramm and Lewis, 1987 and
P. enderleini Wagner, 1933 by the lack of basally notched helmet spines and a
genal comb consisting of 6 spines (the genal spine being smaller than the others
and separated by a space roughly the width of the spines). Schramm and Lewis
(1988) assigned these two species to Species Group A, Subgroup A-3. The males
of P. pallas can now be assigned to this grouping. The male can immediately be
distinguished from those of P. viracocha by the presence of a slender mesal lobe
projecting well beyond the dorsal margin of the basimere (Fig. 2E) and the apex
of St. IX bent caudad at a 90° angle (Fig. 2D). The absence of an acutely angled
dorso-anterior projection of the telomere (Fig. 2E-F) separates it from P. ender-
leini. Separable from males of both species by the presence of a conspicuous
dorsally expanded wing-like sclerite that extends upwards between t. VII and
sensilium (Fig. 2B). Females may be separated from P. enderleini in having a
prectenidial helmet width that is more than one and one-half the length of the
longest helmet spine and from P. enderleini (5-6 well-separated bristles) and P.
viracocha (4 well- separated bristles) in having only 3 well-separated marginal
bristles below the lobe of t. VIIL
Description, Male. — Head: Anterior helmet consisting of 14 spatulate spines without notched bases.
Prectenidial width of helmet more than 1.5 times length of longest helmet spine. Line of minute
marginal setae along anterior margin of helmet; single large seta near base of ventral-most helmet
spine. Two placoid pits along dorsal margin of helmet. Helmet striations absent above dorsal spine.
Gena with 8-9 minute setae along anterior margin; two large lateral setae in vertical row. Genal comb
of 6 spatulate spines; dorsal-most spine less than Vi length of others. Genal lobe extending beyond
dorsal genal spine and truncate. Maxillary palpus of 5 segments; basal segment very short. Maxilla
slightly rounded apically; labial palpus of 5 segments, apex reaching % length of fore coxa. Postan-
tennal region with anterior group of 18-20 setae per side and 4 rows (4, 5, 8 and 8-9 setae per side);
posterior row with intercalaries. Occiput with dorsal tuber. Antennal scape with 5 apical setae extend-
ing onto pedicel; pedicel with 10 long slender setae extending beyond 9-segmented clavus. The latter
appears compressed and fused, extending far short of antennal fossa. Thorax (Fig. 2A): Pronotum with
two anterior irregular rows of setae and a third posterior row; comb composed of 22 spines. Large
dorsal notch in prosternum for cervical link plate. Mesonotum with anterior group of setae; posterior
row of 6 setae per side with intercalaries and single pseudoseta per side under mesonotal collar.
Metapleuron with 9 lateral setae. Mesosternum lobed ventrally downward between pro- and meso-
coxae. Metanotum with irregularly arranged anterior row of setae and posterior row of 6 setae per
side with intercalaries. The dorso-posterior margin of metanotum barely discernible. Lateral metanotal
area with 3 setae. Pleural arch well developed. Metepisternum with single lateral seta; squamulum
present. Metasternum lobed antero-ventrally extending downward between mid and hind coxae. Me-
tepimeron with two vertical rows of 4 setae each; atrium of spiracle larger than abdominal spiracles.
Legs: Numerous lateral setae on fore coxa. Fore femur with 9-10 lateral setae, no mesal setae. Setae
guarding femoral-tibial joint subequal. Dorsal margin of fore tibia with 5 notches bearing setae (2, 2,
3, 3, 3), apical setae arranged in a false comb of 7 setae. Lateral surface of fore tibia with vertical
row of 7 setae. Lateral sulcus of mid coxa complete. Anterior femoral pit of mid femur with long
outer seta and shorter inner seta; 6 setae each on posterior lateral margin and single mesal seta. Lateral
seta guarding femoral-tibial joint of mid femur larger than mesal seta; mesal seta larger than lateral
seta on hind femur. Hind tibia with 10 lateral setae, 6 dorsal notches (2, 2, 3, 3, 3, 3 setae), and apical
false comb of 9-10 setae. Mid and hind coxae, each with fringe of setae extending about % from
apex to base. Dorsal margin of mid and hind femora with short, stout setae. Hind femur with 8-9
lateral setae arranged toward posterior. Hind tibia with 15 lateral setae, 7 dorsal notches (2, 2, 3, 3,
3, 3, and 4 setae); apical setae interrupted and not forming a false comb as on fore and mid tibiae.
The lowest seta in each notch bearing 3 setae separated from adjacent 2 setae by space less than width
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Table 1. — A listing of fleas of Peru by political departments (A = new record for Ancash Department;
C = species found in current study; P = new record for Peru; X = species recorded in literature;
and ? = doubtful occurrence).
Caja-
Huan-
Flea species
Ancash
Apurimac Arequipa Ayacucho
marca
Cuzco cavelica
Adoratopsylla intermedia copha
Adoratopsylla intermedia intermedia
Agastopsylla hirsutior
Agastopsylla nilotica nilotica
Agastopsylla pearsoni
Ayeshaepsylla thurmani
AC
Cediopsylla spillmanni
Ceratophyllus titicacensis
Cleopsylla townsendi
AC
X
Craneopsylla minerva minerva
Craneopsylla minerva wolffhuegeli?
AC
X
Ctenidiosomus spillmanni
Ctenocephalides canis
X
X
Ctenocephalides felis felis
cx
Dasypsyllus plumosissimus
Dasypsyllus stejnegeri
Delostichus phyllotis
PC
X
X
Delostichus xenurus
X
Echidnophaga gallinacea
AC
Ectinorus alejoi n. sp.
PC
Ectinoriis claviger
AC
X
Ectinorus disjugis
X
Ectinorus hecate
Ectinorus hertigi
Ectinorus ineptus
Ectinorus pearsoni
Ectinorus sentus
X
Ectinorus simonsi
X
Ectinorus viscachae
X
Euhoplopsyllus andensis
Euhoplopsyllus manconis
Gephyropsylla klagesi klagesi
Gephyropsylla klagesi samuelis
Hectopsylla eskeyi
Hectopsylla psittaci
Hectopsylla pulex
Hectopsylla suarezi
Hormopsylla egena
Hormopsylla trux
CX
X
X
Leptopsylla segnis
Myodopsylla setosa
Neotyphloceras crassispina chilensis?
CX
X
Neotyphloceras crassispina crassispina
X
X
Neotyphloceras crassispina hemisus
Neotyphloceras rosenbergi
Nonnapsylla rothchildi
AC
X
Nosopsyllus fasciatus
Nosopsyllus londiniensis londiniensis
Parapsyllus cedei
Plocopsylla achilles
Plocopsylla enderlieini
Plocopsylla hector
Plocopsylla inti
Plocopsylla kilya
CP
X
X
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101
Table 1. — Extended.
La Lamba- Moque-
Ica Junin Libertad yeque Lima Loreta gua
X
X
X
XX X
X X X X
San Peru
Piura Puno Martin Tacna Tambo Tumbes Islands
X
X
X
X
X
X
X
XX X
X
X
X X
X
X X
X
X
X
X
X
X
X
X
X
X
X
X XX
X X
XXX X
X
X
XXX X
X
X
X
X
X
X
X
X
X
X
X
X X
X X
X
X
X
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Table 1. — Continued.
Caja-
Huan-
Flea species
Ancash
Apurimac Arequipa Ayacucho marca
Cuzco
cavelica
Plocopsylla pallas
Plocopsylla viracocha
AC
X
Plusaetis equatoris?
Phisaetis dolens quitanus
Polygenis bohlsi bohlsi
Poly gen is brachimis
Polygenis impavidus
AC
X
X
Polygenis litargus
Polygenis litus?
cx
X
Polygenis rimatus?
Polygenis roberti beebei
X
X
X
Polygenis roberti roberti
Pulex irritans
cx
X
X
Rhopalopsyllus australis tupinus
Rhopalopsylliis cacicus
Rhopalopsyllus lugubris lugiibris
Sphinctopsylla inca
Sternopsylla distincta speciosa
AC
X
Tetrapsyllus bleptus
Tetrapsyllus comis
Tetrapsyllus elutus
Tetrapsyllus tristis
AC
Tiamastiis cavicola
CX
X X
Tiarapsylla bella
Tiarapsylla titschacki
AC
Timga penetrans
CX
X
Xenopsylla cheopis
X
of base of same setae. Femoral pits on all three femora with long outer seta and shorter inner seta.
The fifth segment of each tarsus with 5 pairs of plantar bristles, the proximal pair placed on plantar
surface between second pair, preapical plantar bristles small, paired, and broadly spaced on apex of
plantar surface. Unmodified abdominal segments: Number of marginal spinelets variable on t. I-VII
(5-6, 3-4, 4, 2-3, 0, 0, and 0 spinelets per side). Tergites I-VI with 2 rows of setae (anterior short
setae, posterior long setae with intercalaries). Tergum VII with single row of setae, antesensilial bristles
lacking. Single seta of main row below level of pointed spiracular fossa. Notable sclerotized apodemes
on basal st. II, bearing single seta per side, st. III-VI with 3 setae per side, st. VII with 1-2 small
setae per side. Modified abdominal segments (Fig. 2B, D-F): Tergum VIII greatly expanded into wing-
like processes that extend dorsad between t. VII and sensilium, mesal surface spiculose (Fig. 2B).
Basimere of t. IX with mesal process along dorsal margin bearing 2 setae, apex with row of 8-9 long
marginal setae (Fig. 2E). Apex of telomere nearly twice as wide as near point of articulation with
basimere; bearing a scattered group of minute setae on the mesal surface and a large spiniform on the
apical-posterior mesal surface. Membranous dorsal anal lobe with single dorsal seta; ventral anal lobe
with two ventral setae. Sensilium with 16 sensilial pits per side. Sternum VII extending upward
covering st, VIII and aedeagus. Apex of st. IX strongly bent caudad, bearing a single hair-like seta
and two highly modified spiniform bristles (Fig. 2D). Aedeagus (Fig. 2C): Lateral lamina narrow at
neck broadening toward apex; median lamina broader than lateral lamina. Lateral lamina projecting
dorso-caudally above fulcrum. Apodemal rod extending from well differentiated ventral floor of the
girdle. Penis rods slightly longer than lateral/median lamina. Dorsal median lobes membranous and
somewhat squared apically with sclerotized crochet. Sclerotized inner tube covered laterally by dorsal
armature. Modified abdominal segments, female (Fig. 2G-H): Dorsal portion of t. VIII with two stout
setae just above spiracular fossa; dorsal margin beneath sensilium strongly concave with well devel-
oped lobe along upper caudal margin lacking setae. The dorsal caudal lobe subtended by a shallow
sinus with three long marginal bristles equally spaced; anterior and nearly contiguous to these is a
shorter subspiniform seta extending obliquely upward. Among a lateral group of 36-38 setae are 2
stout setae. Caudal margin of St. VII with slight concavity, group of 16-18 setae per side of which 2
2002
103
Table 1. — Continued. Extended.
Ica Junin
La
Libertad
Laniba-
yeque
Lima
Moque-
Loreta gua
Pasco Piura
Puno
San Peru
Martin Tacna Tambo Tumbes Islands
X
X
X
X
X
X
X
X
X X
X
X
X
X X
X
X
X
X
X
X
X
X
X
X
X
X
X
X X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
are strongly curved ventrad (Fig.
2 H). Sternum VIII without setae. Dorsal anal lobe dorsally enclosed
with fringe of 7-8 long setae; anal stylet more than three times as long as wide with 2 short and 1
long setae. Ventral anal lobe inconspicuous with several fine setae along ventral portion. Bulga of
spermatheca about twice length of hilla; lumen deeply inserted into bulga and distinctly narrowed at
apex of internal orifice. Spermatheca strongly concave ventrally and convex dorsally; lumen of duct
enters bulga ventrally (Fig. 2G).
Length (slide mounted specimens). — Male: 2.9 mm (n=20, range=2. 6-3.1
mm); Female: 2.9 mm (n=20, range=2.4-3.2 mm).
Sphinctopsylla inca (Rothschild, 1914)
Specimens Examined. — Chiquian Provinces E of Chiquian (10°08'06"S, 77°10'22"W), —3660 m,
ex P. andium, 27 March 2000, Hastriter Team, 1 $ . Huaylas Province: Parque Nacional Huascaran
(9°30'04"S, 77°25'3T'W), 3630 m, ex Akodon spl cf. mollis, 25 March 2000, Hastriter Team, 2$;
Parque Nacional Huascaran (9°30'27"S, 77°26'’ 13"W), 3630 m, ex Akodon spl cf. mollis, 25 March
2000, Hastriter Team, 1 S .
Tiarapsylla titschacki Wagner, 1937
Specimens Examined. — Recuay Province: W of junction highways 3 and 109 (— 10°03'S, 77°22'W),
—4115 m, ex L. peruanum, 29 March 2000, T. Alejo, 33, 7$.
Remarks. — Three type specimens (lc5^, 2$) were collected on “vizcacha” in
the adjacent Department of La Libertad (4000 m) (Wagner, 1937). Additional
records include one female each from L. puruanum. Department of Puno (ele-
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Annals of Carnegie Museum
VOL. 71
vation unknown) and Department of Junin (—4875 m) (Johnson, 1956). A female
was also reported from a feline host in the Department of Puno (Hopkins and
Rothschild, 1956). The latter record is undoubtedly a predator-related association.
Lagidium puruanum seems a preferred host for this species, occurring only at
high elevations from central to southern Peru.
Discussion
Very common species (Pulex irritans complex, L. segnis, E. gallinacea and T.
cavicola) found on peridomestic animals were not aggressively sought except to
establish records for Ancash Department. Trapping statistics were kept only for
the March 2000 collecting period and an analysis follows. During eight trap
nights, 844 traps were set yielding 67 small mammals [Akodon spl cf. mollis
(n=16, 23.9%), Akodon sp2 cf. mollis (n=5, 7.5%), M. musculus (n~5, 7.5%),
P. andium (n=38, 56.7%) and P. amicus (n=3, 4.5%)] for a catch rate of 7.9%.
Phyllotis andium harbored 57% of the total fleas collected. Plocopsylla pallas was
found on 13 of the 38 (34%) P. andium surveyed. The average number of P,
pallas occurring on P. andium that were positive for this flea was 3.4. Phyllotis
andium was trapped at the same locations as P. amicus and Akodon sp2 cf. mollis,
but P. pallas was found only once on each of these hosts. One can only speculate
as to why P. pallas was not collected in provinces other than Recuay, since P.
andium was present in all provinces at similar elevations and habitats. Fleas most
commonly collected on sylvatic mammals during all collection periods included
the following: Plocopsylla pallas (16%), Neotyphloceras crassispina hemisus
(14%), and Cleopsylla townsendi (9%). These three species comprised 39% of
the total number of fleas collected.
The total number of species of fleas currently reported in Peru is 81 (Table 1);
however, this may be misleading. Several species listed in Table 1 are probable
misidentifications. They include the following: C. m. wolffhuegeli, N. c. chilensis,
P. equitoris, Polygenis rimatus (Jordan, 1932), and Polygenis litus (Jordan and
Rothschild, 1908). The latter species reported from Piura Department by Mac-
chiavello (1948) is doubtful, since only three females (localities unknown) of P.
litus have been reported in the literature (Jordan and Rothschild, 1908, 1923) and
can easily be confused with P. litargus. Populations of C. m. wolffhuegeli flourish
much farther south in northern Argentina and southern Bolivia. Neotyphloceras
c. chilensis, difficult to distinguish from other subspecies, occurs along montane
and coastal Chile from Santiago south. Plusaetis equatoris may be found in the
Andean valleys of northwestern South America as far south as Ecuador. The latter
three species probably represent C. m. minerva, N. c. crassispina or N. c. hemisus,
and P. d. quitanus, respectively. Literature records of P. rimatus were reported
on the same hosts {O. xantheolus and A. mollis) on which we found P. litargus
and likely represents this species. Polygenis rimatus is common in lowland areas
of Argentina, Brazil, and Paraguay, but not in montane regions. Considering these
five species as extralimital to Peru, the total number of recognized taxa that occur
in Peru is 76.
The distribution of several species reported in the literature are of special in-
terest. Populations of Gephyropsylla klagesi samuelis (Jordan and Rothschild,
1923) and P. litargus on the Isla Metapalo (Table 1, Peru Islands) were possibly
transported by commerce on their common peridomestic host, O. xantheolus, or
associated commensal rodents. Although both subspecies of Adoratopsylla inter-
2002
105
media have been reported in Junin and Piura Departments, neither were collected
during our studies. This species is harbored by various marsupial species of Di~
delphis Linnaeus, 1758 and Monodelphis Burnett, 1830. The distribution of these
hosts in diverse habitats throughout the Andean valleys and the lowlands of Peru
should support a much broader distribution than the records would indicate for
this flea species. We were surprised that Neotyphloceras rosenbergi (Rothschild,
1904) was not collected, since it was reported in the departments of Cajamarca
and Piura from hosts we commonly collected (A. mollis and O. xantheoius). Three
fleas are reported from Peru for the first time (D. stejnegeri, P. achilles, and E.
alejoi n. sp.). Eleven of the 21 species herein reported are new records for Ancash
Department (A. thurmani.C. townsendi, C. m. minerva, E. gailinacea, E. clavigep
N. c. hemisus, P. pallas, P. d. quitanus, S. inca, T. bleptus, and T. titschacki).
Future emphasis for flea faunal studies should include departments that have
received little attention. Most notably are the departments of the eastern slope of
the Andes (Amazonas, Cuzco, Huanuco, Junin, Pasco, and San Martin), the low-
land departments (Loreto and Madres de Dios), the southern coastal departments
(Arequipa, Moquegua, and Tacna), and the south central montane departments
(Apurimac, Ayacucho, Huancavelica).
ACKN'OWLEDGMENTS
Grateful appreciation is extended to Dr. Paul Pachas Chavez and Dr. Carlos Enrique Ponce Garcia,
Ministerio de Salud, Hospital de Caraz, and Jorge Lucero Tamayo, Ministerio de Saiud, Laboratorio
Referencial Regional de Salud, Hauaraz for providing coordination and facilities support for part of
this research. For the loan of specimens we are grateful to Theresa Howard and the Trustees of the
Natural History Museum, London. The logistical support and assistance of Pat Carney, Uniformed
Services University of the Health Sciences, Bethesda, MD, and the staff of the U.S. Naval Medical
Research Center Detachment Unit 3800, Lima, Peru, were also essential to these studies, and we are
much indebted. To Dr. Oswaldo Ramirez, Universidad Peruana Cayetano Heredia, Lima, we acknowl-
edge support in identifying the host mammals. The senior author expresses special thanks to the staff
of the Monte L. Bean Life Science Museum for their continued support in providing work space,
equipment and materials necessary for conducting systematic studies. Funding for this research was
partially provided by a grant from the National Science Foundation, Award Number DEB--9983195.
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ANNALS OF CARNEGIE MUSEUM
VoL. 71, Number 2, Pp. 107-129
28 May 2002
REVIEW OF LEPTOMERYX (ARTIODACTYLA, LEPTOMERYICIDAE)
FROM THE ORELLAN (OLIGOCENE) OF NEBRASKA
William W Korth’ ^
Research Associate, Section of Vertebrate Paleontology
Margaret E. Diamond'
Abstract
Several hundred specimens were examined of the artiodactyl Leptomeryx from the Orellan of North
America. The dental morphology used to distinguish Chadronian species of the genus were examined
for consistency in the Orellan sample. Based on the consistent dental characters established, three
distinct species are recognized from the Orellan: L. evansi Leidy, L. exilis Cook, and Leptomeryx
elissae new species. Additional material of Leptomeryx from the Whitneyan was also examined and
suggests that L. minimus Frick and L. lenis Cook may be junior synonyms of L. exilis and L. evansi,
respectively.
Two distinct lineages of Leptomeryx are recognized that range from the Chadronian through the
Whitneyan and possibly Arikareean, the L. yoderi-L. mammifer-L. exilis— L. obliquidens lineage and
the L. speciosus-L. evansi lineage. It is likely that the Arikareean leptomerycid Pronodens Koerner
is derived directly from the L. evansi lineage.
Key Words: Leptomeryx, Chadronian, Orellan, Oligocene, Artiodactyla
Introduction
Historical Review
Leidy (1853) first named Leptomeryx evansi as a new genus and species from
what is now known to be the Orellan Brule Formation in South Dakota. Since
that time, 16 species of Leptomeryx have been named, ranging in age from Du-
chesnean to Hemingfordian (Cope, 1886, 1889; Douglass, 1903; Lambe, 1908;
Lull, 1922; Cook, 1934; Frick, 1937; Wilson, 1974). The majority of these species
were Chadronian in age (see Webb, 1998 for review). The only Orellan species
named, other than the type species, was L. exilis from Sioux County, Nebraska
(Cook, 1934). In his review of the White River artiodactyls, Scott (1940) cited
only a single species from the Orellan, the type species, L. evansi.
Heaton and Emry (1996) reviewed the Chadronian species of Leptomeryx and
followed Scott (1940) in concluding that there was only a single species of lep-
tomerycid from the Orellan, L. evansi, and that the other nominal species from
the Orellan were merely synonyms of L. evansi. Galbreath (1953) recognized only
L. evansi in his Orellan fauna from northeastern Colorado, but cited a single
specimen as Leptomeryx sp. The latter specimen was a maxillary fragment with
upper cheek teeth that were smaller than referred specimens of upper dentitions
of L. evansi. In the most recent review of the Leptomerycidae, Webb (1998) also
agreed that only a single species of Leptomeryx existed in the Orellan and sug-
' Rochester Institute of Vertebrate Paleontology, 265 Carling Road, Rochester, New York 14610.
^ Earth Sciences and Science Education Department, Buffalo State College, Buffalo, New York 14222.
Submitted 29 November 2001.
107
108
Annals of Carnegie Museum
VOL. 71
gested several synonymies for the known Chadronian to Arikareean species of
Leptomeryx.
Black (1978) named a new Duchesnean genus of leptomeryicid, Hendry omeryx.
He distinguished Hendryomeryx from Leptomeryx by its more primitive, lower
crowned cheek teeth. He referred a species previously identified as Leptomeryx
to it, H. defordi, from Texas (Wilson, 1974). Storer (1981) also referred one of
Cope’s Chadronian species, L. esulcatus, to Hendryomeryx. Webb (1998:fig. 31.6)
listed Hendryomeryx from the Orellan of Montana without a citation and figured
Hendryomeryx as surviving into the Whitneyan. However, several authors have
considered Hendryomeryx as a junior synonym of Leptomeryx (Heaton and Emry,
1996; McKenna and Bell, 1997).
In all, as many as three species of leptomerycid or as few as one species may
be present in the Orellan of North America. No review of Orellan Leptomeryx
has ever been made at the species level. With the large samples of Leptomeryx
now available, such a study is warranted.
Fossil Material
Extensive collections of Orellan Leptomeryx are available in several institutions
in North America. The bulk of the material studied was from the Orella Member
of the Brule Eormation, Sioux County, Nebraska. However, comparative samples
from the Orellan of North Dakota, South Dakota, Colorado and Wyoming were
also studied (see Appendices 1 and 2). The institutions providing the sample were
as follows: American Museum of Natural History (AMNH), Carnegie Museum
of Natural History (CM), National Museum of Natural History, Smithsonian
(USNM), University of Nebraska State Museum (UNSM) and Yale-Peabody Mu-
seum (YPM). Many of these samples were collected as early as the early 1900s
and no distinction was made as to horizon other than “Oreodon beds” or “Lower
Brule” (=Orella Member, Brule Eormation). A large, stratigraphically controlled
sample was available from the UNSM. The stratigraphic divisions of this sample
were based on the divisions of the Orella Member defined by Schultz and Stout
(1955): Orella A, Orella B, Orella C, and Orella D (from lowest to highest). A
smaller stratigraphically controlled sample from the White River Formation of
Niobrara County, Wyoming, from AMNH was also included in this study. Small
samples of Whitneyan Leptomeryx from the collections of the AMNH and UNSM
were also examined and measured for comparison.
Methods
Morphology. — Heaton and Emry (1996) used several features of the lower
cheek teeth to distinguish the Chadronian species of Leptomeryx. Each of these
is explained below and was identified on the Orellan sample of Leptomeryx in an
attempt to establish its usefulness in separating species (Fig. 1).
1) entoconulid on M3 — The posterolingual cusp on the talonid of M3 (ento-
conulid) is a cylindrical cusp with a deep valley posterior to it, separating it from
the hypeonulid on specimens of L. speciosus and L. evansi (abbreviated S/E). The
entoconulid on these teeth is also subequal in height to the hypoconulid. On
specimens of L. yoderi and L. mammifer this cusp is more conical (triangular in
lingual view) and slopes more gently posteriorly, eliminating the deep valley
between it and the hypoconulid (Y/M). The entoconulid on these teeth is lower
in height than the hypoconulid.
2002
Korth and Diamond — Orellan Leptomeryx from Nebraska
109
Buccal Column
Entoconulid
Palaeomeryx fold
Entoconulid
L yoderimys/mammifer morphology ^ specious/evansi morphology
Fig. 1 — Schematic diagrams of left M3 of Leptomeryx (occlusal view above, lingual view below)
indicating features discussed in text and variation of morphology of entoconulid. Modified from Heaton
and Emry (1996:fig. 9).
2) Palaeomeryx fold — A small loph that originates from the protoconid on the
lower molars and runs down the posterior slope of that cusp is lacking in Chad-
ronian species of Leptomeryx but is present on L. evansi.
3) enamel wrinkling — Primitively, the enamel of the cheek teeth is smooth as
in the Chadronian species of Leptomeryx', however, in L. evansi it is highly rugose.
4) position of protoconid loph on P3 — Heaton and Emry (1996) found the
major loph running posteriorly from the apex of the protoconid on P3 (although
highly variable) generally ran along the lingual side of the tooth in Chadronian
species, but along the buccal edge of the tooth in L. evansi.
5) buccal column — The stylar cusp between the buccal cusps of the lower
molars, if present, is usually a tall column. Its height and presence is variable.
Measurements. — Size has been shown to distinguish species of Leptomeryx
with similar dental morphology (Heaton and Emry, 1996). Measurements of all
cheek teeth of the specimens examined in this study were taken. Maximum widths
and lengths of each tooth and alveolar lengths of the molar tooth rows were taken
on all specimens. All measurements were taken to the nearest 0.1 mm.
Results
Characters Used to Distinguish Species
Of the characters observed, the most reliable is the morphology of the ento-
conulid on the talonid of M3. The difference between the L. evansi/L. speciousus
morphology and the L. yoderi/L. mammifer morphology (Heaton and Emry, 1996:
hg. 9) is consistently distinguishable. In addition to the differences in the ento-
conulid cited and figured by Heaton and Emry (1996), it is also evident that in
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Annals of Carnegie Museum
VOL. 71
the Y/M morphology the cusp is lower than both the hypoconulid and the ento-
conid and does not show signs of wear until the latter two cusps are moderately
worn. In specimens with the S/E morphology, the entoconulid is as tall as the
adjacent cusps.
The presence of the Palaeomeryx-fold on the lower molars is fairly consistent.
The Palaeomeryx-fold is absent from all specimens with the Y/M entoconulid
morphology. Specimens referred to Leptomeryx evansi nearly always have this
feature present, but there is some variability. In specimens of Leptomeryx evansi
containing all of the molars, this feature is present on all MjS, 90% of the M2S,
and 83% of the M3S. Therefore, the absence of this character was less diagnostic
on isolated M3S.
The degree of enamel crenulation on the molars is generally consistent between
the recognized species of Leptomeryx, but the range of variability has nearly a
complete overlap. The amount of crenulation of the enamel was subjectively re=
corded numerically from 0 (smooth enamel) to 2 (most crenulated) in increments
of 0.5. Specimens of L. evansi ranged from 0.5 to 2.0, with a mean of 1.5,
meaning that the teeth were generally fairly strongly wrinkled. The specimens
with the Y/M entoconulid were generally much smoother, ranging from 0 to 1.5
with a mean of 0.5. It is evident, looking at a large population, that there is a
general trend for more highly wrinkled enamel in L. evansi, but the overlap is
quite large and could not easily be applied to a very small sample of specimens.
Similar results were found with the upper cheek teeth as well.
As indicated in Tables 1-4, the teeth of these species differ in size. With ap-
proximately 40% overlap, however, large samples with good provenience are
needed to distinguish species. Lower dentitions referred to L. evansi were gen-
erally larger than those with the Y/M entoconulid. Similar results were found with
the upper dentitions.
The remainder of the characters analyzed by Heaton and Emry (1996) for the
lower dentitions did not appear to be consistent enough to separate species of
Leptomeryx. For example, presence and size of buccal column on lower molars
and posterior protoconid loph on P3 were extremely variable. The posterior lophid
from the protoconid on P3 of all specimens was directed along the buccal side of
the tooth; none of the Orellan specimens has this lophid on the lingual side of
the tooth. The buccal column is present on nearly all the specimens from the
Orellan and is highly variable in size, ranging from a small cuspule to a distinct
column. No general trend in this morphology can be used to separate the Orellan
species of Leptomeryx.
No features of the upper cheek teeth were examined by Heaton and Emry
(1996). However, in the present study there are two features of the upper dentition
that appear distinguishable between species. First is the amount of enamel wrin-
kling, which is reflective of this character in the lower dentition, and the second
is the development of a mid-lingual column (protostyle). This cusp is generally
large and columnar on most specimens, but is entirely lacking in others. Species
are also separable on size of the upper dentitions as noted above. In general, the
upper molars of the smaller species are also distinguishable from those of L.
evansi by their proportions. In L. evansi, the upper molars are nearly square (width
equal to length), whereas in the smaller species, the upper molars are generally
wider than long except M^ (Tables 1 and 4).
There appear to be differences between species based on morphology of the
mandible, but the features are known only on a few specimens, so their consis-
Fig. 2 — Mandible and lower cheek teeth of Leptomeryx evansi, CM 512A. A. Occlusal view, Pj— M3. B, Lateral view of mandible. Bar scale
Annals of Carnegie Museum
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Korth and Diamond — Orellan Leptomeryx from Nebraska
113
tency is not known. On mandibles of Leptomeryx evansi (Fig. 2) there are one or
more accessory mental foramina. They are always below P4, This is also true of
an intermediate-sized species with the L. evansi-iype of M3 morphology (de-
scribed below). In the known mandibles of the smallest species, there are also
accessory mental foramina. One is always ventral to P3, and if there is more than
one, the second is below P4 (Fig. 6), as in L. evansi.
Another mandibular difference is in the position of P,. In all species of Lep-
tomeryx there is a diastema between P, and P2. The main mental foramen is below
this diastema. In L. evansi Pj is dorsal to the anterior margin of the mental foramen
(Fig. 2). In the smallest species, the alveolus for P, is 2.4 mm anterior to the
mental foramen. Because of the low number of complete mandibles in the fossil
sample, these differences are not included in the diagnoses of the species listed
below.
Species Recognized
Three species of Leptomeryx can be recognized from the Orellan of Nebraska
and the northern Great Plains. Leptomeryx evansi, the type species of the genus,
is separable from the other species by its larger size, more highly crenulated
enamel on the cheek teeth, presence of a protostyle on the upper molars, and S/E
morphology of the entoconulid on M3.
A second species with the S/E-type of entoconulid morphology of M3 is also
recognizable. It is generally smaller than L, evansi with less wrinkling of the
enamel and lacks the Palaeomeryx-io\d on the lower molars.
The third species is the smallest. It lacks a protostyle on the upper molars, has
less crenulated enamel on the cheek teeth, has the Y/M morphology of the en-
toconulid, and lacks a Palae ornery x- fold on the lower molars.
Systematic Paleontology
Leptomeryx Leidy, 1853
Leptomeryx evansi Leidy, 1853
(Eig. 2, 3; Tables 1 and 2)
Type Specimen. — USNM 157, partial skull and associated mandible.
Referred Specimens. — See Appendix 1.
Horizon and Locality. — Holotype from Orellan White River Group, South Da-
kota. Referred specimens from the Orellan of the northern Great Plains.
Emended Diagnosis. — Intermediate-sized species; enamel on cheek teeth highly
crenulated; Palaeomeryx- fold present on lower molars; entoconulid on M3 cylin-
drical with deep posterior valley, separating it from the hypoconulid; protostyle
present on upper molars.
Discussion. — Leptomeryx evansi is, by far, the most common of any of the
Orellan species of the genus, outnumbering the other species as much as 50 to 1
in some collections. This species shows little variation throughout its chronolog-
ical and geographic range. There is virtually no difference in size of the samples
Fig. 3 — Cheek teeth of Leptomeryx evansi. A. Occlusal (above) and lingual view of M,-M3, CM 175.
B. Occlusal view of M'-M^ UNSM 119726. Bar scale = 1 cm.
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Table 1. — Dental measurements o/ Leptomeryx evansi the Orellan (all horizons and localities
combined). Abbreviations: L, length; W, width; N, number of specimens; M, mean; OR, observed
range of size; sd, standard deviation; CV, coefficient of variation. All measurements in mm.
Mi-Mj
LM,
WM,
LM2
WMj
LM3
WM3
N
53
51
48
80
79
129
128
M
22.0
6.2
4.5
6.6
4.9
9.3
4.7
OR
19.8-24.2
5.3-7.2
3. 8-5.2
4.2-7.5
4.1-5.4
8.0-10.7
3.1-5.5
sd
I.l
0.4
0.3
0.5
0.3
0.6
0.3
CV
5.0
6.5
6.0
7.5
6.0
6.4
7.1
M'-M-’
LM‘
WM‘
LM2
WM2
LM’
WM’
N
24
66
66
60
59
41
41
M
19.0
6.3
6.3
6.7
6.8
6.7
6.4
OR
17.8-20.6
5.4-7.0
5. 1-7.5
5. 8-7. 5
5.2-8.4
6.0-7.7
5.4-7.6
sd
0.8
0.3
0.5
0.4
0.7
0.4
0.6
CV
4.2
5.2
8.1
5.4
9.9
6.1
8.8
from Orella A through Orella D (Table 2). Geographically, the only sample that
shows any variation is that from Wyoming. The Wyoming sample of L. evansi
has a slightly longer M3 than all of the other samples. However, the overlap of
the size ranges with other samples is nearly 100% and the difference in the mean
is less than 10% of the length of the tooth. This larger size of M3 is also reflected
in the measurement of the molar row (M,-M3). Heaton and Emry (1996) also
measured a sample from the lower part of the Orellan in Wyoming and produced
measurements similar to those of our sample, slightly larger than the remainder
of the samples. This is not viewed as enough difference to suggest a separate
species with the Wyoming material, only a slight geographic variation in the
Wyoming population.
Leptomeryx elissae, new species
(Fig. 4, 5; Table 3)
Type Specimen. — UNSM 119638, mandible with left P3-M3.
Referred Specimens.— CM 73416; UNSM 1 19636, 1 19637, 1 19639, 1 19640; AMNH 53626A, 606-
25868, 606-25873 (all mandibles with cheek teeth).
Horizon and Locality. — Holotype from UNSM locality Sx-8, Orella Member
(Orella C), Brule Formation, Sioux County, Nebraska. All refeiTed specimens
from the Orella Member (Orella C and D), Brule Formation, Sioux County, Ne-
braska (Orellan).
Diagnosis. — Smaller than L. evansi; enamel of cheek teeth smooth (less cren-
ulated than in L. evansi); S/F-type of entoconulid on M3; Palaeomeryx-fo\d lack-
ing on lower cheek teeth.
Etymology. — Named for a daughter of one of the authors (MED).
Discussion. — A small group of specimens that share the S/F-type of entocon-
ulid on M3 with Leptomeryx evansi are distinguishable from the latter by their
slightly smaller size, less crenulated enamel on the molars, and lack of a Palaeo-
meryx-fo\d on the lower molars. This species is intermediate in size between the
two other Orellan species of Leptomeryx (Table 3), and slightly closer to the
smaller species. However, the amount of overlap in the size range with L. evansi
is large, nearly 50%. The average amount of wrinkling on the molars is 0.7, more
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Korth and Diamond — Orellan Leptomeryx from Nebraska
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Table 2. — Dental measurements of lower molars of Orellan Leptomeryx evansi separated stratigraph-
ically and geographically. Abbreviations as in Table /.
M,-M.,
LM,
WM,
LM2
WMj
LM,
WM,
Nebraska (all levels)
N
41
39
38
66
65
1 12
1 12
M
21.8
6.2
4.5
6.6
4.9
9.2
4.7
OR
19.8-24.2
5.3-7.0
3. 8-4.9
4.2-7.5
4. 1-5.4
8.0-10.7
3.7-5.5
sd
1.0
0.4
0.2
0.5
0.3
0.6
0.3
CV
4.8
6.5
5.5
7.8
6.1
6.3
6.9
Nebraska (Orella A)
N
1
1
1
3
3
14
14
M
20.8
5.7
3.8
6.4
4.4
9.0
4.6
OR
—
—
—
6.0-6.9
4. 1-4.7
8.1-9.6
3.7-5.3
sd
—
—
—
0.5
0.3
0.4
0.5
CV
—
—
—
7.0
7.0
4.7
10.1
Nebraska (Orella C)
N
3
3
3
6
6
15
15
M
21.7
6.2
4.4
6.8
4.9
9.0
4.7
OR
20.5-22.5
6. 1-6.4
4. 1-4.6
6.4-7.2
4.4-5. 1
8. 1-9.9
4. 1-5.2
sd
1.1
0.2
0.3
0.4
0.3
0.6
0.3
CV
5.0
.2.5
6.0
5.3
5.6
6.2
6.4
Nebraska (Orella D)
N
—
—
—
2
2
10
10
M
—
—
—
6.7
4.5
9.2
4.6
OR
—
—
—
6.5-6.8
4.2-4.7
8. 1-9.9
4.4-5. 2
sd
—
—
—
—
—
0.5
0.2
CV
—
—
—
—
—
5.5
5.1
North
Dakota
N
4
4
4
5
5
7
7
M
21.7
6.1
4.5
6.5
4.8
9.3
4.6
OR
20.7-23.7
5.7-6.4
4.0-5.2
6.2-6.9
4.4-5.3
8.5-10.6
4.1-5.3
sd
1.4
0.3
0.6
0.3
0.4
0.8
0.5
CV
6.4
4.9
12.4
4.2
7.7
8.1
10.4
Wyoming
N
7
7
5
8
8
8
7
M
22.8
6.4
4.4
6.9
5.0
9.7
4.7
OR
21.6-24.2
5. 8-7. 2
4. 1-4.6
6.5-7.4
4.6-5. 4
9.0-10.1
4.6-5. 4
sd
1.1
0.4
0.2
0.4
0.2
0.4
0.2
CV
4.6
6.9
4.4
5.2
4.9
4.5
5.0
than the smaller
species.
less than in
L. evansi.
but with
a range of from 0 to 1,
less than in the smaller species (0 to 1.5).
Leptomeryx exilis Cook, 1934
(Fig. 6, 7, 8; Table 4)
Type Specimen. — AMNH 81012, partial right mandible with M2-M3.
Referred Specimens. — See Appendix 2.
Horizon and Locality. — Holotype from “the Brule (Middle Oligocene) beds
about two miles north of Chadron, Nebraska” (Cook, 1934:154). Referred ma-
terial from Orellan of the White River Group, northern Great Plains.
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Fig. 4 — Occlusal (above) and lingual view of M,-M3 of Leptomeryx elissae, UNSM 119638 (holo-
type). Bar scale = 1 cm.
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Fig. 5 — Mandible and lower cheek teeth of Leptomeryx elissae, CM 73416. A. Occlusal view, P3-M3 (alveolus for P2). B. Lateral view of mandible. Bar
scale = 1 cm.
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Table 3. — Dental measurements of Orellan Leptomeryx elissae. Abbreviations as in Table 1.
Mi-Mj
LM,
WM,
LM2
WM2
LMj
WM3
N
4
3
3
5
5
6
6
M
19.8
5.6
4.3
6.4
4.6
8.6
4.4
OR
19.4-20.4
5. 1-6.0
4.0-4.6
6.0-6.7
4.4-4.9
8.0-9.2
4.2-4.6
sd
0.5
0.5
0.3
0.3
0.2
0.4
0.1
CV
2.6
8.4
7.0
4.3
4.6
4.5
3.4
Emended Diagnosis. — Smallest species of the genus; cheek teeth with little or
no wrinkling of the enamel; P alaeomery x-fold lacking on lower molars; Y/M type
of morphology of entoconulid on M3; upper molars wider than long (subequal in
dimensions in L. evansi) and lacking a protostyle.
Discussion. — Specimens of Leptomeryx exilis are easily distinguishable from
those of the other Orellan species by their smaller size and construction of the
entoconulid on M3 (Y/M-type). This species differs from L. evansi in lacking
crenulations on the enamel of the molars, and lacking a Palaeomeryx-fold on the
lower molars. Specimens referred to Leptomeryx elissae also have the S/E-type
of entoconulid on M3. The upper molars are proportioned slightly differently from
those of L. evansi and lack a protostyle.
There appears to be no difference in size or morphology between samples of
L. exilis from the various horizons within the Orella Member in Nebraska.
Conclusions
Orellan Leptomeryx
Contrary to recent authors (Heaton and Emry, 1996; Webb, 1998), three distinct
species of Leptomeryx are recognizable from the Orellan of the Great Plains: L.
evansi, L. exilis and Leptomeryx elissae. They are easily distinguishable by size
and morphology of the lower third molar. All of the species of Leptomeryx from
the Orellan are markedly smaller than the species reported from the Chadronian
(see Heaton and Emry, 1996). The Chadronian species also lack crenulations of
the enamel on the cheek teeth and the P alaeomery x-fo\d found on Orellan L.
evansi.
Heaton and Emry (1996) recognized two lineages of Leptomeryx in the Chad^
ronian based predominantly on the morphology of the lower molars. One lineage
consisted of L. mammifer and L. yoderi that were relatively large species with
the primitive, conical entoconulid on M3 (Y/M-type). The other lineage consisted
only of L. speciosus in the Chadronian that was distinguished from the other
species by the columnar entoconulid on M3 (S/E-type) and slightly different size.
Heaton and Emry (1996) viewed the L. mammifer-L. yoderi lineage as ending in
the Chadronian, whereas the lineage represented by L. speciosus continued into
the Orellan with L. evansi.
The recognition of three Orellan species demonstrates that the radiation of
Leptomeryx in the Orellan is not as simple as previously believed. The morphoh
ogy of M3 of Orellan L. exilis is the same as in L. yoderi and L. mammifer,
making it a likely member of the same lineage, and thus extending this lineage
into the Orellan. Leptomeryx exilis differs from the former Chadronian species
only in its much smaller size. L. exilis could easily be derived from late Chad-
ronian L. mammifer simply by reducing its size. There appears to be no mor-
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119
Fig. 6 — Mandible and lower cheek teeth of Leptomeryx exilis, CM 9109. A. Occlusal view, P3-M3 (alveolus for P2). B. Lateral view of mandible (alveolus
for Pi). Bar scale = 1 cm.
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Korth and Diamond — Orellan Leptomeryx from Nebraska
121
phology that would bar L. mammifer from an ancestral position with respect to
L. exilis.
Heaton and Emry (1996) viewed the Chadronian L. speciosus as ancestral to
the Orellan L. evansi, the latter differing in its smaller size, more crenulate enamel
on the molars, and presence of a Palaeomeryx-fo\d on the lower molars. The only
difference suggested here in this L. speciosus— L. evansi lineage proposed by Hea-
ton and Emry is the addition of another species from the Orellan. Specimens
referred here to Leptomeryx elissae are closer to the Chadronian L. speciosus in
morphology than to Orellan L. evansi, lacking the crenulations and Palaeomeryx-
fold of the latter (primitive characters shared with L. speciosus). Leptomeryx elis-
sae is even smaller than L. evansi, and thus easily distinguishable from L. spe-
ciosus. It appears that this lineage, characterized by the S/E-type of entoconulid
on M3, splits in the Orellan into a smaller species with simpler cheek tooth mor-
phology {Leptomeryx elissae) and a larger species with more complex cheek tooth
morphology (L. evansi). As with the other lineage of Leptomeryx, it is quite likely
that the Chadronian L. speciosus is ancestral to both of the Orellan species of the
genus recognized here.
Whitneyan Leptomeryx
Several species of Leptomeryx have been named from the Whitneyan: L. lenis
Cook (1934), L. minimus Erick (1937), and L. obliquidens Lull (1922). Small
samples of Whitneyan Leptomeryx are present in the collections of the AMNH,
mainly from South Dakota, and were compared to the holotypes of these species.
The holotype of L. lenis (AMNH 87013) contains both upper and lower dental
elements. The cheek teeth of L. lenis are highly crenulated, have distinct Palaeo-
meryx-folds on the lower molars and have the S/E-type of entoconulid on M3.
They are also within the size range of L. evansi. The holotype of L. lenis is
indistinguishable from specimens of the Orellan L. evansi. Several specimens of
similar size and morphology from the Whitneyan were present in the AMNH
collections, and likewise, could not be separated from the Orellan samples of L.
evansi. It appears likely that L. lenis is a junior synonym of L. evansi. This
synonymy would extend the known record of L. evansi into the Whitneyan.
Among the Whitneyan specimens present in the AMNH collections are several
that are similar to L. evansi in morphology (Fig. 9), but are beyond its range of
size (both smaller and larger). These specimens may well represent additional
species of Leptomeryx from this horizon.
The holotype (AMNH 1347a) and paratype (AMNH 1347) of L. minimus from
the Whitneyan of South Dakota are small in size, have the Y/M-type of M3, lack
crenulations on the cheek teeth, and lack the Palaeomeryx-fold on the lower
molars (Fig. 10). In size and morphology, these specimens do not differ from
those referred to L. exilis from the Orellan, The additional Whitneyan specimens
of similar size and morphology in the AMNH collections are also indistinguish-
able from those of L. exilis (Fig. 11). As with L. evansi and L. lenis, it appears
that L. minimus is a junior synonym of L. exilis, thus extending the record of the
latter from the Orellan into the Whitneyan.
Fig. 7 — Cheek teeth of Leptomeryx exilis. A. Occlusal (above) and lingual view of M2-M3, AMNH
81012 (holotype). B. Occlusal view, CM 73412. Bar scale = 1 cm.
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Fig. 8 — Occlusal (above) and lingual views of M3S of Leptomeryx exilis. A. UNSM 1 1605. B. UNSM
1 19606. Bar scale = 1 cm.
Leptomeryx obliquidens was also described from the Whitneyan of South Da-
kota (Lull, 1922). It is much larger than any of the other species of the genus,
nearly twice the size of L. exilis. It is known only from the holotype (YPM 10541)
and one referred specimen (YPM 2248). It has a unique combination of mor-
phologies. The lower molars lack the Palae ornery x-io\d and have the Y/M-type
of entoconulid on M3 (as in L. exilis), but the cheek teeth are highly crenulated,
as in L. evansi (Fig. 12). The upper molars, although heavily worn, show no signs
of a protostyle. No additional specimens referable to this species could be found
Table 4. — Dental measurements of Orellan Leptomeryx exilis. Abbreviations as in Table 1.
Mi-Mj
LM,
WM,
LM2
WM,
LMj
WM.,
N
19
17
17
41
40
68
68
M
19.8
5.4
4.1
6.0
4.5
8.3
4.4
OR
18.0-21.0
4.8-5. 8
3.6-4.5
5. 1-7.0
3.9-5.2
6.1-93
3.4-5.3
sd
0.7
0.3
0.2
0.4
0.3
0.5
0.4
CV
3.7
4.8
6.0
6.4
7.1
6.3
8.0
M'-M^
LM'
WM'
LM"
WM"
LM"
WM"
N
3
4
4
5
5
4
4
M
16.9
5.7
6.0
6.0
6.5
6.1
6.1
OR
16.6-17.2
5.4-5.9
5. 8-6.4
5.9-6.2
5.9-7. 1
5.9-6.4
5.4-6.5
sd
0.3
0.3
0.3
0.1
0.5
0.2
0.5
CV
1.8
3.9 ,
4.5
2.0
7.3
3.6
7.9
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Korth and Diamond — Orellan Leptomeryx from Nebraska
123
mS
7 8
9 10 11
length
Fig. 9 — Scatter diagram of length vs. width of M3 of Leptomyerx evansi-Mke specimens (S/E-type
entoconuiid) from the Whitneyan. Crossed lines indicate range of size of Orellan L. evansi. Darkened
square represents holotype of L. lenis (AMNH 81013). Scales in mm.
in any collections. It appears most likely that L. obliquidens is part of the L.
yoderi—L. mammifer—L. exilis lineage, but that it has attained crenulated molars
separately from those of L. evansi.
Arikareean Leptomeryx
Although several isolated and fragmentary specimens of Leptomeryx have been
reported from the Arikareean (see Webb, 1998:471), only a few species have been
named. Douglass (1903) named L. transmontanus from Montana. However, in an
unpublished dissertation, Rasmussen (1977) demonstrated that this species was
referable to the more derived genus of leptomerycid, Pronodens Koerner, 1940
(also see Taylor and Webb, 1976).
The only other Arikareean species referred to Leptomeryx is L. agatensis from
Sioux County, Nebraska (Cook, 1934). This species is known only from the ho^
lotype, a maxilla with upper cheek teeth (AMNH 81505). In size and morphology,
this specimen is not separable from upper dentitions of L. evansi. Due to the poor
record of this species it is impossible at present to recommend the synonymy of
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Korth and Diamond — Orellan Leptomeryx from Nebraska
125
m3
6 -
length
Fig. 1 1 — Scatter diagram of length vs. width of M3 of Leptomeryx exilis-like specimens (Y/M-type
entoconulid) from the Whitneyan. Crossed lines indicate range of size of Orellan L. exilis. Darkened
circles represent holotype and paratype of L. minimus (AMNH 1347a and AMNH 1347). Scales in mm.
these two species. It is equally as likely that L. agatensis is referable to a species
of the Arikareean leptomerycid Pronodens, the latter differing from Leptomeryx
mainly in the morphology of the anterior dentition which is not preserved in the
holotype of L. agatensis. Webb (1998) listed L. agatensis as a synonym of L.
obliquidens. The marked difference in size of these species makes it very unlikely
that they are synonyms.
Relationships of North American Leptomeryx
Two distinct lineages of Leptomeryx can be recognized from the Chadronian
to the Whitneyan of North America. One contains the Chadronian species L.
yoderi and L. mammifer; the Orellan species L. exilis, and the Whitneyan L.
Fig. 10 — Occlusal (above) and lingual views of M1-M3 Leptomeryx minimus, AMNH 1347 A (holo-
type). Bar scale = 1 cm.
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VOL. 71
Fig. 12 — Cheek teeth of Leptomeryx obliqiiidens . A. P^— M-\ YPM 10541 (holotype). B. P3— M3, YPM 2248. C. Occlusal view (left) and lingual view M3,
YPM 10541 (holotype). Bar scale below each figure = 1 cm.
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Korth and Diamond — Orellan Leptomeryx from Nebraska
127
obliquidens and L. miniumus {1=L. exilis). The other lineage includes the Chad-
ronian L. speciosus, Orellan L. elisscie and L. evansi, the Whitney an L. lenis (?=L.
evansi) and additional species similar to L. evansi, and possibly the Arikareean
L. agatensis.
The only consistent character that separates these two groups throughout their
known record is the morphology of the entoconulid on M3, The L. yoderi-L.
mammifer-L. exilis-L. obliquidens lineage is characterized by a conical entocon-
ulid that is triangular in lingual view, and is lower than the hypconulid and strong-
ly connected to it. In the L. speciosus— L. evansi— L. elissae lineage, the entoconulid
on M3 is cylindrical, equal in height to the hypoconulid, and separated from it by
a deep, distinct valley (see Heaton and Emry, 1996:fig. 9).
In the first lineage, there is little morphological change through time except in
size. There is increase in size during the Chadronian, from L. yoderi to L. mam-
mifer, then a marked reduction in size at the Orellan boundary with L. exilis
which persists into the Whitneyan (?=L. minimus). Also in the Whitneyan there
is a marked increase in size with the appearance of L. obliquidens. The latter also
shows the first change in morphology with the crenulation of the enamel on the
cheek teeth.
The L. speciosus— L. evansi— L. elissae lineage shows an increase in the occur-
rence of the Palaeomeryx-fo\d on the lower molars and increased crenulations on
the cheek teeth from L. speciosus to the Orellan L. evansi along with a reduction
in size. However, the second Orellan species, L. elissae, lacks these derived fea-
tures of L. evansi, and is nearly identical to L. speciosus except for its smaller
size (even smaller than L. evansi). The Whitneyan and later species of this group
appear closely related to L. evansi with the presence of the Palaeomeryx-iold and
crenulations on the cheek teeth.
The only Arikareean or later leptomerycids that may have been derived from
either of these lineages is Pronodens. The latter has slightly higher crowned cheek
teeth than Leptomeryx and a reduction and modification of the anterior dentition
(Taylor and Webb, 1976; Webb, 1998). However, the P alaeomery x-fo\d is more
prominent than in L. evansi and the entoconulid has the S/E-type of morphology
of the L. evansi lineage as well. It is likely that L. evansi or a similar species in
that lineage was directly ancestral to Pronodens.
Acknowledgments
Access to collections and comparative material was generously provided by the following: R. H.
Tedford (AMNH), M. R. Dawson (CM), M. A. Turner (YPM), M. R. Voorhies and R. G. Corner
(UNSM), R. J. Emry (USNM) and T. Heaton of South Dakota State University. Drs. Emry, Heaton,
and Dawson also assisted with discussions of leptomerycid phylogeny. Photographic equipment was
provided by R. Snider of the Department of Biology, Buffalo State College. Figures 2, 5, 6, and 12
were prepared by A. Heyden. This research was funded in part by the Rochester Institute of Vertebrate
Paleontology. Earlier versions of this paper were critically reviewed by S. D. Webb, R. J. Emry, and
M. R. Dawson.
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Cope, E. D. 1 886. The Vertebrata of Swift Current Creek region of the Cypress Hills. Annual Report
of the Geology and Natural History Survey of Canada, 1885:79-85
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Patch Formation, central-western Montana. Unpublished Ph.D. Dissert., University of Kansas,
Lawrence, Kansas.
Schultz, C. B., and T M. Stout. 1955. Classification of Oligocene sediments in Nebraska. Bulletin
of the University of Nebraska State Museum, 4:17-51.
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actions of the American Philosophical Society, 28:363-746.
Storer, j. E. 1981. Leptomerycid artiodactyls of the Calf Creek local fauna (Cypress Hills Formation,
Oligocene, Chadronian) Saskatchewan. Natural History Contributions, Saskatchewan Museum of
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Taylor, B. E., and S. D. Webb. 1976. Miocene Leptomerycidae (Artiodactyla, Ruminantia) and their
relationships. American Museum Novitates, 2596:1-22.
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America (C. M. Janus, K. M. Scott, and L. L, Jacobs, eds.). Cambridge University Press, New
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Appendix 1
Referred specimens of Orellan Leptomeryx evansi examined in this study.
Nebraska (unspecified level in Orella Member)
CM 175 (3 specimens), 185 (2 specimens), 205 (3 specimens), 226 (5 specimens), 362, 442 (3
specimens), 455 (2 specimens), 472, 458, 2497, 3674 (2 specimens), 3679, 3680 (4 specimens). AMNH
53572 (2 specimens), 53573 (9 specimens), 53621, 53627 (3 specimens), 53626 (3 specimens), 330-
726, 330-729, 546-24643, 570-24857, 602-25441, 602-25485, 606-25836, 606-25861, 606-25869,
606-25874, 606-25879, 606-25882, 606-25937, 606-25946 (3 specimens), 61 1-26636, 61 1-26638, 611-
26642, 611-26653, 611-26658, 611-26670, 611-26676, 611-26677, 616-27214 [Lower dental ele-
ments]. CM 205 (7 specimens), 226 (6 specimens), 542, 2497, 3670, 3678, 3682, 3683, 3674 (2
specimens), 3675 (2 specimens), 3676, 3824, 9143, 9144, 9148, 9689 [Upper dental element].
Orella A (Nebraska)
UNSM 119625 through 119630, and 119641 through 119648 [Lower dental elements]. UNSM
1 19649 through 1 19660 [Upper dental elements],
Orella C (Nebraska)
UNSM 119634 and 119661 through 119677 [Lower dental elements!. UNSM 119678 through
1 19698 [Upper dental elementsl.
2002
Korth and Diamond — Orellan Leptomeryx from Nebraska
129
Orella D (Nebraska)
UNSM 119699 through 119706 and 119712, 119713 [Lower dental elements], UNSM 119714
through 1 19726 [Upper denta[ elements].
North Dakota
CM 2010a, 28800, 28803a; USNM 78-5, 78-8, 78-22 [Lower dental elements], CM 2010, 28645
[Upper dental elements].
South Dakota
USNM 9426c (holotype).
Wyoming
AMNH 0-75-1525, 0-81-1611, 0-94-1883, 0-120-2228, 0-442-2044, 058-1379, 088-1691a, 088-
1699 [Lower dental elements].
Appendix 2
Referred specimens of Orellan Leptomeryx exit is examined in this study.
Nebraska (unspecified level in Orella Member)
CM 1289, 73403, 73405, 73407, 7341 1, 73415, 73419, 73420, 73421; AMNH 53572 (2 specimens),
042-973q, 540-24592a, 606-25946 (3 specimens), 606-25875, 611-26678, 615-27163, 616-27220, 616-
27252 [Lower dental elements], CM 73412 [Upper denta[ element],
Orella A (Nebraska)
UNSM 119617, 119631, 119632 [Lower dental elements], UNSM 119621 [Upper dental elements],
Orella C (Nebraska)
UNSM 119600 through 119615, 119618 through 119620, and 119633 [Lower dental elements],
UNSM 119623, 119624 [Upper dental elements],
Orella D (Nebraska)
UNSM 119616 [Lower denta[ e[ement], UNSM 119622 [Upper dental element].
South Dakota
AMNH 38838a, 90281, 208-3596-3, 208-3596-7, 230-4183, 230-4276, 230-4277 [Lower dental
elements], CM 9121; AMNH 25858, 53582a, 53596, 606-25840, 606-25857, 611-26647 [Upper dental
elements].
Colorado
AMNH 6652, 6664, 9002 [Lower dental elements],
Wyoming
AMNH 0-142-2983g [Lower dental element].
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ANNALS OF CARNEGIE MUSEUM
VoL. 71, Number 2, Pp. 131-141
28 May 2002
FROM THE ARCHIVES AND COLEECTIONS
W. J. HOLLAND’S SPEECH AT THE INTERNATIONAL CONGRESS OF
AMERICANISTS, 13TH SESSION, IN 1902
David R. Watters^
The Context of the Address
W, J. Holland, Director of Carnegie Museum, attended the 13th Session of the
International Congress of Americanists (13th ICA) which convened at the Amer-
ican Museum of Natural History in New York City from October 20-25, 1902.
Holland’s participation in the 13th ICA can be tracked through entries in its
Proceedings volume published three years later (International Congress of Amer-
icanists, 1905). The volume indicates he was a Member of the General Commis-
sion of Organization (p. xi). Delegate of the United States (p. xiv), Subscriber (p.
xvii), and Member of the Council (p. xxii).
Holland twice addressed the Congress on Monday, October 20, the day the
13th Session opened. In his initial address, which is the subject of this paper, he
spoke to the Congress regarding the status of Carnegie Museum. Later that af-
ternoon he read his paper entitled “The Petroglyphs at Smith’s Ferry, Pennsyl-
vania.” His speech and paper (Holland, 1905<7, 1905/?) were published in the
Proceedings.
Holland was invited to the dais by the preeminent American anthropologist of
the time, Frederic Ward Putnam, who was chairing the Monday session in his
role as a Vice President (representing the United States) of the 13th ICA. Holland
was the second speaker. He was preceded by W. J. McGee of the Bureau of
American Ethnology, Smithsonian Institution, and succeeded by Stewart Culin
(Archaeological Museum of the University of Pennsylvania), George A. Dorsey
(Field Columbian Museum), Edward S. Morse (Peabody Academy of Science),
Frederick Starr (University of Chicago and Davenport Academy of Sciences), and
Putnam (Peabody Museum of Harvard University and the American Museum of
Natural History), each of whom reviewed the anthropological work conducted by
his institution (International Congress of Americanists, 1905:xxvi-xliv). Although
Holland’s scientific work was in entomology, not anthropology, he shared the
lectern that day with some of the major figures in the emerging discipline of
anthropology. Holland became acquainted with many prominent North American,
Latin American, and European anthropologists during the 13th ICA.
The Content of the Address
Holland’s speech makes it clear that he regarded the 13th ICA as a fine venue
for informing Americanists, anthropologists, and museologists about the efforts
being made in the field of anthropology by the “infant” Carnegie Museum.
Reproduced below in their entirety are Putnam’s introductory remarks, Hol-
‘ Curator, Section of Anthropology
131
132
Annals of Carnegie Museum
VOL. 71
land’s speech, and Putnam’s closing comments made in 1902, as published in
1905 in the 13th ICA Proceedings:
[Vice President F. W. Putnam speaking] I call upon Dr. W. J. Holland,
the director of one of the younger museums (that is, younger in the
sense that it has only recently done anything in anthropology) to briefly
outline the work of the Carnegie Institute of Pittsburg.
W. J. Holland, of the Carnegie Museum, in responding to the call of
the Chair, said:
Mr. President, Ladies and Gentlemen. — The Carnegie Museum is in
its infancy, but it is a strong and vigorous infancy. It has already out-
grown its swaddling clothes and the manger in which it has been lying.
It is now waiting patiently for a decision from the Supreme Court of
Pennsylvania, a decision which will undoubtedly allow the manger to
be transformed into a bed. There are four or five millions of dollars,
thanks to the generosity of the founder, at the service of the Museum
for enlarged buildings, but the money cannot be expended until the
ground is obtained upon which to put these buildings, and a lawsuit
now pending prevents this. Meanwhile we are at work gathering up
collections which are to be placed in the halls of the future edifice. We
are hiring storerooms. We are, to a certain extent, in the “cold storage”
business. Good storage is cold, and should be kept cold for the pres-
ervation of collections which are made. Like some very religious peo-
ple, we are afraid of fire.
Our archaeological work has largely been thus far confined to the
valleys of the Monongahela and the Allegheny, which unite at Pittsburg
to form the Ohio. We here find mounds, shell-heaps, sites of ancient
villages, and sculptured rocks, and we are attempting to map out the
location of these memorials of the primitive peoples who once inhabited
these portions of western Pennsylvania, and to preserve, so far as pos-
sible, the scanty records which relate to the aboriginal inhabitants.
We are also receiving from other places a great deal of interesting
material. We are under obligations to the Honorary President of the
present meeting [13th ICA], the Duke of Loubat, for notable gifts, and
we are under obligations to Mr. [Andrew] Carnegie, the founder, for
grants of money which have enabled us to purchase things that are
desirable for exhibition in order to illustrate the archaeology of the
country at large. We have recently secured considerable sums of money
to aid the work of exploration in Egypt, and we have received in return
from the Egypt Expedition Eund some very interesting collections. From
Colombia, Mexico, India, Japan, and indeed from all parts of the world
we have obtained material, which we are classifying. Ultimately we
hope to have a large and representative museum in which the student
of archaeology will be able to work with satisfaction. We wish to make
our work thorough, scholarly, and complete.
Referring again to our immediate neighborhood, I may say that there
is not very much of surpassing archaeological interest left for us there.
The forefathers shot the Indians, or poisoned them with bad whiskey.
They were poor people, who did not have much in the way of goods
2002
Watters — From the Archives and Collections
133
and chattels, and all that remains of them are the few tools and stone
implements which they carried, and which we find in their graves.
I have the honor, sir, on behalf of the Carnegie Museum, to extend
to this Congress an invitation to visit the institution a week from the
coming Wednesday. I understand that you will be with us at that time.
We have provided for the accommodation of the delegates. We will give
you your breakfast, your luncheon, and your dinner. We will speed you
as parting guests to “Fort Ancient,” somewhere in the “howling wil-
derness” of southern Ohio, to which point Professor Putnam will prob-
ably conduct you. When you come to visit us you shall see for your-
selves what we are doing in our museum, the youngest institution of its
kind in America. The great works of the United States Steel Corpora-
tion, through the courtesy of its president, will be open for inspection,
and a train will be provided to take you there and bring you back to
dinner.
Vice-President Putnam: We thank Dr. Holland for the information he
has given us, and for the cordial invitation to the delegates to meet at
Pittsburg and make that one of the stopping-places on the excursion
which will follow the meeting of the Congress. (International Congress
of Americanists, 1905:xxxi— xxxii).
Putnam referred to Carnegie Institute in his introduction, whereas Holland
spoke of Carnegie Museum. In 1902 Carnegie Museum was one of the compo-
nents of Carnegie Institute, the parent organization; the other components were
the Department of Fine Arts and the Music Hall. Also, Pittsburg was the officially
sanctioned spelling, the “h” having been dropped by the U.S. Geographic Names
Board in 1890 (a decision rescinded in 1911 after protest), but both spellings were
in use at the time (e.g., Carnegie Institute, 1907). Holland’s allusions to “. . .
swaddling clothes and the manger ...” and “. . . very religious people ...” in
the first paragraph seem peculiar until one realizes that he was an ordained min-
ister, having served as pastor of the Bellefield Presbyterian Church in the Oakland
section of Pittsburgh from 1874 until 1891 (Van Trump, 1966:219-220). Holland
subsequently served as Chancellor of the Western University of Pennsylvania
(now the University of Pittsburgh) from 1891 to 1901.
The Significance of the Address
At the time W. J. Holland (Fig. 1) attended the 13th ICA, he had been Director
of Carnegie Museum for only four years, having accepted that position in 1898
at the behest of his friend Andrew Carnegie (Avinoff, 1933). Between 1898 and
1901, he concurrently served as the Director of Carnegie Museum and Chancellor
of the Western University of Pennsylvania. He succeeded the first director, ar-
chaeologist Frank H. Gerrodette, who had resigned in 1896 following a public
brouhaha over his excavation of the McKee’s Rocks Mound near Pittsburgh (Rich-
ardson, 1980:13). A letter in the Carnegie Museum of Natural History Archives,
written by Herbert H. Smith (1897), who identified himself as “Curator” but
seems to have been serving as the acting director, states that Gerrodette had been
elected to the directorship in May, had excavated the McKee’s Rocks Mound in
July, August, and early September, and resigned in September 1896. Gerrodette,
who had been F. W. Putnam’s student at Harvard University, arranged for Putnam
to visit the excavation at McKee’s Rocks Mound during the controversy in 1896.
134
Annals of Carnegie Museum
VOL. 71
Fig. 1 — William Jacob Holland, Director of Carnegie Museum, c. 1910. (Carnegie Museum of Natural
History Archives)
Ironically, it was Putnam who introduced Holland when he spoke to the 13th ICA
six years later.
Holland’s comment about ‘k . . enlarged buildings ...” in the first paragraph
of his speech refers to the planned addition to the original 1895 building. The
settling of the lawsuit to which Holland refers opened the way for the completion
2002
Watters — From the Archives and Collections
135
of the addition, with the new building being dedicated in 1 907 (Carnegie Institute,
1907). Carnegie Museum collections held formerly in '7 . . cold storage ...” thus
finally were displayed in fitting exhibition spaces.
The second and fourth paragraphs relate to the archaeological work carried out
in the vicinity of Pittsburgh. Holland’s comment regarding Carnegie Museum’s
efforts to map the “. . . mounds, shell-heaps, sites of ancient villages, and sculp-
tured rocks ...” was a fairly accurate representation, for the time, of the state of
knowledge about the range of archaeological sites in the region. Despite Holland’s
allusion to Carnegie Museum’s early research efforts, almost a half-century would
pass before an intensive program of systematic archaeological research was in-
augurated in 1950 through the Upper Ohio Valley Archaeological Survey, directed
by Curator James L. Swauger with the fieldwork being carried out initially by
William Mayer-Oakes and Donald W. Dragoo (Richardson, 1980:15-16). Hol-
land’s characterization of the local Native Americans as “. . . primitive peoples
. . and “. . . poor people ...” who left behind “. . . not very much of surpassing
archaeological interest . . .” reflects the prevailing attitude of his times. He ac-
knowledges the forefathers’ role in the decimation of these peoples, although his
comment seems to be simply a factual statement without any clear indication of
regret.
The third paragraph discloses the areas of the world from which anthropological
collections then were being obtained. Acquisition of collections for eventual ex-
hibition in the enlarged building clearly was a priority for Holland, and it was a
task that was very much in line with the needs of the “infant” Carnegie Museum.
His comment about Carnegie Museum having received Egyptian collections from
the Egypt Exploration Fund (EEF) in return for “. . . sums of money ...” refers
to that organization’s Pittsburgh chapter, whose formal charter dates to 1901,
although it existed even earlier (Watters and Patch, 1986:32-33). Interest in cre-
ating this chapter can be tracked to 1898, when the Museum Committee passed
the following resolution:
EGYPTIAN EXPLORATION. Resolved: That the Trustees express their
gratification that it is contemplated to establish a centre [sic] of the
Egyptian Exploration Fund, here in Pittsburgh, and that so far as prac-
ticable the authorities of the Museum will cooperate and offer the Mu-
seum as a depository for such objects as may be obtained by this center.
(Carnegie Museum, Minutes of the Museum Committee [for October
31, 1898], voL II, January 3, 1898 to April 1, 1905).
Holland documents his involvement in the founding, recruitment of new members,
and fund-raising efforts of the Pittsburgh EEF chapter in six of his monthly reports
to the Museum Committee (Holland 1899, 1901 <2, 1901fi, 1901c, 1901 <7, 1902fi).
Other collections that Holland mentions include H. H. Smith’s archaeological
collection from Santa Marta, Colombia; a robe donated by the Maharajah of
Benares, India; casts of Mexican sculptures donated by Andrew Carnegie, who
arranged for their reproduction through the Duke of Loubat; and loans of various
Japanese items. Facsimile publications of Mexican codices were the “. . . notable
gifts ...” presented by the Duke of Loubat.
The speech’s final paragraph invites the Congress participants to visit Carnegie
Museum. The 13th ICA Proceedings volume (p. Ixvii) discloses that the post-
Congress excursion included visits to Philadelphia, Washington, D.C., Pittsburgh,
136
Annals of Carnegie Museu. .
VOL. 71
Fort Ancient in southern Ohio, Cincinnati, and Chicago. Holland tendered this
invitation with the approval of the Museum Committee:
INTERNATIONAL CONGRESS OF AMERICANISTS. The Director
presented a letter from M. H. Saville, General Secretary of the Inter-
national Congress of Americanists, stating that about forty members of
this society desired to come to Pittsburgh about October 29th and that
they would like to be invited to visit the Carnegie Museum and Institute.
The Director was, on motion, requested to invite the members accord-
ingly and to anange for their entertainment. (Carnegie Museum, Min-
utes of the Museum Committee [for September 29, 1902], vol. II, Jan-
uary 3, 1898 to April 1, 1905).
The delegation toured Carnegie Institute and other points of interest in Pittsburgh
on October 29, and Holland took advantage of that opportunity to once again
expound upon his ideas for a new Carnegie Museum. Holland’s invitation to visit
the United States Steel Corporation works (they visited the Homestead plant)
seems somewhat incongruous until one realizes that this formerly was the Car-
negie Steel Company, the foundation of Andrew Carnegie’s wealth and his finan-
cial support of Carnegie Museum. Pittsburgh newspaper accounts of the event
provide a means of identifying the foreign delegates and some of the institutions
and countries they represented (Table 1). One newspaper even published a pho-
tograph of the delegates (Fig. 2). According to the newspapers, the delegation’s
itinerary, after the stop in Chicago, had been expanded to include visits to St.
Louis (hosted by the Louisiana Purchase Commission), New York City, and fi-
nally Boston and Cambridge.
Holland’s speech to the 13th ICA is a valuable document because it is one of
the few records yet discovered on the status of anthropology at Carnegie Museum
during the early years of his directorship. Although brief, the speech nonetheless
provides important insight into Holland’s perception of local archaeological re-
sources, intentional acquisition of anthropological collections from around the
world in preparation for their installation in the new building, and reliance upon
Andrew Carnegie for the financial means to obtain collections and expand the
facility. Holland readily used opportunities afforded him by his speeches at the
1 3th ICA and to the visiting delegation the following week to promote his vision
of a new Carnegie Museum to international, national, and local audiences.
The Significance of Holland’s Attendance
Holland attended the 13th Session of the International Congress of Americanists
for several reasons. The Museum Committee had formally endorsed his partici-
pation:
CONGRESS OF AMERICANISTS. On motion. Dr. W. J. Holland was
appointed to represent the Carnegie Museum at the Thirteenth Annual
Meeting of the International Congress of Americanists, to be held in
New York in 1902. (Carnegie Museum, Minutes of the Museum Com-
mittee [for July 30, 1901], vol. II, January 3, 1898 to April 1, 1905).
The 13th ICA was the first session convened in the United States; all previous
sessions had been held at European locations (except for the 1 1th Session, a
special meeting in Mexico in 1895). At the 12th ICA in Paris in 1900, it was
2002
Watters — From the Archives and Collections
137
Table 1. — 13th IC A foreign delegates visiting Carnegie Museum on October 29, 1902.
Argentine Republic
Senor Juan B. Ambrosetti
Costa Rica
Professor Henri Pittier de Fabrega
Professor Juan Fernandez Ferraz [incorrectly listed as Juan H. Ferraz]
France
Professor Leon Lejeal, College de France, Paris
Senor M. Gonzalez de la Rosa, Paris [misspelled as Gongalez]
Germany
Dr. Eduard Seler, German Government [misspelled as Edward]
Professor Karl von den Steinen, University of Berlin
Great Britain
Professor John H. Biles, University of Glasgow, Scotland
A. P. Maudslay, Anthropological Institute of Great Britain and Ireland [misspelled as Mandslay]
Honduras
Senor N. Bolet Peraza
Italy
Signor Giovanni Branchi
Mexico
Senor Alfredo Chavero
Senor Francisco Belmar, State of Oaxaca
The Netherlands
Professor J. L. van Panhuys
Paraguay
Senor Alonzo M. Criado
Russia
Professor Waldemar Bogarras
Sweden
Professor C. V. Hartman, Swedish Anthropological Society, Stockholm
Professor Hjalmar Stolpe, Royal Ethnological Museum, Stockholm
Venezuela
Senor Genera Garcia
Notes;
(1) The 19 foreign delegates are listed identically in six local newspapers {Leader, Times, Dispatch,
and Chronicle-Telegraph from October 29, and Post and Gazette from October 30); the list prob-
ably was derived from a Carnegie Museum press release since errors in spelling are consistent in
the newspapers.
(2) Most newspapers state that 31 persons formed the delegation; the twelve individuals not named
may have been spouses of delegates and/or American delegates.
(3) The Leader of October 29 mentions five women among the delegation; at least two women are
visible in the photograph; the Times of October 30 mentions Miss Adela Breton (an Englishwom-
an) and the unnamed wife of Dr. Seler.
decided to alternate future sessions between venues in Europe and the Americas,
and the 13th ICA in New York City inaugurated that scheme. The International
Congress of Americanists, the premier anthropological meeting of the times,
brought together participants from throughout the Americas and Europe, and the
13th ICA thus afforded Holland an opportunity to meet many eminent anthro-
pologists. Holland also expanded his contacts in the museum field since many of
the 13th ICA participants were employed by or otherwise associated with major
museums of the day, serving either as anthropologists or administrators on their
staffs.
Holland was acquainted with some major figures in American anthropology
even before he attended the 13th ICA. In June 1902, only four months before the
138
Annals of Carnegie Museum
VOL. 71
Fig. 2 — Photograph from the Dispatch (October 30, 1902 edition, p. 8) of the visiting delegation of
the International Congress of Americanists at the Schenley Hotel in Oakland. (Carnegie Museum of
Natural History Archives, Holland newspaper clipping scrapbook)
13th ICA, the founding meeting of the American Anthropological Association
(AAA) convened in the Oakland section of Pittsburgh (McGee, 1902^, \902b).
The founding meeting was held in conjunction with the American Association
for the Advancement of Science’s (AAAS) annual meeting, which was being
attended by anthropologists from its Section H (Anthropology). It was announced
beforehand that the AAA founding meeting would be held in the . audience
room of the Bellefield Church, on Monday, June 30, at 2 o’clock” (McGee,
1902<2:353), which must have refen'ed to the Bellefield Presbyterian Church (there
being no other Bellefield church in Pittsburgh), the same church where Holland
had served as pastor until 1891 and remained a member. However, McGee (1902/?:
464) afterwards reported the “. . . Oakland Church ...” to have been the venue
of the June 30 meeting. He did not mention the denomination of the church in
either article.
I initially thought McGee’s use of “Oakland Church” simply was a generalized
reference to the Bellefield Presbyterian Church located in Pittsburgh’s Oakland
section, an assumption that seemed logical and warranted given Holland’s per-
sonal ties to that church. However, a newly discovered document brings the va-
lidity of that assumption into question. Holland, when referring to room arrange-
ments being made for the entire AAAS meeting (not specifically the AAA found-
ing meeting), stated:
The Trustees of the Bellefield Presbyterian Church and of the Oakland
Methodist Episcopal Church have kindly tendered the use of their re-
spective buildings for the coming [AAAS] meeting, and I have secured
an option upon the Music Hall and the small lecture room of the Insti-
tute. (Holland, 1902c/).
McGee’s use of the term “Oakland Church” has an equally plausible chance of
having referred to the Oakland Methodist Episcopal Church as the venue for the
AAA founding meeting.
2002
Watters — From the Archives and Collections
139
McGee (1902l?:477) did establish one definite link between Holland and the
visiting anthropologists when he acknowledged Holland's courtesy in arranging
for Carnegie Museum to exhibit a Mexican sculpture discussed by archaeologist
Marshall Saville during the Section H scientific sessions. That Holland is iden-
tified as one of the founders of the American Anthropological Association (Anon-
ymous, 1903:191) provides additional evidence of his early connections with an-
thropologists. Thus, in the audience listening to Holland’s speech and paper at
the 13th ICA were some of the same anthropologists with whom he had interacted
in Pittsburgh only four months before.
Holland attended the 13th ICA for another reason. He was interested in ex-
ploring the possibility of hiring an anthropologist to become Curator of the Sec-
tion of Ethnology and Archaeology at Carnegie Museum, and the meeting pro-
vided him the opportunity to meet possible candidates. In late January 1903, three
months after the 13th ICA ended, Holland offered the curatorship to Stewart
Culin. Culin, representing the Archaeological Museum of the University of Penn-
sylvania, had spoken immediately after Holland at the 13th ICA. He declined
Holland’s offer in early February.
Holland then entered into negotiations with Carl Vilhelm Hartman for the cu-
rator position, and Hartman accepted the offer on February 28 (Watters and Fon-
seca Zamora, 2001). Hartman had attended the 13th ICA as a representative of
the Swedish Society for Anthropology and Geography, and he was one of the
delegates visiting Pittsburgh on the post-Congress excursion (Table 1). Having
twice heard Holland expound upon his vision for Carnegie Museum, Hartman
was aware of the Director’s desire to acquire collections. Hartman made it a point
in his letters to Holland to emphasize his personal knowledge about collections
of antiquities available for purchase in Costa Rica, where he previously had con-
ducted archaeological research in 1896-1897 on a Swedish expedition (Brunius,
1984). Hartman reported for duty on March 17, 1903. Two weeks later, Holland
sent him to Costa Rica on his second expedition. Two Costa Rican delegates,
Henri Pittier de Fabrega and Juan Fernandez Ferraz, with whom Holland became
acquainted at the 13th ICA through Hartman and during their subsequent visit to
Pittsburgh (Table 1), facilitated the Carnegie Museum expedition. Hartman pre-
sented letters of introduction, written by Holland, to both of them when he arrived
in Costa Rica. Thus, Holland’s attendance at the 13th ICA led not only to the
hiring of an anthropology curator, but also resulted in the acquisition of collections
from Costa Rica for the “infant” Carnegie Museum.
There are other connections between Carnegie Museum and the 13th ICA. C. C.
Mellor, Chairman of the Museum Committee, is listed in its Proceedings volume
(p. xviii) as a “Subscriber,” although whether he actually attended the Congress
is uncertain. Four months later, Mellor’s committee approved Holland’s recom-
mendation to hire Hartman (Watters and Fonseca Zamora, 2001). In his Monthly
Report to the Museum Committee, delivered at its February 28, 1903 meeting
(the same day Hartman was hired), Holland wrote:
Dr. C. V. Hartmann [sic], whose acquaintance I formed at the recent
Congress of Americanists, is in the building, and I shall take pleasure
in presenting him to you at sometime this evening. He has had expe-
rience in Museum work, having been associated with Dr. Stolpe, the
celebrated Swedish ethnologist, whose assistant he was at Stockholm.
He was for a long time with Dr. Fumholtz in Mexico and has written
140
Annals of Carnegie Museum
VOL. 71
extensively upon the ethnology of Costa Rica. It may be that he is the
man, for whom we are looking. He produces excellent testimonials.
(Holland, 1903).
Holland refers to Dr. Hjalmar Stolpe, the person responsible for mentoring Hark
man in museum work (Watters and Fonseca Zamora, 2001). Stolpe had attended
the 13th ICA, serving as the Vice President representing Sweden (as Putnam did
for the United States), and during the excursion thereafter he visited Carnegie
Museum (Table 1). Stolpe was Director of the Ethnological Section of the Mu-
seum of Natural History of Sweden, and he was one of the contacts made by
Holland among the museologists at the Congress.
The 13th ICA was the only session Holland ever attended. However, he retained
an interest in the organization and is listed as a subscriber in the Proceedings of
some of its later sessions. The final Proceedings volume in which his name ap-
pears is from the 23rd Session, which lists him as Director Emeritus of Carnegie
Museum but specifies he was “Not in attendance” (International Congress of
Americanists, 1930:xxvii). Holland died December 13, 1932 (Avinoff, 1933). The
Library of the Carnegie Museum of Natural History holds the Proceedings from
most sessions of the International Congress of Americanists convened between
the 12th ICA in Paris in 1900 and the 23rd ICA in 1928 in New York City.
Appropriately, the Duke of Loubat donated the 12th Proceedings volume while
the 23rd Proceedings came from the Estate of W. J. Holland. The Library’s pres-
ervation of the ICA Proceedings serves as a fitting tribute to Holland’s partici-
pation in the 13th Session of the International Congress of Americanists one
hundred years ago, when the “infant” Carnegie Museum was beginning its vig-
orous growth under his directorship.
Acknowledgments: Throughout Holland’s tenure as Director, the institution was
known as Carnegie Museum, only being changed in 1973 to the Carnegie Museum
of Natural History, I discovered Holland’s association with the 13th ICA during
my research on the life of C. V. Hartman, supported by a grant graciously awarded
by the Adrienne and Milton Porter Charitable Foundation. I am indebted to
CMNH Research Associates Hazel Johnson for obtaining newspaper accounts
about the delegation visiting Pittsburgh, researching the churches related to the
AAA founding meeting, and tracking biographical information about W. J. Hol-
land, and Oscar Fonseca Zamora (University of Costa Rica) for providing infor-
mation about Henri Pittier de Fabrega and Juan Fernandez Ferraz, I am grateful
to CMNH Librarian Bernadette Callery for access to archival resources, Anthro-
pology Collection Manager Deborah Harding for information about the collections
mentioned by Holland, Anthropology Secretary Charmaine Steinberg for assisting
in manuscript preparation, and Staff Photographer Mindy McNaugher for pho-
tographing the newspaper clipping of the visiting delegates.
Literature Cited
Anonymous. 1903. The American Anthropological Association. American Anthropologist, 5:178-
192.
Avinoff, A. 1933. Obituary: William Jacob Holland. Annals of Carnegie Museum, 21:i-iv.
Brunius, S. 1984. Carl V. Hartman: svensk arkeolog-etnograf i Centralamerika. Institute of Latin
America Studies Occasional Papers, Stockholm.
Carnegie Institute. 1907. Guide to the Dedication of the New Buildings of the Carnegie Institute
at Pittsburgh, Pennsylvania, April 11, 12, 13, 1907. Carnegie Institute, Pittsburgh.
Holland, W. J. 1899. Monthly Report [for September] of the Director to the Museum Committee,
2002
Watters — From the Archives and Collections
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October 2, 1899. Monthly Reports of the Director, vol. I, January 1897-May 31, 1905. Carnegie
Museum of Natural History Archives, Pittsburgh.
. 1901<:/. Monthly Report [for December 1900J of the Director to the Museum Committee,
January 5, 1901. Monthly Reports of the Director, vol. I, January 1897-May 31, 1905. Carnegie
Museum of Natural History Archives, Pittsburgh.
. \90\h. Monthly Report of the Director to the Museum Committee, April 30, 1901. Monthly
Reports of the Director, vol. I, January 1897-May 31, 1905. Carnegie Museum of Natural History
Archives, Pittsburgh.
. 1901c. Monthly Report of the Director to the Museum Committee, May 31, 1901. Monthly
Reports of the Director, vol. I, January 1897-May 31, 1905. Carnegie Museum of Natural History
Archives, Pittsburgh.
. 1901r/. Monthly Report of the Director to the Museum Committee, November 30, 1901.
Monthly Reports of the Director, vol. I, January 1897-May 31, 1905. Carnegie Museum of Natural
History Archives, Pittsburgh.
. 1902fl. Monthly Report [for January] of the Director to the Museum Committee, February
I, 1902. Monthly Reports of the Director, vol. I, January 1897-May 31, 1905. Carnegie Museum
of Natural History Archives, Pittsburgh.
-. 1902b. Monthly Report [for February] of the Director to the Museum Committee, March 3,
1902. Monthly Reports of the Director, vol. I, January 1897-May 31, 1905. Carnegie Museum of
Natural History Archives, Pittsburgh.
. 1903. Monthly Report [for February] of the Director to the Museum Committee, March 1,
1903. Monthly Reports of the Director, vol. 1, January 1897-May 31, 1905. Carnegie Museum
of Natural History Archives, Pittsburgh.
. 1905a. [Address about Carnegie Museum to the 13th ICA], Proceedings of the International
Congress of Americanists, 13th Session, 1902:xxxi-xxxii. Eschenbach Printing Co., Easton, Penn-
sylvania.
. 1905Z?. The Petroglyphs at Smith’s Perry, Pennsylvania. Proceedings of the International
Congress of Americanists, 13th Session, 1902:1-4. Eschenbach Printing Co., Easton, Pennsyl-
vania.
International Congress of Americanists. 1905. Proceedings of the International Congress of Amer-
icanists, 13th Session, 1902. Eschenbach Printing Co., Easton, Pennsylvania.
-. 1930. Proceedings of the Twenty-third International Congress of Americanists, New York,
1928. Science Press Printing Co., Lancaster, Pennsylvania.
McGee, W. J. 1902a. Proposed American Anthropologic Association. American Anthropologist, 4:
352-353.
-. 1902Z?. Anthropology at Pittsburg. American Anthropologist, 4:464-481.
Richardson, J. B., III. 1980. The Section of Man. Carnegie Magazine, 54(9): 13-20.
Smith, H. H. 1897. Report to C. C. Mellor, Chairman of the Museum Committee, January 13, 1897.
Monthly Reports of the Director, vol. 1, January 1897-May 31, 1905. Carnegie Museum of Nat-
ural History Archives, Pittsburgh.
Van Trump, J. D. 1966. Bellefield’s Tower: The Centenary of the Bellefield Presbyterian Church.
The Western Pennsylvania Historical Magazine, 49:213-225.
Watters, D. R., and O. Fonseca Zamora. 2001. C. V. Hartman’s Letter of Pebruary 20, 1903 to W.
J. Holland. Annals of Carnegie Museum, 70:263-268.
Watters, D. R., and D. C. Patch. 1986. Pittsburgh Discovers Ancient Egypt. Carnegie Magazine,
58(2):30-37.
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4:
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VOLUME 71 16 AUGUST 2002 NUMBER 3
CONTENTS
ARTICLE
Revision of the New World abariform genera Neotalus n.gen. and Abaris
Dejean (Coleoptera: Carabidae: Pterostichini (Auctorum))
Kipling W. Will 143
FROM THE ARCHIVES AND COLLECTIONS
W. J. Holland’s roles in the 1902 meetings of the American Association for the
Advancement of Science and the American Anthropological Association
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ANNALS OF CARNEGIE MUSEUM
VoL. 71, Number 3, Pp. 143-213 16 August 2002
REVISION OF THE NEW WORLD ABARIFORM GENERA
NEOTALUS N.GEN. AND ABARIS DEJEAN
(COLEOPTERA: CARABIDAE: PTEROSTICHINI (AUCTORUM))
Kipling W Will^
Research Associate, Section of Invertebrate Zoology
Abstract
New World ground beetles (Coleoptera: Carabidae) in the abariform genus-group are classified into
three genera, Pseudabarys Chaudoir, Neotalus new genus and Abaris Dejean. Species of the clade
Neotalus + Abaris are revised. Based on the cladistic analysis of adult characteristics, a preferred
hypothesis groups species of Abaris into clades that are designated as two subgenera and four species-
groups: Abaris sensu stricto containing three clades designated as species-groups; A. striolata, A.
aenea, and A. picipes-groups and subgenus Abaridius Chaudoir (type species Abaris tachypoides)
comprised of the A. tacky poides-gvonp and two species placed sedis mutabilis within the subgenus.
Seventeen new species are described in the genus Abaris: A. napoensis, A. bicolor, A. nitida, A. nigra,
from Ecuador; A. franiai, A. inflata, A. convexa, A. inaequaloides, A. wardi, from Bolivia; A. im-
punctata, A. nobilis, from Brazil; A. metallica from Venezuela; A. retiaria, from Colombia and Ve-
nezuela; A. erwini from Peru and Bolivia; A. opaca, from Peru, Bolivia and Brazil; A. mina, from
Argentina, Brazil, Bolivia, and Paraguay; and A. aquilonaria from Central America and Mexico.
Abaris darlingtoni Straneo and A. aenea Dejean are synonymized, the latter is the senior, and hence
valid name of the species. The genus Neotalus is described to include N. portai (Straneo); the species
was originally included in Bothynoproctus Tschitscherine. Descriptions, distributional information,
illustrations of morphological structures, and a key to Neotalus and Abaris species are provided.
Distributional patterns show differences between the A. picipes-group species and their sister clade
the A. aenea-group, with the former being more eurytopic and generally adapted to drier habitats.
Biogeographical analysis of biotic zones, using items of error as the optimality criterion, relative to
the phylogenetic hypothesis for Neotalus + Abaris, suggests South American diversification and re-
striction until at least the Pliocene. Abaris species have probably moved into Central America and
Mexico four to five times since the development of the land connection with South America in the
Pliocene. This contrasts against the biogeographical pattern for Coptodera Dejean species (Lebiini),
which appear to have basaily diverged among the northern areas of Mexico versus South America.
Key Words: New species, systematics, biogeography, classification
Introduction
Members of the Neotropical genera Neotalus new genus and Abaris Dejean are
small ground beetles (Coleoptera: Carabidae), most individuals are brilliantly me-
tallic with very prominent eyes (Fig. 1). Abaris species are uniquely characterized
among all Pterostichine grade taxa by pectinate claws (Fig. 2A). These charac-
teristics give them a unique look that is approached in Pseudabarys Chaudoir,
1873 and Prosopogmus Chaudoir, 1865, both presumed to be closely related gen-
era. I discovered from examination of museum specimens that many more species
than the nine named forms of Abaris existed. The captivating form and unstudied
diversity were the starting points for an interest in revising this group.
A small body of literature treats the abariform species. Straneo (1939) published
a key to the eight then named species of Abaris and Bousquet (1984) added a
' Department of ESPM, Division of Insect Biology, University of California, Berkeley, CA 94720-
3112, e-mail: kiplingw@nature.berkeley.edu.
Submitted 10 October 2001.
143
144
Annals of Carnegie Museum
VOL. 71
Fig. 1. — Photograph of living Abaris splendidula (LeConte). Collected in Cochise Country, Arizona.
ninth species when he discovered that LeConte’s Pterostichus splendidulus be^
longed in the genus Abaris. Bousquet and Liebherr (1994) summarized what little
had been published on the relationships of the genus Abaris and covered nearly
all the genera included herein. The complex of genera thought to be closely related
and/or associated in published classifications (Bates, 1882; Blackwelder, 1944;
Straneo, 1977, 1979; Csiki, 1930; Reichardt, 1977) includes the Nearctic-Neo-
tropical genera A Abaridius Chaudoir, Ophryogaster ChAudoix, 1878, Pseu-
dabarys; the northern African=Palearctic genus Orthomus Chaudoir, 1838; and the
Australian genus Prosopogmus. My previous analysis of both adult and larval
characters (Will, 2000) supports a close relationship of these genera, as well as
Blennidus Motschulsky, 1866, Oribazus Chaudoir, 1874, Dyschromus Chaudoir,
1835, Abacillius Straneo, 1949 (referred to in Will, 2000 as “genus E”), and
Argutoridius Chaudoir, 1876. These genera form, in part, a clade centering on the
subtribe Euchroina (auctorum) and the setalis series (Moore, 1965).
An exceedingly short or absent coronal suture in the larva was considered by
Bousquet and Liebherr (1994) as a potential synapomorphy for Abaris and Or-
thomus species. This condition is also found in Pseudabarys (Will, 2000) and is
a significant character for a clade including some or all of the taxa above.
The combination of large eyes, pectinate claws, and metallic luster suggests
that Abaris species may have peculiar life histories or behaviors relative to related
genera that have characteristics more typical for pterostichine grade taxa. How-
ever, the limited field observations for these species do not provide incontrovert-
ible evidence for any adaptive significance for this suite of characteristics. In fact,
observations of any sort exist for very few of these beetles. Specimens of these
species are uncommon in collections. In addition to my own field work, some
recent efforts by carabid collectors such as T. L. Erwin (USNM) and G. E. Ball
(UASM) have resulted the in collection of large numbers of specimens from a
few locations in Central and South America. I hope this monograph will provide
researchers and students, particularly those in South America, with the tools to
2002
Will — Revision of Abaris
145
identify and study species of Abaris so that a greater understanding of their life
history and ecology can be obtained.
Materials, Methods, and Terms
Taxonomic Materials
Material examined in the course of this study included 1025 adults specimens
of Abaris and Neotalus species. A number of these were examined in detail in-
cluding dissection of various internal systems. Material was taken on loan from
collections listed below. Museum codens are used in the text and the names listed
here are the individuals I corresponded with to obtain material and/or those who
processed the loans for me.
Museums and Codens. — AMNH: Department of Entomology, American Mu-
seum of Natural History, New York, New York (L. H. Herman).
BMNH: Department of Entomology, British Museum, London (Natural His-
tory) (S. Hine).
BORD: C. Bordon collection, Maracay, Venezuela (C. Bordon).
BPBM: Bishop Museum, Honolulu, Hawaii (G, A. Samuelson).
CASC: Department of Entomology, California Academy of Sciences, San Fran-
cisco, California (D. H. Kavanaugh, R. Brett).
CMNC: Entomology, Canadian Museum of Nature, Ottawa, Ontario.
CMNH: Section of Invertebrate Zoology, Carnegie Museum of Natural History,
Pittsburgh, Pennsylvania (R. L. Davidson).
CNCI: Canadian National Collection, Biosystematics Research Institute, Agri-
culture and Agri-Food Canada, Ottawa, Canada (Y. Bousquet).
CUIC: Department of Entomology, Cornell University, Ithaca, New York (J. K.
Liebherr).
EMEC: Essig Museum of Entomology, Berkeley, University of California,
Berkeley, California (C. Barr).
INBC: Instituto Nacional de Biodiversidad (INBio), Costa Rica, Santo Domin-
go de Heredia.
IZWP: Instytut Zoologii, Polska Akademia Nauk, Warszawa, Poland (T. Hu-
flejt).
KWWC: Kipling Will Collection, Berkeley, California.
MSNM: Museo di Storia Naturale di Milano, Milan, Italy (M. Pavesi, C. Leo-
nard!).
MCZC: Department of Entomology, Museum of Comparative Zoology, Har-
vard University, Cambridge, Massachusetts (P. Perkins).
MHNP: Entomologie, Museum National d’Histoire Naturelle, Paris, France (T.
Deuve).
MIZA: Museo del Instituto de Zoologia Agricola Francisco Fernandez Yepez,
Maracay, Venezuela (L. Joly, J. Clavio).
MNHB: Zoologisches Museum, Museum fiir Naturkunde der Humboldt-Univ-
ersitat zu Berlin, Berlin, Germany.
NMW: Zweite Zoologische Abteilung, Naturhistorisches Museum Wien, Wien,
Austria (H. Schonmann, E. Kirschenhofer).
OSUC: Department of Entomology, Ohio State University, Columbus, Ohio
(N. Johnson).
QCAZ: Catholic Zoology Museum, Pontificia Universidad Catolica del Ecua-
dor, Quito, Ecuador (G. Onore).
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RSCI: Riccardo Sciaky Collection, Milan, Italy.
SEMC: Snow Entomological Division, The Natural History Museum of the
University of Kansas, Lawrence, Kansas (R. Brooks, J. S. Ashe).
STOC: H. Stockwell Collection.
UASM: Strickland Museum, Dept, of Biological Sciences, University of AL
berta, Edmonton, Alberta, Canada (D. Shpeley, G. E. Ball).
UATU: Department of Entomology, University of Arizona, Tucson, Arizona
(C. A. Olson, D. R. Maddison).
UMMZ: University of Michigan, Museum of Zoology, Ann Arbor, Michigan
(M. O’Brien).
UVGC: Universidad del Valle de Guatemala, Guatemala City (J. Schuster).
USNM: Department of Entomology, United States National Museum of Natural
History, Smithsonian Institution (T. L. Erwin, D. Furth)
Taxonomic Methods
Species Recognition. — Species were recognized by a set of defining character-
istics that permit grouping like individuals from other such individuals with a
different set of characteristics. This is essentially the species definition of Nixon
and Wheeler (1990).
When multiple specimens were available, variation in characteristics for the
species could be assessed and the limits of the defining characters understood.
However, when only a single, notably different specimen is available it was nec-
essary to decide if the difference was of specific value. For species based on one
or very few specimens, I relied on characters that seemed to be beyond individual
variation observed in species known from larger samples. In general these unique
specimens were not dissected, so external characteristics that were found to be
correlated with genitalic differences in better represented species were relied on.
In a few cases a specimen was included within my interpretation of a species,
though not in the type series, because it exhibited slight differences that may
ultimately prove to define it as a member of a distinct species. I felt it was best
not to multiply names in these marginal cases until more material is collected,
permitting a robust test of stability of species definitions based on observed char-
acter differences.
Species of these genera, like nearly all of life, do not have any single charac-
teristic or complex of characteristics that we can examine to inform us with cer-
tainty on their reproductive isolation, or any other possible mode of speciation.
Entities named here may be explained, after they are recognized, as real biological
and evolutionary individuals. But regardless whether the process thought to un-
derlie species generation is correct or not, species recognition remains the same
as it has for all of recorded history, based on empirical observation of character-
istics augmented by knowledge of reproduction and development in the rare cases
where those data are known. Of course, hypotheses may be adjusted, if necessary,
in light of observations of reproductive isolation or interbreeding.
Ranking Criteria and Classification. — No general standard exists for recogniz-
ing supraspecific groups in pterostichine ground beetles. The repeated failure to
support a monophyletic Pterostichini (auctorum) demonstrates that this taxon is
a grade (Will, 2000). My use of the term pterostichine grade emphasizes the lack
of monophyly among the presently included taxa.
The Holarctic pterostichine grade fauna, primarily Pterostichus Bonelli, 1810,
2002
Will — Revision of Abaris
147
is superficially rather uniform in appearance and some recent authors have sep-
arated supraspecific groups by very small character differences. The application
of similar criteria to the South American and Australian taxa has resulted in what
I feel is an extreme emphasis on differences with less regard for grouping simi-
larities. This has led to the recognition of too many supraspecific taxa. With this
in mind I have attempted to minimize the number of formal names (genera and
subgeoera) proposed here, but have named clades (species-groups) that, based on
the characters, seem to represent biologically interesting groups and groups of
similar form. These names are provided for biologists who may need to discuss
the ecology, behavior, etc., of a monophyletic group. When the higher elements
of the pterostichine grade are established, some clades (genus-groups) may come
to have a formal position in the Linnaean hierarchy.
It is neither practical nor desirable to have a name affixed to every branching
point of the cladogram. The classification here is completely consistent with the
phylogenetic hypothesis but is not identical to it. These two constructs, cladogram
and classification, have slightly different but interconnected purposes. The former
provides a complete summary of characters and relationships, and is the basis for
discussion of hypotheses of common descent. The latter is the primary means of
information retrieval for the taxa, and so should be a useful mnemonic device. In
order to fulfill this function, the classification necessarily ignores some of the
details of the cladogram and focuses on the aspects deemed likely to be biolog-
ically important.
I use the term sedis mutabilis as proposed by Wiley (1979; 1981) for the
polytomous relationship of basal Abaridius species.
Phylogeny Reconstruction.— matrix was constructed with the computer
program WinClada (Nixon, 1999) and submitted to NONA (Goloboff, 1994) for
analysis. NONA’S default settings were used for a heuristic search for most par-
simonious trees (see NONA documentation). The search was done by submitting
the matrix via WinClada using number of replications = 100 (Mult* 100), starting
trees per replication = 15 (hold/15), random seed = Time (rsO) and the search
strategy was multiple TBR+TBR (mult* 100; Max*). Aspects of cladistic methods
and subsequent interpretation of the pattern as a phylogenetic hypothesis em-
ployed here are discussed in my analysis of the genus Lophoglossus LeConte,
1852 (Will, 1999).
Preparation and Imaging Techniques. — External structures were examined us-
ing a dissecting stereo-microscope at magnification of lOOX or less. Some smaller
structures and microsculpture of legs, mouthparts, and elytra were examined and
photographed using a Hitachi 4500 scanning electron microscope. Disarticulated
cuticular structures, including all external parts of the body and various internal
systems, were cleared in 10% KOH and mounted on glycerine slides, then ex-
amined using a phase-contrast compound microscope.
Methods generally used for preparation of male genitalia follow Allen (1972).
However, the small size of most species did not allow for the successful eversion of
the endophallus using Allen’s methods. Two alternatives were tried with some success
for each. First fluid pressure was applied to the foramen using the Phalloblaster
machine (vesica everter, Matthews, 1998) for Neotalus portai (Straneo). The small
size of Abaris specimens precluded the use of this machine as the needle diameter
exceeds the diameter of the median lobe. Instead of direct fluid pressure, a rapid
change in osmotic pressure was used to evert the endophallus in some species. The
median lobe was softened in KOH and then dehydrated in 95% EtOH. In order to
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increase pressure pushing out the endophallus, a small drop of water soluble glue
was placed on the foramen to prevent equalization through the foramen. When the
dehydrated median lobe was placed in distilled water, pressure from the flow of water
into the lobe via the gonopore forced out the endophallus.
Preparation of female reproductive tracts and genitalia followed procedures
outlined by Liebherr and Will (1998).
Initially, drawings were made as pencil line-drawings using an ocular grid or
camera lucida. These line drawings were digitized using a flatbed scanner, “inked”
and shaded on a personal computer using the Corel Photo-Paint® version 7.467.
Some cases digital images of whole or slide-mounted portions of specimens were
captured using a video microscope and a Snappy Video grabber version 3.0 con-
nected to a personal computer. These images were enhanced in Corel Photo-Paint®
version 7.467 and either printed from a desktop printer or black and white print film.
Locality Information. — Locality information is provided for each species and
is divided between type and non-type information. Data from type specimens that
are verbatim from the labels are enclosed in double quotation marks with the
description of the kind of label and a note if handwritten. Notes not on the spec-
imen labels and the description of the labels are added for clarity and this infor-
mation placed between square brackets ( [ . . . ] ). Unless otherwise noted, labels
are white, rectangular with black ink. In as much detail as could be clearly con-
firmed locality data for other material examined is listed for each species by
country in descending order from largest political unit to smallest site information,
including latitude and longitude. Altitude and seasonal information are summa-
rized in the section “Notes on Life History” and are listed under each species if
any additional data were available beyond that found on the type specimens.
Maps show the location of collections for each of the species either from spec-
imen data labels or from published records. A symbol on the map primarily
represents a single collecting site. Multiple sites are represented by a single sym-
bol in cases where collecting sites are very close and multiple symbols on the
map would be impossible to discern or be confusing. If label data were unclear
or the locality not definitively identifiable, the data are listed but not plotted on
the map. In cases where this was the only record for a country a symbol was
place on the map with a “?” to identify it as a questionable record.
Terms
Measurements. — Overall length was measured using an ocular reticle and gen-
erally was the entire distance from the base of the labrum to the tip of the left
elytron. In specimens where it was not possible to measure this way accurately
because of the head position, length was standardized using the sum of lengths:
1. base of labrum to the cervical collar, 2. apex to base of pronotum along the
midline, 3. base of scutellum to apex of left elytron.
The ratio of elytral interval widths is the width of interval 3 divided by the
width of interval 2. Measurements were taken by placing an ocular reticle across
the intervals at the level of the dorsal setigerous puncture.
Ocular ratio is a measurement of the relative size of the eyes or “bug eyed-
ness” of the beetles. This ratio is the width over the widest point of the eyes
divided by the width between the eyes at the level of the anterior supraorbital
setae.
Body Parts. — Terms for adult external structures and segmentation follow
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Will — Revision of Abaris
149
Lawrence and Britton (1994). One exception is the use of sterna II-VII for nor-
mally exposed ventral abdominal segments rather than stereites II-VII as used by
Lawrence and Britton. This follows the definition of sternum as a single ventral
division per segment, as is found in adult beetles, and sternites as subdivisions
of a sternum as is found in many larvae (Snodgrass, 1935; Torre-Bueno, 1937).
Terms for the female reproductive tract and external genital structures follow
Liebherr and Will (1998). Nomenclature of endophallic structures follows Noonan
(1991).
The median lobe of the aedeagus is discussed using the following: basal bulb,
generally expanded region of the foramen from the base to the parameres; blade,
remaining section from the basal bulb to the tip; apex, apical portion of blade
from the level of the ostium to the tip; and tip, portion of the apex of the median
lobe beyond the ostium. Terms used for colors follow Torre-Bueno (1937).
The term scutellar stria has been used to refer to the short striae on the elytra
near the scutellum even though they may not be homologous structures across all
groups. The term has been used in reference to the parascutellar striae, which are
directly adjacent to the scutellum and continuous with the basal marginal border
of the elytra when the border is evident and striae are sulciform. Additionally,
the term has been used for the short basal section of stria 1 when that stria is
interrupted, even though this section is not adjacent to the scutellum. In order to
be consistent with my hypotheses of primary homology I abandon the term scu-
tellar stria but retain the use of parascutellar stria as restricted above. In all
abariform taxa elytral stria 1 is continuous with the parascutellar stria. Stria 1
may end at its junction with the parascutellar stria or have a short, disconnected
basal section arising from near the basal setigerous puncture. When stria 1 is
interrupted and the disconnected basal section is present, it is referred to as the
basal section of stria 1.
Discussion of Clades and Taxa. — Cladistic diagnoses are summaries of syna-
pomorphic characteristics of a monophyletic group of taxa based on the cladogram
present herein. The purpose of this section is to allow for quick assessment of
group membership. The character states of a clade that apply to all included taxa,
are not discussed in subordinate clades as they are plesiomorphic and are not
useful for assessing group membership at that level.
Recognitory diagnoses correspond to the more traditional diagnosis found in
m.ost taxonomic works. The characteristics provided under this heading are in-
tended to permit accurate recognition of specimens. This includes any general
attributes and comparative aspects, without regard to phylogenetic implications
of the characteristic.
Taxonomic and Cladistic Accounts
Systematic Entomology
Order Coleoptera
Family Carabidae
Subfamily Harpalinae
Tribe Pterostichini (Auctorum)
Key to Adult Neotalus new genus and Abaris Dejean species
1 . Metacoxal anterior sulcus incomplete, ended medially on coxa, broadly arcuate in most
species, not approaching anterior margin (Fig. 3A); basal section of stria 1 present or
absent ........................................................ 2
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Fig. 2. — Structures of abariform species. A. Scanning electron micrograph, pro tarsal claw of Abaris
aenea Dejean showing pectination. B. Digitized video-capture image, ventral view, pro and mesotho-
racic area. Neotalus portal (Straneo), mesepimeral tubercle indicated by arrow.
1'. Metacoxal anterior sulcus complete, ended at or near apex, straight, not broadly arcuate,
approaching anterior coxal margin, (Fig. 3B); basal section of stria 1 absent. Abaris
tachypoides-group 21
2 (1). Basal section of stria 1 absent 3
2'. Basal section of stria 1 present 8
3 (2). Sterna V-VII without sulci (Fig. 4C); tarsal claws pectinate (Fig. 2A) 4
3'. Sterna V-VII with sulci (Fig. 4A); tarsal claws smooth Neotalus portal (Straneo)
4 (3). Pronotum widest just before middle, lateral margin reflexed in basal third or not; most
species shinier with faint or obsolete microsculpture; if microsculpture obvious, then
pronotum in basal third laterally distinctly reflexed (Fig. 24D-E) 5
4'. Pronotum widest slightly behind middle, lateral margin not reflexed in basal third (Fig.
25D); dull species, microlines forming reticulate microsculpture obvious. Argentina
[12] Abaris basistriata Chaudoir (in part)
5 (4). Microsculpture faint but visible on head and pronotum 6
5'. Microsculpture obsolete on head and pronotum [4] Abaris aenea Dejean
6 (5). Elytral interval 3 much broader than 2; width ratio 3/2 >1.4 7
6'. Elytral interval 3 only slightly wider than 2; width ratio 3/2 <1.4
[5] Abaris erwini new species
7 (6). Pronotum punctate basally and laterally, base laterally sinuate with median base dis-
tinctly extended posterad of hind angles (Pig. 24H) South America
[7] Abaris notiophiloides Bates
7'. Pronotum impunctate laterally, base in some with a few shallow punctulae, base not
Fig. 3. — Line drawing illustrating diagrammatically abariform left metacoxae. A. With incomplete/
arcuate metacoxal anterior sulcus (ms). B. With straight/appressed metacoxal anterior sulcus (ms).
2002
Will — Revision of Abaris
151
Fig. 4. — Line drawing illustrating diagrammatically abariform abdominal sterna. A. With complete
transverse sulci. B. Incomplete/laterally present sulci. C. Sulci absent.
sinuate; hind angles only slightly anterior to median base (Fig. 24G) Central America/
Mexico ................................ [6] Abaris aquilonaria new species
8 (2'). Sterna V-VII with lateral sulci, medially complete or interrupted (Fig. 4A-B); sulci
shallowly to deeply impressed, best developed on sternum VI (variable species handled
in both halves of couplet) .......................................... 9
8'. Sterna V-VII without a trace of lateral or medial sulci (Fig. 4C) .............. 19
9 (8). Sulci on sterna V-VII complete (Fig. 4A) 10
9'. Sulci on sterna V-VII broadly interrupted medially and slightly developed laterally
(Fig. 4B) ..................................................... 12
10 (9). Pronotum constricted at base, widest in anterior half; both basal foveae distinct ... 11
10'. Pronotum widest at base; outer basal foveae absent (Fig. 25E) .............
[13] Abaris metalUca new species
1 1 (10). Dorsum of head and pronotum metallic green, elytra cupreous; lateral pronotal margins
sinuate to base, hind angles clearly denticulate (Fig. 25F) .................
......................................... [14] Abaris bicolor new species
11'. Entire dorsum cupreous; lateral pronotal margins convexly arcuate to base, hind angles
minutely denticulate (Fig. 25G) .................. [15] Abaris nobilis new species
12 (9'). Disc of pronotum shiny, without obvious microsculpture .................... 13
12'. Disc of pronotum dull, with obvious mesh microsculpture either reticulate or somewhat
transversely elongate ............................................. 18
13 (12). Pronotal lateral margin sinuate anterior of hind angles, reflexed in basal third (Fig.
24A-B); narrowly convex immediately lateral of basal fovea ................ 14
13'. Pronotal margin laterally arcuate and straight or subsinuate anterior of hind angles, not
reflexed laterally; convex from basal fovea to lateral margin. (Fig. 25B-C) ....... 17
14 (13). Pronotum with outer basal foveae present, punctate or rugose along base and/or punc-
tulate in foveae (Fig. 24A-B) ....................................... 16
14'. Pronotum with outer foveae absent, impunctate (Fig. 24C) .................. 15
15 (14). Mediae lobe tip asymmetrical in dorsal view, left edge emarginate (Fig. 21E) ......
.......................................... Abaris striolata Bates (in part).
15'. Median lobe tip nearly symmetrical in dorsal view, left edge convex (Fig. 21G) ....
....................................... [3] Abaris robustula Tschitscherine
16 (14). Pronotal base distinctly rugose medially and around basal foveae; aedeagus median
lobe tip very broad in dorsal view (Fig. 21A-C) .... [1] Abaris napoensis new species
16'. Pronotal base impunctate or faintly punctured medially and around basal foveae; ae-
deagus median lobe tip acuminate in dorsal view (Fig. 21D-E) .............
....................................... [2] Abaris striolata Bates (in part)
17 (13'). Pronotal base impunctate (Fig. 25C); endophallus of aedeagus without spine patch
visible at bend of mediae lobe (Fig. 23D); spermathecal duct tightly twisted .......
........................................... [1 1] Abaris mina new species
17'. Pronotal base punctate (Fig. 25B); endophallus of aedeagus with spine patch visible at
bend of median lobe (Fig. 23 A); spermathecal duct broad, not twisted (Fig. 19E) . . .
.............................................. [10] Abaris picipes Bates
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18 (12').
18'.
19 (8').
19'.
20 (19).
20'.
21 (1').
21'.
22 (21).
22'.
23 (22').
23'.
24 (23).
24'.
25 (21').
25'.
26 (25).
26'.
27 (25').
27'.
28 (27).
28'.
29 (27').
29'.
30 (29).
30'.
31 (30').
31'.
Pronotal base with coarse punctulae in basal foveae (Fig. 25A); dorsum black; obvious
reticulate microsculpture on pronotum, transversely elongate mesh on elytra, elytra
shiny. Mexico . • • • [9] Abaris bigenera Bates
Pronotal base without coarse punctulae (Fig. 241); dorsum aeneous; reticulate micro-
sculpture obvious throughout, dull species. South America ................
[8] Abaris impunctata new species
Pronotal lateral margins not reflexed, arcuate, straight just anterior to slightly obtuse
hind angles (Fig. 25B-D) 20
Pronotal lateral margins distinctly reflexed basally and subsinuate to right angled hind
angles (Fig. 24B) [2] Abaris striolata Bates (in part)
Reticulate microsculpture obvious on head and pronotum; dorsal surface duller
[12] Abaris basistriata Chaudoir
Microsculpture obsolete on head and pronotum; dorsal surface very shiny .....
[10] Abaris picipes Bates
Sterna V-VII with lateral sulci, medially complete or interrupted (Fig. 4A-B) .... 22
Sterna V-VIl without sulci (Fig. 4C) 25
Pronotal base without punctulae; if punctulae present then outer basal foveae clearly
round (Fig. 26A) [17] Abaris franiai new species
Pronotal base with coarse punctulae; outer basal foveae elongate 23
Apex of pronotum wider, tip of front angles distant from occiput and slightly produced
(Fig. 25H, 26B) . 24
Apex of pronotum narrow, tightly rounded and appressed to occiput, front angles not
produced (Fig. 9) [19] Abaris tachypoides Bates
Pronotum medially punctate between inner foveae (Fig. 25H); mental tooth broadly
flattened at apex [16] Abaris nitida new species
Pronotum medially impunctate between inner foveae (Fig. 26B); mental tooth rounded
at apex [18] Abaris inflata new species
Pronotum relatively small, lateral margins sinuate and narrowly constricted basally (Fig.
26D-E); width of pronotum at base less than width across eyes, or if approximately
equal then legs bicolored 25
Pronotum relatively larger, lateral margins sinuate or not; width of base greater than or
equal to width across eyes, legs not distinctly bicolored 26
Coxae, trochanters and base of femora darkly infuscated, contrasted with flavous apices
of femora, tibiae and tarsi; dorsum black; pronotal base punctulate; front angles of
pronotum slightly produced (Fig. 26D) [21] Abaris nigra new species
Legs concolorous, flavous; dorsum bronzed; pronotal base smooth, only foveae im-
pressed; front angles of pronotum round and tightly appressed to occiput (Fig. 26E)
[22] Abaris inaequaloides new species
Pronotal base with irregular, coarse punctulae at least in area of basal fovea; if punctulae
restricted to fovea then head dull from reticulate microsculpture 28
Pronotal base without punctulae or with only a few shallow punctulae in basal fovea;
if punctulae present then head quite shiny, micro-lines scarcely visible .......... 29
Mental tooth form simple, sagittiform [25] Abaris wardi new species
Mental tooth apex blunt and slightly emarginate ...... [23] Abaris opaca new species
Elytral intervals convex, more prominently so in apical third; eastern and southern
Mexico, Central or South America 30
Elytral intervals flat throughout their length; northwestern Mexico, Baja California,
Arizona, southern California [24] Abaris splendidula (LeConte)
Dorsum uniformly shinier, though not brilliantly so, microsculpture of pronotal disc
transverse, somewhat irregular mesh, transverse mesh on elytra. Central America, Mex-
ico . [26] Abaris aequinoctialis Chaudoir
Pronotal disc very dull from reticulate microsculpture; elytral microsculpture reticulate,
shinier than pronotal disc. South America . 31
Pronotal base not bordered laterally (Fig. 26C) [20] Abaris convexa new species
Pronotal base with distinct lateral border (Fig. 27C) .... \21\Abaris retiaria new species
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Will — Revision of Abaris
153
Classification and Checklist of Abariform Genera, Subgenera and Species
(Pseudabarys {Neotalus 5- Abaris))
Neotalus Will, new genus (one species)
Neotalus portal (Straneo)
Abaris Dejeae (27 species)
Abaris sensu stricto
[A. striolata-gmwpl
1. A. napoensis Will, new species
2. A. striolata Bates
[A. aenea-gmwpl
3. A. robustula Tschitscherine
4. A. aenea Dejean
5. A. erwini Will, new species
6. A. aquilonaria Will, new species
7. A. notiophiloides Bates
[A. picipes-gmup\
8. A. impunctata Will, new species
9. A. bigenera Bates
10. A. picipes Bates
IL A. mina Will, new species
12. A. basistriata Chaudoir
13. A. metallica Will, new species
Abaridius Chaudoir
{sedis mutabilis}
14. A. bicolor Will, new species
15. A. nobilis Will, new species
[A. tachypoides-gmup]
16. A. nitida Will, new species
17. A. franiai Will, new species
18. A, inflata Will, new species
19. A. tachypoides Bates
20. A. convexa Will, new species
21. A. nigra Will, new species
22. A. inaequaloides Will, new species
23. A. opaca Will, new species
24. A. splendidula (LeConte)
25. A. wardi Will, new species
26. A. aequinoctialis Chaudoir
27. A. retiaria Will, new species
Accounts of Clades
Abariform Clade (Pseudabarys (Neotalus + Abaris jj
Cladistic Diagnosis. — All members of the abariform clade share the following
non-homoplasious synapomorphies (Fig. 31: characters 2, 8, 12, 26): eyes large,
ocular ratio greater than 1.60; humeral umbilicate puncture 2 displaced medially
from 1 and 3; ventral setae of tarsomere 5 of unequal length and paired; dorsal
bursal glands present (Fig. 5A-B).
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Fig. 5. — Abaris aenea Dejean digitized video-capture images of female reproductive tract, showing
paired dorsal glands. A. Ventral view of bursa laid flat, arrow indicates gland. B. Single gland enlarged.
Clade fNeotalus + Abaris j
Cladistic Diagnosis. — All members of this clade share the following synapo-
morphies (Fig. 31: characters 1, 25): maxillary palpomere 3 much smaller than
palpomere 4 (Fig. 28B), diverticulum of the spermatheca appended gland absent.
Neotalus, new genus
Type Species. — Bothynoproctus portai Straneo; here designated.
Etymology of Generic Name. — Noun in apposition, masculine. Neotalus is a genus-group name
formed from the Greek neos (veoa) — new and Talos (TaXoa) — the name of the bronze giant faced by
the Argonauts in Crete. The single New World species placed in this genus is larger than species in
the genus Abaris, has a distinctive metallic luster and lacks the pectinate claws found in Abaris species.
Nomenclatural and Taxonomic Notes. — Straneo (1941) described Bothyno-
proctus portai placing it with B. mattoensis Tschitscherine, 1900, the type species
for the genus. No doubt as a result of World War II, Stefano Straneo did not have
the benefit of seeing the type of B. mattoensis, and therefore he misplaced his
new species. The characteristic deep punctures of the submentum and sternum
VII in B. mattoensis are lacking in N. portai. The former is not closely related
to the abariform genera and is more closely related to Dyschromus Chaudoir and
Tichonilla Strand, 1942 (Will, 2000), whereas the latter has several significant
synapomorphies in common with Abaris species.
Based on these characteristics and my analysis of a broader set of characters
(Will, 2000), Straneo’s species is removed from Bothynoproctus and a new genus,
Neotalus, is proposed to hold this species. Therefore, Neotalus portai (Straneo)
is the type species for this monotypic genus.
Cladistic Diagnosis. — As this is a monotypic genus, its synapomorphic char-
acters cannot be specified. If a second species is found that groups with N. portai
it will likely share some of the apomorphic characteristics described for the spe-
cies below.
Recognitory Diagnosis of Adults. — See species below and characters in the key.
Description of Adults. — None required because the genus is monotypic. Its characters are as in the
type species described below.
Distribution. — See species account below and figure 10.
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Fig. 6. — Dorsal habitus of Neotalus portal (Straneo).
Species Account
Neotalus portal (Straneo, 1941), new combination
Fig. 2B, 6, 10
Bothynoproctus portal Straneo, 1941:28.
Type Material. — HOLOTYPE. Female. Labeled: “Foret Vierge, au bord de Riv. Paraguay, S An-
tonio” [handwritten]/“Holotypus portai” [red label]/“portai Stran.” [handwritten, red ink], [MSNM].
ALLOTYPE. Male, labeled: “S. Antonio” [handwritten], “Allotypus” [red label]/[dissected genitalia
and mouthparts glued to card] [MSNM], PARATYPES. 2 Males, same data as Allotype, both labeled
with “M.C.Z. Paratype 28422”, MCZC.
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Type Locality. — Paraguay, S Antonio, Staz. Entomol. Fabre., from original de-
scription.
Range. — Bolivia, Brazil, Paraguay (Fig. 10)
Recognitory Diagnosis of Adults. — Recognized from all other Carabidae by the
combination of form of mouthparts — labial palpi short and broad, lacinia large
with a thick, curved apical digitus, maxillary palpi broadly fusiform — and me-
sepi sterna with large tubercle (Fig. 2B) and a corresponding protuberance of pro-
notal base, tarsal claws smooth, and female reproductive tract with paired bursal
glands (Fig. 5A-B).
Description. — In addition to characteristics of the Neotalus + Abaris clade, large sized, 7. 0-8. 5
mm. Elytra aeneous or cupreous to vinous, head and pronotum darker, shiny. Ventral surface black,
tarsi and palpi paler brunneous.
Head. Large, very broad, frontal impressions deep, divergent. Fronto-clypeal suture impressed, clyp-
eus emarginate, membrane at base of labrum exposed. Eyes very large and prominent, hemispherical,
ocular ratio 1.62-1.72. Submentum separated from mentum by distinct suture. Mentum very broad,
with epilobes slightly produced beyond broad, apically emarginate medial tooth. Ligula with glossal
sclerite broad and convexly rounded at apex with two large setae. Paraglossae very long and free with
cristate microspicules, especially laterally and near apex. Labial palpi short and broad, palpomere 2
with two large medial setae and 2-3 small apical setae, palpomere 3 with small scattered setae.
Maxillae with lacinia large and with a thick, curved apical digitus plus a medial field of large spines
with scattered finer setae. Cardo distinctly developed, stipes with two large setae, one apical one basal.
Palpifer simple, maxillary palpi fusiform but broad; palpomere 1 broad with a few small setae at apex,
palpomere 3 very short relative to palpomere 4 and with several small setae at its apex, palpomere 4
expanded at tip with scattered small setae, 2-3 lateral setae larger. Galea with two small terminal
setae. Mandibles with retinaculum discrete, premolar and molar teeth small.
Thorax. Ventral thoracic segments impunctate and moderately shiny, microsculpture faint mesh.
Pronotum broad, lateral margins straight before hind angles, apical margin straight, front angles very
slightly produced, basal margin straight, inner basal foveae present, broadly impressed, outer basal
foveae shallow round depressions. Prosternal process apically margined. Metepisternum quadrate.
Flight wings small scales. Elytra broad, depressed, oblong-ovate, humeri angulate, equal to width of
pronotal base, striae deep, smooth (Fig. 6); stria 1 continuous with parascutellar stria, basal section of
stria 1 absent, elytral intervals slightly convex; each elytron with a single setigerous dorsal puncture
in apical half of third interval; second and third intervals of approximately equal width. Legs of
moderate length; metacoxal anterior sulcus divergent, and arcuately ended medially in coxa, coxae
each with two large setae; pro and meso-trochanters each with a single large seta, metatrochanter
apically rounded without setae; femora slender, setation as follows; profemora — anterior face, 1 apical,
1 medial; dorsal face, 1 apical and 2 subapical; posterior face, 1 apical, 1 medial and 1 basal; meso-
femora — anterior face, 1 apical, 1 medial and 1 basal; dorsal face with row of 4 setae, 2-4 apical;
posterior face, 0-2 setae and occasional scattered setae laterally; metafemora — anterior face, 1 apical,
1 medial; dorsal face with row of 2-4 setae; posterior face glabrous. Tibiae slender; protibia with a
row of 3-4 fine setae on medial face and a row of 3-5 coarser setae laterally, two clip setae, anterior
brush of fine setae ends at enlarged medial seta; meso- and metatibiae with four rows of prominent
setae, medial row very fine, others much stouter. Male protarsomeres expanded, first to third with
articulo-setae ventrally (Stork, 1980). Four basal protarsomeres of female and four basal meso- and
metatarsomeres in both sexes with two, more or less distinctly defined rows of ventral setae and
numerous scattered setae, most scattered setae smaller than the row setae. Fifth tarsomere with two
pairs of fine ventral setae, apical nearly twice length of subapical. Tarsal claws smooth.
Abdomen. Sterna moderately shiny, microsculpture obscure. Transverse sulcus of sterna V-VII
present and entire. Male genitalia simple, endophallus minutely spinose and scrotiform. Median lobe
of aedeagus straight, smooth. Female reproductive tract with spermatheca not discrete from duct, duct
broad. Gonocoxite-1 with 3 small ensiform setae near apex, gonocoxite-2 arcuate and laterally exca-
vated, apex round, 1 dorsal and 1 lateral large ensiform seta, 2 nematiform setae in elongate furrow.
Paired bursal glands dorsally present.
Additional character information is provided in original description.
Notes on Life History. — The only information beyond data given for the types
is on a specimen from Sara, Bolivia, collected at 450 m in November.
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Material Examined. — Twelve specimens, including the type series, were examined. Nontype ma-
terial from: BOLIVIA. La Paz^ Sud. Yungas. Ocbaya [MSNM], Sara: [CMNH, MNHB].
BRAZIL. Chapada [CMNH],
PARAGUAY. Central: [MHNP].
UNKNOWN: [MSNM].
Genus Abaris Dejean, 1831
Abaris Dejean, 1831:780. TYPE SPECIES (by monotypy): Abaris aenea Dejean, 1831:781. Dejean
and Boisduval, 1832:211. Laporte, 1840:118. Chaudoir, 1852:76-77. Chenu, 1851:161. Lacor-
daire, 1854:347. Bates, 1871:218-220. Bates, 1882:85. Csiki, 1930:576. Straneo, 1939:38. Black-
welder, 1944:35. Reichardt, 1977:409. Straneo, 1977:111,115. Straneo, 1979:350,352. Bousquet,
1984:384-389. Bousquet and Larochelle, 1993:165. Bousquet and Liebherr, 1994:435-441. Lo-
renz, 1998:243. Bousquet, 1999:48-51. Ball and Bousquet, 2001:85.
Abarys Agassiz, 1846:1 (unjustified emendation). Gemminger and Harold, 1868:366. Chaudoir, 1873:
96. Tschitscherine, 1898:83.
Abaridius Chaudoir, 1873:97. Reichardt, 1977:409. Erwin, 1991:40. New Synonymy: Type species
(original designation): Abaris tachypoides Bates, 1871:220.
Etymology. — Dejean constructed the generic name from the Greek using a, absence of, and (Bapua,
heavy, in reference to the lightly constructed form and fleetness of foot for the included species.
Dejean transcribed jSapw with u = i. Most Greek lexicons transcribe n = y, which led to the unjustified
emendation of Abaris by Agassiz (1846) to Abarys. Indeed, a strict transcription of Abaris back to
the Greek would mean “not boat shaped,” whereas abarys would translate to “not heavy” as described
by Dejean. Nonetheless, the ICZN (1999, Art. 33.2.3) does not allow emendation of a name based on
changes in character transcription.
Abarys (apapw) is a masculine word in Greek, but Abaris as used by Dejean is a feminine form.
He did this in conjunction with the single species he named aenea, which is clearly feminine. There-
fore, various specific epithets that have been changed to, or originally formed in the masculine must
be made to agree in gender with Abaris as a feminine Latinized noun.
Synonymy. — Bates (1871) described five Abaris species. These plus A. aequin-
octialis described by Chaudoir (1852) increased the total number of Abaris species
to seven. Bates does not mention the presence of pectinate tarsal claws in his
discussion of the characteristics of the genus, but does note the transverse sulci
on the sterna of some species. Chaudoir (1873) subsequently newly defined two
genera, Abaridius and Pseudabarys that included two of Bates’ species. A, tach-
ypoides and A. robustus, respectively. The three genera were diagnosed as follows:
Abaris species with pectinate claws and smooth sterna, Abaridius species with
both pectinate claws and ventral sulci, and Pseudabarys with smooth claws and
a sulcate venter. My study of these characteristics and additional characters shows
that Abaris and Abaridius are congeneric. Abaris is distinct from Pseudabarys,
but the latter is in need of revision and its monophyly has not been tested.
Cladistic Diagnosis. — Abaris is considered a monophyletic group within the
larger abariform clade and is defined by the synapomorphous presence of pectinate
claws (Fig. 2, Fig. 31: character 11).
Recognitory Diagnosis. — Recognized from all other Carabidae by the combi-
nation of pectinate claws, distinctly developed internal and external elytral plica
(sensu Liebherr, 1986:90-91) and paired bursal glands in the female reproductive
tract (Fig. 5).
Description of Adults. — In addition to characteristics of the Neotalus + Abaris clade, small to
medium sized ground beetles, 4. 8-8. 9 mm. Nearly all individuals with an aeneous or virescent luster,
sometimes brilliantly so (only A. nigra and some individuals of A. bigenera black without metallic
luster). Nearly all individuals shiny or very shiny. Body ventrally piceous, rufous or ferruginous, legs
and mouthparts paler than venter in most, some individuals variously infuscated.
Head. Head large, relatively broad. Frontal impressions present, deep in most species, shallowest
in members of the A. picipes-gvowp. Eyes very large and prominent, hemispherical, ocular ratio 1.63-
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VOL. 71
E
E
if>
o
A
C
1.0 mm
Fig. 7. — Structures of Abaris splendidula (LeConte). A. Pygidial gland system, ventral view of right
gland. B. Extracted gut. C. Right hind wing, dorsal view. Legend: aal+2, anterior anal vein 1+2;
aa3+4, anterior anal vein 3+4; cc, collecting canal; crp, crop; ct, crypt ; e, esophagus; eff, efferent
duct; FG, foregut; HG, hindgut; MG, midgut; mt, Malpigian tubles; mp3, posterior medial vein 3;
mp4, posterior medial vein 4; o, oblongum cell; pv, proventriculus; ra4, anterior radial vein 4; re,
rectum; rp2, posterior radial vein 2; rp3+4, posterior radial vein 3+4; rs, gland reservoir; rtp, rectal
pad; sc, secretory cells; tVIII, tergite VIII.
1 ,9 1 , most greater than 1 .70, eyes least developed in A. nobilis. Submentum separated from mentum
by well-marked suture. Mentum broad, with epilobes produced anterad medial tooth. Mentum tooth
may be entire or emarginate at the apex, broad or slightly acuminate in form but not sharply pointed.
Ligula with glossal sclerite broad at apex, with 2 large apical setae. Paraglossae long and free with
cristate microspicules, especially near apex and medially. Labial palpi fusiform, palpomere 2 with 2
large medial setae and 2-3 small apical setae, palpomere 3 with small scattered setae. Maxillae with
lacinia large and with thick, curved apical digitus and medial field of large spines. Cardo distinctly
developed, stipes with 2 large setae, 1 apical 1 basal. Palpifer simple, maxillary palpi fusiform. Pal-
pomere 1 broad with few small setae at apex, palpomere 3 very short relative to palpomere 4 and
with several small setae at its apex, palpomere 4 with scattered small setae, 2 lateral setae larger in
most individuals. Galea with 2 small terminal setae. Mandibles with retinaculum discrete, premolar
and molar teeth small.
Thorax. Ventral thoracic segments impunctate except for a few punctures on mesepisternum in some
individuals, shiny, microsculpture obsolete or faint, irregular mesh, or slightly transverse. Pronotum
variously formed, lateral margins sinuate or subsinuate in most species, apical margin straight or front
angles slightly produced, basal margin straight or bi-sinuate, inner basal foveae present, sharply im-
pressed in most species, outer basal foveae quite variable among species and somewhat variable among
individuals. Prosternal process apically margined. Metepisternum longer than wide. Flight wings full
(Fig. 7C) in all individuals investigated. Elytra oblong-ovate, free at suture, broad across humerus and
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Will — Revision of Abaris
159
humeri angulate in most species, slightly wider than pronotum; striae deep, smooth; stria 1 continuous
with parascutellar stria; intervals convex to flat; third interval broader than other intervals in some
species, notably in species of the A. aenea-group; each elytron with a single setigerous dorsal puncture
in third interval behind middle, in contact with second stria in most individual. Legs slender and of
moderate length. Metacoxal anterior sulcus appressed along anterior margin (Fig. 3B), or divergent,
and arcuate/sinuate ended medially in coxa (Fig. 3A), coxae with two setae. Pro- and mesotrochanters
with a single large seta, metatrochanter apically rounded or slightly pointed, without setae. Femora
slender, setation various between species and individuals but in most individuals as follows: profe-
mora— anterior face, 1 apical, 1 medial; dorsal face, 1 apical, 1 subapical; posterior face, 1 apical, 1
medial and 1 basal, presence of basal seta quite variable; mesofemora — anterior face, 1 apical, 1 basal
and 2 medial; dorsal face with row of 8-16 setae, 2-4 setae apical; posterior face, 0-2 setae; meta-
femora— anterior face, 1 apical,! medial; dorsal face with row of 3-4 setae; posterior face glabrous.
Tibiae slender; protibia with row of 3-4 fine setae on medial face and row of 3-5 stouter setae laterally,
2 clip setae and anterior brush of fine setae ended at enlarged medial seta; meso- and metatibiae with
four rows of prominent setae, medial row with very fine, others rows with much stouter seta. Male
protarsomeres expanded, 1-3 with articulo-setae ventrally (Stork, 1980). Female protarsomeres 1-4
as well as meso- and metatarsomeres 1-4 in both sexes with two more or less well-defined rows of
ventral setae and numerous, scattered setae that are mostly smaller than row setae; fifth tarsomere
with two pairs of fine ventral setae, apical nearly twice length of subapical. Tarsal claws pectinate,
with 4-6 small denticles on each claw (Fig. 2A).
Abdomen. Sterna in most species shiny, microsculpture in all species obscure medially, more prom-
inent laterally. Transverse sulcus of sterna V-VI present and entire or medially absent, or completely
absent. Male genitalia (Fig. 21-23) either simple, median lobe relatively straight, endophallus minutely
spinose and scrotiform, or variously modified with median lobe slightly curved and endophallus with
fields of large spines and sacculi. Median lobe smooth in most species, strigose in some species of
the A. picipes-group. Female gonocoxite-1 with 0-4 small ensiform setae near apex, gonocoxite-2
arcuate and laterally excavated, apex round, 1 dorsal and 1 lateral large ensiform seta, 2 nematiform
setae in elongate apical furrow (e.g.. Fig. 19A). The gut of a single specimen of A. splendidula was
dissected (Fig. 7B). Crop (crp) pouched ventrally; proventriculus (pv) with 4 compact, fibrous pads;
mid-gut (MG) covered by short similarly shaped crypts (ct) and 4 Malpighian tubules (mt) intertwined
along its length; hind-gut (HG) with 5 oval rectal pads (rtp). Pygidial gland reservoir (rs) simple, oval
(Fig. 7A), efferent duct (eff) broad, subequal in length to reservoir. Collecting canal (cc) short, ramified
10-12 times, each ended with secretory cell (sc^ Collecting canal 1 mm long approximately 2X
length of efferent duct, efferent duct joined by canal just above midpoint.
Defensive chemical compounds. Methacrylic and tiglic acids, undecane, dodecane and tridecane
produced by pygidial glands (Will et al., 2000, 2001).
Species Accounts
Subgenus Abaris sensu stricto
Cladistic Diagnosis. — All members of this clade share the synapomorphic lat-
erally sulcate basal bulb of the aedeagal median lobe (Fig. 2 IB, Fig. 31: character
19).
A. striolata- group
Cladistic Diagnosis. — Individuals of this group have the tip of the aedeagus
very thin, blade-like in the lateral view (Fig. 21A-B,D, Fig. 31: character 22)
and the appended gland of the spermatheca (sg) elongate (Fig. 19B, Fig. 31:
character 30).
{l)Abaris napoensis, new species
Fig. 11, 18A, 19A--B, 21A-C, 24A
Type Material. — HOLOTYPE. Male, labeled: “Ecuador, Napo: Onkone Gare Camp, 00°39'10"S,
76°26'00"W, 220 m, terra firma forest, flowerfall-leaf litter; at night; 5&8.X.1995; 07-95. TL. Erwin
Ecuador Expedition 1995, G.E. Ball & D. Shpeley”, USNM. ALLOTYPE. Female, same data as
holotype, USNM. PARATYPES. 2 females, same data as holotype, UASM, CMNH. 7 males, same
data as holotype, UASM(4), CMNH(l), QCAZ(l), KWWC(l).
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Type Locality. — As given for holotype.
Range. — Ecuador (Fig. 11).
Recognitory Diagnosis. — The prominent baso-lateral punctation of the prono-
tum (Fig. 24 A) and form of the tip of the aedeagal median lobe (Fig. 21A-C)
are distinctive. Photograph of dorsal view figure 18A.
Description. — Medium sized, overall length 6. 5-6. 8 mm. Very shiny, aeneous head and pronotum
darker than virescent elytra. Ventral body surface, legs, mouthparts and antennae ferrugino-testaceous;
legs and abdominal sterna slightly paler than other areas.
Head. Ocular ratio 1.80-1.84. Head very shiny, reticulate microsculpture scarcely visible, frontal
impressions deep, short, slightly convergent, not clearly delimited medially, clypeus shallowly de-
pressed apically and laterally. Mentum tooth form simple, broad.
Thorax. Pronotum (Fig. 24A) sinuate laterally; basolateral region reflexed in basal third of pronotum
but slightly convex near hind angles; front angles scarcely produced; apex not narrowly constricted;
hind angles denticulate; lateral bead broad in basal third, then sharply narrowed to hind angles; ba-
solateral margin not bordered; microsculpture on disc very faint, transverse mesh. Elytra with trans-
verse microsculpture; basal section of stria 1 present; intervals 2-3 equal width, intervals flat; humeral
tooth produced anterad interval 8. Mesepisternal angle produced as small rounded tubercle. Metacoxal
sulcus arcuate, ended medially in coxa (in some specimens sulcus very short and so not arcuate, but
still divergent from apical margin).
Abdomen. Sterna V-VI with lateral sulci. Basal bulb of male median lobe laterally sulcate (Fig.
21A-C), endophallus folding pattern visible in cleared lobe; blade smooth, sinuate; tip thin, reflexed,
with broad asymmetrically expanded area. Female reproductive tract (Fig. 19B) seminal canal broad
with few loose twists, receptaculum not discrete from canal, expanded base of seminal canal very
large and continuous with bursa, appended gland elongate; gonocoxite-1 with 3 apicolateral ensiform
setae.
Etymology. — Specific epithet is Latinized adjective based the type region.
{2)Abaris striolata Bates, 1871
Fig. 16, 21D-E, 24B
Abaris striolata Bates, 1871:220.
Type Material. — LECTOTYPE (here designated). Male, labeled: “Ega” [handwritten]; “Abaris
striolata type, Bates” [handwritten]; “Ex Musaeo H.W. Bates, 1892”, MHNP.
Type Locality. — Brazil, Tefe [Ega]. From original description.
Range. — Brazil, Peru (Fig. 16).
Recognitory Diagnosis. — Individuals small, with the pronotal base smooth in
most individuals, outer basal foveae shallow, and baso-lateral region reflexed. Best
separated from other species by the characteristic pronotum (Fig. 24B) and char-
acter combinations in the key.
Description. — Small sized, overall length 5. 3-6. 2 mm. Elytra cupreous, head and pronotum black
with slight metallic reflection. Ventral body surface, mandibles and labrum piceous; legs, palpi and
antennae ferruginous to brunneous.
Head. Ocular ratio 1.71-1.90. Mentum tooth emarginate at apex. Microsculpture of head obsolete;
frontal impressions short, deep, parallel.
Thorax. Pronotal basolateral region convex, apex broad, front angles not produced, hind angles
denticulate, lateral bead narrow, basolateral margin not bordered (Fig. 24B); microsculpture on disc
transverse. Microsculpture of elytra transverse, basal section of stria 1 present, elytral humeral tooth
anterad interval 8; intervals flat on disc, more convex laterally, intervals 2-3 of equal width. Metacoxal
sulcus arcuate, ended medially in coxa. Mesepisternal angle produced as small sharp tubercle.
Abdomen. Abdominal sulci present laterally on sterna V-VI. Basal bulb of male median lobe
laterally sulcate; endophallus folding pattern visible; tip thin in lateral profile, asymmetrical in dorsal/
ventral views (Fig. 21D-E). Female tract with seminal canal broad, straight; receptaculum not discrete
from canal; spermatheca without basal bulb, expanded base of seminal canal large and continuous
with bursa; appended gland elongate; gonocoxite-1 with 2 apicolateral ensiform setae.
Notes on Life History. — Adults have been collected in July and September-
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Will — Revision of Abaris
161
November at elevations of 290-840 m. A single specimen was collected at mer-
cury vapor light and one at a “Fig fall.” Other notes include collections in sub-
tropical moist forest, leaf litter in a dry stream or simply “on ground.”
Material Examined. — In addition to type, 12 specimens. PERU. Cusco: Quincemil [MCZC]. Junin:
San Beni Valley 8 km E Satipo [SEMC]. Madre de Dios: Rfo Tambopata Reserve, 30 km (air) SW
of Puerto Maldonado [USNM]; Rfo Tambopata Reserve, 30 km (air) SW of Puerto Maldonado
12°50'S, 69°20'W [BMNH, DHK]. Tambopata Reserve, 30 km (Air) SW of Puerto Maldonado, main
trail -tf, 12°50'S, 69°20W [USNMJ.
A. aenea-gxowp
Cladistic Diagnosis. — All members of this clade have the lateral area in the
basal third of the pronotum reflexed and flat laterad the outer basal foveae (Fig.
31: character 13). A similar condition is known in species of Prosopogmus and
other Carabidae. These other occurrences of this condition are considered analo-
gous.
{3)Abaris robustula Tschitscherine, 1898
Fig. 15, 21F-G, 24C
Abarys robustulus Tschitscherine, 1898:83.
Type Material. — HOLOTYPE. Female, labeled: “Mih’om” [questionable interpretation of hand-
writing], “Abarys robustulus m. typ. Tschitscherin det” [handwritten name on printed determination
label], “Co]l. Mus. Vindob, TYPUS” [red paper], NMW.
Type Locality. — Brazil, as given in original description.
Range. — Bolivia, Brazil, Peru (Fig. 15).
Recognitory Diagnosis. — Pronotum distinctive with outer basal foveae absent
and a nearly flat, reflexed area along the margin (Fig. 24C). Very similar in general
facies to A. striolata but male median lobe is straighter and thicker at the tip
(compare Fig. 21D— E to Fig. 21F-G).
Description. — Small sized, overall length 6.4-6. 9 mm. Black with aeneous luster, less pronounced
on head and pronotum. Ventral body surface, mandibles and labrum piceous; legs, palpi and antennae
paler brunneous.
Head. Ocular ratio 1.62-1.88. Mentum tooth shallowly emarginate at apex. Microsculpture of head
faint, nearly effaced on disc, reticulate, micropunctulae present in some individuals (visible at 25 X).
Frontal impressions short, shallow, not clearly delimited medially.
Thorax. Pronotal (Fig. 24C) basolateral region reflexed, apex broad, front angles produced, hind
angles denticulate, lateral bead narrow, margin sinuate to base, basolateral margin not bordered, outer
basal foveae absent, inner punctiform; microsculpture on disc transverse, faint and shiny. Microsculp-
ture of elytra transverse; basal section of stria 1 present, intervals 2-3 of equal width, intervals flat to
slightly convex; elytral humeral tooth anterad interval 8. Metacoxal sulcus arcuate.
Abdomen. Shallowly sulcate laterally on sterna V-VI. Basal bulb of male median lobe laterally
sulcate, median lobe with subapical expansion visible in dorsal view (Fig. 21F-G); endophallus folding
pattern visible. Female reproductive tract with seminal canal narrow with many tight twists, recepta-
culum distinct, spermatheca without basal bulb, appended gland spherical; gonocoxite-1 with 2-3
apicolateral ensiform setae.
Notes on Life History. — Collected in September, October and November at
elevations of 400-750 m. The Pakitza specimen was collected in the leaf litter in
a dry, stony stream and the Bolivia specimen was taken at mercury vapor light.
Material Examined. — In addition to type, 4 specimens. BRAZIL, unknown [NMW].
PERU. Madre de Dios: Pakitza, Zone-3, 12°07'S, 70°58 W; In leaf litter of stony dry stream. Trail
Castanal [USMN]. Cuzco: Quincmil, Pena. [MCZC].
BOLIVIA. Santa Cruz: Ichilo Province, Buena Vista [CMNH].
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{A)Abaris aenea Dejean, 1831
Fig. 5, 8, 14, 18C, 21H~I, 24D-E
Abaris aenea Dejean, 1831:781.
Type Material. — LECTOTYPE, (here designated). Male, labeled: “d” [drawn on green paper];
“aenea. mihi, Carthagena” [handwritten on green paper]; “ex Museo, Chaudoir” [red ink], MHNP.
Abaris darlingtoni Straneo, 1939:38. [New Synonymy].
Type Material. — HOLOTYPE. Male, labeled: “Barro Colorado, Canal Zone, May 17 1929, Dar-
lington; 23393, MCZC. Type, darlingtoni” [red paper]; “Holotype” [red paper]; “Abaris darlingtoni,
Holotypus n.sp., det. ing. Straneo; ADP, 03439; 9 winged”; MCZC. ALLOTYPE. Female labeled:
“Barro Colorado, V. 18.26, C.Z. Van Tyne” [handwritten]; Allotype [red paper], “77” [red], “Dar-
lingtoni Straneo”, MSNM.
Type Locality. — Cartagena, Bolivar, Colombia.
Range. — Colombia, Ecuador, Panama, Peru, Venezuela (Pig. 14).
Recognitory Diagnosis. — Small and very shiny. Similar to A. erwini but distin-
guished by the obsolete microsculpture of the head and pronotum. Habitus figure
18C.
Description. — Small sized, overall length 5. 0-6.0 mm. Bronze or virescent, head and often pron-
otum darker and greener than elytra. Ventral body surface brunneous, legs and mouthparts paler rufous
or ferrugino-testaceous.
Head. Ocular ratio 1.71-1.83. Clypeus medially tumescent, apically and laterally depressed and
finely strigate or rugose in region near hind setigerous puncture, frontal impressions parallel. Micro-
sculpture absent except as faint mesh on depressed portion of clypeus; scattered micropunctulae on
dorsum. Mentum tooth slightly emarginate at apex.
Thorax. Pronotum (Fig. 24D-E) flat and reflexed in basolateral region, apex broad, front angles
slightly produced, margins subsinuate to base and broadly reflexed in basal half; lateral bead of uniform
width throughout, irregularly punctulate-rugose in basal quarter. Some specimens with faint rugosities
and punctulae along lateral margin (Fig. 24E) (This is constant and more developed in specimens
from Ecuador, only slightly punctate in specimens from Peru and mixed in specimens from Panama.
In all specimens pronotum is smoother than in A. notiophiloides, though approaching the condition in
that species in Ecuadorian populations). Hind angles slightly denticulate. Elytral microsculpture trans-
verse; basal section of stria 1 absent, intervals slightly convex, interval 3 width 1. 2-1.57 X interval 2
(see synonymy below). Metepisternal angle produced as small round tubercle. Metacoxal anterior
sulcus arcuate, ended medially in coxa.
Abdomen. Sterna without sulci. Basal bulb of male median lobe laterally sulcate, endophallus
structure visible in repose as a darker medial area in cleared lobe (Attempts to evert endophallus only
succeeded to expose it half way. From this, it appears to be scrotiform, uniformly covered with spines,
without sacculi.) Female tract with seminal canal long and narrow, with numerous tight twists, recep-
taculum distinct, appended gland spherical; gonocoxite-1 with 1-2 apicolateral ensiform setae.
Chemical data. Defensive chemicals secreted from the pygidial glands are methacrylic and tiglic
acids, undecane, dodecane and tridecane (Will et ah, 2000, 2001).
Notes on Life History. — Adults have been collected in all months of the year
except April and September, most commonly at 150-200 m elevation. Specimens
have been collected mostly at night and in thin, relatively dry leaf litter. Several
Ecuadorian specimens were collected in primary tropical forest on a well-drained
ridge in red clay soil by raking leaf litter and scraping the humus and root layer
beneath ferns at night. Although rotten Ficus fruits were found nearby — and in
association with the fruit a species of Pseudabarys Chaudoir and several species
of Harpalini — A. aenea was not found to be associated with the fruit. Specimens
have been collected in leaf litter and at flower falls both during the day and night
at several sites. A few specimens have been taken at ultraviolet or white lights.
Synonymy. — Straneo (1939) described A. darlingtoni from Barro Colorado Is.,
Panama. Given the small amount of material at his disposal, Straneo determined
2002
Will — Revision of Abaris
163
12i
▼
11.5^
▼ B3
£
1
▼ fl T
€0
ro 10.5^
ffl ffl
.1 10
m ▼
9.5
▼
Q-
6 6.5 7 7.5 8 8.5 9
interval 2 width
▼ Panamanian specimens ffl mainland specimens
Fig. 8. — Graph of width ratio of elytral intervals for specimens of Abaris aenea Dejean. Average ratio
of interval 3(Y axis)/2(X axis). Units on axes in ticks of ocular reticle. Standard deviation and number
of individuals measured: “darlingtoni” Panamanian form, 1.57 ± 0.04, n = 12; Columbia, 1.45 ±
0.31, n = 6; Venezuela, 1.47 ± 0.27, n = 3; Ecuador, 1.50 ± 0. 13, n = 4.
that specimens with a relatively wider third elytral interval, as compared to A.
aenea, combined with a lack of lateral pronotal punctures, as in A. notiophiloides,
belonged to a distinct species. Based on the larger amount material I have at hand,
it is clear that interval width variation is not a sufficient character for species
recognition in this complex.
Individuals were measured for the ratio of interval 3/interval 2 from all col-
lecting sites, representing both South American A. aenea and the Panamanian
''darlingtonf’ form (Fig. 8). No discrete difference can been seen when all data
are compared. Individuals from the island population are very similar to each
other, but are within the variation seen in the Venezuelan, Ecuadorian and Col-
ombian populations. Similarly, all other characteristics show no difference among
populations of A. aenea that are not subsumed by variation between individuals.
Likewise, the Panamanian ''darlingtoni” form is not distinguishable from South
American specimens of A. aenea.
Material Examined. — 133 specimens. COLOMBIA: [IZWP]; Cartagena [BMNH, MHNP]; Rio
Frio, Magdalena [MCZC].
ECUADOR. Napo; Onkone Gare Camp, 0°39'10"S, 76°26'0"W [UASM]; Limoncocha [UASM];
Yasuni Scientific Station, 00°40'36"S, 76°24'02"W [KWWC, CUIC].
PANAMA. Canal Zone: Barro Colorado Island [CMNH, MCZC], 9°9'15"N, 79°15'0"W, [USNM];
Chepo, Altos de Maje [STOC].
PERU. Madre de Dios: Rio Tambopata Reserve, 30 km (air) SW Pto., Maldonado, 12°50'S,
06°20'W [BMNH]; Rio, Tambopata Reserve, 30 km, (air) SW of Puerto Maldonado, 12°50'S, 69°20'W
[DHK].
VENEZUELA. [MHNP]; San Esteban [MHNP].
{5)Abaris erwini, new species
Fig. 11, 18B, 21J-M, 24F
Type Material. — HOLOTYPE. Male, labeled: “Peru: MADRE DE DIOS, Pakitza, Zone 02, 9&12
Feb90 T L Erwin, 70°58W 12°07'S; Under leaves and, fallen fruit, Tr. Aguajal 107; BIOLAT,COLE,
000007170”, USNM. ALLOTYPE. Female with same data except last label, “BIOLAT,COLE,
000007153”, USNM. PARATYPES. 5 males with the same data as the Holotype except the following
labels; BIOLAT,COLE, 000007156; BIOLAT,COLE, 000007152; BIOLAT,COLE, 000007154; BIOL-
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AT,COLE, 000007155; BIOLAT,COLE, 000002472, USNM, and 5 females with the same data as the
holotype except the following labels; BIOLAT,COLE, 000007157; BIOLAT,COLE, 000007169; BIOL-
AT,COLE, 000007164; BIOLAT,COLE, 000007156; BIOLAT,COLE, 000007154, USNM. Female,
Chambireyacu, pres Yurimaguas, (Huallaga-Perou), M. de Mathan, Juin-Aout 1885, MHNR Female,
Peru, Madre de Dios, Rio Tambopata Res., 30 km (air) sw Pto., Maldonado, 12°50'S 069°20W;
B.M. 1983.455, N.E. Stork, 3.x.-15.xi.l983, on ground, BMNH. Male, Peru: Tambopata Prov., Madre
de Dios Dplo.l5 km NE Puerto; Maldonado, Reserva, Cuzco Amazonica, 12°33'S, 69°03'W, 200 m.
Plot #Z1E9; 17 June 1989, J.S Ashe, R.A.Leschen #136, ex. Flight intercept trap, SEMC. Male, Peru,
Junm San Ramon de Pangoa 40 km SE Satipo, Schuh, R.T.& J.C., 7 June 1972, Soils-litter layer in
primary forest, AMNH.
Type Locality. — As given for holotype.
Range. — Peru, Bolivia (Fig. 11).
Recognitory Diagnosis. — Small and shiny, somewhat variable species, separat-
ed from A. aenea by the more evident microsculpture on the head and generally
less punctate pronotum. Separated from other similar species by characters in the
key. Photograph of dorsal view figure 18B.
Description. — Small sized, overall length 5. 3-5. 9 mm. Shiny, aeneous, head and pronotum darker,
virescent. Ventral body surface, labrum and mandibles dark brunneous; legs either brunneous or sightly
paler than venter of body; tibiae, tarsi palpi and distal antennomeres paler.
Head. Ocular Ratio 1.73-1.78. Frontal impressions short, parallel, shallow, sharply delimited exter-
nally. Mentum tooth slightly emarginate at apex. Microsculpture of head very faint, reticulate.
Thorax. Pronotal basolateral region slightly convex though obscured in some individuals by punc-
tures; apex broad, front angles little produced, margins subsinuate; hind angles not denticulate, lateral
bead uniform thickness throughout or narrowed in apical quarter, basolateral margin not bordered;
microsculpture transverse, very faint, shiny. Elytral microsculpture transverse; basal section of stria 1
absent, intervals flat, interval 3 width 1.1-1.4X interval 2, mesepisternal angle produced as low flat
tubercle; elytral humeral tooth anterad interval 8. Metacoxal sulcus arcuate, ended medially in coxa.
Abdomen. Sterna without sulci. Male aedeagus (Fig. 21J-M) with basal bulb of median lobe lat-
erally sulcate (Fig. 21J, bs); endophallus folding pattern visible in cleared lobe (I attempted to evert
the endophallus of several specimens but the internal sac only everted half way. A basal right sacculus
covered in medium length spines was apparent on everted portion); median lobe blade smooth. Female
tract with seminal canal thin with many tight twists, receptaculum distinct, appended gland spherical;
gonocoxite-1 with 3 apicolateral ensiform setae.
Variation. — In regard to pronotal form, this species is rather variable and may
prove to be a complex of species rather than one polymorphic species.
Notes on Life History. — Adults have been collected in January, February, June,
July and September-November at elevations of 200-750 m. Specimens were col-
lected in leaf litter and in areas with fallen rotten fruits, such as Ficus fruits.
Material Examined. — 29 specimens examined, including types. BOLIVIA. Chapare [MSNM].
PERU. Huallaga: Yurimaguas, Chamireyacu [MHNP]. Junm: San Ramon de Pangoa 40 km SE
Satipo [AMNH], Madre de Dios: Pakitza, 12°7'0"S, 70°0'0"W [USNM]; Pakitza, Rio Manu, BIOLAT
Sta. 11°56'47"S, 71°17'0"W [USNM]; Pakitza, Zone 04, 12°7'0"S, 70°0'0"W [USNM]; Rio Tambopata
Reserve, 30 km (air) SW of Puerto Maldonado, 12°50'0"S, 69°20'0"W [CASC]; Rio Tambopata Re-
serve, Explorer’s Inn 12°50'0"S, 69°17'0"W [USNM].
Etymology. — Eponym based on the surname of the collector and carabidologist Terry L. Erwin.
{6)Abaris aquilonaria, new species
Fig. 12A, 20A, 21N-0, 24G
Type Material. — HOLOTYPE. Female, labeled: “Mexico, Chiapas, 3200'. 1 1.6 mi. N Ocozocuautla,
vi. 10-13. 1966; George E. Ball, D.R. Whitehead collectors”, USNM. ALLOTYPE. Male, labeled:
“Coyame, Lake Catemaco, Veracruz, Mexico, 10-1 8. VII. 63-Black Its. D.R. Whitehead”, USNM.
PARATYPES. Female, label data same as holotype, UASM. Female, labeled: “Mexico, Chiapas 11.6
mi.N Ocozocoautla, 3200', May 25 & June 20 ’72, A. Meyer, G.E.& K.E. Ball, Collectors”, UASM.
Male, same label data as allotype, KWWC. Male, genitalia dissected, labeled: “Mexico, Chiapas,
2002
Will — Revision of Abaris
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Parque Laguna Belgica 19 km N.Ocozocoautla, 2 June 1991, 970 m, J.S.Ashe Coll#46, ex; flight
intercept trap”, SEMC.
Type Locality. — As given for holotype.
Range. — Costa Rica, Honduras, Mexico (Fig. 12A).
Recognitory Diagnosis. — Similar to A. aenea but the presence of visible mi-
crosculpture on the pronotum and the lack of punctation along the base imme-
diately distinguishes A. aquilonaria.
Description. — Small sized, overall length 5.4-6. 1 mm. Shiny, aeneous, with head and pronotum
slightly darker. Ventral body surface dark brunneous; legs, antennae and mouthparts paler brunneous
to ferrugino-testaceous.
Head. Ocular ratio 1.74-1.85. Mentum tooth slightly emarginate at apex, microsculpture of head
faint, reticulate. Frontal impressions short, deep, not clearly delimited medially. Clypeus smooth or
with a few shallow lines.
Thorax. Pronotum (Fig. 24G) with obvious, slightly transverse mesh microsculpture; region laterad
of basal foveae flat and reflexed, apex broad, front angles not produced, hind angles denticulate, lateral
beads uniform or slightly narrower at hind angles, lateral margins sinuate to base. Metacoxal sulcus
arcuate, ended medially in coxa. Basal section of elytral stria 1 absent; intervals slightly convex. Elytra
with interval 3 width 1.5-2. IX interval 2, humeral tooth anterad interval 8; microsculpture transverse.
Abdomen. Sterna without sulci. Basal bulb of male median lobe laterally sulcate, blade smooth
(Fig. 2 IN). Female reproductive tract (Fig. 20A) with seminal canal long, narrow with a few (4-5)
tight twists, receptaculum distinct, appended gland (sg) spherical; gonocoxite-1 with 2 apicolateral
ensiform setae.
Notes on Life History. — Adults collected in May-July at elevations of 700-
970 m. Collections have been made using flight intercept traps and ultraviolet
lights.
Material Examined. — 6 specimens in addition to types. COSTA RICA. Guanacaste: Guanacaste
Conservation Area, Martinez Biological Station [CASC]. This specimen was not included in the type
series as it differs in color (bright green) and the pronotum has longer straight sections in front of the
hind angles. However, no other differences could be found in the external characteristics or the median
lobe of the aedeagus. Additional material of A. aquilonaria will either show population level variation
that includes this form, or the necessity of describing a new species.
HONDURAS. San Juan Pueblo [USNM]. This specimen is all black, much duller, and the pronotum
is somewhat flatter without the sharp denticles of the hind angles of the pronotum found in the typical
A. aquilonaria. Unfortunately, genitalia of this male specimen are lost. In general form it is quite
similar to typical specimens but, like the Costa Rican form, may represent another species. Again,
additional material is needed. [USNM],
MEXICO. Chiapas: 11.6 mi.N Ocozocoautla. Veracruz: Coyame, Lake Catemaco [UASM, teneral
and damaged specimens not included in type series]
Etymology. — Latin adjective aquilonaris (northern). This refers to the fact that this species ranges
farther north than any other A. aenea-group species.
{l)Abaris notiophiloides Bates, 1871
Fig. 11, 19C, 21P-Q, 24H
Abaris notiophiloides Bates, 1871:220. Abarys notiophiloides (Chaudoir, 1873:98).
Type Material. — LECTOTYPE (here designated). Female: Labeled: “Ega” [Handwritten]; “Ex.
Museao H.W.Bates, 1882”, “Abaris notiophiloides Bates type” [Handwritten]; “Lectotype Abaris
notiophiloides Bates, K.W.Will 1997” [red paper label], MHNP.
Type Locality. — Brazil, Tefe [Ega], from original description.
Range. — Brazil (Fig. 11).
Recognitory Diagnosis. — Immediately recognizable by the greatly widened ely-
tral interval 3 in combination with the punctate pronotal margins.
Description. — Small sized, overall length 4.8-5. 1 mm. Brunneous with a cupreous luster, head and
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pronotum in most individuals, some with only head, slightly darker. Ventral body surface somewhat
paler; legs, palpi and antennae paler.
Head. Ocular ratio 1.79-1.86. Mentum tooth shallowly emarginate at apex. Microsculpture of head
reticulate, frontal impressions short, divergent, not clearly delimited medially.
Thorax. Pronotal (Fig. 24H) basolateral region flat and reflexed, apex broad, front angles not pro-
duced, punctate along margin anterior to outer foveae; margins sinuate, hind angles denticulate, lateral
bead narrow and of uniform thickness, basolateral margin not bordered; microsculpture transverse on
disc, more irregular at base and near foveae. Elytra with basal section of stria 1 absent, intervals
moderately convex, flatter dorsally, interval 3 width 2X width of interval 2, elytral humeral tooth
anterad interval 8; microsculpture transverse, faint. Mesepisternal angle produced as small flat tubercle.
Metacoxal sulcus arcuate, ended medially in coxa.
Abdomen. Sterna without sulci. Basal bulb of male median lobe laterally sulcate, endophallus
folding pattern visible, blade smooth (Fig. 21 P-Q). Female reproductive tract (Fig. 19C) with seminal
canal short, broad, straight, receptaculum not discrete from seminal canal, without basal bulb, fibrous
region at base of spermatheca, appended gland spherical, gonocoxite-1 with 2 apicolateral ensiform
setae.
Notes on Life History. — Found together with Pseudabarys robustus (Bates)
(Bates, 1871:220).
Material Examined. — Including types, 12 specimens examined. BRAZIL. Chapada [MSNM]; Ega
[MHNP]; Itaituba, Amazones (Hahnel) [MHNP]; Santarem [CMNH]; Unknown; [MHNP].
A. picipes-group
Cladistic Diagnosis. — Members of this clade have either faint or obsolete trans-
verse microsculpture on the pronotal disc and the region near the basal foveae
(Fig. 31: characters 16, 17). This clade includes some of the most widespread
and variable species in the genus. Although the general form and internal struc-
tures of the male genitalia are similar among all species in the clade, it is not
possible to define any non-homoplasious synapomorphies of the male genitalia
that characterize the clade at the level of the species-group. Within the group,
strigose ventral surface of the median lobe (character 21) found in A. mina, A.
picipes, and A. basistriata is a synapomorphy for those species. The median lobe
of A. metallica is not known, but is predicted to be strigose as well (Fig. 31).
{^)Abaris impunctata, new species
Fig. 15, 21R-S, 241
Type Material. — HOLOTYPE. Male, labeled: “Brazil, Santa Catarina, Nova Teutonia, 27°1F0"S,
52°23'0"W, 300-500 m, F.Plaumann”, MCZC.
Type Locality. — As given for holotype.
Range. — Brazil (Fig. 15).
Recognitory Diagnosis. — Recognizable, dark species with dull surface from
reticulate and somewhat granulate microsculpture and impunctate pronotal base
(Fig. 241).
Description. — Medium sized, overall length 7.0 mm. Black with a slight aeneous luster. Ventral
body surface, legs and mouthparts brunneo-piceous.
Head. Ocular ratio 1.67. Mentum tooth form simple. Microsculpture of head reticulate; frontal
impressions short, punctiform.
Thorax. Pronotal basolateral region convex, apex broad, front angles very slightly produced, hind
angles minutely denticulate; lateral bead of uniform width through most of its length narrowed just
before hind angles; basolateral margin not bordered (Fig. 241); microsculpture reticulate and surface
dull. Microsculpture of elytra obvious, reticulate; basal section of stria 1 present, intervals flat, intervals
2-3 equal width, mesepisternal angle produced as small flat tubercle; humeral tooth anterad interval
8. Metacoxal sulcus arcuate, ended medially in coxa.
Abdomen. Sterna V-VI shallowly sulcate laterally, deeper on VI where it is oblique to base of
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sternum. Male median lobe (Fig. 21R-S) basal bulb laterally sulcate, endophallus folding pattern
visible in cleared median lobe, blade smooth, tip symmetrical. Female genitalia and reproductive tract
not studied.
Etymology. — Specific epithet is a Latin adjective referring to the lack of obvious punctures along
the base of the pronotum.
(9)Abaris bigenera Bates, 1882
Fig. 12B, 19D, 22A-D, 25A
Abaris bigenera Bates, 1882:86. Larval description by Bousquet and Liebherr (1994).
Type Material. — LECTOTYPE, (here designated). Male, labeled: “Oaxaca, Mexico. Hoege”,
BMNH.
Type Locality. — Mexico, Oaxaca, from original description.
Range. — Guatemala, Honduras, Mexico (Fig. 12B).
Recognitory Diagnosis. — One of the most common Mexican species. Easily
separated from the other common Mexican species, A. aequinoctialis, by the larg-
er size and more somber color. Relatively large and dark species, frontal impres-
sions effaced, rather variable but recognizable using characters in the key.
Description. — Variable in size but most relatively large sized, overall length 6.4-7. 9 mm. Dark,
moderately shiny, metallic luster faint, cupreous in most individuals, but striking blue or green in a
few individuals; head and pronotum concolorous with elytra in most, darker in some individuals.
Ventral body surface brunneous; legs, mouthparts and antennae slightly paler but infuscated in some
individuals, especially mandibles, antennae and femora.
Head. Ocular ratio 1.68-1.80. Mentum tooth shallowly emarginate at apex. Microsculpture of head
reticulate; frontal impressions short, shallow and convergent or effaced.
Thorax. Pronotal margins smoothly arcuate or subsinuate to hind angles that are either minutely
denticulate or not; apex wide, front angles slightly produced, lateral bead narrow and uniform through-
out its length, basal margin not bordered, basolateral region slightly convex; microsculpture reticulate,
surface dull. Basal section of elytral stria 1 present, intervals flat, intervals 2-3 of equal width, elytral
humeral tooth anterad interval 8; microsculpture transverse; surface moderately shiny. Metacoxal sul-
cus arcuate, ended medially in coxa. Mesepisternal angle very small, produced as low round tubercle.
Abdomen. Abdominal sulci present laterally on sterna V-VI. Median lobe basal bulb laterally
sulcate, endophallus (Fig. 22C-D) with ventral field (vs) of large spines and left and right basal-lateral
sacculi (11s, rls), right sacculus recurved, partially spine covered with its position corresponding to
spined ridge in A. basistriata; median lobe (Fig. 22 A) ventrally smooth, lobe tip asymmetrically
produced to right (Fig. 22B). Female reproductive tract (Fig. 19D) with seminal canal narrow with
many tight twists, receptaculum distinct, appended gland (sg) spherical, gonocoxite-1 with 2, 3 or 4
apicolateral ensiform setae.
Notes on Life History. — Adult specimens have been collected in April-Septem-
ber at elevations of 200-1680 m. When habitat was noted, most specimens were
collected in the leaf litter of relatively dry oak-pine forest, e.g., “oak-pine zone,
Alnus near stream in litter.” Specimens were also collected in oak forest, wet oak-
pine forest and montane tropical forest. Other habitat associations include coffee
finca, crustose fungi and crustose fungi on logs or under logs and gilled mush-
rooms. In addition to collecting from leaf litter, pan traps were reported as a
collection method for some specimens.
Material Examined. — Including type, 223 specimens. GUATEMALA. Baja Verapaz, 7.6 km W
Chilasco [SEMC]; Baja Verapaz, 1.6 km S Pantin 15°16'0"N, 90°14'0"W [USNM]; Baja Verapaz, 8
km S Purulha [UASM]; El Quiche, 7.3 m S Chichicastenango 14°54'0"N, 91°7'0"W [USNM]; S
Geronimo [MHNP]; Sacatepequez Florencia [UVGC]; El Progreso 21 km NE San Augustine Acasa-
guastlan [UASM]; Zacapa Dept., San Lorenzo [CNC]; Sierra de las Minas [CNC].
HONDURAS. Los Pinos, 14°32'0"N, 87°53'0"W [UASM]; Ocotepeque 12.7 km E & 10.6 km S
Ocotepeque lower slopes El Pital, 14°25'0"N, 89°4'0"W [SEMC]. Morazan: Francisco 21.3 km N
Teguicigalpa, la Tigra, 12°14'0"N, 86°6'0"W [SEMC].
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MEXICO. Chiapas: 6 km SE Tollmen [UASM]; 18.7 km W Rizo de Oro, Chiapas/Oaxaca border
[UASM]; 3.1 mi. N Pueblo Nuevo [UASM];3.1 mi. N Pueblo Nuevo, Rtel95 [UASM]; 8.6 mi. E San
Cristobal, rte 190 [UASM]; El Rincon [CNC]; Lagos de Monte Bello National Park [UASM]; Mun-
icipio de Ocosingo 2nd ridge NE of Las Margaritas above La Soledad [CASC]. Guerrero: 12.1 mi.
W Chilpancingo [UASM]; 13.9 mi. W Chilpancingo [UASM]; 143.1 km NE Atoyac de Alvarez
[UASM]; 2 km SW Yerba Santa 39 km S Filo de Caballo [CMNH]; 52.9 km NE Atoyac de Alvarez
[UASM]; 78.5 km N junction Rt200 on rtl34 to Ciudad Altamirano [UASM]; B. Vista de Cuellar,
“El Pochote” [UASM]; Omiltemi [MHNP, UASM]. Jalisco: 10.8 mi. S Talpa de Allende [UASM];
12.4 mi. S Tecalitan [UASM]; 21.4 mi. S Tecalitan [UASM]; 42.4 km NW Cuatla [UASM]; 5.5 mi.
NW Cuatla [UASM]; 7 km S Tequila rd to Microondas [UASM]; 8.8 km NW Cuatla [UASM]; 9.6
km S Tequila on rd to Microondas [UASM]; El Ricon 30.5 mi. NW Los Volcanes [UASM]; Mexico,
2.7 km NE Temazcaltepec, rte 130 [UASM]. Michoacan: 97.7 km W Apatzingan on road to Dos
Aguas [UASM]. Oaxaca: 11.4 mi. E Sta. Catarina Juquila [CNC, UASM]; 12.8 mi. E Sta. Catarina
Juquila [UASM]; 14.3 km E Ixtlan de Juarez [UASM]; 14.9 km N Sola de Vega [UASM]; 14 km E
Mitla (road to Zacatepec) [CUIC]; 15.7 km E Rte 190 road to Ojo de Agua [UASM]; 20 mi. S
Juchatengo, rtl31 [UASM]; 2 km W Capulalpam [UASM]; 4.9 mi. E Sta. Catarina Juquila [UASM],
6.6 mi. E Sta. Catarina Juquila [UASM], 7.3 mi. E Sta. Catarina Juquila [UASM], 11.4 mi. E Sta.
Catarina Juquila [UASM], 12.8 mi. E Sta. Catarina Juquila [UASM]; Hoege [MHNP]; NW Sta. Maria
Nizavaguiti 16°4rO"N, 95°50'0"W [UASM]; Hwyl35, 23.6 km N Jctl90 at Huitzo [CUIC]; Hwyl75,
18.3 km S Guelatao (7.5 km N El Punto) [CNC, CUIC]; Rte 131, 21.8 mi. N Juchatengo [UASM];
Queretaro, 17.8 mi. E Landa de Matamoros [UASM]; 6.4 mi. E Pinal de Amoles [UASM], San Luis
Potosi: 24.7 mi. S Landa de Matamoros [UASM]; 4.3-17.9 mi. E Ciudad del Maiz rteSO [UASM],
Sinaloa: 60.9 km E Concordia rte 40. Veracruz: 16.4 km S Orizaba on road to Tlaquilpan [UASM];
2.3 km S Jalapa on rd to Coatepec [UASM].
{\(y)Abaris picipes Bates, 1871
Fig. 13, 18D, 19E, 23A-C, 25B
Abaris picipes Bates, 1871:220.
Type Material. — LECTOTYPE (here designated). Female, labeled: “Tapajos” [handwritten];
“Abaris picipes type, Bates” [handwritten]; “Ex Musaeo H.W. Bates, 1892” [printed], MHNP.
Type Locality. — Brazil, Tapajos, from original description.
Range. — Argentina, Bolivia, Brazil, Paraguay (Fig, 13).
Recognitory Diagnosis. — Very similar in general form to A. mina (see descrip-
tion of that species) but can be separated from that species by the punctate pro-
notal base. Photograph of dorsum figure 18D.
Description. — Small to medium sized, overall length 5. 5-6.4 mm. Shiny, black with cupreous luster;
less metallic on head and pronotum, head and pronotum slightly darker in most individuals. Ventral
body surface brunneous to piceous; legs, mouthparts and antennae concolorous or slightly paler than
ventral body surface.
Head. Ocular ratio 1.67-1.74. Mentum tooth shallowly emarginate at apex. Microsculpture of head
obsolete, dorsal micropunctulae present; frontal impressions very short, shallow almost effaced.
Thorax. Pronotal (Fig. 25 B) basolateral region convex, punctate in region of basal foveae (single
female from Bolivia lacks punctures), apex broad, front angles produced, hind angles with or without
a small denticle, lateral bead narrow and of uniform thickness, basolateral margin in some specimens
with a hne engraved line in form of border at margin beneath lateral foveae; microsculpture of pron-
otum obsolete; very shiny. Microsculpture of elytra transverse, faint; basal section of stria 1 present,
intervals flat, intervals 2-3 of equal width, humeral tooth anterad interval 8. Mesepisternal angle
produced as small tubercle. Metacoxal sulcus arcuate.
Abdomen. Abdominal sulci absent in most individulas, but in some present laterally as faintly
impressed, irregular lines. Median lobe (Fig. 23A-C), aedeagal basal bulb laterally sulcate, endophallus
folding pattern (fp) visible as a large dark region in cleared lobe, spine patch visible in bend of median
lobe; blade strigate (Fig. 23A), tip (Fig. 23B) asymmetrically produced to right. Female reproductive
tract (Fig. 19E) with seminal canal broad, simple; receptaculum distinct, slightly enlarged relative to
canal; appended gland (sg) spherical; gonocoxite-1 with 0-2 apicolateral ensiform setae.
Notes on Life History. — Adults have been collected in January, February, June,
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Will — Revision of Abaris
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October-December at elevations of 450-1000 m. Habitat associations include
forested areas, in leaf litter and in a litter filled ravine.
Material Examined. — In addition to type, 65 specimens. ARGENTINA. Mendoza [MSNM]; Tar-
tagal Salta, Dio San Martin [MSNM]; Tucuman [MHNP]; Jujuy Calilegua National Park, Aguas Ne-
gras [CNC]; Jujuy Calilegua National Park, Estaca El Cero [CMNC].
BOLIVIA. Santa Cruz: Alto de Chaves Ascencion [MHNP]; Andres Ibanez Potrerillo de Guenda,
Terebinto 17°40'20"S, 63°27'0"W [CMNH]; Prov. Andres Ibanez, Vallecito [CMNH]; Sara [CMNH,
MSNM]; Villa Montes [UASM],
BRAZIL. Chapada [CMNH,MSNM]; Mtt. Grosso, Corumba [MCZC]; Santarem [CMNH]; Tapajos
[MHNP]; Bahia Bon “jiue-garbe” [illegible handwriting] [MHNP]; Bahia, Encruzilhada [MHNP];
Distrito Federal, 20 km E Brasilia [CNC], Goias, Cuyaba [MHNP]; Rio Verde [MHNP]; Jatahy
[MHNP]; Minas Gerais, Aquas Vermelhas [CMNH]; Sao Paulo, Teodoro Sampaio [CMNH].
PARAGUAY. [MNHB]; Dralze [MCZC]; Puerto Bertoni [MCZC]; Sa. Trinidad [MNHB]; Villar-
rica [MSNM,MCZC]; vie. Horqueta [MSNM, MCZC]; Boqueron Medina-cue [MCZC]; Paraguari,
Parc. Nac. Ybycuf [USNM].
UNKNOWN: Molinasque [MCZC]; Salta Urundel [MHNP].
{\\)Abaris mina, new species
Fig. 16, 23D-F, 25C
Type Material. — HOLOTYPE. Female, labeled; “Brazil, Chapada, Acc. No 2966”, CMNH. AL-
LOTYPE. Male, same label data as holotype, CMNH. PARATYPES. Female, same data as holotype
and “Oct”, CMNH, Female, same data as holotype and “Aug”, CMNH. Male, same data as holotype
and “Sept”, USNM. Male, labeled: “Brazil, Goyaz, Jatahy, Museum Paris, Ex. Coll. M. Maindron,
Coll. G. Babault 1930”, MHNP. Male, labeled: “Goyaz, Rio Verde, Museum Paris, Ex. Coll. M.
Maindron, Coll. G. Babault 1930”, MHNP. 2 Females, labeled: “Bresil, Jatahy, Prov. De Goyaz,
Donchier march 1903, Museum Paris, Ex. Coll. M. Maindron, Coll. G. Babault 1930”, MHNP. Female,
labeled: “Brazil, 20 km E. Brasilia, DF, 111-4-1970,1000 m, JM& BA Campbell”, CNC. Female,
labeled: “Paraguay, Stapua Cantera” [handwritten], MNHP. Female, labeled: “Paraguay”, MSNM.
Female, labeled; “vie. Horqueta, Paraguay”, MCZC. Female, labeled: “Villarrica, Paraguay, VI- 1932,
Zool. Mus. Berlin”, MNHB. Female, labeled: “Paraguay, leg. Piebrig, Zool. Mus. Berlin”, MNHB.
Female, labeled: “Bolivia, Santa Cruz, Ichilo Province, Buena Vista, 400 m, R.Ward, mercury vapor
light”, CMNH.
Type Locality. — Brazil, Chapada.
Range. — Brazil, Bolivia, Paraguay (Fig. 16).
Recognitory Diagnosis. — Very similar in form to A. picipes but distinguished
from that species by the smooth pronotal base.
Description. — Medium sized, overall length 6. 0-6. 7 mm. Shiny, black, aeneous, or virescent in
some individuals. Ventral body surface, legs, mouthparts and antennae brunneous.
Head. Ocular ratio 1.67-1.74. Mentum tooth shallowly emarginate at apex. Microsculpture of head
obsolete, surface shiny; frontal impressions shallow or effaced.
Thorax. Pronotal (Fig. 25C) basolateral region convex, apex broad, front angles produced, lateral
margins arcuate almost to hind angles or slightly anterad base; hind angles with or without small
denticle, lateral bead narrow, of uniform thickness throughout; basolateral margin without border in
most, few specimens with finely engraved line in form of border along base posterad foveae; micro-
sculpture obsolete on disc, very shiny. Microsculpture of elytra transverse; basal section of stria 1
present, elytral humeral tooth anterad interval 8. Mesepisternal angle produced as a very small round
tubercle. Metacoxal sulcus arcuate.
Abdomen. Abdominal sulci present as very shallow irregular lateral lines on sterna V-VI, best
developed on V. Median lobe (Fig. 23D-F) basal bulb laterally sulcate, endophallus folding (fp) pattern
visible as large dark area in cleared median lobe, blade (Fig. 23D) strigate ventrally, tip (Fig. 23E)
symmetrically produced. Female tract with seminal canal narrow with many tight twists, receptaculum
distinct, appended gland spherical; gonocoxites-1 with 2-3 apicolateral ensiform setae.
Notes on Life History. — Adults have been collected in March, June, and Au-
gust-November. Only one specimen was labeled with elevation data, a collection
at 1000 m.
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Material Examined. — Including types, 22 specimens. ARGENTINA. Unknown [MCZC],
BRAZIL. Goias, Jatahy [MHNP].
UNKNOWN: [MNHB].
Etymology. — Specific epithet is a Latin adjective that refers to the smoothness of the pronotum of
this species.
{\2)Abaris basistriata Chaudoir, 1873
Fig. 14, 18E, 22E-I, 25D
Abarys basistriatus Chaudoir, 1873:98.
Type Material. — LECTOTYPE, (here designated). Female, labeled: “ex Museo, Chaudoir” [red
ink], “Brezil, Schlb-jun” [? Handwritten, not legible], “Lectotype, Abarys basistriatus Chd., K.W.Will
1997” [Red label], MHNP.
Type Locality. — Nova Friburgo, Brazil, from original description.
Range. — Argentina, Bolivia, Brazil, Colombia, Paraguay, Peru, Venezuela (Fig.
14).
Recognitory Diagnosis. — Similar to A. mina and A. picipes in general form but
recognizable by the reticulate microsculpture on the dorsum. Photograph of dor-
sum figure 18E.
Description. — Medium to small sized, overall length 5.4-6. 6 mm. Black with aeneous luster, only
moderately shiny. Ventral body surface, legs, mouthparts and antennae brunneous.
Head. Ocular ratio 1.74-1.86. Mentum tooth emarginate at apex. Microsculpture of head reticulate;
frontal impressions shallow, not clearly delimited medially.
Thorax. Pronotal (Fig. 25D) basolateral region slightly convex, apex broad, front angles produced,
hind angles not denticulate; basolateral bead narrow, uniform thickness; basolateral margin not bor-
dered, margins subsinuate or straight to base; microsculpture reticulate, surface dull. Basal section of
elytral stria 1 present, (a single specimen from Rio Salado, Argentina lacks the basal section of stria
1, except for a slight trace on the right elytron); mesepisternal angles produced as small sharp tubercles,
elytral humeral tooth anterad interval 8; microsculpture reticulate, intervals flat, intervals 2-3 of equal
width, striae fine. Metacoxal sulcus arcuate ended medially in coxa.
Abdomen. Sterna without sulci. Basal bulb of male median lobe laterally sulcate, endophallus (Fig.
22H-I) with apico-ventral field (als) of large spines extended dorsally near base, prominent ventro-
medial spined row (sr), a long right lateral scale (Is) that is apically free, and a short dorso-basal scale
(dbs) that is spine tipped and held appressed to tip of median lobe when endophallus is everted,
median lobe ventrally strigate (Fig. 22E), lobe tip asymmetrical, produced to right. Female reproductive
tract with seminal canal long, thin and with many tight twists, receptaculum distinct, appended gland
spherical; gonocoxite-1 with 2-3 apicolateral ensiform setae.
Notes on Life History. — Adults have been collected in all months of the year
at elevations of 100—3600 m. The two specimens from Huanuco, Peru, labeled
2500-3600 m are from a much higher elevation than known for any other
Abaris species. It is likely that these represent a chance collection or are mis-
labeled specimens, as all other records for A. basistriata are for elevations of
1550 m or less. The only additional information is from a specimen collected
using a flight intercept trap in subtropical humid forest, one marked as col-
lected “under stones” and ten specimens collected at Mercury vapor or ultra-
violet lights.
Material Examined. — In addition to type 166 specimens examined. ARGENTINA. Eldorado Mi-
siones [AMNH]; Gargas [MHNP]; LaPampa (Pico) [MCZC]; Rio Salado [MSNM, MHNP]; Cata-
marca, S of La Vina, 28°3'0"S, 65°35'0"W [AMNH]; Salta, La Caldera, El Ucuma, 1550 m [CMNC];
San Martin, Tatagal Salt. [MSNM].
BOLIVIA. Santa Cruz: Alto de Chaves Ascencion [MSNM]; Ichilo, Bueno Vista [CMNH]; Andres
Ibanez Potrerillo de Guenda, Terebinto, 17°40'20"S, 63°27'0W [CMNH]; Sara [CMNH, MNHB].
BRAZIL. [IZWP, MCZC]; Chapada [CMNH]; Guapi [USNM]; Sao Paulo Curityba [MNHB]; San-
tarem [AMNH, CMNH]; Bahia, Encruzilhada [MHNP]; Distrito Federal, Estacao Florestal Cabeca do
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Veado [CNC]; Goias, Cuyaba [MHNP]; Rio Verde [MHNP]; Jatahy [MHNP, MNHB]; Mato Grosso
do Sul, R. Caraguata [MCZC]; Minas Gerais Aquas Vermelhas [MSNM]; Belo Horizonte [MHNP];
Lambary [USNM]; Parana, Bocaiuvu [UASM]; Caviuna [AMNH]; nr. Pato Branco [UASM]; Rio
Grande do Sul, S.Rosa [MCZC]; Sta. Catarina, Nova Teutonia, [CASC, MCZC, USNM]; Nova Teu-
tonia 27°1 L0"S, 52°23'0"W [MSNM, MCZC, MHNP, UASM, USNM]; Nova Teutonia, Sa Catharina
[MCZC]; nr Chapaco [UASM]; Sao Paulo, Campos do Jordao [CMNH].
COLOIVIBIA. NW Sierra N de Sta. Maria [MCZC].
PARAGUAY. 40 km E Villarrica [MSNM]; vie. Horqueta [MCZC]; Amabay Cerro Cora [USNM];
Central San Lorenzo [MHNM, CMNH],
PERU. Chanchamayo [MSNM]; Rio Pampas (Hyw 7) [CASC]; Huanuco [MCZC]; Piedras Grandes
[SEMC].
VENEZUELA. [MHNP]; Caracas [MHNP, MNHB]; Aragua, El Limon [MIZA]; Bolivar, Guri Rio
Caroni [MIZA].
UNKNOWN: [MHNP], [?]Araras-Sp. [USNM]; [?]Delbrando Leme-Sp.[USNM]; [?]Lagoa santa
[IZWP]; [?]Rio Jan. [MHNP]; [?]Tlupinangu [MHNP].
{\3)Abaris metallica, new species
Fig. 13, 25E
Type Material. — HOLOTYPE. Female, labeled: “Venezuela, Bolivar, Kavanayen, Aug 8 1970;
R.E. Dietz IV, 1000 m.”, USNM. ALLOTYPE. Male, labeled: “Caracas” [handwritten on green paper
other writing is not legible], MNHB.
Type Locality. — As given for holotype.
Range. — Venezuela (Fig. 13).
Recognitory Diagnosis. — Distinctly amariform and so set apart from all other
Abaris species. Shiny virescent green with a proportionally large head and pron-
otum that is broadest at the base.
Description. — Medium sized, overall length 5. 8-6. 3 mm. brilliant green luster. Ventral body surface,
mouthparts, antennae and legs ferruginous to ferrugino-testaceous.
Head. Relatively large, ocular ratio 1.71-1.72. Mentum tooth emarginate at apex. Microsculpture
of head reticulate; frontal impressions shallow and broad, clypeus anterior margin slightly emarginate.
Thorax. Pronotal (Fig. 25E) front angles produced, hind angles obtuse, not denticulate, lateral bead
very narrow and uniform throughout, basolateral margin not bordered although obscure impression
along margin suggests border; microsculpture distinct, reticulate, shiny. Elytra broad, humeral angles
prominent, microsculpture on disc reticulate and surface shiny, basal section of stria 1 present, intervals
flat dorsally, 2-4 very wide; striae 3-5 almost effaced dorsally; elytral humeral tooth anterad interval
8. Mesepisternal angle produced as prominent, sharp tubercle. Metacoxal sulcus arcuate, ended me-
dially in coxa.
Abdomen. Abdominal sulci complete, finely engraved, very shallow medially. Male genitalia and
female genitalia and reproductive tract not studied.
Etymology. — Specific epithet is a Latinized adjective referring to the metallic luster of this species.
Subgenus Abaridius Chaudoir, 1873:97
Type Species. — Abaris tachypoides Bates.
Cladistic Diagnosis. — Support for this clade comes from two homoplasious
characters; form of the pronotal bead (character 15) and the microsculpture in and
near the lateral foveae (character 17). A wide bead that sharply narrows before
the hind angles is characteristic of the basal grade species and a more uniform
bead is characteristic of species in the clade including A. tachypoides and the
remaining species (Fig. 32: character 15). The wider/narrowing lateral bead is
also known from A. napoensis but is hypothesized as a convergent occuiTence.
The second character, microsculpture in and near the lateral foveae, is highly
homoplasious in regard to the selected phylogenetic hypothesis for the genus. The
basal species of Abaridius have a generally shiny surface, and very faint and
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VOL. 71
Fig. 9. — Dorsal habitus, Abaris tachypoides Bates.
transverse microsculpture in this region, \vhereas most derived members have
reticulate and more or less obvious microsculpture.
Two species, A. bicolor and A. nobilis, included in the subgenus Abaridius
lack the more convincing synapomorphies of the A. tachypoides-group (see be-
low) and are considered sedis mutabilis within the subgenus. Both are large and
brilliantly metallic, but otherwise not particularly similar to each other. The form
of the pronotum is quite different between the two, with A. bicolor (Fig. 25F)
more similar to A. inflata (Fig. 26B) or A. nitida (Fig. 25H), whereas A. nobilis
has a pronotum that is unique in the genus. Neither male nor female genitalia
have been studied for either species as each is represented by only a single
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Will — Revision of Abaris
173
Fig. 10. — Locations of collecting sites for Neolalus portai (Straneo) specimens in South America.
specimen. Characteristics of these systems may resolve the relationships of these
taxa.
{\A)Abaris bicolor, new species
Fig. 15, 25F
Type Material. — HOLOTYPE. Male, labeled: “Ecuador, Napo, Onkone Gare Camp, 00°39'10"S,
76°26'00"W, 220 m; terra firma forest, flowerfall, litter, at night; T.L. Erwin, G.E. Ball & D. Shpeley;
5&8.X.1995; 07-95, T.L. Erwin Ecuador Expedition 1995”, USNM.
Type Locality. — As given for holotype.
Range. — Ecuador (Fig. 15).
Recognitory Diagnosis. — Distinguished by its large size, smooth pronotum and
subdepressed form.
Description. — Large sized, overall length 8.5 mm. Very shiny with brilliant metallic luster, head
and pronotum dark virescent, elytra cupreous. Ventral body surface dark brunneous; legs, mouthparts
and antennae slightly paler.
Head. Ocular ratio 1.63. Mentum tooth emarginate at apex. Microsculpture of head faint, reticulate;
frontal impressions sharp, parallel.
Thorax. Pronotum (Pig. 25P) sinuate to base, basolateral region convex, apex broad, front angles
not produced, hind angles denticulate; pronotal basolateral bead broad, sharply narrowed near base;
margins paler than disc; basolateral margin not bordered; pronotal microsculpture mesh pattern trans-
verse, very faint, shiny. Basal section of elytral stria 1 present; mesepisternal angle slightly produced
as small rounded tubercle; elytral humeral tooth anterad interval 8; microsculpture faint, transverse;
intervals slightly convex, intervals 2-3 of equal width. Metacoxal sulcus arcuate, ended medially in
coxa.
Abdomen. Abdominal sulci complete, deeply and finely engraved. Male genitalia and female gen-
italia and reproductive tract not studied.
Etymology. — Specific epithet is a Latin adjective referring to the color contrast between the elytra
and forebody.
{l5)Abaris nobilis, new species
Fig. 15, 25G
Type Material. — HOLOTYPE. Pemale, labeled: “Brazil, Para: 5 km E Belem, Soil-litter layer in
primary “terra firma” forest. June 6:1973, R.T.Schuh”, AMNH.
Type Locality. — As given for holotype.
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VOL. 71
Fig. 11. — Locations of collecting sites for Abaris species specimens in South America.
Range. — Brazil (Fig. 15).
Recognitory Diagnosis. — Distinctive species. Largest member of the genus,
bright coppery color and arcuate pronotal margins readily distinguishes this spe-
cies from all others.
Description. — Large size, overall length 8.9 mm. Cupreous throughout; ventral body surface, labrum
and mandibles piceous; legs antennae and palpi brunneous.
Head. Eyes relatively small, ocular ratio 1.65. Frontal impressions deep, short, divergent. Mentum
tooth deeply emarginate at apex. Microsculpture of head very faint, reticulate.
Thorax. Pronotal (Fig. 25G) basolateral region convex, apex broad, front angles slightly produced,
hind angles denticulate, lateral bead broad narrowed to base, basolateral margin not bordered; pronotal
microsculpture transverse, very faint, surface shiny. Microsculpture of elytra transverse; basal section
of stria 1 present, intervals convex, intervals 2-3 of equal width; humeral tooth anterad interval 8.
Mesepisternal angle produced as rounded tubercle. Metacoxal sulcus arcuate.
Abdomen. Sterna V-VI with complete transverse sulci. Male genitalia and female genitalia and
reproductive track not studied.
Etymology. — The specific epithet is the Latin adjective nohilis (noble) referring to the large size
and brilliant copper color of holotype, making it a noble looking animal.
A. tachypoides-group
Cladistic Diagnosis. — All members of this clade share a derived form of the
metacoxal sulcus. The metacoxal sulcus is straight and appressed to the apical
coxal margin (Fig. 3B, Fig. 32: character 6). The basal grade species, A. nitida,
A. franiai and A. inflata, are relatively large and somewhat or quite convex in
form. They are similar to A. bicolor but have the distinctive straight metacoxal
sulcus, not the arcuate condition of A. bicolor. The male and female reproductive
structures have not been studied for these species. All three species have complete
transverse sternal sulci, as does the more apically placed A. tachypoides. However,
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Will — Revision of Abaris
175
Fig. 12. — Locations of collecting sites in Central America ior Abaris species specimens. A. A. aqui-
lonaria n.sp. B. A. bigenera Bates.
A. tachypoides is much smaller, and has a very different general form than any
of these species.
The mid^grade species, A. tachypoides, A. convexa, A. nigra and A. inaequa-
loides, are small, have rather narrow pronota with constricted apices. In general,
their form somewhat resembles ants or lachnophorine ground beetles.
{\6)Abaris nitida, new species
^ Fig. 13, 25H
Type Material. — HOLOTYPE. Male, labeled; “Ecuador, Napo, Limoncocha, 100 m, H. Frania, Apr
13-24:1979, Swamp Forest, leaf litter”, USNM. ALLOTYPE. Eemale, labeled: “Ecuador, Napo, Ya-
suni Scientific Station, 0°40'36"S, 76°24'2"W, 210 m, K.Will, 24:IV:1998”, CUIC.
Type Locality. — As given for holotype.
Range. — Ecuador (Fig. 13).
Recognitory Diagnosis. — Distinguished from other member of the group by
convex form, deep basal foveae and general form of the pronotum (Fig. 25 H).
Description. — Medium sized, overall length 6.7-7. 1 mm. Shiny bright aeneous elytra; head and
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Annals of Carnegie Museum
VOL. 71
/
pa /
p...-
/ XSt p
'Px ' propic Of Capncom
\ &Y
Abaris picipes: □
Abaris opaca: ^
Abaris metallica: •
Abaris nitida: A
J — — Ei-Q
Fig. 13. — Locations of collecting sites fov Abaris species specimens in South America.
pronotum darker, slightly virescent. Ventral body surface brunneous; mouthparts, antennae and legs
paler, ferrugino-testaceous; femora either concolorous with ventral body surface or paler.
Head. Relatively large, ocular ratio 1.67-1.72. Mentum tooth flat and broad at apex, form simple.
Head smooth throughout, microsculpture obsolete. Frontal impressions moderately deep, slightly di-
vergent, not delimited medially.
Thorax. Pronotum (Fig. 25H) microsculpture faint, transverse; form very convex, narrow at apex,
front angles scarcely produced, lateral bead thick, widened in basal third and narrowed just before
denticulate hind angles. Basal section of elytral stria 1 absent; humeral tooth anterad interval 8;
microsculpture transverse mesh. Mesepisternal tubercle large. Metacoxal sulcus linear, appressed.
Abdomen. Abdominal sulci complete. Male genitalia and female genitalia and reproductive tract
unstudied.
Notes on Life History. — Collected in April at 100-210 m elevation. The Li-
moncocha specimen was collected in a swamp forest from leaf litter and the
Yasuni specimen was collected while using a headlamp and searching leaf litter
along a trail in terra firme forest.
Etymology. — The specific epithet is a Latin adjective referring to the shiny appearance of these
beetles.
(n)Abaris franiai, new species
Fig. 16, 26A
Type Material. — HOLOTYPE. Male, labeled: “Bolivia, Cochabamba, Villa Tunari, For. leaf litter,
430 m. May 9:1979, H. Frania”, USNM.
Type Locality. — As given for holotype.
Range. — Bolivia (Fig. 16).
Recognitory Diagnosis. — Relatively large and robust form with a relatively
broad head. Similar to A. inflata and A. nitida and must be separated from these
by the characters given in the key, e.g., the impunctate pronotal base and sulcate
sterna.
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Will — Revision of Abaris
177
yAtyg-'--"' /
y-'A a I
Epystof
W\
V®
m
W
nrapfe ef Cipfieom
\ fij
fV J
} Abaris aonea:^
•
j ADam oasismata:^
Fig. 14. — Locations of collecting sites for Abaris species specimens in South America.
Description. — Large sized, overall length 7.9 mm. Aeneous, head and pronotum viresceet. Ventral
body surface, labram and mandibles brunneous; legs, palpi and antennae paler.
Head. Ocular ratio 1.89. Meetum tooth form simple. Microsculpture of head faint, reticulate; frontal
impressions short, sharply divergent; clypeus with setigerous punctures surrounded laterally by de-
pressions.
Thorax. Pronotal (Fig. 26A) basolateral region slightly convex, apex broad, front angles slightly
produced, hind angles with small denticle, lateral bead broad and narrowed to base, basolateral margin
not bordered; microsculpture transverse, faint mesh, surface shiny. Microscuipture of elytra transverse;
basal section of stria 1 absent, intervals convex, intervals 2-3 of equal width; elytral humeral tooth
anterad interval 7. Mesepisternal angle produced as small sharp tubercle. Metacoxal sulcus linear,
appressed.
Abdomen. Sternal sulci complete. Male genitalia and female genitalia and reproductive tract un-
studied.
Etymology. — Specific epithet is an eponym based on the surname of the collector H. Frania.
{l^)Abaris inflata, new species
Fig. 16, 26B
Type Material. — HOLOTYPE. Male, labeled: “Bolivia, Cochabamba, Chapare Alto Pol. mar, XI:
1960” [handwritten], MSNM [Straneo Collection],
Type Locality,— given for holotype.
Range.- — -Bolivia (Fig. 16).
Recognitory Diagnosis.— KelatiYely large species, very convex with the pron-
otum broad.
Description. — Large sized, overall length 7.5 mm. Moderately shiny; elytra with aeneous luster,
head and pronotum slightly darker. Ventral body surface, coxae and trochanters brunneous; remaining
leg segments, mouthparts and antennae paler.
Head, Relatively broad, ocular ratio 1.78. Meetum tooth form simple, apex broadly rounded. Mi-
croscuipture of head obsolete, frontal impressions short, not delimited medially. Clypeus tumescent
near frontal suture and depressed near apex.
Thorax. Pronotum (Fig, 26B) convex, front angles scarcely produced, hind angles about right,
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VOL. 71
Fig. 15. — Locations of collecting sites for Abaris species specimens in South America.
denticulate, lateral bead broad in basal third and sharply narrowed to hind angles, without basolateral
border; microsculpture faint and transverse. Elytra basal section of stria 1 absent; mesepisternal angle
prominently produced as sharp tubercle, elytral humeral tooth anterad interval 7; microsculpture trans-
verse mesh, intervals 2-3 of equal width. Metacoxal sulcus linear, appressed.
Abdomen. Male genitalia and female genitalia and reproductive tract unstudied.
Etymology. — The Specific epithet is a feminized Latin adjectival of inflatus (to blow into) referring
to the generally tumescent or robust form of the body.
{\9)Abaris tachypoides Bates, 1871, revised combination
Fig. 9, 15, 21T-U
Abaris tachypoides Bates, 1871:220.
Type Material. — LECTOTYPE (here designated). Male, labeled: “Ega” [handwritten]; “Abaris
tachypoides type. Bates” [handwritten]; “Ex Musaeo H.W. Bates, 1892”; “Lectotype, Abaris tachy-
poides Bates, K.W. Will”, MHNR
Type Locality. — Brazil, Tefe [Ega], from original description.
Range. — Brazil (Fig. 15).
Recognitory Diagnosis. — The relatively large head and narrow pronotum are
distinctive (Fig. 9). Similar to A. inaequaloides, but distinguished from that spe-
cies by the straight striae, equal width of elytral intervals and darker, more cu-
preous body color.
Description. — Small sized, overall length 5. 7-6.4 mm. Cupreous, head and pronotum black with
slight metallic luster. Ventral body surface, mandibles and labrum brunneous to piceous; legs, palpi
and antennae paler ferrugino-testaceous.
Head. Head relatively large, ocular ratio 1.84-1.91. Mentum tooth form simple. Microsculpture of
head obscure, reticulate; frontal impressions long, deep, parallel.
Thorax. Pronotal basolateral region convex, apex narrowly constricted, front angles not produced,
appressed to occiput, hind angles not denticulate, lateral bead narrow, basolateral margin not bordered;
microsculpture on disc reticulate. Elytral microsculpture transverse; basal section of stria 1 absent,
intervals convex, intervals 2-3 of equal width; elytral humeral tooth anterad interval 7. Mesepisternal
angle produced as small, flat tubercle. Metacoxal sulcus linear, appressed.
Abdomen. Sternal sulci complete, very finely engraved. Basal bulb of male median lobe unmodified
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Fig. 16. — Locations of collecting sites fov Abaris species specimens in South America.
(Fig. 21T-U), endophallus folding pattern not visible, blade smooth (Fig. 2 IT). Female genitalia and
reproductive tract not studied.
Material Examined. — Five specimens examined. BRAZIL. Ega [MHNP]; Amazonus [BMNH],
UNKNOWN: [BMNH].
{2Q)Abaris convexa, new species
Fig. 15, 23G-H, 26C
Type Material. — HOLOTYPE. Male, labeled: “Bolivia, Cochabamba, Villa Tunari, 430 m,
H.Frania, inundation for., 9-10:V:1979, rotting flood debris”, USNM.
Type Locality. — As given for holotype.
Range. — Bolivia (Fig. 15).
Recognitory Diagnosis. — A small species dorsal surface somewhat dull, more
convex in form and with relatively larger eyes than other species of similar size.
Description. — Small sized, overall length 5.4 mm.
Head. — Ocular ratio 1.90. Mentum tooth form simple. Microsculpture of head reticulate; frontal
impressions moderately deep, parallel.
Thorax. Pronotal (Fig. 26C) basolateral region convex, apex narrow; front angles not appressed to
occiput, not produced; lateral margins subsinuate, hind angles not denticulate, lateral bead uniformly
thick, basolateral margin not bordered; microsculpture obvious, reticulate mesh, surface dull. Micro-
sculpture of elytra distinct, transverse; basal section of stria 1 absent, mesepisternal angle produced
as small flat tubercle, elytral humeral tooth anterad interval 7. Metacoxal sulcus linear, appressed.
Abdomen. Sterna without sulci. Median lobe (Fig. 23G-H) basal bulb unmodified, endophallus
folding pattern not visible; lobe tip (Fig. 23H) bluntly rounded, symmetrical. Female genitalia and
reproductive tract unstudied.
Etymology. — Specific epithet is a Latin adjective referring to the convex form of this species.
{2l)Abaris nigra, new species
Fig. 16, 20B, 23I~J, 26D
Type Material. — HOLOTYPE. Female, labeled: “Ecuador, Napo Prov., Tena, Misahualli Hotel Jun-
gle Lodge. 30:1:1999. Italo Tapia”, CUIC. ALLOTYPE. Male, same data as holotype, CUIC. PARA-
TYPES: Female, same data as holotype, UASM; Male same data as holotype, QCAZ; Female, labeled:
“Ecuador, Napo, Anaconda Lodge, Napo River, D. Brzoska, 14:11:1991”, SEMC.
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Fig. 17. — Locations of collecting sites for Abaris species specimens in southern North America and
Central America.
Type Locality. — As given for holotype.
Range. — Ecuador (Fig, 16).
Recognitory Diagnosis. — The dark bronze-black color, bicolored legs, and
rounded humeral angles set this species apart from all others. Somewhat similar
to A. inaequaloides, but front angles not tight to occiput as in that species.
Description.—Small sized slightly built, overall length 6. 0-6. 2 mm. Black with bronze luster. Ven-
tral body surface black; legs, except femora, mouthparts and antennae brunneous with darker infus-
cation; femora flavotestaceous, distinctly contrasted with ventral surface, only darker near base.
Head. Ocular ratio 1.74-1.80. Mentum tooth form simple. Microsculpture of head reticulate; frontal
impressions deep, divergent, not clearly delimited medially.
Thorax. Pronotal (Fig. 26D) basolateral region convex, apex narrow, front angles not produced,
lateral margins sinuate, hind angles not denticulate, lateral bead narrow and uniformly thick, basolateral
margin not bordered; microsculpture on disc reticulate, distinct mesh, surface dull. Elytral microsculp-
ture transverse; basal section of stria 1 absent; intervals slightly convex, flatter on disc; intervals 2-3
of equal width, both widened medially; stria 3 sinuate in basal quarter; elytral humeral tooth anterad
interval 7. Mesepisternal angle produced as very small, sharp tubercle. Metacoxal sulcus linear, ap-
pressed.
Abdomen. Sterna without sulci. Median lobe (Fig. 23I-J) basal bulb unmodified, endophallus fold-
ing pattern visible in cleared lobe, blade (Fig. 231) smooth. Female reproductive tract with broad,
straight seminal canal, receptaculum large, distinct, spermatheca with a large basal expansion, ap-
pended gland spherical; gonocoxite-1 with 2 apicolateral ensiform setae.
Notes on Life //Atory.— Specimens have been collected in January and Feb-
ruary. The Misahualli site was sampled in April, May and October as well, and
no A. nigra were found during these months. The habitat in the area of the type
locality is terra firme forest that is mostly secondary growth mixed with small
plots of primary forest.
Material Examined. — Six specimens examined. In addition to the type material, a single male with
the same data as the holotype was preserved in 100% EtOH, KWWC.
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ABC
D E
Fig. 18. — Photographs of dorsal view of Abaris species. A. A. napoensis n.sp. B. A. erwini n.sp. C.
A. aenea Dejean. D. A. picipes Bates. E. A. basistriata Chaudoir.
Etymology. — Specific epithet is a Latin adjective referring to black body color in this species.
(22)Abaris inaequaloides, new species
Fig. 15, 26E
Type Material. — HOLOTYPE. Female, labeled: “Bolivia, Beni, Cercado, 1 km N Ballivian, Rio
Ibare, 14°47'S, 64°59'W, 124 m, 3 Apr 1994, R.Ward”, CMNH.
Type Locality. — As given for holotype.
Range. — Bolivia (Fig. 15).
Recognitory Diagnosis. — Very distinctive species, at once recognized by the
reticulate microsculpture, dull dorsal surface, bronze color and narrow pronotum
relative to the large head.
Description. — Medium sized, overall length 6.3 mm. Head and pronotum dull, elytra moderately
shiny, bronze luster throughout. Ventral body surface, coxae and trochanters brunneous; remaining leg
segments, mouthparts and antennae paler, flavotestaceous.
Head. Relatively large, ocular ratio 1.85. Mentum tooth form simple. Microsculpture of head pro-
nounced, reticulate; surface finely strigate laterad frontal impressions. Frontal impressions short, par-
allel and moderately deep.
Thorax. Pronotum (Fig. 26E) relatively narrow, basolateral region convex, front angles rounded,
not at all produced, hind angles obtuse, not denticulate, lateral bead narrow uniformly thick, laterobasal
margin not bordered; microsculpture reticulate mesh; surface dull. Elytra shiny, microsculpture retic-
ulate mesh; basal section of stria 1 absent, interval 3 width 1.7X interval 2, dorsal punctures foveate,
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striae 4-5 slightly sinuate in basal third; humeral tooth anterad interval 7. Mesepisternal angle slightly
produced, broad and rounded. Metacoxal sulcus linear, appressed.
Abdomen. Sterna without sulci. Male genitalia and female genitalia and reproductive tract unstudied.
Etymology. — Specific epithet is Latinized adjective refering to this species’ general similarity to
species of the B. inaequale-group of the genus Bembidion Latreille, 1802.
(23)Abaris opaca, new species
Fig. 13, 26F
Type Material. — HOLOTYPE. Female, labeled: “Peru, Madre de Dios, Tambopata Res. Zone, Ex-
plorer’s Inn, 12°50'S, 69°17'W, Fig fall, at night, Rfo La Torre to Sunset Point trail, 1 Nov 1982, 290
m, T.L.Erwin and L.Sims”, USNM.
Type Locality. — As given for holotype.
Range. — Peru (Fig. 13).
Recognitory Diagnosis. — Black species with reticulate microsculpture on dor-
sum. Very similar to A. wardi but with the mentum tooth form broad and slightly
emarginate at the apex. Pronotal apex narrow but angles not appressed to occiput
as in A. inaequaloides.
Description. — Medium sized, overall length 6.5 mm. Black with aeneous luster, less pronounced on
head and pronotum. Ventral body surface piceous; legs, mouthparts and antennae brunneous.
Head. Ocular ratio 1.89. Mentum tooth form broad, and emarginate at apex. Microsculpture on head
reticulate mesh; frontal impressions shallow, sharp, slightly divergent, not clearly delimited medially.
Thorax. Pronotal basolateral region convex, apex narrow; hind angles very slightly obtuse to right
angled, not denticulate; lateral bead uniformly thick, basolateral margin not bordered; obvious micro-
sculpture on disc, reticulate; surface dull. Microsculpture of elytra reticulate; basal section of stria 1
absent, intervals slightly convex, interval 3 width L4X interval 2, humeral tooth anterad interval 8.
Mesepisternal angle barely produced, not tuberculate. Metacoxal sulcus linear, appressed.
Abdomen. Sterna without sulci. Male genitalia and female genitalia and reproductive tract unstudied.
Etymology. — Specific epithet is a Latin adjective referring to the dull of the dorsal surface.
{2A)Abaris splendidula (LeConte, 1863)
Fig. 1, 7A-C, 17, 23K-L, 27B
Pterostichus spendidulus LeConte, 1863:10. Abaris splendidula: redescription (Bousquet 1984:384).
Type Material. — HOLOTYPE. Female, labeled: [round label with yellow= western states]; “979”;
“Type 5648”; “Pterostichus splendidulus Lee.”; “J.LeConte collection; Abaris splendidula (LeC.)
Det. 1982, YBousquet”, MCZC. Redescribed by Bousquet (1984).
Type Locality. — Fort Yuma, California, from original description.
Range. — Mexico, southwestern USA (Fig. 17).
Recognitory Diagnosis. — Moderately shiny with a convex form and flat elytral
intervals. No other Abaris species is known from the region inhabited by A.
splendidula.
Description. — Small sized, overall length 5.4-6.5 mm. Black with a cupreous or virescent luster.
Ventral body surface, mandibles and labrum rufopiceous; legs, palpi and antennae ferruginous.
Head. Ocular ratio 1.75-1.86. Mentum tooth form simple. Microsculpture of head reticulate; frontal
impressions deep, sharply delimited laterally, obscurely so medially.
Thorax. Pronotal (Fig. 27B) basolateral region convex, apex broad, front angles not produced,
margins subsinuate, hind angles not denticulate, lateral bead narrow and uniformly thick, basolateral
margin bordered; microsculpture transverse. Microsculpture of elytra reticulate, obvious; basal section
of stria 1 absent, intervals flat, intervals 2-3 of equal width, humeral tooth anterad interval 8. Mese-
pisternal angle produced as a very small round tubercle. Metacoxal sulcus linear, appressed. Hind
wing as in figure 7C.
Abdomen. Sterna without sulci. Pygidial gland system as in figure 7A. Alimentary tract as in figure
7B. Basal bulb of male median lobe (Fig. 23K-L) unmodified, endophallus folding pattern not visible;
blade smooth (Fig. 23K). Female reproductive tract seminal canal narrow with few tight twists along
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Fig. 19. — Abaris species female reproductive tract and genitalic features of A. napoensis n. sp. A.
Ventral view of right gonocoxite. B. Ventral view of reproductive tract and genitalia, right gonocoxite
removed. Same for C. A. notiophiloides Bates. D. A. bigenera Bates. E. A. picipes Bates. Scale for
B, C, D and E at left. Legend: be, bursa copulatrix; co, common oviduct; gel, basal gonocoxite; gc2,
apical gonocoxite; It, laterotergite IX; sg, appended gland of spermatheca; sp, spermatheca.
length, receptaculum distinct, appended gland spherical; gonocoxite- 1 with 1 apicolateral ensiform
seta.
Notes on Life History. — Bousquet (1984) cited comments by G. E. Ball on the
situation in which A. splendidula was collected in Mexico. Specimens were col-
lected “in thin leaf litter, on damp sandy-clay spots in open woodland dominated
by acacias and on the flood plain of a riparian forest.” Two specimens I collected
in Arizona, USA in November were found by raking Salix leaf litter on sandy
soil along the Santa Cruz River. I collected many additional specimens in various
sites in Arizona in July and August in similar situations along washes with or
without water present. Individuals were generally 1-3 meters back from the main
erosion channel in semi-shaded areas with ample leaf litter. Some activity was
observed in late afternoon but most beetles were found at night while searching
with a headlamp. Two specimens collected at light in Tucson were found at large
fluorescent lights at the edge of the city on evenings when it rained either before
or just after collecting the beetles. This suggests some relationship between timing
of dispersal activity and rain patterns. Adults have been collected in January, July,
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Fig. 20. — Abariform species female reproductive tract and genitalic features of A. Abaris aquilonaria
n.sp., ventral view of reproductive tract and genitalia, right gonocoxite removed. Same for B. A. nigra
n.sp. C. A. aequinoctialis Chaudoir. D-E. A. retiaria n.sp. F. Pseudabarys undescribed sp. Ecuador,
both gonocoxites shown. Scale for A, B, and C at upper left. Legend: be, bursa copulatrix; co, common
oviduct; gel, basal gonocoxite; gc2, apical gonocoxite; It, laterotergite IX; sg, appended gland of
spermatheca; sgd, spermathecal gland duct diverticulum; sp, spermatheca.
August and November at elevations from sea level to 1100 m. Data labels include
collections in several similar forest types; dry tropical scrub, gallery, riparian,
riparian tropical scrub forests. Specimens have been taken in leaf litter and rarely
at lights (n = 3).
The male and female collected in Arizona in November were dissected and
reproductive organs examined. Neither specimen was in breeding condition, i.e,
ovaries were not enlarged and male accessory glands were not turgid. Beetles
collected from July to August were kept alive and actively mated producing eggs.
Material Examined. — More than 100 specimens examined. MEXICO. Baja California Sur Notri:
18 km S Loreto [UASM]. Sonora: 17 km sw Moctezuma rte 21[UASM]; 19.3 km n Imuris Rte 15
[UASM]; 55 km sw Moctezuma rte 21 [UASM]; Euerte, 18 km n Los Mochis on rte 15 [UASM]; Sta.
Ana [EMEC],
UNITED STATES. Arizona. Cochise Co.: St. David [UASM]; rt 80 at San Pedro River[KWWC].
Pima Co.: 31 km NW Tucson [CMNH]; Arivaca [UASM]; Florida Cyn.Sta Rita Range Res.[UATU];
Green Valley [AMNH]; Waterman Mtns. [UATU]; Baboquivari Mnts, Brown Canyon [KWWC]; Ari-
vaca Wash, Arivaca [KWWC]; Tucson at lights [KWWC]. Sta. Cruz Co.: [USNM]; Patagonia
[USNM]; Tumacacori at Sta. Cruz River, 31°33'46"N, 111°2'43"W [KWWC].
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{25)Abaris wardi, new species
Fig. 11, 23M-N, 27 A
Type Material. — HOLOTYPE. Male, labeled: “Bolivia, Santa Cruz, Ichilo Province, Buena Vista,
400 m R.Ward, 3:X:1994, MV light”, CMNH. ALLOTYPE. Female, labeled: “Bolivia, Cochabamba
Puerto Villarroel”, UASM. PARATYPES. Female, labeled: “Brazil, Cuyaba, Mtt. Grosso”, MCZC;
Male, labeled: “Bolivia, Sta Helena, W.M.Mann, X, Mulford BioEpl 1921-22”, USNM. Male, labeled:
“Cetto Verde (S.Ama) Bechyne, 16:V:1969”, MSNM. Female, labeled: “Bolivia, Santa Cruz, Province
Andres Ibanez Vallecito, 4-5 Oct 1994, R.D.Ward”, CMNH.
Type Locality. — As given for holotype.
Range. — Bolivia, Brazil (Fig. 11).
Recognitory Diagnosis. — Medium to small size with obvious, reticulate micro-
sculpture. Very similar to A. opaca but separable from that species by the simple
form of the mentum tooth. Pronotal apex narrow but front angles not appressed
to occiput as in A. inaequaloides.
Description. — Medium to small sized, overall length 5. 8-7.0 mm. Black with bronze or cupreous
luster. Ventral body surface piceous; legs, mouthparts and antennae paler brunneous or rufous.
Head. Ocular ratio 1.74-1.82. Mentum tooth form simple. Microsculpture of head reticulate; frontal
impressions long, divergent or sinuate, not clearly delimited medially.
Thorax. Pronotal (Fig. 27A) basolateral region convex, apex narrow, front angles not produced,
hind angles not denticulate, lateral bead narrow, basolateral margin not bordered; microsculpture ob-
vious on disc, reticulate; surface dull. Microsculpture of elytra reticulate; basal section of stria 1 absent,
intervals convex to almost flat, interval 3 width equal to or up to L4X interval 2, elytral humeral
tooth anterad interval 8. Mesepisternal angle rounded, not produced. Metacoxal sulcus linear, ap-
pressed.
Abdomen. Sterna without sulci. Basal bulb of male median lobe (Fig. 23M-N) unmodified, endo-
phallus folding pattern not visible, lobe smooth, apical blade uniform, straight, thin. Female tract with
seminal canal broad, straight, receptaculum not distinct from canal, spermatheca without basal bulb,
appended gland spherical; gonocoxites- 1 with 3 apicolateral ensiform setae.
Etymology. — Specific epithet is an eponym based on the surname of the collector Robert D. Ward.
(26)Abaris aequinoctialis Chaudoir, 1852
Fig. 17, 20C, 230~P, 26G-H
Abaris aequinoctialis Chaudoir, 1852:76. Abarys aequinoctialis: Chaudoir, 1873:98.
Type Material. — LECTOTYPE, (here designated). Female, labeled: “ex Museo Chaudoir” [red ink];
“Lectotype” [Handwritten on circular label with purple edge]; “Lectotype, Abaris aequinoctialis Chd.,
Det. G.E.Ball ’72”, MHNP.
Type Locality. — Temax, Yucatan, Mexico, from original description.
Range. — Mexico, Costa Rica, El Salvador, Guatemala, Honduras, Nicaragua
(Fig. 17).
Recognitory Diagnosis. — Most common species from southern Mexico and
Central America. Similar in form to A. splendidula but separable by the transverse
microsculpture of the elytra. The known range of A. aequinoctialis is not over-
lapping and south of A. splendidula (Fig. 17).
Description. — Small sized, overall length 5. 1-6.5 mm. Dorsal surface shiny, virescent or cupreous
throughout, or bicolored with head and pronotum darker green and elytra dark cupreous. Ventral body
surface rufous; legs and mouthparts paler rufous or ferrugino-testaceous; coxae and femora concolo-
rous with ventral body surface; tibiae and tarsi paler in some individuals, nearly flavescent.
Head. Ocular ratio 1.74-1.88. Mentum tooth form simple. Microsculpture of head faint, reticulate.
Clypeus smooth, fronto-clypeal suture evident, frontal impressions short and not clearly delimited
medially.
Thorax. Pronotum convex, margins sinuate anterad base and convex laterad basolateral foveae, hind
angles slightly obtuse, not denticulate, lateral bead narrow, uniformly thick; microsculpture on disc
transverse (microsculpture faint and surface shiny in most specimens from Guatemala and Yucatan,
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Fig. 21. — Features of male genitalia of Abaris species. A, E, G, I, K, O, Q, S, U, dorsal view blade
of median lobe; D, F, H, N, P, R, T, right lateral view of blade; C, J, right lateral view of median lobe;
B, left lateral view of blade; L, left paramere; M, right paramere. A-C. A. napoensis n.sp. D-E. A.
striolata n.sp. F-G. A. robustula Tschitscherine. H-I. A. aenea Dejean. J-M. A. erwini n.sp. N-O. A.
aquilonaria n.sp. P-Q. A. notiophiloides Bates. R-S. A. impunctata n.sp. T-U. A. tachypoides Bates.
Legend: bs, basal bulb sulcus.
Mexico, obvious mesh in most specimens from other areas). Base of pronotum with or without lateral
border (best developed in Yucatan, Mexico specimens). Basal section of elytral stria 1 absent; position
of humeral tooth variable either anterad interval 7, stria 7, or barely attaining interval 8; microsculpture
transverse; intervals 2-3 subequal.
Abdomen. Sterna without sulci. Blade of median lobe (fig. 230-P) thin, smooth; tip (Eig. 23P)
bluntly rounded, endophallus folding pattern not visible, basal bulb unmodified laterally. Eemale re-
productive tract with seminal canal straight, broad, receptaculum not distinct; appended gland spher-
ical; gonocoxite-1 with 2-3 apicolateral ensiform setae.
Notes on Life History. — Specimens have been collected in all months. Habitat
data includes leaf litter of rainforest, coffee finca and palm forest. Specimens
were collected at sea level to 1400 m elevation. One record was associated with
rotting fruit. Nine specimens were taken at “lights” or Ultraviolet light.
Material Examined. — 239 specimens. COSTA RICA, unknown [USNM]; Guanacaste Prov.; 3 km
N Canas Hac. La Pacifica [UASM]; 5 km N Canas [UASM]; Las Canas [UASM]; Santa Rosa National
Park Administ. & Research Center, 150 m [CASC]; 30 km norte de Liberia, Finca Jenny [INBC]; 3
km N de Nacaome, P.N. Barra Honda [INBC]; 8 km S de Caujiniquil, Est. Murielago [INBC]; Barra
Honda, A.C.Tempisque, Los Mesoes [INBC]; Est. Cacao, Lado suroeste del Vol. Cacao [INBC]; Est.
Las Pailas, P.N. Rincon de la Vieja [INBC]; Est.Lomas Barbudal, A.C.T. [INBC]; Est.Maritza, Lado
oeste del Volcan Orosi [INBC]; Est. Palo Verde, P.N. Palo Verde [INBC]; Est.Sta Rosa, P.N. Guanacaste
[INBC]; Ojochal, P.N.Sta. Rosa [INBC]; P.N. Barra Honda [INBC]; Barcelona. Asemtamiento Juan
Santamaria [INBC]; Playa Naranjo, P.N.Sta. Rosa [INBC]; Ref.Nac.Eauna Silvestre R.L. Rodriguez
[INBC].
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Fig. 22. — Features of male genitalia of Abaris species. B, F, dorsal view blade of median lobe; A, E,
right lateral view of blade; G, ventral view of blade; C, H, right lateral view tip of median lobe and
everted endophallus; D, I, left lateral view tip of median lobe and everted endophallus. A-D. A.
bigenera Bates. E-I. A. basistriata Chaudoir. Legend: als, anterior spine field; bsc, basal scale; fp,
folding pattern of endophallus; 11s, left lateral sacculus; Is, lateral scale; rls, right lateral sacculus; sr,
spine row; vs, ventral spine field.
EL SALVADOR. Isalco [USNM]; Sta. Anna, 6 km W Hwy.CAl, above Lago de Coatepeque,
[USNM].
GUATEMALA. 250 km N San Felipe [BPBM]; Tikal [BPBM, RSCI]; Tikal Rainforest, [UASM];
Zapote [MCZC, MHNP]; El Progreso, 6 km E San Cristobal, Acasaguastlan [USNM]; Peten, Tikal
[USNM, UASM].
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Fig. 23. — Features of Abaris species male genitalia. B, E, H, J, L, N, P, R, dorsal view blade of median
lobe; A, D, G, I, O, right lateral view of blade; C, F, ventral view of blade; K, M, Q, right lateral
view of median lobe. A-C. A. picipes Bates. D-F. A. mina n.sp. G— FI. A. convexa n.sp. I— J. A. nigra
n.sp. K-L. A. splendidula (LeConte). M-N. A. wardi n.sp. O-R A. aequinoctialis Chaudoir. Q-R. A.
retiaria n.sp. Legend: esp, endophallus spine patch; fp, endophallus folding pattern.
HONDURAS. [MHNP]; Honduras, Copan, Ruinas de Copan [UMMZ]; Morazan, Esc. Agr. Pan.
Zamorano [UMMZ]; Francisco, Zamorano, 47°0'N-80°0'W [SEMC].
MEXICO, unknown [MHNP]<— Paz, V.San Yicente Finca, J.Bechyne [MHNP]; 18 mi. N San Andre
V [MHNP]. Chiapas: 32.5 mi. E Comitan, Rtel90 [UASM]; 4.9 mi. N Frontera Comalapa, 727 m;
7.7 mi. N Frontera Comalapa [UASM]; 5.3 mi. S Union Juarez; Cinco Cerros; Cinco Cerros, km30
on Hwyl90 [UASM]; El Aguacero,16 km W Ocozocuautla [UASM]; Trinitaria [UASM]; Puente
Macuilapa nr. Los Amates [CASC]. Hildalgo: 7 mi. NE Jacala [OSUC]. Oaxaca: 5.0 mi. E Tapan-
atepec, rtel90 [UASM]; 7.6 mi. W Zanatepec, rtel90 [UASM]; Rio Miltepec 18.4 mi. W Zanatepec,
rtel90 [UASM]; Hy200 km 194 E Puerto Angel [AMNH]. San Luis Potosi: 1.8 mi. N El Naranjo
[UASM]; El Salto de Agura [CNC]; El Salto de Agura, 7 mi. N El Naranjo [UASM], Tamulipas:
1.4 mi. SE Chamal [UASM]. Yucatan: 12 km N Piste [UASM];2 km E Chichen Itza [UASM];
Chuminopolis [AMNH]; Tehuantepec [AMNH]; Temax [MHNP]; Zopilote, 1.5 mi. E nr.Pan Am. Hyw.
[UASM].
NICARAGUA. Chontales [MHNP].
{21)Abaris retiaria, new species
Fig. 11, 20D-E, 23Q-R, 27C
Type Material. — HOLOTYPE. Male, labeled: “Venezuela, Falcon, Paraguana Pen S Jose Cueva
Piedra Honda, 100 m, S.Peck, 3:111:1971”, USNM. ALLOTYPE. Female, labeled: “Colombia, Mag-
dalena, Rio Frio, RJ. Darlington, 1:XI:1928”, MCZC. PARATYPES. Female, label data same as al-
lotype, MSNM. Male, label data same as allotype, MCZC. 3 females, label data same as allotype
except date Sept, XI and VIII respectively, MCZC. Female, labeled: “Venezuela, Buena Vista Para-
guana, Falcon, Bordon, 2:1:1971”, BORD. Male, labeled: “Venezuela, Maracaibo”, USNM.
Type Locality. — As given for holotype.
Range. — Colombia, Venezuela (Fig. 11).
Recognitory Diagnosis. — Easily recognized from all other species by the ob-
vious reticulate mesh microsculpture, square form and bordered base of the pron-
otum.
Description. — Small sized, overall length 5. 1-5.8 mm. Brunneous with a cupreous or virescent
luster. Ventral body surface, legs and mouthparts paler.
Head. Ocular ratio 1.64-1.75. Mentum tooth form simple. Microsculpture obvious on head, retic-
ulate; frontal impressions sharp, divergent, clypeus finely strigate.
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E
E
m
o
Fig. 24. — ^Pronota of Abaris species. A. A. napoensis n.sp. B. A. striolata. C, A. robustula Tschitsch-
erine. D. A. aenea Dejean, smooth form. E. Same, punctate form. F. A. erwini n.sp. G. A. aquilonaria
n.sp. H. A. notiophiloides Bates. I. A. impunctata n.sp.
Thorax. Pronotal (Fig. 27C) basolateral region slightly convex, apex broad, front angles produced,
lateral margins straight, hind angles obtuse, not denticulate; pronotal lateral bead uniformly thick,
continuous laterally on base; microsculpture obvious, reticulate, dull. Elytral microsculpture reticulate;
basal section of stria 1 absent, intervals little convex, intervals 2-3 of equal width, elytral humeral
tooth anterad interval 8. Metacoxal sulcus linear, appressed to apical margin of coxae. Mesepisternal
angles produced as low tubercle.
Abdomen. Sterna without sulci. Basal bulb of male median lobe (Eig. 23Q) unmodified, endophallus
folding pattern not visible; median lobe straight, smooth; tip (Fig. 23R) evenly rounded. Female tract
with seminal canal broad, straight, receptaculum not discrete, spermatheca with basal bulb, appended
gland unknown (Fig. 20D-E); gonocoxite-1 with 2 apicolateral ensiform setae. (Because all female
specimens were collected into very strong EtOH by EJ. Darlington, female tracts could not be removed
without damage. Therefore, this description is based on the fragments of the attempted dissections
and may require subsequent revision.)
Etymology.- — Specific epithet is a feminized noun in apposition based on the Latin Retiarius (type
of Roman gladiator bearing a net to entangle his adversary). This calls attention to the net-like retic-
ulate mesh of microsculpture on the pronotum.
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Fig. 25. — Pronota of Abaris species. A. A. bigenera Bates. B. A. picipes Bates. C. A. mina n.sp. D.
A. basistriata Chaudoir. E. A. metallica n.sp. F. A. bicolor n.sp. G. A. nobilis n.sp. H. A. nitida n.sp.
Cladistic Analysis
Outgroup Selection
Three genera, Orthomus, Prosopogmus, and Pseudabarys, were used as out-
groups in this analysis. They were selected from among 115 genera examined as
the basis of the analysis of pterostichine taxa (Will, 2000). All three outgroups
plus Neotalus and Abaris comprise a taxon characterized by a “gooseneck”
shaped bursa of the female reproductive tract (e.g.. Fig. 19B, 20F). Orthomus,
Pseudabarys and Abaris share a reduced or absent coronal suture in the larvae
(Bousquet and Liebherr, 1994; Will, 2000). Prosopogmus adults are very similar
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Fig. 26. — Pronota of Abaris species. A. A. frania n.sp. B. A. inflata n.sp. C. A. convexa n.sp. D. A.
nigra n.sp. E. A. inaequaloides n.sp. F. A. opaca n.sp. G. A. aequmoctialis Chaudoir, punctate form,
Chiapas, Mexico. H. Same, smooth and basally bordered form, Guatemala.
in general form to some Abaris species and Pseudabarys shares the paired dorsal
bursal glands with Neotalus and Abaris. Although some of the relationships hy-
pothesized in my higher level analysis (Will, 2000) are not found in the consensus
of all most parsimonious cladograms, the set of relationships {Orthomus {Pro-
sopogmus {Pseudabarys {Neotalus + A^arA)))) is supported under every reso-
lution in the full character analysis. Therefore, rooting between Orthomus and the
remaining taxa is clearly justified.
Character List
1. Maxillary palpomere 3: greater than half length of paplpomere 4 (Fig. 28 A)
(0); less than half length of palpomere 4 (Fig. 28B) (1).
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Fig. 27. — Pronota of Abatis species. A. A. ward! n.sp. B. A. splendidula (LeConte). C. A. retiaria
n.sp.
2. Ocular ratio, measured as width across eyes/width between eyes at level of
anterior supraorbital setae (Fig. 29). Less than 1.55 (0); Greater than 1.60 (1).
Measurements of ocular ratio were taken to determine if a discrete gap existed
between clusters of measurements. Speeies with similarly developed eyes could
then be coded as having the same character state. Measurements of three individ-
uals per species were taken if specimens were available. This ratio was plotted
and a gap between 1.55 and 1.60 was found and used to establish states. Figure
29 shows a plot of the measurements and indicates the gap between states.
3. Mentum tooth apex emarginate (0); simple, not emarginate (1).
Fig. 28. — Scanning electron micrograph, ventral view. A. Right maxillary palpi Lophoglossiis scru-
tator (LeConte) Pterostichini. B. Left maxillary palpi Abatis aetiea Dejean.
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2
1.9
1.8
•2 1.7
m
“^1.6
1.5
1.4
Ocular Ratio
^ ■
M I I I I I I I I I I I I I I I I I I I I I I M I I I I I I'
Ort Neo 3 6 9 10 15 18 21 24 27
Pro 1 4 7 12 11 16 19 22 25
Pse 2 5 8 13 14 17 20 23 26
Species
X ml - m2* m3
Fig. 29. — Graph of ocular ratio measurements showing separation of states for the character. Up to
three measurements were taken for each species. Legend: ml, m2, m3, measurements; Ort, Orthomus;
Pro, Prosopogmus; Pse, Pseudabarys; Neo, Neotalus', Numbers 1-27 correspond to Abaris species
numbers in the text.
4. Reticulate microsculpture on vertex of head. Very faint but visible in clean
specimens (0); obsolete (1).
5. Micropuectulae present on vertex of head (0); absent (1). These punctures
are small (visible at 25 X or greater) and irregularly scattered.
6. Metacoxal sulcus arcuate and divergent from apical margin, ended medially
in coxae or, if short, divergent from margin (Fig. 3 A) (0); sulcus linear, complete
or not, appressed to margin; if incomplete then not divergent from margin (Fig.
3B) (1).
7. Basal section of stria 1 present (0); absent (1).
8. Elytral setigerous lateral umbilicate puncture 2 in line with puncture 1 and
3(0); 2 distant from margin, separated from margin by a distinct convexity and
mesad of 1 and 3 (1).
9. Elytral humeral tooth anterad and in line with interval 8, elytra across hu-
meri broad and humeral angles sharp (0); tooth anterad interval 7, elytra with
rounder and narrower humeral angles (1). Only the position of the tooth was
scored. The general shape is not considered discrete.
10. Microsculpture on elytra reticulate and isodiametric (0); transversely elon-
gate mesh (1).
11. Tarsal claws smooth (0); pectinate (Fig. 2A) (1).
12. Ventral setae of fifth tarsomere subequal in length (0); unequal in length,
apical pair much longer (ca 2X) than subapical pair (1).
13. Pronotal basolateral region outside outer basal foveae convex (0); basola-
teral region flat and reflexed (1).
14. Pronotum front margin broad, front angles produced or not, separate from
occiput (0); pronotum narrow, appressed to occiput, front angles not protruding
(1).
15. Pronotal lateral bead narrow and of uniform width throughout, if narrowing
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at all then narrowed just before hind angles (0); bead narrower in apical third,
then broadened near middle and again narrowed near base, basal taper starts well
before hind angles (1).
16. Pronotal disc microsculpture obsolete (0); transverse and faint, surface
shiny (1); transverse and well developed, microlines clearly visible and surface
slightly duller (2); reticulate and isodiametric mesh regular or somewhat irregular,
surface dull (3) [nonadditive].
17. Microlines in and near basal fovea of pronotum obsolete (0); transverse
and faint, surface shiny (1); transverse and well developed, microlines clearly
visible and slightly duller (2); reticulate and isodiametric or somewhat irregular,
surface dull (3) [nonadditive].
18. Transverse sulci on sterna V-VI complete across sternum (0); laterally
present and medially absent (1); completely absent (2). This character was con-
sidered additive to set adjacency of states. State 1, interrupted medially, is equally
similar to completely absent and entire.
19. Median lobe basal bulb unmodified, small and smoothly rounded (0); basal
bulb enlarged and laterally sulcate (1).
20. Endophallus folding pattern and/or spine fields not visible in cleared lobe
(0); endophallus folding and pattern and/or spine fields visible as darker regions
in lobe (1). Only in a few specimens was the endophallus successfully everted.
However, it is presumed that development of larger spines, lobes and scales, that
can be seen through the median lobe, is a character that may group taxa. It is
likely to be less informative than the multiple characters that potentially exist in
the various structures of an everted endophallus, e.g., A. basistriata (Fig. 22E~I).
21. Blade of median lobe smooth ventrally (0); ventrally strigate (1).
22. Tip of median lobe tapered gradually to apex, not noticeably thin in lateral
view (0); tip very thin in lateral view (1).
23. Apex of median lobe approximately symmetrically narrowed in dorsal view
(0) ; apex distinctly asymmetrically produced to right (Fig. 22D) (1).
24. Blade of median lobe smoothly curved and uniformly tapered in lateral
view, slightly curved in dorsal view (Fig. 21M-R) (0); blade uniformly thin from
bend to tip in lateral view and straight and narrow in dorsal view (Fig. 23Q~R)
(1) .
25. Diverticulum of appended gland present (Fig. 20F) (0); diverticulum absent
(Fig. 19B) (1).
26. Paired bursal glands absent (0); present (Fig. 5) (1).
27. Seminal canal straight and simple or few (<3) loose twists, duct relatively
broad (Fig. 19E) (0); duct narrow with a few (4-5) tight twists along its length
(Fig. 20A) (1); duct narrow with many tight twists along its length (Fig. 19D)
(2). This character was set as additive to establish adjacency of similar states.
28. Receptaculum of spermatheca discrete from duct as an expanded, annulated
in most species, reservoir (Fig. 19D) (0); receptaculum not distinct from duct
(Fig. 20C) (1).
29. Spermatheca without basal bulb (0); base of spermathecal duct expanded
as a bulb (Fig. 20B) (1).
30. Appended spermathecal gland spherical (0); elongate (1).
Results of Cladistic Analysis
Of the 100 search replications (mult* 100), 40 found trees of length 88 steps.
Of these 34 were unique arrangements. Swapping on these trees (max*) found
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1
Orthomus
Prosopogmus
Pseudabarys
Neotalus
basistriata ”
bigenera
impunctata
metallica
robustula
aenea
enwini
aquilonaria
notiophiloides
striolata
napoensis
picipes
mina
bicolor
nobilis
nitida
franiai
inflata
tachypoides
convexa
nigra
inaequaloides
opaca
splendidula
aequinoctialis
wardi
retiaria
Fig. 30. — Consensus cladogram of the 36 MPTs of 88 steps.
two additional trees. A total of 36 most parsimonious cladograms was found (RI
= 71, Cl = 40, length 88 steps). These scores show that a reasonable level of
grouping information is present and homoplasy is within the expected range (de
Queiroz and Wimberger, 1993; Sanderson and Donoghue, 1989) for this matrix
of taxa and characters.
The consensus cladogram (Fig. 30) is largely resolved, with monophyly for all
clades named in the classification supported except the generic separation of Neo-
talus and Abaris and grouping of the members of the A. picipes-group.
In some resolutions Neotalus is placed as sister to Abaridius. This placement
is not preferred as it is supported only by the lack of obvious structure of the
endophallus of the median lobe and would require accepting the hypothesis that
pectinate claws gained in the common ancestor of Abaris were lost in Neotalus
portai. The endophallic structure is not known for most of the Abaridius species,
absent in five species and is present in A. nigra. In other groups of Carabidae,
and particularly in pterostichine clade taxa, the endophallus has been best used
for differentiating between species and is assumed a less suitable system for ge-
neric distinction in abariform taxa.
A number of resolutions have the clade of A. mina + A. picipes placed as sister
to the A. striolata-gvonp. This placement is not preferred as the grouping is based
only on the microsculpture of the head and requires parallel gains of a very
similarly shaped and strigate median lobe in both A. mina + A. picipes and A.
basistriata. Grouping A. mina, A. picipes, and A. basistriata minimizes homoplasy
in the characteristic form of the median lobe. This preferred arrangement also
minimizes the hypothesized number of origins of the long, narrow and tightly
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twisted spermathecal canal. It does require a hypothesis that this peculiar form of
spermatheca reversed to a simpler, straight form in A. picipes.
Little support exists for any particular arrangement of Abaridius species and
thus relationships among them are largely unresolved in the consensus.
The selected cladogram (Fig. 31, 32) shows only clades supported by unam-
biguous character state changes. These groups are supported under both acceler-
ated and delayed transformation optimizations. This arrangement is one of three
that are consistent with my preferred placement of Neotalus and the clade of A.
mina + A. picipes. These three differ only in the arrangement of the species within
the A. tachypoides-group.
Interpretation of the Pattern
Based on the selected arrangement, it is possible to discuss the pattern in terms
of evolutionary process. This involves extrapolation from the observed and sum-
marized data to the likely conditions of the hypothetical ancestors and the char-
acter state transformation.
Character Evolution
Of the 30 characters included in the analysis based on my initial hypothesis of
homology (Patterson, 1982; de Pinna, 1991), 12 are perfectly congruent (ci =
1.0) and can be considered to have passed the test of secondary homology (de
Pinna, 1991) (Fig. 31, 32: characters 1, 2, 6, 8, 11, 12, 19, 21, 22, 25, 26, 30).
Other characters show some homoplasy but still provide grouping information for
some taxa as reversals or convergent origins of states, e.g., the loss of the basal
section of stria 1 (Fig. 31: character 7) and the flat-reflexed baso-lateral margin
of the pronotum (Fig. 31: character 13). States of only two characters (Fig. 31,
32: characters 14, 29) were shown to be entirely parallel occurrences of conditions
not otherwise distinguishable.
The female reproductive tract has several interesting characters that define both
generic and species level relationships. The paired dorsal glands of the bursa (Fig.
5) in species of the abariform genera are similar to those found in cicindelines
(Deuve, 1993) and some platynines (Liebherr and Zimmerman, 1998), but are
unique among all pterostichine taxa. The absence of the diverticulum is a return
to the plesiomorphic condition for the pterostichines (Will, 2000), but is clearly
a synapomorphic loss for Neotalus + Abaris. The form of the spermatheca is at
once synapomorphic at some levels, e.g., for A. aenea-gronp + A. picipes-group
clade, and convergent in pairs of different states. Pairing occurs in the A. striolata-
group [state OJ and (A. aenea-groxxp + A. picipes-group) [2, although reversed in
some]; A. mina [0] and A. picipes [2]; A. aquilonaria [1] and A. notiophiloides
[0]; and A. splendidula [1] and the terminal polytomy including A. aequinoctialis
[0]. Since some of the species pairs are entirely sympatric, these strikingly dif-
ferent forms of the spermatheca may be indicative of differences in oviposition
and/or mating behavior. The ovipositors, however, are almost identical in all spe-
cies. Most of the variation in form of the bursa and spermatheca is found in
species of Abaris s.str., e.g., A. striolata- group and A. notiophiloides. These taxa
also have the greatest modifications of the male genitalia.
The male genitalia are quite simple in most species and with a trend toward a
very simple, straight median lobe and unmodified endophallus in species of the
subgenus Abaridius. Paralleling the modifications in the female reproductive tract,
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Will — Revision of Abaris
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Abaris s.str.
Fig. 31. — Portion of selected cladogram showing outgroup genera, Neotalus and Abaris (s.str.) species.
Unambiguous character state changes are mapped on the cladogram, black discs — non homoplasious
change, white — homoplasious change. Numbers above branches are character numbers, below are
character state.
species of Abaris s.str. have smaller parameres, a relatively larger basal bulb of
the median lobe, variously modified median lobe tip, and a spinose and lobed
endophallus in some species.
Microsculpturing shows no clear transformation series and is highly homopla-
sious. From the cladogram one can only generalize that transverse (either faint,
obvious or irregular) is in most clades plesiomorphic, and all the other states are
frequently derived from it.
Biogeographical Overview
Plant and animal distributions in tropical America have been the data source
for many studies on ecological and historical biogeography (see Whitmore and
Prance, 1987). Carabid beetles are well represented in these studies (Shpeley and
Ball, 1993; Noonan, 1985 and included references). The distribution of Abaris
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Fig. 32. — Portion of selected cladogram showing Abaris {Abaridius) species. Unambiguous character
state changes are mapped on the cladogram, black discs — non homoplasious change, white — homo-
plasious change. Numbers above branches are character numbers, below are character state.
species includes the entire breadth of the New World tropics and so investigation
of their pattern of distribution adds to our general understanding of the bioge-
ography of animals in the New World.
Areas of Endemism
The “zoogeographic zones” presented by Shpeley and Ball (1993) for species
of Coptodera Dejean, 1825 (Fig. 33, Table 1) are used in the biogeographic
analysis and discussion of the distribution of Neotalus and Abaris below. The
areas identified for Coptodera species, in general, correspond to the distribution
of Abaris species and contain more or less identifying sets of taxa. Therefore it
is reasonable to use these areas for cladistic biogeographic analysis from which
descriptive patterns can be discussed.
In addition to the similar ranges, both Abaris and Coptodera have species that
are found at fruit falls in tropical forest, many individuals are more or less me-
tallic, and many have large eyes and pectinate claws. These similarities suggest
an overlap in behavior or life histories. Coptodera species, however, are clearly
associated with tree trunks, limbs and logs (Shpeley and Ball, 1993); whereas
Abaris species are only known as ground dwellers. Adults of both genera pre-
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Fig. 33. — Zoogeographic zones in southern North America, the West Indies, Central and South Amer-
ica for Abaris and Coptodera species. From Shpeley and Ball (1993). See text for explanation of
zones.
sumably fly; however, the more arboreal habits and the relatively more numerous
records of flight for Coptodera species suggest its members fly more frequently.
Descriptive Aspects of Distributions
Refer to distribution maps of species for the following discussion (Fig, 10-17)
and to table 1 and figure 33 for zones. Because of the paucity of material and
limited number of collecting sites, the analysis of the biogeographical patterns for
species of Neotalus and Abaris must assume the material in hand represents the
ranges of species, and therefore the areas assigned to them in the analysis. Several
species that have been sampled more adequately have large ranges, and in many
cases are sympatric with one or more other species. The collection of more ma-
terial, especially from the vast and under-collected areas in Brazil, may corrob-
orate or call for revision of hypotheses presented here.
Most Abaris species (13 spp.) live entirely or in part in tropical lowland forests
corresponding to zone C or the drier forests south of the Amazon Basin, zone B
(11 spp.). Only four species, all members of the A. picipes-group, have ranges
that extend southward beyond the tropics. Abaris splendidula is the only species
that ranges north of the tropics. The range of N. portai also extends south of the
tropics.
Members of the A. aenea-group and A. picipes-group show a marked diver-
gence in habitat breadth. The A. aenea-group species are relatively stenotopic and
found in moist tropical lowland forest from Bolivia to southern Mexico at ele-
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Table 1. — Zoogeographic zones from Shpeley and Ball (1993) used as areas for biogeographical
analysis for Abaris and Coptodera species.
Zone Area
A Southern Atlantic South America. Atlantic forest
B cis-Andean South America south of Amazon Basin from eastern slope of
Andes west of zone A
C Amazon Basin
D cis-Andean South America north of the Amazon Basin and west of zone E
E cis-Andean South America north of the Amazon Basin including northern
Brazil and the Guianas
F Choco refugium in trans-Andean Colombia and Ecuador
G Lower Central America north to southern Nicaragua
H Nuclear Central America from Nicaragua to the Isthmus of Tehuantepec in
southern Mexico
I Mexico north of the Isthmus of Tehuantepec, Pacific versant
J Mexico north of the Isthmus of Tehuantepec, Gulf versant
K West Indies
L Southwest United States and northern Mexico
M Eastern United States
vations of 100-970 m, with most specimens collected from between 200-400 m.
In contrast, members of the A. picipes-group are centered in drier forests south
of the Amazon Basin (zone B), are generally more widespread, and apparently
eury topic. The widespread A. picipes-group species have been collected at ele-
vations of 100-1680 m (or as high as 3600 m, see discussion under A. basistriata
above). In the A. aenea- group, A. aenea ranges into lower Central America and
A. aquilonaria is found through nuclear Central America and into southern Mex-
ico at elevations of 200-970 m. Similarly A. bigenera and A. (Abaridius) ae~
quinoctialis are found through nuclear Central America and into southern Mexico.
The former, like other A. picipes-group species, has been collected at a variety of
elevations (200-1680 m) and in several habitat types. Conversely the latter, like
the A. aenea-group species, has been found only at elevations less than 850 m.
Most Abaridius species are known from zones B and C; only A. aequinoctialis
and A. splendidula occur outside of South America. Abaris splendidula lives in
drier and cooler habitats and, like the species of the A. picipes-group, can be
found across a wide range of elevations (0-1100 m).
Biogeographic Analysis
Biogeography is a synthetic field that draws a constellation of data from other
disciplines such as systematics, geology and ecology. Historical biogeography
(Rosen, 1978:160) specifically attempts to reconstruct the origin of taxa and biotas
based on the pattern of relationships found through the systematic study of taxa
and the resulting implied genealogy. If each terminal taxon lived exclusively in
one area then, the area relationships and historical pattern could be read by re-
placement of the taxa with their areas on the cladogram. However, distributional
patterns are complicated by widespread taxa and redundant areas, i.e., sympatric
species distributions (Page, 1990; Nelson, 1984).
Several methods have been proposed to reduce complex taxon-area cladograms
to a fundamental area cladogram (Nelson and Platnick, 1981) in which areas occur
only once. Component analyses include various conversion rules that make as-
sumptions regarding missing areas, widespread taxa and redundant distributions
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(Page, 1990)« The methods include Assumption 0 (= Brooks parsimony analysis
(BPA), Wiley, 1987); Assumptions 1, 2 (Nelson and Platnick, 1981), and three-
area statements (TAS) (Nelson, 1991).
Assumption 0 (BPA) has been criticized as restrictive in its treatment of wide-
spread taxa (Page, 1990: 124). Assumptions 1—2 are increasingly more realistic but
because of memory handling limitations of the only available computer program,
Component L5 (Page, 1989), the large number of hypotheses that can result from
complex data under these assumptions makes analyses under these assumptions
impossible to complete. TAS and three-taxon analysis (3TA) methods and impli-
cations have come under extensive criticism (Farris and Kluge, 1998).
An alternative parsimony criterion to the component based analyses is recon-
ciled tree analysis or tree mapping (Page, 1990, 1994; Nelson and Platnick, 1981)
(referred by some authors as a component analysis, e.g,, Morrone and Carpenter,
1994:101). Although no criteria have been presented that allow for determining
a single best method for biogeographical analysis (Morrone and Carpenter, 1994:
111), reconciled tree analysis as implemented in Component 2.0 is interpretable
and practical means for analyzing biogeographical data and is used herein.
Reconciled tree analysis finds the best fitting pattern(s) of area relationships
for the observed taxon-area cladogram by inserting terms (leaves or components
(Page, 1993)) into a user-tree until the observed pattern is reconciled with the
presumed area relationships. Three measures of fit for the reconciled tree that
have been proposed are (Page, 1994), 1. duplications, 2. leaves added, and 3.
number of losses.
“Real” duplication events manifest as redundant nodes (Page, 1994:65) and
are explained as primitive sympatry or secondary dispersal events subsequently
followed by vicariance (Liebherr and Zimmerman, 1998:157). Minimizing dupli-
cations results in a set of trees that generally includes trees produced by mini-
mizing “leaves added”, and so is equal to or larger than that set. The number of
leaves added is often reported as items of error (lOE), to measure fit between two
trees (Nelson and Platnick, 1981). lOE is readily calculated by doubling the num-
ber of leaves added to the observed tree. The measure is easily derived but bio-
logical interpretation of this measure is unclear (Page, 1994).
In regard to the reconciled tree, losses are supposed ancestral species that did
not disperse into an area or became extinct before a vicariance event resulted in
two areas. Thus, absence of potential descendant taxa is accounted for by a single
event in the ancestor rather than an event for each leaf and component. This is
most clearly thought of in a strictly cospeciatioeal situation when a parasite is
lost from a host through extinction. Counting leaves or using lOE may overes-
timate the number of such events (Page, 1988) by preferring the addition of leaves
to reduce duplications. When a heuristic search is conducted for best fitting user-
trees for a set of areas where dispersal is deemed likely, minimization of dupli-
cations and leaves is appropriate. Such is the case in the Hawaiian Islands (Lieb-
herr and Zimmerman, 1998). In such a system, no preference for ancient versus
recent extinction events is defensible.
Focusing on duplications and leaves can increase the number of losses. In a
hypothetical example shown in figure 34, the single duplication of the user-tree
in 34B requires 3 losses and 10 lOE (5 leaves added) to account for the observed
occurrence of area II and its sister-area relationship with area III. Conversely
figure 34A has the same number of leaves added (5 leaves = 10 lOE) as 34B,
but the paraphyletic relationship of II Till requires one less loss than in 34B. In
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rx ^1 rX-
Fig. 34. — Hypothetical components from two reconciled trees for areas 1-VI. Both represent a single
duplication of the respective user-tree requiring 5 added leaves (dashed lines). Area II represents the
occurrence of the area in the observed tree. Losses are mark with an “X.” A. 2 losses. B. 3 losses.
the user-tree 34A, only 2 events (primitive losses or failure of the ancestor to
disperse) are required.
In a system where vicariance and not dispersal is held to be the primary type
of event, minimizing losses is a more reasonable parsimony criterion. By this
method, more tree duplications are acceptable if they result in fewer losses (ex-
tinction events) during the history of diversification. This is reasonable for the
large continental areas occupied by Abaris and Coptodera species. Using this
criterion does not preclude dispersal as an explanation for parts of the resulting
pattern that include duplications. However, these may be best explained as prim-
itive sympatry and subsequent vicariance.
Fig. 35. — Taxon-area cladogram for Neotalus and Abaris species. See text and figure 33 for explanation
of zones.
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Will — Revision of Abaris
203
By looking at the areas relative to their relationships on the preferred cladogram
(taxon-area cladogram Fig. 35), it is likely that the present distribution is the
result of 4-5 separate dispersal/speciation events into or in Central America. Three
pieces of evidence suggest that these dispersals are relatively recent, probably
occurring no earlier than the early Pliocene when the Central American land
bridge was formed (Stehli and Webb, 1985): 1, few Abaris species are present in
Central America; 2. no species are found in the West Indies; and 3. all are ap-
parently fully winged. This contrasts with other groups whose ancestors are pre-
sumed to have undergone earlier emigration from South America. These groups
have greater numbers of Central American and Mexican species (e.g., Coptodera
(Shpeley and Ball, 1993), Pseudabarys (Will, unpubl.)); species in the West Indies
(e.g., Coptodera, Dyschromus (Erwin, 1979)); and/or species with brachypterous
flight wings in some Central American species (e.g., Dyschromus, Pseudabarys).
When no preferred geological hypothesis is available, a heuristic search for a
best fitting set of area relationships, a fundamental area cladogram, can be con-
ducted using Component 2.0 (Page, 1993, 1994). Options used in heuristic search
and reconciled tree analyses of Coptodera and Neotalus + Abaris included the
following: 1. absence was treated as missing data, 2. widespread associates were
mapped, 3. branch swapping used was nearest- neighbor interchange, 4. optimality
criterion used was minimize losses.
Since the preferred cladogram contains unresolved nodes. Component neces-
sarily resolves them arbitrarily before running the analysis. Allowing these arbi-
trarily resolved cladograms could permit the use of resolutions that are suboptimal
in length. Therefore, a fully resolved cladogram (Fig. 36A) was constructed that
was of length 88 steps and contained nearest geographic neighbors. The phylo-
genetic hypothesis for Coptodera also contained unresolved nodes (Shpeley and
Ball, 1993, their figure 36). These polytomies were resolved following the species
pairs found in the cladistic analysis of Liebherr (presented in appendix B of
Shpeley and Ball, 1993; their figure 41A-B). Taxa and area cladograms were
submitted to Component and a heuristic search conducted as described above.
For the Neotalus + Abaris species, 196 rearrangements produced 14 area clado-
grams that equally minimized losses (47 losses, 24 duplications, 164 leaves
added), and for Coptodera species, 66 rearrangements produced 3 area cladograms
that equally minimized losses (99 losses, 36 duplications, 380 leaves added) (Fig.
37A-C). The two sets of area relationships were compared by using Component’s
quartet dissimilarity measure. Two of the 14 Neotalus + Abaris species clado-
grams were found to be more similar to the three Coptodera species cladograms
than the others (Fig. 37B-C). These two cladograms were used to develop the
general historical relationships of the areas and the hypotheses for the process
leading to the current distribution of Abaris. The strict consensus was also found
for each set of data (Fig. 36D, 37D).
Based on the consensus cladograms for the areas. Neotalus + Abaris diversified
in South America first with vicariance of the Atlantic forest (zone A), followed
by isolation of the Amazon Basin (zone C), and finally separation of the cis-
Andean region (zone B). The latter may have remained in contact with zone D
by way of habitats along the lower-mid elevation corridor west of the Amazon
Basin. This hypothesized corridor would have species distributions like those seen
today from Bolivia, Peru, Ecuador, and Colombia, e.g., A. erwini and A. aenea
(Fig. 11, 14). Plausibly, the common ancestor of Neotalus + Abaris originated in
southern South America and species were restricted there until the Pliocene. This
204
Annals of Carnegie Museum
VOL. 71
A
” Neotalus
_r napoensis
^ striolata
robustula
erwini
aenea
aquilonaria
notiophiioides
““ impunctata
r” bigenera
r basistriata
I ^ metaica
y- mina
^ picipes
nobiiis
bicolor
nitida
franiai
infiata
tachypoides
nigra
convexa
inaequaloides
splendidula
aequinoctiaiis
retiaria
Fig. 36. — Cladograms for Neotalus + Abaris species and resulting relationships from biogeographical
analyses. A, Fully resolved cladogram submitted to Component. B, C. Two of the 14 resulting fun-
damental area cladograms that were most compatible with the fundamental area cladogram derived
from the Coptodera analysis. D. Consensus of the 14 fundamental area cladograms.
pattern and timing is similar to the hypothesis presented for ground beetles of the
subgenus Anisotarsus Chaudoir, 1837 (Notiobia Perty, 1830) by Noonan (1981).
However, Abaris would probably have been found in the subtropical and tropical
habitats, as it is today, and not in the warm temperate habitats favorable to Anb
so tars us.
The consensus cladogram (Fig. 36D) is not resolved for any of the Central
American zones. This is a result of having only a few, primarily widespread
species in these areas. If the two cladograms that are maximally similar to the
results based on Coptodera. are considered (Fig. 36B-C), the Central American
and southern Mexican zones have separated sequentially from south to north.
Unlike the results from the analysis of the Coptodera data, zones D, northern cis-
Andean and zone L, northern Mexico + southwestern United States, are shown
to have separated from the Central American zones. This suggest additional taxa
may remain to be sampled in Central America that would provide evidence for
area relationships similar to those resulting from the analysis of Coptodera species
data.
By comparison, the pattern resulting from analysis of the Coptodera species
shows a general pattern involving an early divergence of the northern and southern
regions and then parallel differentiation proceeding within both sets of areas (Fig.
37A-D). Again, this is consistent with a group reaching proto-Middle America
and North America earlier and more frequently than observed for Abaris. Such
an early divergence (late Cretaceous or early tertiary) and frequent dispersals (10
events south to north) was proposed for Coptodera species by Shpeley and Ball
(1993:162, 167).
2002
Will — Revision of Abaris
205
Fig. 37.— Resulting cladograms from the biogeographical analysis of Coptodera species. A, B, C.
Three possible area relationships produced. D. Consensus cladogram of the three cladograms above.
It is surprising, given the apparent dispersal abilities of Abaris species (full
flight wings and broad distributions of some species) that none have been found
west of the Andes in the Choco region (zone F) or on any island in the Carribean
(zone K). Other carabid groups found in tropical lowland habitats, such as pla-
tynines, have established and diversified on all the Antilles (Liebherr, 1997). Pla-
tynus Bonelli, 1810 is species rich in the islands, Glyptolenus Bates, 1878 some-
what less diverse (Liebherr, 1997) and Abaris and Pseudabarys are completely
absent, even from the island of Trinidad. Since Abaris is apparently no less vagile
than these platynine taxa (some abariform taxa have been collected at lights and
are known to fly). The difference may be related to biotic factors (likely for the
islands of the West Indies) or inadequate collecting (probable for western South
America).
Relative to apparently older elements (e.g., Dyschromus), the hypothesis that
Abaris species simply did not reach northern South America until relatively re-
cently may be sufficient explanation for their absence from the West Indies.
Conclusions
The monophyletic Neotalus + Abaris is revised. After necessary nomenclatural
changes and the description of 17 newly recognized species (nearly tripling the
species number of Abaris), the species of Neotalus and Abaris are now available
to students of carabids and tropical biology. It is hoped that the ability to identify
species will lead to more extensive study of their habits and perhaps studies that
lead to a functional explanation for structures such as the paired bursal glands
and pectinate claws.
The remaining abariform genus Pseudabarys stands, like Abaris stood, with
few named species and many recognizable unnamed forms. Once these species
are treated to the level of Abaris, a broader synthesis will be possible and hope-
fully another level of understanding reached in Neotropical pterostichine ground
beetles.
206
Annals of Carnegie Museum
VOL. 71
Acknowledgments
This project, which was part of my dissertation project at Cornell University, could not have been
accomplished without the assistance of many people whose contributions, small or large, have been
graciously accepted. For continuous support I thank Jim Liebherr and my other excellent and sup-
portive committee members Quentin Wheeler and Melissa Luckow. I thank David Maddison, Uni-
versity of Arizona for support during preparation of the manuscript for publication. Numerous collec-
tion managers and carabidologists as listed in the materials section provided me with specimens and
assistance along the way. Three individuals deserve special thanks. George Ball (UASM) made the
monograph of Abaris possible by taking type specimens on loan from MHNP, providing copious notes
regarding the specimens and reviewing the manuscript with great care. Bob Davidson (CMNH) was
always a source of knowledge, specimens, solid review of my manuscript and not the least an excellent
host. Yves Bousquet (CNCI) kindly provided valuable information on carabids in general and was
particularly generous in allowing me to see data in his unpublished manuscripts and notes. He also
provided a sound review of the manuscript. My successful fieldwork in Ecuador could not have been
so, except for the assistance provided by Giovanni Onore (QCAZ), a man of great heart and benefactor
of entomologists from throughout the world. It was Giovanni who wisely assigned Italo Tapia to assist
me during my stay in Ecuador. Italo’s impressive fortitude, sharp wit and willingness greatly enhanced
my time in the field. At Cornell, Bryan Danforth and Tom Eisner both substantially assisted me with
time and materials. An earlier version of the manuscript received an exceptionally thorough review
by John Rawlins (CMNH). His efforts greatly improved this paper and his sage advice benefitted me
personally. The project received generous financial support from National Science Foundation Doctoral
Dissertation Improvement grant (DEB-9700764); A.C. Rawlins Endowment, Cornell University; Ernst
Mayr research grant. Harvard University; G. Griswold Fund, Cornell University; and Palmer Assis-
tantship, Cornell University.
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5
Orthomus
00010
Prosopogmus
00000
Pseudabarys
01001
Neotalus
11001
aenea
11010
aequinoctialis
11101
aquilonaria
11001
basistriata
11001
bicolor
11001
bigenera
11001
convexa
11101
erwini
11001
franiai
11101
impunctata
11101
inaequaloides
11101
inflata
mil
metallica
11001
mina
11011
napoensis
11110
nigra
11101
nitida
mil
nobilis
11001
notiophiloides
11001
opaca
11001
picipes
11010
retiaria
11101
robustula
1100*
splendidula
11101
striolata
11011
tachypoides
11101
Appendix 1
Characters 1-
-30
10
1
15
1
00000
00000
030
00000
00100
220
00101
01000
120
01101
01000
220
01101
moo
022
111*1
11000
222
01101
moo
222
00100
11000
332
00101
11001
110
00101
11000
331
mil
11000
332
01101
moo
122
mil
11001
110
00100
11000
331
11110
11010
312
mil
11001
130
00100
11000
330
00101
11000
001
00101
11001
111
mil
11000
332
11101
11001
110
00101
11001
110
01101
moo
222
moo
11000
332
00101
11000
00 (1
moo
11000
332
00101
moo
121
moo
11000
232
00101
11000
221
mil
11010
330
20
1
25
1
30
1
00
00000
00000
01
00000
00000
01
00000
10000
00
00011
10000
11
00001
12000
00
00011
10100
11
00001
11000
11
10101
12000
? ?
9 9 9 9 9
9 9 9 9 9
11
00101
12000
00
0000?
9 9 9 9 9
11
00001
12000
? ?
? ? ? ? 9
? ? ? ? ?
11
0000?
? ? ? ? ?
? ?
9 9 9 9 9
9 9 9 9 9
? ?
9 9 9 9 9
9 9 9 9 9
? ?
9 9 9 9 9
? ? ? ? ?
11
10001
12000
11
01001
10101
01
00001
10010
? ?
9 9 9 9 9
? ? ? ? ?
? ?
9 9 9 9 9
9 9 9 9 9
11
00001
10100
? ?
? ? ? ? ?
9 9 9 9 9
2) 11
10101
10000
00
00011
10?1?
11
00001
12000
00
00001
11000
11
01001
10101
00
0000?
9 9 9 9 9
210
Annals of Carnegie Museum
VOL. 71
wardi 11101 11100 11000 332 00 00011 10100
State unknown = ?, Polymorphic =
Appendix 2
The Nexus File Submitted to Component 2.0 for Neotalus + Abaris
#NEXUS
BEGIN TAXA;
DIMENSIONS NTAX=9;
TAXLABELS
A
B
C
D
G
H
I
J
L
ENDBLOCK;
BEGIN DISTRIBUTION;
TITLE = ‘BLOCK L;
NTAX=28;
RANGE
nap
3,
str
3,
rob
2 3,
erw
3,
aen
3 4 5,
aqu
5 6 8,
not
3,
imp
1,
big
6 7 8,
bas
1 2 3 4,
met
4,
min
1 2,
pic
1 2 3,
nob
3,
bic
3,
nit
3,
fra
2,
inf
2,
tac
3,
nig
3,
con
2,
ine
2,
spl
9,
aeq
5 6 7 8,
2002
Will — Revision of Abaris
21 1
ret
: 4,
opa
: 2,
war
: 2,
Neo
: 1 2
TREE T1==(28,(((1,2),((3,(4,(5,(6,7)))),(8,(9,((10,11),(12,13)))))),(14,(15,
(16X17,(18,(19,(20,(21,(22,(23,(24,(25,(26,27)))))))))))))));
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BEGIN TREES;
[!>Heuristic search settings:
> Trees fitted to block BLOCK 1
> Absence is treated as missing data
> Widespread associates mapped
> Nearest neigbor interchange (NNI) branch swapping performed
> Total number of rearrangments tried =196
> Criterion minimised = number of losses
> Trees found = 14
> Minimal value = 45
]
TRANSLATE
1 A,
2 B,
3 C,
4 D,
5 G,
6 H,
7 I,
8 J,
9 L
TREE T1 =(1,(3, (2,(4, (5,(9, (8, (6,7))))))));
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TREE T6=(l,(3,(2,(4,(9,(5,(8,(6,7))))))));
TREE T7 = (l,(3,(2,(4,((5,9),(7,(6,8)))))));
TREE T8 = (l,(3,(2,(4,(9,(5,(7,(6,8))))))));
TREE T9 = (l,(3,(2,(4,((5,(6,8)),(7,9))))));
TREE T10=(l,(3,(2,(4,((5,9),(6,(7,8)))))));
TREE Tll = (l,(3,(2,(4,(9,(5,(6,(7,8))))))));
TREE T12=(l,(3,(2,((4,9),(5,(8,(6,7)))))));
TREE T13 = (l,(3,(2,((4,9),(5,(7,(6,8)))))));
TREE T14=(l,(3,(2,((4,9),(5,(6,(7,8)))))));
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The Nexus File Submitted to Component 2.0 for Coptodera
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BEGIN TAXA;
212
Annals of Carnegie Museum
VOL. 71
DIMENSIONS NTAX=I3;
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3 4 7 8,
rel
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7 8 10,
nvir
8 10,
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1 2 3,
meg
3,
ruf
1 5,
cham
6 7,
dial
3,
pic
1 2 3 4 5 7 8 10 11 13,
fov
10,
poe
7,
aen
3 5,
aer
13,
vir
9,
pak
3,
way
3,
acu
1 3 4 5 6 7 8 10,
apic
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sto
7,
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3 7 8 9 10 11 13,
drom
1 2 3 5,
nit
1 2 3 5 6 7 8 9 10 12,
brun
: 8 10 12,
2002
Will — Revision of Abaris
213
erw
3,
ema
1 2 3 5,
nig
3 5,
teut
1,
braz
1,
aur
8 10,
dep
1 2 4,
cup
1 2 7 8 10,
sig
1,
squ
1,
tran
1 2 3 4 7 8 9 10,
ful
1,
bif
1,
xan
8 9,
tri
1 3;
TREE T1=((1,((7,((5,10),(12,23))),(42,(19,34)))),((2,35),(((26,(3,(40,(31,(4,
24))))), (39, (37, 38))), ((9, ((16, (6, 20)), (25, (15, (8, (11, 32)))))), ((17,
((13,(22,30)),((33,36),(41,43)))),(14,((21,(18,29)),(27,28))))))));
ENDBLOCK;
BEGIN TREES;
[!>Heuristic search settings:
> Trees fitted to block BLOCK 1
> Absence is treated as missing data
> Widespread associates mapped
> Nearest neigbor interchange (NNI) branch swapping performed
> Total number of rearrangments tried = 66
> Criterion minimised = number of losses
> Trees found = 3
> Minimal value = 99
]
TRANSLATE
1 A,
2 B,
3 C,
4 D,
5 E,
6 F,
7 G,
8 H,
9 I,
10 J,
11 K,
12 L,
13 M
TREE Tl=((3,(l,(5,(2,4)))),((7,(9,(6,(12,(ll,13))))),(8,10)));
TREE T2=((3,(l,(5,(2,4)))),((7,((6,12),(9,(ll,13)))),(8,10)));
TREE T3 = ((3, (1,(5, (2, 4)))), ((7,(9, ((6, 12), (11, 13)))), (8, 10)));
ENDBLOCK;
ANNALS OF CARNEGIE MUSEUM
VoL. 71, Number 3, Pp. 215-232 16 August 2002
FROM THE ARCHIVES AND COLLECTIONS
W. J. HOLLAND’S ROLES IN THE 1902 MEETINGS OF THE AMERICAN
ASSOCIATION FOR THE ADVANCEMENT OF SCIENCE AND THE
AMERICAN ANTHROPOLOGICAL ASSOCIATION
David R. Watters ^
The Pittsburgh Venue
One hundred years ago, Pittsburgh was the host city for the 5 1 st Annual Meet-
ing of the American Association for the Advancement of Science (AAAS), from
June 28 to July 3, during which the American Anthropological Association (AAA)
held its founding meeting on June 30. The meetings actually convened in the
Oakland district of the city, although each organization consistently listed Pitts-
burgh as the venue for its meeting (Howard, 1902; MacDougal, 1902a; McGee,
1902c; Smith, 1902). In 1890 the city’s name had begun to be spelled Pittsburg,
the “h” having been dropped following a decision by the U.S. Geographic Names
Board (rescinded in 1911), but in practice both spellings were still commonly
used in 1902 (e.g., MacDougal, 1902a; McGee, 1902c).
William Jacob Holland (Fig. 1) deserves credit both for attracting the AAAS
meeting to Pittsburgh and selecting Oakland as the venue for its activities. Doc-
uments in the Holland Archives at Carnegie Museum of Natural History provide
information on his efforts to promote the city as the convention site, the assis-
tance he enlisted to bring this meeting to fruition, and the obstacles he overcame
to do so. His achievement is even more impressive when one realizes that Hol-
land had become the Director of Carnegie Museum only four years before the
AAAS meeting, had served concurrently as the Chancellor of the Western Uni-
versity of Pennsylvania (now the University of Pittsburgh) for three of those
years, until 1901, and was simultaneously involved in intensive efforts to re-
organize the museum, acquire collections, institute fieldwork, and plan the mu-
seum’s expansion in the enlarged Carnegie Institute facility that opened in 1907
(Watters, 2002).
Holland achieved his goal of bringing the AAAS meeting to Pittsburgh because
of his prior involvement with AAAS, his personal contacts with important persons
in the city, his long-term involvement with the Oakland district, his own orga-
nizational skills, the support given by the Museum Committee, and the assistance
provided by local corporations, scientific associations, and Pittsburgh’s social elite.
His ability to orchestrate the overall organization of the 5 1 st AAAS annual meet-
ing derived from his position as Chairman of the local Executive Committee, one
of nine local planning committees established to ensure the success of the venture
(Table 1).
The purpose of this paper is to present the results of research about the venue
of the founding meeting of the American Anthropological Association within the
context of the AAAS meeting and the activities of its Section H (Anthropology).
' Curator, Section of Anthropology
215
216
Annals of Carnegie Museum
VOL. 71
Fig. 1 — William Jacob Holland, Director Emeritus of Carnegie Museum, c. 1930. (Holland Archives,
Carnegie Museum of Natural History).
That Pittsburgh was the location for the founding of AAA ties directly to W. J.
Holland’s desire to have the city host the 51st Annual Meeting of the AAAS. The
research project makes use of unpublished documents in the Holland Archives
and published data available in the Proceedings of the AAAS 51st Annual Meet-
ing (Howard, 1902) and assorted articles.
Procuring and Planning the AAAS Meeting
Holland wanted to host the AAAS 50th Annual Meeting in 1901, but he had
to settle for the next year. He first broached this idea to the members of the
Museum Committee in June 1900, a mere month after informing them that he
would the accept the Carnegie Museum directorship on a fulltime basis and would
tender his resignation as Chancellor of the Western University of Pennsylvania,
as soon as a suitable replacement could be found (which happened in 1901).
Holland’s rationale for proposing Pittsburgh is preserved:
2002
Watters — From the Archives and Collections
217
Table 1. — Local committees and supporting organizations for the 1902 AAAS Meeting.
Local Committees and Chairs for the Pittsburgh Meeting
Honorary President — George Westinghouse, Jr.
Executive Committee — W. J. Holland
Finance Committee — John B. Jackson
Committee for the Reception and Entertainment of Visiting Ladies — Miss Julia Morgan Harding
Reception and Entertainment Committee — W. N. Frew
Committee on Accommodations, Hotels, etc. — R. D. McGonnigle
Committee on Room, Halls, and Equipment — C. R. Cunningham
Committee on Excursions — Charles F. Scott
Committee on Transportation — Samuel Moody
Committee on Press and Printing — Rev. P. A. McDermott
Supporting Organizations for the Pittsburgh Meeting
The Trustees of the Carnegie Institute
The Chamber of Commerce of the City of Pittsburgh
The Trustees and Faculty of the Western University of Pennsylvania
The Faculty of the Pittsburgh College of the Holy Ghost
The Engineers' Society of Western Pennsylvania
The Academy of Science & Art of Pittsburgh
The Western Pennsylvania Botanical Society
The Twentieth Century Club
The Pittsburgh Chapter of the Daughters of the American Revolution
The Dolly Madison Chapter of the Sons and Daughters of the War of 1812
The University Club and several other organizations had “announced their intention”
Sources: Howard, L. O. (ed.), 1902, Proceedings of the 51st AAAS Meeting (pp. 8-9); Monthly Report
of the Director [for February] to the Museum Committee, March 3, 1902, pp. 4-5.
INVITATION TO AMERICAN ASSOCIATION FOR THE
ADVANCEMENT OF SCIENCE
The Director stated that the American Association for the Advance-
ment of Science, which is now about fifty years old, had never held a
meeting in Pittsburgh, for the reason, he believed, that no formal invi-
tation had ever been pressed upon the body. He thought that Pittsburgh
would be a most interesting place for the next meeting of the Associ-
ation, and he recommended that an invitation to meet at the Carnegie
Institute next year be extended, and that a committee of three be ap-
pointed for that purpose. Whereupon the following resolution was
adopted:
Resolved: That a committee of three, consisting of Dr. W. J. Holland,
Chairman, C. C. Mellor and John A. Brashear be appointed with in-
structions to extend an invitation to the American Association for the
Advancemicnt of Science to hold its next annual meeting in the year
1901 in the city of Pittsburgh.
Resolved, further, That this committee be instructed to secure, in the
extending of this invitation, the assistance and cooperation of the mu-
nicipal authorities, the trustees of the Carnegie Institute, and the various
institutions of learning and learned societies of the cities of Pittsburgh
and Allegheny.
Resolved, further, That for this purpose the committee be given full
218
Annals of Carnegie Museum
VOL. 71
power to act. (Carnegie Museum, Minutes of the Museum Committee
[for July 16, 1900], voL II, January 3, 1898 to April 1, 1905).
Mellor was the Chairman and Brashear a member of the Museum Committee, so
their willingness to appoint themselves to the AAAS invitation committee ensured
the close cooperation desired by Holland, as its Chair. Their involvement likewise
enabled and facilitated his contact with important local persons who would play
key roles in planning an AAAS meeting. Samuel H. Church, the only member of
the Museum Committee who did not serve on the invitation committee, became
involved with the AAAS meeting in other ways. Pittsburgh and Allegheny (now
Pittsburgh’s North Side) then were separate cities.
Brashear and Holland had close ties to AAAS. Brashear, a Life Member, joined
AAAS in 1884 at its 33rd meeting, became a Fellow in 1885, belonged to three
scientific Sections, and in 1900 served as Vice-President of Section D (Mechanical
Science and Engineering) at the New York meeting (Howard, 1902:7, 20, 53).
Holland joined in 1888 at the 37th meeting, became a Fellow in 1896, and be-
longed to Section F (Zoology) (Howard, 1902:92). Mellor had been a member of
AAAS since 1889 at the 38th meeting, but he did not join a specific scientific
Section (Howard, 1902:1 14). Church became a member at the Pittsburgh meeting,
joining two Sections (Howard, 1902:60).
The Minutes of the Museum Committee contain nothing more about the hoped
for 1901 AAAS meeting (it went to Denver), and there are only two entries
concerning the 1902 meeting. That Pittsburgh had been selected is evident in the
Minutes from November 30, 1901, which contain a resolution on the need for a
special meeting “. . . to discuss matters relating to the coming meeting of the
American Association for the Advancement of Science.” The February 3, 1902
Minutes direct Holland to look into organizing a “. . . committee for the enter-
tainment of the American Association for the Advancement of Science. ...”
There is another set of documents that provides better background on Holland’s
efforts to organize the 1902 AAAS meeting. These efforts are outlined in his
Monthly Reports of the Director to the Museum Committee, with the reports of
April 30 and November 30, 1901, and February 1, March 3, April 30, May 31,
and June 30, 1902, pertaining to the AAAS meeting. The April 30, 1901 Monthly
Report mentions his effort to “. . . increase the [AAAS] membership in Pittsburgh
... in view of the fact that that distinguished body is to hold its meeting in 1902
in this city ...” which confirms that Pittsburgh by then had been selected as the
location. The November 30 Monthly Report references a letter received from Dr.
Charles S. Minot, AAAS President, asking that '‘. . . makers of scientific apparatus
...” be invited to exhibit at the meeting, and the report also includes an appeal
by Holland to the Museum Committee stating that the preliminary arrangements
“. . . should be taken up by us at once and with energy.”
Holland’s comments in the Monthly Report for February 1, 1902 are especially
interesting in terms of identifying the range of activities involved in preparing for
the meeting, for which “. . . not another moment ought to be allowed to elapse.”
He says the department of paleontology’s entire force was “. . . busily engaged
in extracting material from the matrix and in preparing the same for exhibition
in view of the meeting.” He had received valuable information from Denver, the
site of the preceding AAAS meeting, and mentions a “. . . gentleman who might
perhaps be secured to act as a press agent.” He had secured use of the Bellefield
Presbyterian Church and the Oakland Methodist Episcopal Church buildings, ob-
2002
Watters — From the Archives and Collections
219
tained options on the Music Hall and small lecture room of Carnegie Institute,
and notified the proprietor of the Hotel Schenley that his hotel would be the
headquarters. Holland had to . . take these steps promptly . . to forestall use
of the buildings by the State Teachers Association of Pennsylvania, which planned
to meet in Pittsburgh at the same time as the A A AS.
The March 3 Monthly Report lists ten local groups which already had appointed
arrangement committees for the A A AS meeting (Table 1). One might well have
empathy for Holland who was to . convene these committees to meet in the
Lecture Room of the institution on Friday next at three o’clock.” However, events
were “. . . progressing favorably . . .” by the time he wrote his April 30 Monthly
Report, to the extent that he had turned his attention to raising “. . . a considerable
sum of money ...” with the expectation that “. . . a generous response will be
rendered by our fellow-citizens to the appeal that is about to be made to them
for financial assistance.”
The Museum Committee was informed in Holland’s May 31 Monthly Report
only that “. . . we have been exceedingly busy.” On June 30 he reported . .
arrangements are well nigh satisfactorily completed” (one would hope so since
the AAAS opening general session was held that morning), and he announced
that almost “nine thousand dollars has been contributed by the citizens towards
the entertainment of the Association. ...” It is of more than passing interest that
there was no Monthly Report for July, and in his August 30 report Holland says
“After an absence of between three and four weeks I returned to the Museum on
August the 11th.” Even the indomitable Holland deserved a respite.
Oakland Facilities and AAAS Activities
The facilities Holland reserved initially, the Bellefield Presbyterian Church, the
Oakland Methodist Episcopal Church, and Carnegie Institute, had insufficient
space for the numerous meetings planned by AAAS and its Affiliated Societies.
He expanded the available venues by securing the use of the Eirst United Pres-
byterian Church, the Bellefield School House, and the Botanical Lecture Hall of
Phipps Conservatory, the latter located in nearby Schenley Park (Anonymous,
1902:802). All facilities were within five minutes’ walk of the Hotel Schenley,
AAAS headquarters (Pig. 2). Carnegie Institute’s Music Hall was the venue for
general sessions of AAAS, whereas individual Sections and Affiliated Societies
held their sessions in the other buildings.
Carnegie Institute, of which Carnegie Museum was one component, was built
on the edge of Schenley Park in 1895, one of the first cultural institutions con-
structed in the developing Oakland district. Holland had close ties to this area
long before he assumed the directorship of Carnegie Museum. He had lived in
Oakland ever since becoming the pastor of the Bellefield Presbyterian Church,
where he served for seventeen years (1874-1891), before becoming Chancellor
of the Western University of Pennsylvania, and his home, the former manse ad-
jacent to that church, was but a brief walk from Carnegie Museum. Use of meeting
rooms in the Oakland churches by the AAAS was due, in large part, to Holland
being a member of the Bellefield Presbyterian Church and to his personal ties to
the pastors and trustees of the other churches.
The scientific sessions were the principal reason for attending the AAAS annual
meeting. However, members also could avail themselves of excursions from Oak-
land to many of the manufacturing establishments in the region, which appear to
220
Annals of Carnegie Museum
VOL. 71
Fig. 2 — The Oakland district of Pittsburgh, c. 1905. Carnegie Institute is the building on the left (with
the two towers later removed during the 1907 expansion), Hotel Schenley is in the right foreground,
and Phipps Conservatory is the domed structure (beyond the hotel) in Schenley Park in the back-
ground. (Holland Archives, Carnegie Museum of Natural History).
be the reason the 51st annual meeting was especially well attended by persons
representing physics, chemistry, and mechanics and engineering (MacDougal,
1902^:41). The excursions and social activities are documented in Pittsburgh
newspapers (e.g.. The Pittsburg Press, July 1 edition, p, 11) but are mentioned
only in passing in the 51st AAAS Proceedings. These extracurricular activities
were well received:
The arrangements for the excursions and for the general entertainment of
the members were on a larger scale than anything attempted at recent
meetings of the Association, the local committee having collected and at
its disposal a fund of $9,000.00 for this purpose. (MacDougal, 1902<2:4).
A general schedule for daily activities was outlined in a report in Science about
five weeks before the meeting:
It is expected that the Council will meet daily at 9 a.m., and that the
usual brief general session will assemble at 10:00 a.m., the meetings of
the scientific sections following, with a brief interruption for lunches,
until 4 o’clock p.m. (Anonymous, 1902:802).
The American Association for the Advancement of Science
The American Association for the Advancement of Science, at the time it met
in Pittsburgh, included ten scientific Sections (A-I and K) and numerous Affiliated
Societies (Table 2). A President headed AAAS; Vice-Presidents chaired the Sec-
tions. The AAAS Council was comprised of the Past Presidents, Vice-Presidents
from the two previous meetings (in New York and Denver), Officers of the Pitts-
2002
Watters — From the Archives and Collections
221
Table 2. — Scientific Sections and Affiliated Societies of A A AS.
Scientific Sections at the Pittsburgh Meeting
Section A.
Section B.
Section C.
Section D.
Section E.
Section F.
Section G.
Section H.
Section I.
Section J.
Section K.
Mathematics and Astronomy
Physics
Chemistry
Mechanical Science and Engineering
Geology and Geography
Zoology
Botany
Anthropology
Social and Economic Science
[None listed]
Physiology and Experimental Medicine
Affiliated Societies at the Pittsburgh Meeting
American Anthropological Association (affiliated in Pittsburgh)
American Chemical Society
American Folk-Lore Society
American Microscopical Society
American Psychological Association
American Physical Society
American Society of Naturalists
Association of Economic Entomologists
Botanical Society of America
Geological Society of America
National Geographic Society (affiliated in Pittsburgh)
Society for the Promotion of Agricultural Science
Society for the Promotion of Engineering Education
Sources: Howard, L. O. (ed.), 1902, Proceedings of the 51st AAAS Meeting (pp. 3-4, 7); Anonymous,
1902, Science, 15:803; and MacDougal, D. T., 1902a, Science, 16:43.
burgh Meeting (the incoming Vice-Presidents and Secretaries of the Sections),
Members from the Association at Large, and Members from the Affiliated Soci-
eties (Howard, 1902:7).
The AAAS met as a body in general sessions, of which the first was held in
the Music Hall at 10:00 a.m., Monday, June 30, when retiring President Charles
S. Minot relinquished the position to President-Elect Asaph Hall. W. J. Holland,
Chair of the local Executive Committee then welcomed members on behalf of
Pittsburgh and Allegheny, followed by remarks by Samuel H. Church, Secretary
of Carnegie Institute, and George H. Anderson. Their speeches are included in
the Proceedings of the 51st Annual Meeting (MacDougal, 1902Z?:55 1-558). The
Music Hall was the location of three other general sessions, for speeches presented
by Leonard P. Kinnicutt, Minot, and Robert T Hill in the evenings of June 30
and July 1 and 3, respectively (MacDougal, 1902(3:44). The Pittsburgh Gazette
(July 2 edition) reported that more than 1,000 persons attended Minot’s speech.
According to a report in The Pittsburg Press (July 2 edition), the entire Museum
Committee — W. J. Holland, C. C. Mellor, S. H. Church, and J. A. Brashear —
stood in the receiving line at Minot’s reception.
MacDougal (1902(3:41-42), AAAS General Secretary, reported 431 members
in attendance and 320 papers and addresses read before scientific Sections and
Affiliated Societies, with special lectures and evening sessions raising total pre-
sentations to nearly 350. MacDougal (1902(3:47) lists by name the 73 members
222
Annals of Carnegie Museum
VOL. 71
elected AAAS fellows at the Pittsburgh meeting (oddly, MacDougal, 1902Z?:569,
says eighty, but provides no names). Smith (1902:201-202) identified nine new
fellows as being . . interested in anthropology . . Livingston Farrand, Wil-
liam C. Mills, Charles L. Owen, A. E. Jenks, A. H. Thompson, J. D. McGuire,
Frank W. Blackmar, William Wallace Tooker, and William Henry Goodyear.
George Westinghouse, Jr., the Honorary President of the Pittsburgh Meeting
(Table 1), was elected an Honorary Fellow, only the ninth person so honored by
AAAS (Howard, 1902:40-41; MacDougal, 1902:47). Holland emphasized that
Westinghouse was deserving of this great honor because of his “. . . knowledge
of physics and of mechanics as sciences ...” (1902:502), not merely for having
been the Honorary President. Holland was honored by colleagues in Section F
(Zoology), who elected him to the AAAS Council as their Member at Large
(Howard, 1902:7, 426). The AAAS selected Washington, D.C., as the venue for
its winter meeting six months later, from December 29, 1902, to January 3, 1903.
AAAS Section H — Anthropology
Anthropology as a discipline was first represented in AAAS in 1875 at the 24th
meeting, when the Subsection of Anthropology was organized under then Section
B, the Section of Natural History. Anthropology was assigned to Section H in
1882, at the 31st meeting when AAAS reorganized into nine sections (Howard,
1902:18, 21). Frederick Ward Putnam was the key individual promoting this dis-
cipline within AAAS, and he did so during a quarter-century-tenure (1873-1897)
as its Permanent Secretary (Kohlstedt, 1999:39-49). Section H had existed for
twenty years at the time of the Pittsburgh meeting.
The activities of Section H at the 5 1 st AAAS Annual Meeting are well reported
by Harlan 1. Smith (1902), the Section’s Secretary. The officers, standing com-
mittees, and resolutions (Table 3) are extracted from Smith’s report, augmented
by additional sources. J. Walter Fewkes, the retiring Vice-President, was suc-
ceeded by Steward Gulin; Smith replaced George Grant MacCurdy, the retiring
Secretary. When and where the transition occurred are documented:
Section H was organized, in the audience room of the Bellefield [Pres-
byterian] Church, on Monday morning, June 30, immediately after the
adjournment of the general session, and, with two later mentioned ex-
ceptions, held all of its sessions in the same place. (Smith, 1902: 201).
The general session to which Smith refers was the first session, convened at 10:00
a.m. in the Carnegie Institute Music Hall, when Asaph succeeded Minot as AAAS
President and where Holland, Church, and Anderson welcomed the members.
Therefore, Section H’s organizing meeting would have occurred in the late morn-
ing, probably around 1 1 :00 a.m. on June 30.
The retiring Vice-President traditionally gave a keynote address to the scientific
section he had chaired. J. Walter Fewkes spoke on “Prehistoric Porto Rico” to
Section H on Monday afternoon, June 30, at the Oakland Methodist Episcopal
Church, referred to by Smith (1902:202) as the Oakland church. The Pittsburgh
Gazette (July 1 edition) confirms that Fewkes had given his address the day before
at the Oakland Methodist Episcopal Church. Fewkes’ (1902<3, 1902Z?) address was
published in Science and the 51st Proceedings.
The next two days, July 1 and 2, were devoted to Section H scientific sessions.
The Bellefield Presbyterian Church (Fig. 3) was the main venue for anthropology
papers, based on Smith’s statement that Section H had “. . . held all of its sessions
2002
Watters — From the Archives and Collections
223
Table 3. — AAAS Section H (Anthropology) at Pittsburgh.
Officers of Section H
Vice-President: J. Walter Fewkes, 1901 (retiring); Stewart Culin, 1902 (incoming); George A. Dor-
sey, 1903 (elected)
Secretary: George Grant MacCurdy, 1901 (retiring); Harlan I. Smith, 1902 (incoming); Roland H.
Dixon, 1903 (elected)
Member of Council: W J McGee (incoming)
Sectional Committee: Culin and Smith; ex-officio, Fewkes and MacCurdy; at large, Franz Boas, G.
A. Dorsey, and W. H. Holmes
Member of the General Committee: Walter Hough
Standing Committees of Section H
Committee on Anthropometric Measurements: J. McK. Cattell, W. W. Newell, W J McGee, and
Franz Boas
Committee on the Teaching of Anthropology in America: W J McGee (Chairman), G. G. Mac-
Curdy, Frank Russell, Franz Boas, and W. H. Holmes
Committee on the Protection and Preservation of Objects of Archaeological Interest: Chairman
(open due to the death of Thomas Wilson), F W. Putnam, N. H. Winchell, G. K. Gilbert, A. W.
Butler, and G. A. Dorsey
Resolutions
On the American International Archaeological Commission: Stewart Culin, Chairman; Harlan L
Smith, Secretary
On the death of Dr. Thomas Wilson: Warren K. Moorehead, Stewart Culin, Harlan 1. Smith, and J.
Walter Fewkes
On the American Anthropological Association: Franz Boas
Sources: Howard, L. O. (ed.), 1902, Proceedings of the 51st AAAS Meeting (pp. 5-14); MacDougal,
D. T, 1902n, Science, 16:45-46; Smith, H. I., 1902, Science, 16:201-202; McGee, W J, 1902c,
American Anthropologist, 4:467-471.
Fig. 3 — The Bellefield Presbyterian Church, c. 1900. (Courtesy of the Carnegie Library of Pittsburgh,
Pennsylvania Department).
224
Annals of Carnegie Museum
VOL. 71
in the same place” (1902:210), with only two exceptions. The first was Fewkes’
speech on June 30 at the Oakland Methodist Episcopal Church. The second ex-
ception was a brief change of venue from the Bellefield Presbyterian Church to
the Bellefield School, during the Tuesday, July 1 afternoon session, for two papers
requiring facilities for lantern-slides (Table 4).
A total of 30 papers on anthropological topics were presented at the scientific
sessions on July 1 and 2 (MacDougal, 1902<3:42). The order of presentation and
the venues in Table 4 are derived from Smith’s (1902:202-212) report, which
also abstracts salient points from most papers and mentions persons discussing
certain papers. Four papers were read by title only. The Section H sessions on
July 2 were held jointly with the American Folk-Lore Society in the morning and
the American Anthropological Association in the afternoon. Howard (1902:513-
516) lists the same 30 titles, but his ordering differs somewhat from Smith’s (cf.
McGee, 1902c:47 1-481 for other comments on the papers). Smith (1902:210)
and McGee (1902c:477) acknowledge the role played by W. J. Holland in ar-
ranging for Carnegie Museum’s loan of a sculpture, exhibited in conjunction with
Marshall H. Saville’s lecture on “A Rare Form of Sculpture from Eastern Mex-
ico” on July 2 (Table 4), This sculpture probably was one of the casts of Mexican
sculptures recently donated by Andrew Carnegie (Watters, 2002).
The American Anthropological Association
The founding meeting of the American Anthropological Association was held
in the Oakland district of Pittsburgh at 2:00 p.m. on Monday, June 30, 1902, the
same day as the opening general session of the 51st Annual Meeting of the
American Association for the Advancement of Science. The AAA founding meet-
ing’s time, day, and date and the reason it happened in Pittsburgh are known.
What has been uncertain for a century is the meeting’s venue; that is, the building
in which the AAA founding meeting convened — either the Bellefield Church or
the Oakland Church.
Two pre-founding meeting reports by McGee announced the day, date, hour,
and place; his post-meeting report corroborated the day, date, and hour but dif-
fered in place. McGee (1902a:353; 1902Z?:1035) beforehand said the venue would
be “. . . the audience room of Bellefield Church ...” but afterwards stated the
. . meeting was held in Oakland Church . . (McGee, 1902c:464). He omitted
the denominations of the churches in all three reports.
Smith (1902:201) confirmed that the AAA founding meeting had occurred but
did not specify its date or location. Smith wrote this report in his capacity as
Secretary of Section H, and it therefore logically focuses on Section H activities.
Nevertheless, the report is a valuable document with respect to the AAA, because
it clarified one confusing point and provided a lead that was key to determining
the venue. Smith (1902:201) places the Section H organizing meeting, at which
its new officers were installed (Table 3), at the Bellefield Church in the morning
of June 30, and this meeting most likely began around 11:00 a.m., since it fol-
lowed the AAAS opening general session at 10:00 a.m. Smith’s record allows us
to discount the notion that the Section H meeting and the AAA founding meeting
were held jointly. Even though they happened on the same day and involved
many of the same anthropologists. Section H’s organizing meeting was in the
morning and AAA’s founding meeting in the afternoon.
Smith (1902:202) names the . . Oakland church . . .” as the location of Vice-
2002
Watters — From the Archives and Collections
225
Table 4. — Anthropological papers presented in Pittsburgh.
Retiring Vice-President’s Speech to Section H
Monday afternoon, June 30 (at Oakland Methodist Episcopal Church)
Fewkes, J. Walter, “Prehistoric Porto Rico”
Section H Scientific Sessions
Tuesday, July I, morning session (at the Bellefield Presbyterian Church)
Mills, William C., “The Human Effigy Pipe, taken from the Adena Mound, Ross Co., Ohio”
Moorehead, Warren K., “Gravel Kame Burials in Ohio”
Mills, William C., “Microscopical Sections of Flint from Flint Ridge, Licking Co., Ohio”
Hitchcock, C. H., “The Hernandes Shell-heap, Ormond, Florida”
Moorehead, Warren K., “The Late Dr. Thomas Wilson”
Dorsey, George A., “An Osage Mourning — War Ceremony”
Tuesday, July 1, afternoon session (at the Bellefield Presbyterian Church)
Wright, G. Frederick, “Anthropological Museums in Central Asia”
Wright, G. Frederick, “Climatic Changes in Central Asia traced to their Probable Causes, and
discussed with Reference to their Bearing upon the Early Migrations of Mankind”
McGee, W J, “Mortuary Ceremonies of the Cocopa Indians”
Tuesday, July 1, afternoon session (at the Bellefield School where the session adjourned to avail itself
of lantern facilities for the next two papers)
MacCurdy, George Grant, “A Collection of Crania from Gazelle Peninsula, New Pomerania”
Mills, William C., “Burials of Adena Mound”
Wednesday, July 2, morning session (at the Bellefield Presbyterian Church; joint meeting with Amer-
ican Folk-Lore Society)
Houghton, Frederick, “Cooperation Between the Anthropological Museum and the Public School”
Smith, Lee H., “Uses of Archaeological museums in Education in the Public Schools” (read by
title only)
Hough, Walter, “Explorations of 1901 in Arizona”
Pepper, George H., “The Throwing-stick of Prehistoric People of the Southwest”
Wednesday, July 2, afternoon session (at the Bellefield Presbyterian Church; joint meeting with Amer-
ican Anthropological Association)
Fewkes, J. Walter, “A War Festival of the Hopi Indians”
Saville, Marshall H., “A Rare Form of Sculpture from Eastern Mexico”
Surface, H. A., “The Possible Origin of the Folk-Lore about Various Animals” (read by title only)
McGee, W J, “The Place of Anthropology among the Sciences”
Culin, Stewart, “Anthropological Museums and Museum Economy”
Holmes, William H. “Classification and Arrangement of the Collections of an Anthropological
Museum”
Smith, Harlan L, “Methods of Collecting Anthropological Material”
Hough, Walter, “The Preservation of Museum Specimens”
Fowler, J. A., “The Australian Native” (read by title only)
Boas, F, “The Growth of Children” (read by title only)
Moseley, E. L., “Charcoal Covered by Stalagmite in Put-in-Bay”
Moseley, E. L., “The Sandusky Engraved Slates”
Smith, Harlan I., “Exhibition of a Modern Clay Tablet from Michigan”
Leon, Nicolas, “Square Occipital in the Cranium of a Modern Othomi Mestizo”
Wardle, Harriett Newell, “Evanescent Congenital Pigmentation in the Sacro-Lumbar Region”
Sources: Smith, H. L, 1902, Science, 16:202-212; Howard, L. O., 1902, Proceedings of the 51st AAAS
Meeting (pp. 513-516); McGee, W J, 1902c, American Anthropologist, 4:471-481. [Note: the order
of papers, including those read by title only, and the venues are derived from Smith; Howard’s list
verified the titles but his ordering differed; McGee aggregates papers by related topics, not chrono-
logically; Howard lists journals in which some of the papers were published.]
President Fewkes’ address to Section H on Monday afternoon, June 30. He was
the only writer to locate Fewkes’ speech to a specific building. Moreover, Smith
emphasizes the point that the speech at the Oakland church was one of two
exceptions to the normal meeting place, the Bellefield Church, for Section H
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Annals of Carnegie Museum
VOL. 71
Fig. 4 — The Oakland Methodist Episcopal Church one year after it hosted the founding meeting of
the American Anthropological Association (from the Chronicle Telegraph, October 16, 1903 edition).
(Courtesy of the Carnegie Library of Pittsburgh, Pennsylvania Department).
scientific sessions (the other exception was the two lantern-slide talks given on
July 1 at Belleheld School). However, Smith does not mention the time when
Fewkes gave his address. Smith omitted the denominations of the churches, as
had McGee.
A third source, the authorship of which has been attributed to McGee or George
A. Dorsey (Stocking, 1960:1 and fn. 2), contains the most detailed account of the
events surrounding the founding meeting in Pittsburgh (Anonymous, 1903:184-
186). It reiterates the “Oakland Church” as the location for the 2:00 p.m. meeting
and adds that Culin, the newly installed Vice-President of Section H, served as
the founding meeting’s Chair and Dorsey as its Secretary. This source also lists
the thirteen participants: Franz Boas, Stewart Culin, George A. Dorsey, J. Walter
Fewkes, J. N. B. Hewitt, William H. Holmes, Walter Hough, George Grant
MacCurdy, W J McGee, Joseph D. McGuire, F. W. Putnam, Marshall H. Saville,
and Harlan I. Smith.
The evidence better supports the conclusion that the founding meeting of the
American Anthropological Association convened in the Oakland Methodist Epis-
copal Church (Fig. 4) at 2:00 p.m. on Monday, June 30, 1902 (Watters and Fowler,
2002). Three statements support this view. McGee (1902c:464), the newly elected
AAA president, named the Oakland Church as the venue, in contradiction to his
pre-meeting announcements listing the Belleheld Church. The anonymous author
(1903:184) lists the Oakland Church and places Fewkes at the founding meeting
before he gave his speech. Smith (1902:202) locates the Section H anthropologists
in the Oakland church (a venue exceptional to the norm) on that Monday after-
noon to hear Fewkes deliver his address. The sequence of events that afternoon
was the AAA founding meeting at 2:00 p.m. followed by Fewkes’ speech some-
time later, with both events taking place in the Oakland Methodist Episcopal
2002
Watters — From the Archives and Collections
227
Table 5. — American Anthropological Association (founding meeting) at Pittsburgh.
Officers of AAA
President: W J McGee
Vice-President (four years): F. W. Putnam
Vice-President (three years): Franz Boas
Vice-President (two years): W. H. Holmes
Vice-President (one year): J. W. Powell
Secretary: George A. Dorsey
Treasurer: Roland B. Dixon
Editor: F. W. Hodge
Council of AAA
Frank Baker, Charles P. Bowditch, A. F. Chamberlain, Steward Culin, Livingston Farrand, J. Walter
Fewkes, Alice B. Fletcher, J. N. B. Hewitt, Walter Hough, Ales Hrdlicka, A. L. Kroeber, George Grant
MacCurdy, O. T Mason, Washington Matthews, J. D. McGuire, James Mooney, W. W. Newell, Frank
Russell, M. H. Saville, Harlan 1. Smith, Frederick Starr, John R. S wanton, Cyrus Thomas, and E. S.
Wood
Executive Committee
Boas, Culin, Dorsey, Fewkes, Hodge, Holmes, MacCurdy, McGee, and Putnam
Standing Committees
Program: Putnam, Boas, Holmes, Powell, Dorsey, Fewkes, and Russell
Publication: Hodge, Baker, Boas, Chamberlain, Culin, Dorsey, Fletcher, Holmes, Powell, and Putnam
Finance: not then appointed
AAA Representatives to Council of AAAS
W. H. Holmes and Franz Boas
Sources: Anonymous, \903>, American Anthropologist, 5:184-185; McGee, 1902c, American Anthro-
pologist, 4:465-466.
Church. This scenario means that anthropologists attending the AAAS annual
meeting assembled in two different churches on June 30: (1) in the late morning
at the Bellefield Presbyterian Church for the AAAS Section H organizing meeting,
and (2) in the afternoon at the Oakland Methodist Episcopal Church for the AAA
founding meeting and to hear Fewkes’ address.
The officers of AAA and the members of its Council, Executive Committee,
and Standing Committees were elected and installed at the founding meeting in
Pittsburgh (Table 5). The Pittsburgh Gazette (July 1 edition) was the only news-
paper reporting the officers elected the day before. There was much overlap be-
tween AAA and Section FI, both in officers and members (Tables 3 and 5). W J
McGee served as AAA President and as the Section H Associate at Large Member
of the AAAS Council. Section H President Stewart Culin and Secretary Harlan
I. Smith were members of AAA Council and Culin was on its Executive Com-
mittee. Franz Boas and William H. Holmes, two of the Vice-Presidents for AAA,
served on the Sectional Committee of Section H and as the AAA (Affiliated
Society) Representatives to the AAAS Council. There are three examples of spe-
cific activities corroborating this close relationship. Franz Boas proposed and Sec-
tion H adopted the following resolution:
Resolved, That it is the sense of this meeting that it is desirable to bring
about the closest possible correlation between the work of Section H of
the American Association for the Advancement of Science and the
American Anthropological Association. (Smith, 1902:202).
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Annals of Carnegie Museum
VOL. 71
Second, AAAS officially admitted the American Anthropological Association as
an Affiliated Society while the Pittsburgh meeting was still in progress (Mac-
Dougal, 1902a:43). Third, the scientific session in the afternoon of July 2, only
two days after the founding meeting, was a joint meeting of Section H and AAA
(Smith, 1902:209).
The AAA founding meeting was the culmination of a lengthy process to create
a national anthropological organization, the need for which had been discussed
for years (Stocking, 1960; Darnell, 1998:245-254). At a conference in Chicago,
held six months before the Pittsburgh meeting, representatives of AAAS Section
H, the Anthropological Society of Washington, and the American Ethnological
Society met to discuss various issues pertaining to a national organization (Anon-
ymous, 1903; McGee, 1902a, \902b). Three months after the Chicago meeting,
in March 1902, the American Anthropologic [al] Association was incorporated in
Washington, D. C. (Anonymous, 1903:181), thereby providing the legal status
necessary for the election of officers at the AAA founding meeting in Pittsburgh
three months later.
The AAA founding meeting was, in one sense, a pro forma meeting, because
extensive correspondence and discussion had settled many items on the agenda
ahead of time (Stocking, 1960). Details about AAA continued to be worked out
at other meetings held at the Hotel Schenley (Anonymous, 1903:184, 186). The
relationship of Section H to AAA was a point of discussion on July 1, even
before the scientific sessions started at the Bellefield Presbyterian Church (Smith,
1902:202). Thus, the Oakland Methodist Episcopal Church was the building
where the AAA founding meeting formally convened, but there certainly were
other buildings in Oakland where other meetings were held, before and after the
founding meeting, that became instrumental in determining the future directions
of the American Anthropological Association (Watters and Eowler, 2002).
The single most contentious issue in the founding of AAA was whether the
organization should adopt an exclusive or inclusive policy regarding membership.
Boas (1902) and other anthropologists in favor of an exclusionary policy wanted
to restrict membership to professional anthropologists, whereas McGee and many
others were inclined toward including amateurs or non-professionally trained an-
thropologists in the membership of AAA. Although this issue nominally dealt
merely with membership, in reality it was one aspect of a broader debate centering
on the professionalization of the discipline (Stocking, 1960; Darnell, 1998:249-
251). McGee and others involved with the March 1902 incorporation of AAA
proposed a list of 60 anthropologists to be invited to the founding meeting in
Pittsburgh. Boas and his supporters whittled that number down to 40 persons who
were canvassed for their views before the Pittsburgh meeting. If the anonymous
author (1903:184) is accurate in listing the persons who attended, there were only
13 anthropologists who actually participated in the AAA founding meeting in the
Oakland Methodist Episcopal Church.
Not only did the “inclusive” membership policy carry the day, but the period
of time in which an individual could be classed as a “founder” of AAA was also
extended:
. . . one of the most important actions taken at Pittsburg was that pro-
viding that other anthropologists who so desire may, during the re-
maining months of the year [1902], become affiliated with the Associ-
ation and classed as founders. (McGee, 1902c:466).
2002
Watters — From the Archives and Collections
229
Anonymous (1903:191-192) confirms that many persons took advantage of the
chance to become AAA founders. The list of Life Members (5) and Members
(160) covers two pages; of these 165 individuals, 142 (86%) have asterisks beside
their names indicating they were “founders of the Association.’' Ironically, for
reasons that make no sense apart from asterisks having been omitted, inadvertently
or otherwise, five of the 13 participants attending the founding meeting in Pitts-
burgh are not indicated as founders. Franz Boas, J. Walter Fewkes, J. N. B. Hewitt,
and Harlan 1. Smith have no asterisks, and Marshall H. Saville’s name is not on
the list at all.
W. J. Holland and C. C. Mellor, two of the individuals listed as AAA founders,
very likely would not have made the list had the exclusionary policy advocated
by Boas been adopted in Pittsburgh.
W. J. Holland, the A A AS, and the AAA in Retrospect
Holland rightfully was pleased with the success of the 51st Annual Meeting of
AAAS and the roles played by Carnegie Institute and nine local committees in
ensuring its success. He expressed these sentiments in his Editorial appearing in
the Annals of Carnegie Museum later that year. He closed that article by stating:
One of the most gratifying features of the occasion was the universal
recognition on the part of the assembled body of scientific men and
women of the thoroughness and success of the work which is being
done in that department of the Institute which stands for the advance-
ment of science. A becoming modesty forbids us to quote the expres-
sions which were publicly heard, but the appreciation of what has been
done in the Museum in the few years of its life, by men who are most
competent to express a just judgment, was most gratifying and encour-
aging. (Holland, 1902:503).
L. O. Howard, the AAAS Permanent Secretary, wrote an overall favorable
review of the meeting (though he incorrectly referred to it as the “fifty-second
annual meeting”):
In many respects it has been one of the most successful meetings which
the Association ever held. The attendance, while not very large, has
been composed of members of the active working class, many of them
being Fellows, and the meeting may be safely characterized as a work-
ing meeting. The registration has shown four hundred and thirty-five
members in attendance. ... It must be remembered as usual that the
number registered, namely, 435, includes only the active members and
associates of the Association, and that as a matter of fact there are
always a few members in attendance who are so characteristically for-
getful of all things except scientific matters that they entirely fail to
register. . . . [EJleven affiliated societies of a national scope have met
with us and have swelled the gathering of scientific men in Pittsburgh
during the past week to approximately 750 individuals. The meeting
has, therefore, been a scientific congress of great importance. (Howard,
1902:572).
MacDougal (1902^2:41) reported a registration of 431 persons while Howard
(1902:572) stated 435, so one can surmise that four of the forgetful scientists
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VOL. 71
must have remembered eventually to register. Howard’s comment about the “not
very large” attendance is odd, because the same Proceedings volume (Howard,
1902:16) discloses the number of members in attendance at Pittsburgh surpassed
the figures for the two prior meetings in New York (434) and Denver (311).
However, not everything went smoothly: . it must be added, somewhat
ungraciously perhaps, that the agreement with the headquarters hotel [Schenley]
was so loosely made that exorbitant rates were demanded ...” (MacDougal,
1902^:42).
The industrial sector of the economy of Pittsburgh and Allegheny assuredly
was a major factor in drawing members of some of the AAAS scientific Sections
and Affiliated Societies to the 51st Annual Meeting. Holland, working with Brash-
ear and Mellor, the nine local committees, and the supporting organizations was
able to draw heavily on the managers of these industries to facilitate the scientific
excursions and support the social activities (Table 1). Holland, Church, and An-
derson consistently associated science with industry in their speeches (in
MacDougal, 1902Z?:55 1-558). The connection was not lost on AAAS President
Asaph Hall, who responded:
It is fortunate for us to meet in the city of Pittsburg, famous for its
wonderful production of iron and steel, materials which lie at the foun-
dation of modern civilization. We are glad to see the homes of men
who are the munificent benefactors of our libraries and of our scientific
institutions. We shall be interested in visiting the great shops where you
convert the products of a generous nature into articles for our daily use.
(quoted in MacDougal, 1902Z?:558-559).
One of Holland’s earliest concerns, expressed in his April 30, 1901 Monthly
Report of the Director to the Museum Committee, was to increase the membership
of AAAS in Pittsburgh. The AAAS rolls list 12 members from Allegheny and
66 from Pittsburgh (Howard, 1902:240, 246-248), not including other members
in a number of nearby towns. Whether the number of local members increased
overall as a result of Holland’s efforts is uncertain, but his own effort assuredly
paid off in terms of persons he employed at Carnegie Museum. Five Carnegie
Museum employees, John Bell Hatcher, John A. Shafer, Herbert H. Smith, Doug-
las Stewart, and Frederick S. Webster are listed as AAAS members (Howard,
1902:247-248), and most appear to have joined for the Pittsburgh meeting. Join-
ing AAAS proved beneficial for Hatcher, Shafer, and Webster because Holland,
six months later, sent them to the next AAAS meeting, in Washington, D.C.
(Monthly Report of the Director to the Museum Committee, December 31, 1902).
Holland apparently did not present a scientific paper at the Pittsburgh AAAS
meeting, as none is listed for him in the papers read before Section F (Zoology).
John Bell Hatcher presented the only paper attributed to Carnegie Museum,
“Structure of the Pelvic Girdle in the Sauropoda” in Section F (Howard, 1902:
459). However, other papers by Carnegie Museum personnel could have been
given in meetings of the various Affiliated Societies, since those papers are not
always listed in the AAAS Proceedings.
Holland monetarily assisted Carnegie Museum in one instance as a direct result
of the AAAS meeting:
After paying all the expenses of the meeting there remains a balance of
approximately two thousand dollars [of the $9,000 raised], which the
2002
Watters — From the Archives and Collections
231
donors have unanimously consented to allow to be applied to the ac-
quisition of Egyptian collections for the Carnegie Museum. (Holland,
1902:501).
This money went to the Pittsburgh Chapter of the Egypt Exploration Eund, for-
mally chartered at Carnegie Museum in 1901 largely through Holland’s efforts
and through which Carnegie Museum obtained substantial Egyptian collections
(Watters, 2002; Watters and Patch, 1985).
Holland did not attend the American Anthropological Association’s founding
meeting, even though he later was classed as a founder. In fact, the only evidence
of Holland being involved, even indirectly, with anthropologists in Pittsburgh was
the loan he arranged of the Mexican sculpture for Saville’s lecture in the joint
Section H and AAA scientific session. Nevertheless, Holland can be said to have
provided the Oakland Methodist Episcopal Church venue for the AAA founding
meeting, since he arranged for the church buildings used for meetings by the
Affiliated Societies and scientific Sections. Had the founding meeting taken place,
as originally scheduled, in the Bellefield Presbyterian Church, where he formerly
had been pastor, the irony would have been delicious.
Holland must be credited with being the person primarily responsible for bring-
ing the 51st Annual Meeting of the American Association for the Advancement
of Science to the Oakland district of Pittsburgh. It was the first AAAS meeting
held in the city. Pittsburgh twice more hosted AAAS annual meetings (Kohlstedt,
Sokal, and Lewenstein, 1999:179), in December 1917, while Holland was still
Director of Carnegie Museum, and December 1934, two years after he died. It
has not hosted one since.
Acknowledgments
Throughout Holland’s tenure as Director, the institution was known as Carnegie Museum, being
changed to Carnegie Museum of Natural History only in 1973. I discovered Holland’s involvement
with the AAAS 51st Annual Meeting while conducting research into the life of Carl V. Hartman (first
anthropology curator at Carnegie Museum) under a grant awarded by the Adrienne and Milton Porter
Charitable Foundation. Purely by chance, Don D. Fowler of the University of Nevada, Reno, a pro-
fessor of mine when I received my M. A. degree there, contacted me in the late summer of 2001
regarding the AAA founding meeting in Pittsburgh in 1902. Don was researching the event for a
paper he delivered at the American Anthropological Association annual meeting that fall. It is no
small irony that Don and I each received our Ph.D. at the University of Pittsburgh. We decided to
join forces and contribute a short report (Watters and Fowler, 2002) to Anthropology News (then
running a series of reports on the AAA centenary), with Don focusing on historical contexts, while I
explored the church venue issues. I opted to write the present article to document the data obtainable
from unpublished materials reposing in the Holland Archives (a largely untapped resource pertinent
to intriguing scientific and historical issues). I am grateful to CMNH Research Associate Hazel Johnson
for researching newspaper reports concerning Holland and the 1902 meetings; CMNH Librarian Ber-
nadette Gallery for access to archival and library resources vital to this study; Gil Pietrzak of the
Carnegie Library of Pittsburgh for images of the churches; and Don D. Fowler, who many years ago
showed me that an undergraduate degree in history could be transformed into a master’s degree in
anthropology, with interesting results.
A Note on Sources
There are three instances where very similar articles were published by the same author in different
sources. McGee’s (1902a and 1902^?) short articles, both of which announced the upcoming AAA
founding meeting, are essentially identical apart from the final paragraph in 1902a, listing the pro-
spective founders, that is omitted in 1902Z?. W J McGee’s idiosyncrasy of not using periods after his
initials has been respected in this publication. MacDougal’s 1902a report is an abbreviated version of
the 1902Z? report in the 51st Proceedings; the latter contains materials omitted from the former, the
232
Annals of Carnegie Museum
VOL. 71
speeches of Holland, Church, and Anderson being an example. J. Walter Fewkes’ (1902a, 1902Z?)
Vice-Presidential address is published identically in two sources. I have retained his use of “Porto
Rico,” but note that it represented the U.S. Government’s attempt to anglicize spelling of Spanish
words in that newly occupied territory. Authorship of the Anonymous (1903) article, an important
resource about the AAA founding meeting, has been attributed to McGee and to Dorsey (see Stocking,
1960:1 and fn. 2, for discussion). I have opted to retain it as anonymously authored. Howard is listed
as editor of the 51st AAAS Proceedings (1902), and I have attributed all information therein to him,
unless specified otherwise (e.g., MacDougal, 1902^; Fewkes, \9Q2b). Pittsburgh newspapers provided
coverage of the AAAS meeting, and they contain the only information about certain excursions and
most social events. The July 1 and 2 editions of The Pittsburg Press and The Pittsburgh Gazette are
recommended especially (note the newspapers’ inconsistent spelling of the city).
Literature Cited
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ANNALS
0/ CARNEGIE MUSEUM
THE CARNEGIE MUSEUM OF NATURAL HISTORY
4400 FORBES AVENUE • PITTSBURGH, PENNSYLVANIA 15213
VOLUME 71 26 NOVEMBER 2002 NUMBER 4
i
CONTENTS
ARTICLES
A Late Miocene-Early Pliocene population of Trachemys (Testudines:
Emydidae) from East Tennessee
. Paul W. Parmalee, Walter E. Klippel,
Peter A. Meylan, and J. Alan Holman 233
Redescription of Eopelobates grandis, a Late Eocene anuran from the
Chadron Formation of South Dakota Amy C. Henrici 241
i
Expeditions, Expositions, Associations, and Museums in the Anthropological
Career of C. V Hartman
David R. Watters and Oscar Fonseca Zamora r2<
Index to Volume 71
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@ This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper).
ANNALS OF CARNEGIE MUSEUM
VoL. 71, Number 4, Pp. 233-239
26 November 2002
A LATE MIOCENE-EARLY PLIOCENE POPULATION OF TRACHEMYS
(TESTUDINES: EMYDIDAE) FROM EAST TENNESSEE
Paul W. Parmalee*
Research Associate, Section of Vertebrate Paleontology
Walter E. Klippel^
Peter A. Meylan^
J. Alan Holman^
Abstract
A Late Miocene-Early Pliocene (cf. Hemphillian) deposit of plant and vertebrate remains was
exposed during preliminary highway construction in Spring 2000 near the city of Gray, Washington
County, upper East Tennessee. In addition to as yet unidentified fish, frog, salamander, crocodilian,
and rodent species, two taxa of snakes (cf. Sistrurus sp., cf. Regina sp.), rhinoceros {Teleoceras sp.),
tapir (Tapirus, cf. T. polkensis), sloth (cf. Megalonyx sp.), gomphothere (Gomphotheriidae), peccary
(cf. Catagonus sp.), shrew (Soricidae), mustelid (Mustelidae), bear (cf. Ursus sp.), and turkey {Me-
leagris sp.) are represented. Remains of at least eight aquatic turtles of the genus Trachemys were
also recovered, whereas others were lost due to construction and fossil collectors. The pronounced
rugosity of the costal bones, deeply serrated anterior and especially posterior peripherals, and deeply
incised pygal of the carapace, and a pronounced and deeply serrated anterior margin of the plastron
suggest a relationship close to Trachemys inflata, an Early Pliocene turtle previously known only from
Florida. The numerous individuals represented at the Gray Site indicate a well-established population
and may reflect, along with crocodilians and certain terrestial mammals, a warm environment at some
period(s) during the Late Miocene-Early Pliocene.
Key Words: Gray Site, Tennessee, Miocene, Pliocene, Trachemys, Carapace, Plastron
Introduction
In May 2000, Tennessee Department of Transportation (TDOT) construction
crews widening State Highway 75 near the city of Gray, Washington County,
Tennessee unearthed layers of soft gray clay and an approximately two-meter
thick black, ancient lake/swamp deposit rich in plant macrofossils and articulated
(for the most part) skeletons of extinct animals. During the summer highway
construction was discontinued, and Highway 75 was redesigned to bypass the
fossil bed. A chain-link fence was erected around the site, and straw and grass
seed were blown over the exposed surfaces to provide protection against erosion.
Subsequent core drillings indicated that part of the site (lacustrine deposits) is in
excess of 30 meters in depth and covers an area of up to 1.6 hectares. The total
area of the site is estimated to be approximately 3.0 hectacres, of which 1.4
hectacres were exposed during construction.
’ Frank H. McClung Museum, University of Tennessee, Knoxville, TN 37996.
^ Department of Anthropology, University of Tennessee, Knoxville, TN 37996.
^Natural Sciences Collegium, Eckerd College, St. Petersburg, EL 33711.
Michigan State University Museum, Michigan State University, East Lansing, MI 48824.
Submitted 28 September 2001.
233
234
Annals of Carnegie Museum
VOL. 71
Vertebrates identified thus far include a small fish (Osteichthyes), salamander
(Caudata), frogs (Ranidae), crocodilians (Alligatoridae), snakes (cf, Sistrurus sp.,
cf. Regina sp.), turtles (Trachemys sp.), turkey (Meleagris sp.), tapir (Tapirus, cf.
T. polkensis), rhinoceros {Teleoceras sp.), gomphothere (Gomphotheriidae), sloth
{Megalonyx sp. or Pliometanastes sp.), peccary (cf. Catagonus sp.), bear (cf.
Ursus sp.), shrew (Soricidae), small mouse (Rodentia), and a mustelid (Musteli-
dae). The occurrence of Tapirus, cf. T. polkensis, Teleoceras sp., a small Mega-
lonyx sp. or Pliometanastes sp., and cf. Catagonus sp. strongly suggests that the
mammals from the fossil bed belong to the Hemphillian Land Mammal Age and
date to the Late Miocene-Early Pliocene (>4.5 million years ago). Plant remains
are extremely abundant, including tree trunks, limbs, seeds, and leaf impressions
and occur throughout the various layers of sediment. To date, remains of hickory
nuts {Cary a sp.), acorns {Quercus sp.), grape seeds {Vitis sp.), and hazel nuts
(Corylus sp.) have been identified, but a detailed study of the macroflora and
pollen has not yet been undertaken. Thus far, preliminary geological investigations
have included core samples, stratigraphic profiles, and paleomagnetic dating, but
these have yielded inconclusive evidence as to the exact age and formation of the
site. The site, believed to be a large water-filled sinkhole or natural basin, provided
habitat for a diverse aquatic fauna. Although aquatic mollusks were apparently
common during the interval, they were poorly preserved, and most appear as
white stains in the lacustrine sediments. A species of pea clam (Spheriidae) and
gastropod (Planorbidae: Helisoma sp.?) were also identified.
Materials and Methods
Throughout late spring and early summer of 2000, students and faculty from
the University of Tennessee, Knoxville, periodically collected fossils exposed by
earth-moving equipment. Specimens were prepared and catalogued at the Uni-
versity’s Zooarchaeology Laboratory and the Frank H. McClung Museum. With
few exceptions, all vertebrate specimens recovered at the Gray Site were salvaged
after being unearthed during highway construction and/or after the bone-bearing
matrix was removed and deposited on fields adjacent to the site. Consequently,
few elements survived intact; this was especially true of the turtles. The balance
of work at the site in 2000-2001 consisted primarily of salvaging specimens
exposed on the surface. Plans for future excavations are being formulated that
will be carried out by researchers from the University of Tennessee, Knoxville,
and East Tennessee State University, Johnson City, Tennessee.
Well-preserved elements of the plastron and carapace of several individuals
were recovered, although in most cases shells were fragmented or disarticulated
as a result of the unavoidable method of removal. None of the skeletons of in-
dividual turtles encountered were complete. Depending upon the type of matrix
in which they were preserved, some shells could easily be removed, cleaned
(brushing with water, soaking in calgon), and repaired, whereas others were ce-
mented to a conglomerate of sand and pebbles, usually to the inside of the car-
apace. Except for sections of a lower jaw found with one individual and fewer
than a dozen fragmented limb bones, the majority of appendicular skeletal ele-
ments appear either to have been lost during removal of the deposit by earth-
moving equipment or were disassociated from the shell prior to burial.
The most complete sections of plastron and carapace recovered at the Gray
Site, representing several individuals, were compared with the modern skeletons
2002 Parmalee et al. — Late Miocene-Early Pliocene Trachemys from East Tennessee 235
of the genera Chrysemys, Trachemys, and Graptemys housed in the collections of
the Zooarchaeology Section, Department of Anthropology, Universtiy of Tennes-
see, Knoxville and the Michigan State University Museum, East Lansing. On the
basis of descriptions and illustrations of fossil turtles of Hemphillian Age in North
America by Hay (1908) and Weaver and Robertson (1967), and comparison with
those of other repositories, the aquatic turtles from the Gray Site can be assigned
to the genus Trachemys within the Family Emydidae, as possibly a representative
of the Trachemys scripta complex or a closely related species within the genus.
Description and Discussion
The characters used most frequently to distinguish among the closely related
living species of sliders and coolers of the genera Pseudemys and Trachemys are
external. They include general carapace shape and the colors and patterns of the
shell, head, neck and limbs. Because very few skulls of this group have been
described, the systematics of fossil coolers and sliders is based almost entirely on
variation in the shell, including differences in bone and scute arrangement, size
and overlap of certain scutes, surface texture, bone thickness, and within Trach-
emys, degree of carapacial inflation (e.g., Hay, 1908). Within the larger group,
members of the genus Trachemys are easily recognized. They are the only forms
with doubly serrate carapacial margins (Seidel and Jackson, 1990). In fossil and
living Trachemys (except some neotropical forms), there is a notch in the cara-
pacial margin at the sulcus between each pair of marginal scutes and a notch
between each pair of peripheral bones. Together, these features produce a doubly-
serrate posterior carapace margin. Although recognition of the genus is not prob-
lematic, because of the subjective nature of the characters used, and because the
types of several species are fragmentary pieces of shell, the validity of several
fossil Trachemys species have been questioned. Weaver and Robertson (1967) re-
evaluated the nine species of Trachemys listed by Hay (1908) and synonymized
six with Chrysemys {=Trachemys) scripta petrolei, a mid-Pleistocene subspecies.
Moreover, they identified one as Terrapene, another as Deirochelys, and a third
as closer to Chrysemys picta than to T. scripta.
Weaver and Robertson (1967:58) described two new fossil species of Chryse-
mys {=Trachemys) from Florida, T. inflata from the Pliocene, and T. platymar-
giriata from the Pleistocene. They concluded that T. inflata was a member of the
T. scripta complex ‘A . . the basis of the highly sculptured scute areas of the
nuchal bone, the associated double toothed peripheral bones, epiplastra having an
extensive gular scute overlap and nuchal bone with extensive nuchal scute un-
derlap, and neural bones whose architecture suggests the presence of a well de-
fined median keel on the carapace.” In addition, they indicated that Trachemys
inflata differed from the fossil T. scripta petrolei and living T. s. scripta, T s.
troosti, and T. s. elegans by the shape, depth, and width of the anterior nuchal
notch and the smooth medial half of the marginal scute area of the nuchal bone.
There is some doubt that Trachemys inflata is a valid species, as well as un-
certainty as to its evolutionary position in the T. scripta complex. Weaver and
Rose (1967:70) commented that "'Chrysemys {Trachemys} inflata is unquestion-
ably a member of the evolutionary line leading to North American C. scrip-
ta ... C This view was also repeated by Auffenberg (1972). Weaver and Rob-
ertson (1967:65), on the other hand, expressed the- view that “The extent of the
nuchal and peripheral bone notching and the massive, rugose grooves and ridges
236
Annals of Carnegie Museum
VOL. 71
XIPHIPLASTRA PYGAL
Fig. 1 — Plastron TDOT 13.01 A and carapace TDOT 13.02B sections (same individual) of Trachemys,
cf. T. inflata from the Gray Site, Tennessee. Note the characteristic deeply serrated posterior peripheral
bones of the carapace. Catalog Nos.: A. TDOT 13.01. B. TDOT 13.02.
of C. inflata suggest that it was a specialized or aberrant species characterized by
an extreme development of Trachemys features and not representative of the main
evolutionary sequence leading to recent C. scriptaT
Specimens from the Gray Site tend to exhibit characters of Trachemys scripta,
and particularly those of T inflata. The general shape, sculpture, and deep notch-
ing of the nuchal bone, the exaggerated rugosity of the costals, and especially the
deeply notched, sharp-angled posterior peripheral bones (Fig. IB) are suggestive
of T. inflata. As in most extant subspecies of Trachemys scripta, and the few
known specimens of T. inflata, neural bones of the Gray Site specimens are
smooth and lack any evidence of a median keel on the carapace. Although in-
complete, plastra (Fig. 2) from the Gray Site compare closely with those of taxa
belonging to the T. scripta complex except for the deeper, narrower anal notch,
and the anterior margin of the epiplastra, which exhibits a more pronounced and
deeper serrated edge than is found in living Trachemys.
Representative plastra and carapace sections of Trachemys from the Gray Site
were compared with the type and other material of T. inflata, a large series of T.
platymarginata, and undescribed specimens of the T scripta group from the Moss
Acres Racetrack Site (Hulbert, 2001), all from Florida fossil sites and housed in
the Florida Museum of Natural History, Gainesville. In the holotype of T. inflata
the bone under each scute area on the nuchal is greatly inflated, as are the adjacent
peripherals. In the Gray Site specimens the scute areas are thick but not noticeably
2002 Parmalee et al. — Late Miocene-Early Pliocene Trachemys from East Tennessee 237
Fig. 2 — A-D. Selected examples of plastra of Trachemys, cf. T. injiata from the Gray Site, Tennessee,
exhibiting a pronounced and deeply serrated antierior margin. Catalog Nos.: A. UTK 65.02. B. DTK
64.02. C. UTK 63.01. D. UTK 66.01.
inflated. However, it seems possible that this character, involving the degree of
inflation, may vary with size and/or age.
Of the eight individuals of Trachemys represented in the Gray Site emydid
material recovered so far, only one carapace (Fig. IB) and one plastron (Fig, 2C)
of different individuals could be restored to permit a total length measurement.
238
Annals of Carnegie Museum
VOL. 7]
Both measured 210 mm and compare well in size to the Moss Acres Racetrack
Site T. scripta specimens, but are much smaller than those of T. inflata and T.
platymarginata (Late Pliocene). In addition, Trachemys from the Gray Site differs
from T. platymarginata in having a much more rugose carapace, especially an=
teriorly, and in the anterior margin being deeply serrate.
As already noted, the carapace of the Gray Site Trachemys is extremely serrate,
especially the posterior peripheral bones, and the serrations are deeper than those
observed in T. platymarginata, T. inflata, and the undescribed specimens of T.
scripta from the Moss Acres Racetrack Site. On the other hand, the plastron of
the Gray Site specimens are comparable to the Moss Acres Racetrack Site T.
scripta in having projecting epiplastra with a finely serrate anterior margin. How-
ever, the pygal of the Gray Site specimens is more like T. inflata in being deeply
incised.
Taking into account the variability of certain characters used to distinguish
among the species of Trachemys from the Late Miocene-Early Pliocene, we refer
the Gray Site specimens to Trachemys, cf. T. inflata. The presence of this species
of aquatic turtle, as well as the crocodilians, tapirs, rhinoceros and peccaries,
suggests that the climate during which the Gray Site (sinkhole?) sediments ac-
cumulated was warm. Farlow et al. (2001) described a diverse fauna of Hem-
phillian Age (in part) from the Pipe Creek Sinkhole Site in north-central Indiana
as also indicating a warm environment with possibly an adjacent dry, open grass-
land or savannah habitat. In the case of the Gray Site, however, the vertebrate
taxa identified thus far in association with the macroflora constitute a unique
assemblage that will permit reconstruction of a Late Miocene-Early Pliocene biota
unequalled in most other regions of eastern North America.
Acknowledgments
We thank Han'y Moore, Larry Bolt, Rick Noseworthy, George Danker and several other members
of the Tennessee Department of Transportation, Knoxville, TN, for their initial awareness of the Gray
Site fossils and their efforts in salvaging specimens during and following highway construction. Martin
S. Kohl, Tennessee Division of Geology, Knoxville, TN, contributed numerous fossil specimens from
the Gray Site to the Frank H. McClung Museum collections. A special note of appreciation is extended
to Marta Adams and Tony Underwood, Dandridge, TN, for their continuing work at the Gray Site
that has inovlved field recovery of fossils, and the cleaning, preservation and cataloguing of specimens
in the laboratory. We are grateful to Lindsay Kromer and Robert Pennington for the preparation of
figures 1 and 2. Richard C. Hulbert, Jr. is acknowledged with a special note of appreciation for making
available for comparative study fossil turtle specimens housed in the vertebrate paleontological col-
lections in the Florida Museum of Natural History, Gainesville, Florida. We thank Ruth Huppert,
Frank H. McClung Museum, for typing drafts of the manuscript, and two outside reviewers for their
helpful editorial comments.
Literature Cited
Auffenberg, W. 1972. Fossil turtles. Plaster Jacket, 16.
Farlow, J. O., J. A. Sunderman, J. J. Havens, A. L. Swinehart, J. A. Holman, D. L. Richards,
N. G. Miller, R. A. Martin, R. M. Hunt, Jr., G. W. Storrs, B. B. Curry, R. H. Fluegeman,
M. R. Dawson, and M. E. T. Flint. 2001. The Pipe Creek Sinkhole biota, a diverse late tertiary
continental fossil assemblage from Grant County, Indiana. American Midland Naturalist, 145:
367-378.
Hay, O. P. 1908. The fossil turtles of North America. Carnegie Institution of Washington, Washington,
D.C., Publ. 75.
Hulbert, R. C., Jr. [Editor.] 2001. The fossil vertebrates of Florida. University Press of Florida,
Gainesville, Florida.
Seidel, M. E., and D. R. Jackson. 1990. Evolution and fossil relationships of slider turtles. Pp. 68-
2002 Parmalee et al. — Late Miocene-Early Pliocene Trachemys from East Tennessee 239
73, in Life History and Ecology of the Slider Turtle (J. W. Gibbons, ed.). Smithsonian Institution
Press, Washington, D.C.
Weaver, W. G., Jr., and J. S. Robertson. 1967. A re-evalutation of fossil turtles of the Chrysemys
scripta group. Tulane Studies in Geology, 5:53-66.
Weaver, W. G., Jr., and F. L. Rose. 1967. Systematics, fossil history, and evolution of the genus
Chrysemys. Tulane Studies in Zoology, 14:63-73.
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ANNALS OF CARNEGIE MUSEUM
VoL. 71, Number 4, Pp. 261-299
26 November 2002
EXPEDITIONS, EXPOSITIONS, ASSOCIATIONS, AND MUSEUMS IN THE
ANTHROPOLOGICAL CAREER OF C. V. HARTMAN
David R. Watters
Curator, Section of Anthropology
Oscar Fonseca Zamora
Research Associate, Section of Anthropology
Abstract
Within a span of twenty years (1890-1910), Carl Vilhelm Hartman changed careers from botany
to anthropology, achieved recognition as a professional colleague by peers on both sides of the At-
lantic, pioneered systematic archaeological research in Costa Rica, and produced two widely acclaimed
monographs about his fieldwork. Hartman’s transition to anthropology ensued from four principal
entities with which he became engaged — expeditions, expositions, associations, and museums, and
through which he developed a broad network of contacts among European, Central American, and
North American anthropologists and museologists. He effectively used these trans-Atlantic links to
advance his anthropological career. Hartman, a Swede by birth, learned his field methods in the Old
World but conducted research exclusively in the New World, where he incorporated high standards
for archaeological fieldwork. His career, always firmly rooted in museum anthropology, included five
years (1903-1908) as Carnegie Museum’s first Curator of Ethnology and Archaeology. Hartman’s term
of employment at Carnegie Museum coincided with the onset of a gradual shift in the training of
anthropologists, moving from the museum to the university setting, under the Boasian paradigm for
professionalization of the discipline.
Key Words: Costa Rica, museum anthropology. International Congress of Americanists, Carnegie
Museum, Hjalmar Stolpe, W. J. Holland
Introduction
Carl Vilhelm Hartman, born August 19, 1862 in Orebro, Sweden, the son of a
distinguished botanist initially pursued a career following in his father’s footsteps.
His university training was in botany, his initial work was in applied botany for
the Swedish Academy of Agriculture, and he spent five years studying at major
botanical gardens in Europe under a grant awarded by the Swedish Academy of
Science (Brunius, 1984; Eranzen, 1969).
In 1890, at the age of 28, Hartman joined an expedition that profoundly affected
his life, stimulating a total change of career to the emerging discipline of anthro=
pology. Within the span of a single decade, the 1890s, Hartman had established
his professional credentials sufficiently to obtain employment as a museum an-
thropologist, a profession to which he would devote the remainder of his life.
How did it come to pass that a Swede, trained as a botanist, proficiently ex-
cavated Costa Rican archaeological sites and purchased collections of antiquities
for a museum in the United States a century ago? This intriguing question, the
impetus for this paper, implicitly acknowledges the inter-hemispheric links that
characterize Hartman’s career, from his training in the Old World and research in
the New World, to his employment by museums on both sides of the Atlantic,
Submitted 3 May 2002.
261
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Fig. 1. — C. V. Hartman (standing) and unidentified visitor at the Chinchilla site. Central Highlands,
Costa Rica, in 1903. (Section of Anthropology glass-plate negative G998).
and to his collegial relations with anthropologists in Europe, Central America,
and North America.
This paper’s central thesis is that “internationalism,” in the sense of inter-
hemispheric linkages, was the driving force in the transformation of Hartman’s
career. We follow the theoretical orientation of Givens (1992:64): “The role of
biography in explaining archaeology’s past is to delineate and assess the place of
the individual worker within the science.” The purposes of this study are to
document the four entities — expeditions, expositions, associations, and muse-
ums— in which this internationalism is manifested and, in turn, to analyze their
relationships to Hartman’s career transformation and his development as a pro-
fessional anthropologist. The two decades (1890-1910) spanning Hartman’s tran-
sition to and most active involvement with anthropology coincide with a period
of significant developments in the discipline in the Americas and Europe.
In the Americas, Hartman is known as an archaeologist, most notably for his
introduction of systematic field methods developed in Europe into Costa Rican
archaeology (Fig. 1). The field methods he learned in Sweden have been discussed
in depth by Rowe (1959) and commented upon briefly by Willey and Sabloff
(1993:85). His pioneering research in Costa Rica was addressed by Fonseca Za-
mora (1984), Jones (1998), Ohlsson de Formoso (1991), and Skirboll (1984b);
2002 Watters and Fonseca Zamora — C. V. Hartman’s Anthropological Career 263
his contributions were recognized by a symposium held in his honor at Carnegie
Museum of Natural History (Skirboll and Creamer, 1984).
Hartman’s anthropological research extended well beyond archaeology, a fact
known in Sweden (Brunius 1984) but rarely acknowledged in the Americas (cf.
Lutz, 2001). He wrote ethnographic (1895, 1897, 1901Z?, 1907J, 1907^) and eth-
nobotanical (1906, 1910c) articles and recorded but only sparsely published his
linguistic and anthropometric data. Hartman’s work as an anthropology curator
and museum administrator likewise is also little appreciated in the Americas.
Carnegie Museum (it became Carnegie Museum of Natural History in 1973)
employed Hartman as its first Curator of Ethnology and Archaeology in 1903,
largely because of the anthropological experience he had gained in the previous
decade (Richardson, 1980). His tenure coincided with the implementation of Di=
rector W. J. Holland’s plan for a new Carnegie Museum, and he became a key
player in fulfilling Holland’s vision through various curatorial duties ranging from
acquisition of collections, to installation of exhibits, to publication of scientific
research. Viewed from one hundred years later, however, Hartman’s primary leg-
acy lies in the knowledge generated by his 1903 expedition to Costa Rica. The
research resulted in his three publications (Hartman, 1907a, 1907Z?, 1910a) and
subsequent studies based on collections he brought back, by Fonseca Zamora and
Richardson (1978), Fonseca Zamora and Scaglion (1978), Heckenberger and Wat-
ters (1993), Skirboll (1981, 1984a), Swauger and Mayer-Oakes (1952), and Wat-
ters and Fonseca Zamora (2001a, 2001b).
Materials and Methods
The present study makes liberal use of another of Hartman’s legacies, the doc-
uments and photographs from his tenure at Carnegie Museum, which currently
reside in the Hartman Archives at Carnegie Museum of Natural History, as well
as Hartman-related documents contained in the Holland Archives of the same
institution. These unpublished sources contain a wealth of biographical informa-
tion about Hartman, insights into his views on the four entities stimulating his
career change, and his perspectives on the colleagues and activities fostering his
professional development in anthropology. The Holland Archives, especially his
Reports of the Director to the Museum Committee and the Minutes of the Museum
Committee’s monthly meetings, document the interplay between Holland and
Hartman during his five-year term of employment.
Published materials provide a second source of information. These include
Hartman’s articles and monographs and, where applicable, publications in which
he is mentioned. The former category is particularly valuable for establishing
Hartman’s own perspectives on individuals who assisted his career (e.g., the pref-
aces to his two monographs). The latter category provides other persons’ assess-
ments of him, such as Carl Lumholtz’s comments about Hartman’s roles on the
expedition to Mexico. These publications also provide the historical context for
the four entities that facilitated his transition to anthropology, such as the pro-
ceedings of the various Sessions of the International Congress of Americanists he
attended. Pittsburgh newspapers provide another source of published information.
We present initially the data from the four entities — expeditions, expositions,
associations, and museums — that compose the structure or framework of C. V.
Hartman’s transition from botanist to anthropologist. In doing so, the chronolog-
ical ordering of the events becomes of secondary significance. In the final section
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of this paper, we restructure the primary events chronologically, discuss certain
individuals who played key roles in Hartman’s anthropological career, and place
the twenty-year span (1890-1910) of his most active involvement with anthro-
pology in historical context.
Expeditions
Hartman began his anthropological career during his first expedition to the New
World and enhanced his standing as an anthropologist during two later ones. He
was a crewmember on the first expedition but headed the second and third. The
first expedition was the initial factor stimulating his transition to anthropology.
Lumholtz Expedition to Mexico
Hartman’s career took a major turn in 1890 when he was selected as expedition
botanist by Norwegian explorer and ethnographer Carl Lumholtz who, sponsored
by the American Geographical Society of New York, led an expedition to the
SieiTa Madre mountains of northwestern Mexico. Lumholtz’s expedition also was
linked to the American Museum of Natural History, being allied loosely with its
Hyde Expedition (Fowler, 2000:235). In all, Lumholtz spent five years in Mexico
conducting field research on four expeditions between 1890 and 1898. Hartman
served from September 1890 until summer 1892, and then rejoined Lumholtz for
a few months in 1893.
Lumholtz (1902 I:vii-xix) provides information on the origin, support, staffing,
and timing of the four Sierra Madre expeditions (cf. Lumholtz, 1891). He distin-
guishes between the first expedition (September 1890-April 1891) and the second
(January 1892-August 1893) because in the intervening months he had returned
to the United States to raise funds. Hartman stayed on in Mexico in charge of
the expedition camp at San Diego and from his perspective the first expedition
continued uninterrupted, from when he joined in September 1890 until he departed
in the summer of 1892. He was the only member of the “scientific corps” carried
over from the first to the second expedition, leaving in the midst of the second,
seemingly the last of the “scientific corps” to depart. However, he resumed field-
work with Lumholtz for several months in 1893 (Lumholtz, 1902 1:186, 444),
and dated locality records for some of the plants he collected (Robinson and
Fernald, 1895) verify that he had rejoined the second expedition in May. Lumholtz
conducted fieldwork by himself for the remainder of the second expedition (sum-
mer 1892-August 1893), apart from the brief time he was rejoined by Hartman,
and for all of the third (March 1894-March 1897). He was joined by physical
anthropologist Ales Hrdlicka for the brief (four month) fourth expedition in 1898.
Lumholtz (1891:389; 1902 Lx, xii) identifies the members of his “scientific
corps” from the first and second expeditions (Table 1). The first expedition’s
archaeologist, A. M. Stephen (Lumholtz just used initials and variably spelled the
last name Stephens and Steven) is mentioned only three times in the 1902 book,
in the first two chapters covering the expedition’s early months; Stephen’s assis-
tant, R. Abbott, is mentioned only in the preface. It appears that Stephen and
Abbott left the expedition early on, thus opening the way for Hartman to assume
responsibility for archaeological projects. Hartman carried out archaeological ex-
cavations at mounds on the Piedras Verdes river (Lumholtz, 1902 I: xi, 93), at
the time he headed the camp at San Diego while Lumholtz was away (April-
winter 1891) between the first and second expeditions.
2002 Watters and Fonseca Zamora — C. V. Hartman’s Anthropological Career 265
Table 1. — The Scientific Corps of Lumholtz’s first and second Sierra Madre expeditions.
First expedition (September 1890-ApriI 1891)
Physical geographer — Professor W. Libbey (of Princeton, NJ) and his unnamed laboratory man
(possibly G. Haviland, as in Lumholtz, 1891:389)
Archaeologists — A. M. Stephen (also spelled Steven and Stevens by Lumholtz; almost certainly
Alexander McGregor Stephen), assisted by R. Abbott
Botanists — C. V. Hartman and C. E. Lloyd (listed as F. Lloyd in Lumholtz, 1891:389)
Zoological collector — F. Robinette
Mineralogist — H. White
Second expedition (January 1892-August 1893)
Botanist — C. V. Hartman (only member of the scientific corps on both expeditions)
Civil engineer and photographer — C. H. Taylor
Mineralogist and zoological collector — A. E. Meade (also spelled Meeds)
Notes:
(1) Lumholtz (1891:389; 1902 I:x) twice lists members of the first expedition’s scientific corps, but
with some inconsistencies
(2) Second expedition members are from Lumholtz (1902 L.xii); none of the scientific corps members
served for the entire second expedition
(3) Hartman served from September 1890 until the summer of 1892, and then rejoined Lumholtz for
several months in 1893; he remained in Mexico in charge of the camp, succeeding H. White, between
the first and second expeditions ( April- v/inter 1891)
A. M. Stephen assuredly was Alexander McGregor Stephen, “ . . . one of the
more fascinating and least-known characters in the coterie of nineteenth-century
Southwestern anthropologists” (Fowler, 2000:138). By 1890, when the Lumholtz
expedition began, Stephen had a decade of experience helping organize expedi-
tions for anthropologists from the Smithsonian Institution’s Bureau of American
Ethnology. Stephen had married a Navajo woman, lived at First Mesa, spoke
Navajo and Hopi, and was friends with Thomas Varker Keam and John Gregory
Bourke, two other persons involved with mounting expeditions (Fowler, 2000:
133-139; Parsons, 1936). The notion that this person was Alexander McGregor
Stephen is supported further by Hartman: “The thee archaeologist of the expe-
dition, Mr. A. M. Stephen, who had spent some spent fifteen years studying tribes
of the South-west ...” (Hartman, 1897:120). Stephen’s participation in the Lum-
holtz expedition and his link to Hartman, albeit brief, seem not to have been
recognized previously. Stephen’s archaeological assistant, R. Abbott, remains a
complete mystery.
Hartman’s exposure to and experience with anthropological fieldwork during
the Lumholtz expedition dramatically changed his life. His botanical studies were
important scientifically and resulted in some new species being identified (Rob-
inson and Fernald, 1895), but his interests thereafter were directed more toward
anthropology than botany. He confirmed the impact of this expedition in a 1903
letter to W. J. Holland, Director of Carnegie Museum, requesting employment as
a curator:
When Dr. Carl Lumholtz (who is a Norwegian) organized his expedition
for the exploration of Sierra Madre he engaged my services as a bota-
nist. Soon however I became greatly interested in the archaeological
and ethnological features and as Dr. Lumholtz needed a special assistant,
I turned my attentions wholly to this line of investigation. (Hartman
Archives, letter, February 20, 1903, C. V. Hartman to W. J. Holland).
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VOL. 71
As special assistant, Hartman gained experience in managing the expedition
and Lumholtz apparently placed considerable trust in his abilities. Hartman had
been kept on when the scientific corps was being reduced in number, assumed
charge of the camp when Lumholtz returned to the United States, and he con^
ducted excavations at various archaeological sites (Lumholtz, 1902 1:93, 186,
444). Hartman’s name appears far more frequently in the index of this two- volume
work than does any other member of the scientific corps.
The Lumholtz expedition resulted in Hartman’s first anthropological paper, giv-
en at the 10th International Congress of Americanists in Stockholm in 1894, and
his first anthropological publications (Hartman, 1895, 1897). He had published
botanical papers before the Lumholtz expedition and a few later in his life, but
ever after' the expedition he published mostly on anthropological topics (Franzen,
1969). He combined his botanical and anthropological interests in his “tree cal-
abash” publications, including an article In German in the Boas Anniversary
Volume (Hartman, 1906), later translated into French (1910c), and two articles
based on his Salvadoran research (1907£f, 1907c).
Swedish Expedition to Central America
In May 1896, Hartman began a three-year expedition to Central America that
was broad in anthropological scope, encompassing archaeological, ethnological,
linguistic, and anthropometric research, and in geographic scale with fieldwork in
Costa Rica, El Salvador, and Guatemala. He was the expedition’s only full-time
staff member, although he hired local workers as needed. He did not return to
Sweden until the autumn of 1899.
The expedition, nominally sponsored by the Swedish Society for Anthropology
and Geography, really was organized and funded by Ake Sjogren, a Swedish
geologist and mining engineer who, while working in Costa Rica in the early
1890s, had become interested in the country’s archaeology. Stolpe (1905) credits
Sjogren with conceiving the idea and drawing up the overall plan for archaeo-
logical research implemented by Hartman. Sjogren’s geological background led
to his fascination with archaeology, which focused on jade artifacts found in Costa
Rican sites and issues surrounding their origin (Wilson, 1902). In the years before
he funded Hartman’s expedition, Sjogren had visited the Smithsonian Institution
and provided Costa Rican jade samples for microscopic and thin section analysis
to curators at the United State National Museum (Wilson, 1898:458-459).
Hartman spent 17 months (May 1 896-September 1897) excavating sites in
Costa Rica’s Atlantic plain, Central Highlands, and Pacific slope before moving
on to El Salvador for ethnographic and linguistic studies of the Pipiles, and to
Guatemala with the Xiecas, through the spring of 1899 (Hartman, 1901«:l-2).
He amassed a sizeable amount of Costa Rican antiquities that he presented to
Ake Sjogren who, in turn, donated the collection to the Royal Ethnographical
Museum in Stockholm (Stolpe, 1905).
He invoked this expedition when applying for the position at Carnegie Muse-
um:
Returning to Sweden, I received a favorable offer to carry out explo-
rations of my own in Central- America under the auspices of the An-
thropo. Geogr, Society. This position I accepted and carried on explo-
rations, which extended over a period of three years. (Hartman
Archives, letter, February 20, 1903, C. V. Hartman to W J. Holland).
2002 Watters and Fonseca Zamora — C. V. Hartman's Anthropological Career 267
The Central American expedition resulted in Hartman’s first monograph on
Costa Rican archaeology (190 la), published in English and summarized in Swed-
ish (1902), and three ethnographic publications (19011?, I901d, 1907^). Major
figures in anthropology favorably reviewed his 1901 archaeological monograph
in leading journals in Europe and North America (Gordon, 1906; MacCurdy,
1905; Peet, 1904; Seler, 1904). Such acclaim was a important step in his career
advancement. His Swedish ethnographic article on El Salvador was translated
recently into Spanish (Hartman, 2001).
Carnegie Museum Expedition to Costa Rica
Hartman reported for duty at Carnegie Museum on March 17, 1903 and de-
parted Pittsburgh two weeks later on his second expedition to Costa Rica. In that
brief period, Director Holland had obtained letters of introduction for Hartman
from the Costa Rican embassy in Washington, written personal letters to influ-
ential individuals he knew in Costa Rica, arranged for wire transfer of monies,
purchased necessary field equipment, and obtained a camera and photographic
supplies for Hartman. Holland’s two-page letter of instruction for the expedition
directs Hartman to conduct archaeological research and “ . . . incidentally if you
are able to make collections of botanical, entomological and other specimens”
(Holland Archives, letter, March 24, 1903, W. J. Holland to C. V. Hartman).
Hartman returned to Pittsburgh in mid November, having spent about seven
months excavating sites, purchasing collections, and conducting museum research.
He also brought back zoological and botanical specimens.
Hartman concentrated his excavations in the Central Highlands (Fig. 2) and the
Nicoya peninsula on the Pacific side, with only a brief excursion to the Atlantic
plain. He studied and photographed artifacts in the National Museum of Costa
Rica and expended a great deal of time, effort, and money in purchasing collec-
tions of antiquities from private collectors, an assignment given to him by Holland
who clearly regarded that task as most important for Carnegie Museum (Watters,
2002a; Watters and Fonseca Zamora, 2001^). Hartman succeeded in that task. In
the Annual Report of the Director it is stated: “All of the collections acquired by
Mr. Hartman in Costa Rica, either by purchase or as the result of excavations,
filling more than eighty large cases, have been brought to the Museum ...”
(Holland, 1904:27).
The Carnegie expedition resulted in Hartman’s second monograph on Costa
Rican archaeology (1907a), an American Anthropologist article on the alligator
motif on pottery (19071?), and a comparative study of four grave forms (1910a).
Leading anthropologists in Europe (Beuchat, 1909) and North America (Holmes,
1908) again favorably reviewed his monograph. Hartman had produced two im-
portant monographic works on his Costa' Rican research in just six years.
Expositions
Hartman’s anthropological career coincided with a period during the late nine-
teenth and early twentieth centuries when international expositions (“world’s
fairs”) were being staged at an accelerated rate in Europe (Greeehalgh, 1988) and
the United States (Rydell, Findling, and Pelle, 2000). His personal involvement
with such expositions was limited, but it nonetheless directly affected his career
by expanding his knowledge of New World indigenous cultures and providing
him with the opportunity to meet anthropologists from the Americas and Europe.
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Annals of Carnegie Museum
VOL. 71
Fig. 2. — Hartman’s 1903 excavations at the Chinchilla site on the slope of Irazu volcano. (Section of
Anthropology glass-plate negative G979).
Hartman’s enduring link to Costa Rican archaeology is tied to that country’s
national expositions and its subsequent involvement in international expositions.
Costa Rica's National Expositions and the Exposicion Historico-Americana
Activities sanctioned and sponsored by the Government of Costa Rica in the
latter half of the 1880s were crucial for encouraging national and international
interest in the country’s archaeological patrimony. Kandler (1987) indicates Costa
Rica’s first National Exposition was held in 1885 and that others followed yearly,
in preparation for the Universal Exposition at Paris in 1889, and subsequently the
Exposicion Historico-Americana at Madrid in 1892, and the World’s Columbian
Exposition at Chicago in 1893. One of the reasons for incorporating artifacts in
the national expositions was to develop and refine a systematic way of presenting
Costa Rica’s archaeological heritage to the international audiences (Garron de
Doryan, 1974:23, 26). The national expositions also promoted the creation of the
Museo Nacional in 1887, an idea first proposed in 1862 (Peralta and Alfaro, 1893:
xxix-xxx). Archaeology unquestionably was part of the nationalistic agenda of
the times (Corrales Ulloa, 1999:6; Lines, 1934:5; Viales Hurtado, 1997:102-103).
Costa Rica disseminated information about the country and exhibited a variety
of its products at these events, but
... it was the archeological collection that aroused the greatest interest
on the part of the international public. For many, it was the first time
to be able to view these artefacts and they were admired for their ex-
2002 Watters and Fonseca Zamora — C. V. Hartman’s Anthropological Career 269
ceptional beauty and the mystery which surrounded them. (Kandler,
1987:24).
Hough (1893:273-274), clearly impressed with the Costa Rican antiquities he
observed in Madrid, reported the exhibits occupied two halls and 40 cases, dis-
played pottery, stone carvings, gold objects, jade carvings, and ornamented me-
tates, among other artifacts, and were augmented by paintings of the excavations,
maps, and photographs.
Anastasio Alfaro, head of the National Museum, was largely responsible for
the success of Costa Rica’s involvement at the international expositions, following
up on his efforts with the national expositions. His crucial role in the international
effort is clear: “By government decree, Don Anastasio was appointed to organize
how the country should be represented at these events” (Kandler, 1987:24). Al-
faro’s excavations in 1891 at the Guayabo de Turrialba site secured artifacts for
these expositions, and that fieldwork earned him a silver medal award at Madrid,
while the National Museum received a gold medal (Garron de Doryan, 1974:34;
Stone, 1956:12). He authored the exhibition catalog (Peralta and Alfaro, 1893)
with Costa Rican historian Manuel M. de Peralta. Alfaro is characterized as one
of Costa Rica’s two pioneering professional archaeologists (Hartman is the other)
by Fonseca Zamora (1984). The Madrid Exposition included Costa Rican eth-
nographic materials, obtained from the Talamanca Indians by Swedish naturalist
Carl Bovallius in 1882 (Brunius, 1992:68).
World’s Columbian Exposition
Alfaro accompanied the Costa Rican collection from Madrid to Chicago in
1893 and reinstalled it in the Anthropological Building at the World’s Columbian
Exposition, following the guidelines provided by Frederic Ward Putnam, head of
its Department of Ethnology (Garron de Doryan, 1974:26-38, 96-98). Department
M of the Exposition was often referred to as the Department of Ethnology even
though it included anthropology and history; the Anthropological Building con-
tained exhibit areas devoted to ethnology, archaeology, and ancient religions,
games, and folk-lore, in addition to anthropological laboratories and a library
(Anonymous, l^93a\ Flinn, 1893:52-59).
Much to Alfaro’s displeasure, Chicago had insufficient space for the 7,000
Costa Rican artifacts displayed in Spain and 3,000 had to be shipped back to the
National Museum in San Jose (Stone, 1956:13). Nonetheless, the antiquities ac-
tually exhibited were highly regarded and Costa Rica’s exhibit is twice noted in
the Exposition’s official guide (Flinn, 1893:53, 134). Starr (1893) confirms that
much of what Hough (1893) saw in Madrid had been reinstalled in Chicago and
his report not only lauds the overall exhibit, but also recognizes Alfaro and Peralta
by name.
Costa Rica’s participation at the World’s Columbian Exposition was a fortuitous
circumstance for Hartman:
After the conclusion of Dr. Lumholtz’ first expedition [what Lumholtz
terms the second expedition, ending in 1893] I went with him to the
Columbian Exhibition [sic] at Chicago, where I remained six months in
the Anthropological Department, arranging exhibits and assisting in the
purchase and packing of ethnological collections. (Hartman Archives,
letter, February 20, 1903, C. V. Hartman to W. J. Holland).
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Annals of Carnegie Museum
VOL. 71
It is likely that Lumholtz either arranged for or facilitated Hartman’s employment
at the World’s Columbian Exposition. Hartman’s presence there directly influenced
his career change to anthropology because he met Alfaro and Peralta and observed
Costa Rican antiquities for the first time. Both events would affect his life dra-
matically within a short time.
The Hartman Archives contain an interesting commentary concerning an at-
tempt to secure Costa Rican artifacts for the Columbian Exposition. Fourteen
years after the event, in a letter to the Chairman of the Carnegie Museum Com-
mittee, Hartman wrote:
Other collectors, but with less practical training, have tried the Costa
Rican field but with little success. During the preparations for the
World’s Fair in Chicago a collector was dispatched by Professor Putnam
and provided the means for securing archeological material from Costa
Rica. He spent several months in the country but was able to pick up
a few specimens here and there, altogether two or three small boxes.
(Hartman Archives, letter. May 27, 1907, C. V. Hartman to C. C.
Mellor).
Putnam’s attempt to obtain other Costa Rican artifacts seems unwarranted in view
of the large exhibition Alfaro mounted subsequently in Chicago. His effort makes
more sense, however, when one remembers that many of the artifacts he assem-
bled for the Exposition eventually formed the Field Columbian Museum’s an-
thropological collections. Alfaro’s artifacts would be unavailable to the museum
because they were going back to Costa Rica.
The Columbian Exposition provided Hartman with the chance to become ac-
quainted with major figures in the field, among them Putnam and his assistant
Franz Boas. The concurrent International Congress of Anthropology, convening
in Chicago from August 28 to September 2, afforded an even greater opportunity
to meet anthropologists and hear them lecture on a wide range of topics. Lumholtz
must have willingly acquainted his colleagues with his longest serving field as-
sistant, whom he had just accompanied from Mexico to Chicago. The Mexican
expedition was the subject of Lumholtz’s lecture, “Cave Dwellers of the Sierra
Madre” (Holmes, 1893:427). Among the many North Americans attending the
Congress, aside from Putnam and Boas, were Daniel G. Brinton, Otis T. Mason,
Zelia Nuttall, Frank Cushing, Alice C. Fletcher, George A. Dorsey, W. H. Holmes,
Stewart Culin, and Walter Hough; Latin America was represented by M. A. Muniz
and Emilio Montes (Peru), Emil Hassler (Paraguay), Manuel M. de Peralta (Costa
Rica), and A. Ernst (Venezuela) (Anonymous, 1893^). Fowler (2000:211) men-
tions that A. M. Stephen presented a paper in Chicago, although at the Interna-
tional Folk-Lore Congress, not at the Congress of Anthropology. It thus seems
likely that Hartman, Lumholtz, and Stephen were together again at least one time
after the Lumholtz expedition ended.
Hartman benefited from this Exposition in other ways. He gained practical
experience in the mounting of anthropology exhibits. His previous anthropological
background was restricted to fieldwork and his familiarity with archaeology lim-
ited to northwest Mexico, but by the time he returned to Sweden in 1894, his
anthropological experience was broader and his network of personal contacts wid-
er. A Swedish scholar attests to the central role this played in broadening Hart-
man’s exposure to anthropology, in stating that his “ . . , interest in the region
[Spanish America] was further strengthened at the World’s Fair in 1893 in Chi-
2002 Watters and Fonseca Zamora — C. V. Hartman’s Anthropological Career 271
cago, where New World archaeology and ethnography were given considerable
attention” (Brunius, 1985:152).
Louisiana Purchase Exposition
Hartman attended the Louisiana Purchase International Exposition in St. Louis
in September 1904, while employed as a curator at Carnegie Museum, and the
activities in which he participated are detailed in his report to the Director:
... I spent one week in St. Louis, where I studied the various ethno-
logical exhibits and attended the meetings of the Ethnological and Ar-
chaeological sections of the “Congress of Arts and Science.” At one
of the meetings I read a paper on “Recent archaeological investigations
on the Pacific Coast of Costa Rica.” For the Museum I purchased from
the Coco-Maricopa Indians at the Fair (for $3) a small collection of
their pottery and the peculiar implements they use for its manufacture.
(Hartman Archives, Report from the Ethnological Department for the
month of September 1904, C, V. Hartman to W. J. Holland).
Hartman probably referred to the Cocopa Indians, a group from Baja California
forming “ . . . a part of the anthropological exhibit ...” (Louisiana Purchase Ex-
position Co., 1904:285), although the Maricopas also were present (Fowler, 2000:
215). W J McGee (he did not use periods after his initials), formerly with the
Bureau of American Ethnology and known to Hartman from previous meetings,
had become head of the Anthropology Department at the St, Louis exposition by
1903 (Watters and Fowler, 2002). Rydell, Findling, and Pelle (2000:54) claim that
this exposition had the most anthropological exhibits of any world’s fair and credit
McGee, a believer in living exhibits, with creating an “outdoor laboratory for
anthropological fieldwork.”
Hartman’s standing as an anthropologist was enhanced by the eminent German
anthropologist, Eduard Seler, who lauded his Costa Rican research in an invited
paper, “The Problems of Archeology,” which reviewed the status of archaeology
across the Americas. He stated:
A limited region, including the old settlements on the slopes of the
volcano of Irazu and certain groups of hills which extend down into the
Atlantic lowlands, has lately been investigated in a really exemplary
manner by E. [sic] V, Hartman .... Outside of this, to be sure, we still
lack excavations [in Central America] undertaken in a scientific manner
and authenticated by documents. (Seler, 1906:536).
Hartman had known Seler for at least a decade before the St. Louis meeting. Seler
(1904) reviewed Hartman’s first Costa Rican monograph and provided one of the
“excellent testimonials” that Hartman submitted to Holland while negotiating
employment in 1903. McGee and Boas also were invited speakers.
Our research has been unable to confirm that Hartman actually read his paper
at the Congress. Rogers (1905-1907) produced an eight- volume set of proceed-
ings of the Congress of Arts and Science. The volumes relevant to invited an-
thropological papers, in volume V (Rogers, 1906:513-571), and “short papers,”
in volume I (Rogers, 1905:82-84), do not mention a paper by Hartman; nor does
his name appear in volume VIII (Rogers, 1907), the index compiled for the eight-
volume set. It seems that despite his claim to Holland, he did not “read” his
272
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paper (in a formal, officially sanctioned sense), although he may have informally
discussed or otherwise provided information about his Costa Rican research while
at the Louisiana Purchase Exposition. Nevertheless, he once again effectively used
his involvement with an exposition as a means of interacting with major figures
in anthropology.
Associations
Hartman’s development as a professional anthropologist was enhanced by his
involvement with three scholarly associations, two of which were national orga-
nizations but with extensive links abroad and one that was explicitly international
in scope. His talks at their meetings covered archaeological, ethnographic, and
linguistic topics and his articles, sometimes published in their proceedings and
journals, dealt almost exclusively with his anthropological research in the western
hemisphere.
Swedish Society for Anthropology and Geography
The Svenska Sallskapet for Antropologi och Geografi (SSAG) was the primary
scholarly association promoting the interests and interactions of Americanist re-
searchers in Sweden during Hartman’s career. His involvement with the SSAG
occurred at a time crucial for his transition from botanist to anthropologist in the
mid- 1890s and gave him access to influential Swedish Americanists who would
guide his career. Hjalmar Stolpe, who in 1873 enthusiastically endorsed the found-
ing of this society, then known as the Antropologiska Sallskapet (Brunius, 1990),
played significant roles in mentoring Hartman and advancing his anthropological
career two decades later. The SSAG was involved with the General Ethnographic
Exposition organized by Stolpe in Stockholm in 1878-79 and the Madrid Ex-
position of 1892, along with its antecedent 9th Session of the International Con-
gress of Americanists held in Huelva, Spain (Alvarsson and Brunius, 1994:43;
Brunius, 1990, 1992).
The SSAG served as titular sponsor of the Central American expedition (1896-
99) funded by Ake Sjogren. Hartman served as an SSAG delegate at the 13th,
16th, and 18th Sessions of the International Congress of Americanists; at the 15th
Session he is listed as the Swedish government delegate though not specifically
the SSAG representative.
Hartman published three (1895, 1901/?, 1902) of his first five articles on an-
thropological topics as well as lesser contributions such as book reviews, obitu-
aries, and meeting reports in Ymer, the SSAG journal, (Eranzen, 1969). The SSAG
appointed W. J. Holland a corresponding member within a year of his having
hired Hartman at Carnegie Museum (Anonymous, 1904r/).
International Congress of Americanists
If one factor were to be singled out as the most important influence in ad-
vancing Hartman’s anthropological career, it would be his participation in the
International Congress of Americanists (ICA), a scholarly association with which
he remained involved for thirty years. It seems fitting that he attended his first
and last ICA sessions in Sweden. At his first Session, the 10th ICA in Stockholm
in 1894, he was a relatively young person (32 years old) about to embark on a
new profession. By his final Session, the 21st ICA held in Goteborg in 1924, he
2002 Watters and Fonseca Zamora — C. V. Hartman’s Anthropological Career 273
was 62 years old and had enjoyed a career that was international in scope, inti-
mately linking Sweden with Central and North America.
Hartman’s participation in ICA Sessions, in a variety of roles, was tracked by
reviewing the pertinent proceedings, and his publications therein were checked
against the enormously valuable ICA centenary volume compiled by Comas
(1974). The formal convening of the International Congress of Americanists is
termed a Session. Sessions and their proceedings are numbered sequentially since
the 1st Session was held in Nancy, France in 1875 (see Comas, 1974:14, 117—
132, for lists of the sessions and proceedings). For convenience and brevity, we
have opted in this paper to drop the word Session and use the numerical designator
and the abbreviation ICA when referring to a particular Session (e.g., 13th ICA),
Depending on the roles played, an individual’s name can recur in various cate-
gories in the proceedings — Committee of Organization, Delegates of Govern-
ments, Delegates of Scholarly Societies, Members of Congress, and Subscribers —
although one category sometimes subsumes another in certain proceedings. A
Member who actually attended a Session can be confirmed, but only for selected
proceedings, by the presence of an asterisk beside the name. Most of the earlier
proceedings fortunately include titles of all papers presented at a Session (some-
times even those read “by title” only), irrespective of whether they later were
published in the volumes. With these constraints in mind, it is feasible to docu-
ment fairly completely Hartman’s involvement with the International Congress of
Americanists.
10th International Congress of Americanists (Stockholm, 1894). — The 10th
ICA was held in August of the same year Hartman returned to Sweden, having
completed his first anthropological fieldwork with Lumholtz and participated in
his first international exposition in Chicago. He was fortunate to be in Sweden
because Lumholtz, who was in Mexico on his third expedition, declined an in-
vitation to attend this Session and asked him to be his substitute and speak about
their Sierra Madre fieldwork. His first anthropological paper, “The Indians of
North-Western Mexico,” was presented to an international audience of anthro-
pologists and published in Swedish and English (Hartman 1895, 1897). The 10th
ICA proceedings volume (p. xvi) lists him as a naturaliste voyageur in the roster
of Swedish members, implying he was not yet regarded as an anthropologist.
Personal contacts he made at this ICA Session ultimately became more impor-
tant than his paper for his anthropological career. Costa Rican participants in-
cluded Anastasio Alfaro and Manuel M. de Peralta, whom he had met just the
year before in Chicago, and Bernardo Augusto Thiel, a Bishop who had assembled
a large collection of antiquities while serving in that country. German members
included H. Polakowsky, who had traveled to Costa Rica and written about its
antiquities, and the anthropologists Seler, Adolph Bastian, and Rudloph Virchow.
Americans included Brinton, Nuttall, and the Due de Loubat. However, it was the
Swedish members, especially Hjalmar Stolpe and to a lesser degree Carl Boval-
lius, collector of the Talamanca ethnographic materials exhibited in Madrid in
1892, who most directly influenced Hartman’s transition. Hjalmar Sjogren is listed
as a member from Sweden but his son, Ake Sjogren, is not. However, Ake Sjogren
later was appointed Interim Secretary General and it fell to him to assemble the
proceedings and write its forward. Hartman departed on the Ake Sjogren-funded
Swedish expedition to Central America less than two years after the 10th ICA
ended.
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13th International Congress of Americanists (New York City, 1902). — Hartman
established his stature as a professional anthropologist in the eight years elapsing
between the International Congress of Americanists meetings in Stockholm and
New York. The 13th ICA proceedings reveal that he was no longer regarded as
a naturaliste voyageur. Instead, he and Ake Sjogren are listed as official repre-
sentatives of the Swedish Society of Anthropology (actually SSAG by then);
Hjalmar Stolpe was the delegate of the government of Sweden and a Vice Pres-
ident of the ICA. W. J. Holland was a delegate of the United States and C. C.
Mellor, Chair of the Museum Committee, was listed as a subscriber (Watters,
2002a). Holland spoke about Carnegie Museum, then in its “infancy,” issued an
invitation for delegates to visit Pittsburgh, and presented a paper on petroglyphs
in Pennsylvania (Holland 1905^2, 1905Z?). Delegates or members from Costa Rica
included Henri Pittier de Fabrega and Juan Fernandez Ferraz. Delegates repre-
senting French societies included the Due de Loubat for the Societe des Ameri-
canistes de Paris, and Yale University professor George Grant MacCurdy, for the
Societe d’ Anthropologic de Paris. Seler was a German delegate. Franz Boas,
Stewart Culin, J. Walter Fewkes, Walter Hough, A. L. Kroeber, Otis T. Mason,
W J. McGee, Zelia Nuttall, George Pepper, and F W. Putnam were among the
many members (either delegates or subscribers) representing anthropology in the
United States.
His participation in the 13th ICA strengthened relations with some major figures
in anthropology and introduced him to other persons important to his career.
Along with Boas, Pittier de Fabrega, Fernandez Ferraz, Holland, MacCurdy, and
Seler, among others, he served on the Council of the Commission of Organization,
The Due de Loubat was its Honorary President; Stolpe and Putnam were Vice
Presidents.
The 13th ICA proceedings (pp. Ixivi-lxvii) disclose that he read one paper,
“Archaeological Researches in Costa Rica,” in full, and a second paper, “The
Aztecs of Salvador,” by title only. Neither was published in the proceedings.
Hartman’s status as a professional anthropologist was enhanced greatly when
it was made known that he had been awarded the prestigious Due de Loubat prize
by the Royal Academy of Belles Lettres, History and Antiquities of Sweden.
Hjalmar Stolpe (1905) announced this award in a speech to the 13th ICA mem-
bers, which appears in the proceedings. The Loubat award was bestowed for
Archaeological Researches in Costa Rica, Hartman’s monograph (190 la) based
on the Swedish expedition. This sumptuous volume, the cost of which had been
borne by Hartman’s benefactor, Ake Sjogren, subsequently was “laid upon the
table” for the Congress to examine.
After Stolpe ’s speech, Putnam called for a resolution of congratulations for
Hartman. The resolution’s text is not included in the proceedings but selected
passages were extracted by Hartman and included in his letter to Mellor five years
later:
Resolved, that the Members of the 13th International Congress of Amer-
icanists, assembled in New York, hereby express their hearty appreci-
ation of the results obtained by the Archaeological Expedition to Costa
Rica under the direction of C. V. Hartman . . . and they congratulate Mr.
Sjogren upon the magnificent manner in which the Report has been
published. [They further characterize this volume as] . . . the most pains-
taking and elaborate Report of the exploration of ancient graves in Cen-
2002 Watters and Fonseca Zamora — C. V. Hartman’s Anthropological Career 275
^ AiVlERICAINISTS lINVADE PIT'^BURO * ^
Fig. 3.^ — Photograph from the Dispatch (October 30, 1902 edition, page 8) of the visiting delegation
of the International Congress of Americanists at the Hotel Schenley. (Carnegie Museum of Natural
History Archives, Holland newspaper clipping scrapbook).
tral America which has ever been undertaken . . . [and add] . . . the beau^
tiful volume will always serve as a model for this class of archaeological
work. (Hartman Archives, letter, May 27, 1907, C. V. Hartman to C. C.
Mellor, citing passages extracted from the Putnam resolution).
According to Hartman, the resolution was proposed by Putnam, seconded by
Boas, and adopted unanimously by the Congress. Being recognized personally
and having his research honored in this manner by his fellow anthropologists
surely was enormously satisfying for Hartman who had commenced his transition
to anthropology with the heldwork he conducted on the Lumholtz expedition only
twelve years before.
The 13th ICA proceedings volume discloses that a delegation traveled “ on
an excursion to Chicago, including visits to Philadelphia, Washington, Pitts-
burg[h], and Tort Ancient,’ in southern Ohio and Cincinnati” (p. Ixvii) the week
after the Congress. Hartman, Stolpe, Pittier de Fabrega, Fernandez Ferraz, and
Seler were among the delegation of foreigners (Fig. 3) welcomed by Holland
when they visited Pittsburgh on October 29, 1902. Pittsburgh newspapers disclose
that 3 1 dignitaries arrived in the city, toured Carnegie Institute, and heard Holland
speak of his vision for Carnegie Museum, and further state that the excursion was
to be extended to St. Louis following the visit to Chicago (Watters, 2Q02a).
15th International Congress of Americanists (Quebec City, 1906). — Canada’s
hosting of the 15th ICA resulted from a resolution adopted six years earlier, at
the 12th ICA in Paris, whereby Sessions were to alternate between venues in
Europe and the Americas, with the 13th ICA in New York inaugurating this
scheme in 1902. Hartman’s presence in Pittsburgh caused him to miss the 14th
ICA in Stuttgart, Germany but, conversely, facilitated his attendance at the 15th
ICA in Quebec,
The 15th ICA proceedings list Hartman as the lone delegate for the government
of Sweden, a member of the Council, and a curator of Carnegie Museum. Seler
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again was a delegate of the government of Germany (actually listed as Prussia)
and a Vice President of the ICA. Delegates from scholarly organizations in the
United States included Boas, W. H. Holmes, and Clark Wissler. Fewer interna-
tional members were present and fewer countries represented at Quebec than at
the New York Session of 1902. Costa Rica and many other Latin American coun-
tries had no representation.
At the 15th ICA, Hartman informed the Congress of the death of his mentor
and friend, Hjalmar Stolpe, who had died the year before (Culin, 1906), and it
seems that eulogizing Stolpe was the major reason he attended. He participated
in meetings of the Council and presided over one morning session, but seems not
to have presented a paper since none is listed in the 15th ICA proceedings.
16th International Congress of Americanists (Vienna, 1908). — Hartman’s return
to Sweden in 1 908 following his resignation from Carnegie Museum explains his
attendance at the next European venue of the ICA, at the 16th Session held in
Vienna, Austria in September. After this final trans-Atlantic move, Hartman’s
employment and professional life forever after were centered in Sweden.
His participation in the Vienna Session, as the delegate of the government of
Sweden, representative of the SSAG, an attending Member, and, for the first time,
a Vice President (along with Seler), facilitated his reentry into European anthro-
pology and museology.
He presented two papers. ‘'Some features of Costa Rican Archeology” com-
pared the architectural forms and geographic distributions of four kinds of graves
investigated during his two expeditions. The second paper dealt with a sound
recording device called the photographone, invented by his fellow Swede, Sven
Berglund. Both papers were published in the 16th ICA proceedings (Hartman,
1910<2, \9\0b). His Costa Rica article is listed as “an abstract” and unquestion-
ably was an abbreviated version of a longer lecture illustrated with lantern-slides.
He never published the full version of this paper to the enduring frustration of
persons interested in his Costa Rica research. He also laid upon the table for
examination by the Congress his two recent publications, the second Costa Rican
monograph, published by Carnegie Museum, and his article in American Anthro-
pologist (Hartman, 1907 <2, 1907^).
Other International Congress of Americanists Sessions. — Hartman attended lat-
er Sessions of the International Congress of Americanists but participated less
actively than in the past, possibly due to deteriorating health. He had experienced
health problems while at Carnegie Museum and in several monthly reports to the
Director he mentioned missing work because of illness. The underlying cause of
his impaired health may have been malaria. In his 1907 letter to C. C. Mellor, he
mentions his outlays for bills from doctors resulting from his anemic condition
caused by months of work in malarial regions during the rainy season. Comments
in the 18th and 19th ICA proceedings suggest deteriorating health, and we know
he was granted a medical leave of absence from the Ethnological Section in 1923,
five years before he retired (Lindblom, 1941).
At the next European Session (18th ICA), in London in 1912, he is listed as a
delegate for the government of Sweden and for the SSAG, an attending Member,
and a Council member. However, the 1 8th ICA proceedings, without further ex-
planation, state (p. xl): “Professor C. V, Hartman, Stockholm, was unable to
present his paper ‘Physical Anthropology of the Aztecs of Salvador.’” Immedi-
ately after the ICA, he stayed on in London to attend an International Conference
2002 Watters and Fonseca Zamora — C. V. Hartman’s Anthropological Career 277
Fig. 4. — The Carnegie Institute building (left), which contained Carnegie Museum, and the Hotel
Schenley (right) were situated in the predominantly rural setting of the Oakland district of Pittsburgh
when Hartman was Curator. The twin towers, features of the original (1895) building, were removed
during the 1907 addition. (Carnegie Museum of Natural History Archives).
in which a proposal for an International Anthropological Congress was being
debated. No remarks are attributed to him in the summary of that meeting in the
18th ICA proceedings.
Hartman did not attend the 19th ICA in Washington, D.C., which took place
in 1915 when Europe was embroiled in World War I but before the United States
entered. In the proceedings, the Swedish delegate, Amandus Johnson, extended
his well wishes to the Congress and added (p. xlvi): “Professor Hartman would
have been with us except for his health and the unfortunate state of affairs in
Europe.”
The 21st ICA included two Sessions held in August 1924, first at The Hague
in the Netherlands and then at Goteborg in Sweden. Hartman attended the Go-
teborg Session and was listed as a member of the Commission of Organization,
a member of the Council, and an attending member, but no longer as a delegate
of the government of Sweden or the SSAG. Comas (1974) lists no Hartman papers
or publications for the 21st ICA, at either session. The 1924 Goteborg Session,
the final one he attended, occurred thirty years after his first Congress, the 10th,
in Stockholm.
American Anthropological Association
The American Anthropological Association (AAA) convened its founding meek
ing on June 30, 1902 in the Oakland Methodist Episcopal Church in the Oakland
district of Pittsburgh (Fig. 4), almost exactly four months prior to Hartman’s visit
to Pittsburgh on the excursion following the 13th ICA in New York City. The AAA
founding meeting was held in conjunction with the 51st annual meeting of the
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VOL. 71
American Association for the Advancement of Science (AAAS), which was being
attended by anthropologists from its Section H (Anthropology) (Watters, 2002b;
Watters and Fowler, 2002). W J. Holland’s courtesy to Section H, by arranging for
Carnegie Museum to exhibit a Mexican sculpture discussed by Marshall Saville in
the scientific sessions, was acknowledged by Smith (1902:210) and McGee (1902:
477). Holland did not participate in the founding meeting but soon afterward was
accredited as being an AAA founder (Anonymous, 1903:191).
Hartman became involved with the American Anthropological Association dur-
ing his tenure at Carnegie Museum but the Hartman Archives contain few details
about the extent of his participation. His attendance is documented so far only
for the AAA meeting in 1905, held from December 27-29 at Cornell University
in Ithaca, New York. The meeting provided Hartman with an excellent opportunity
to broaden his network of contacts, because it was held jointly with the Archae-
ological Institute of America, American Folk-Lore Society, and American Phil-
ological Association. Prior to his departure, he informed Holland of his planned
activities:
With the Director’s kind permission, I leave the 25th for Ithaca, where
I at the meeting of the Am. Anthropological Association will read a
paper entitled “Use and ornamentation of the tree-calabash in tropical
America.” I will also attend the sessions of the “Committee for the
preservation of the antiquities of the United States,” which has honored
me with membership. (Hartman Archives, Report from the Section of
Ethnology for the month of December 1905, C. V. Hartman to W J.
Holland).
His paper, one of only nine read in full at the AAA meeting, was published
subsequently in the Boas Anniversary Volume (Hartman, 1906). He illustrated his
talk with the stereopticon and exhibited a number of calabash specimens according
to MacCurdy’s (1906) summary of the meeting. Participants he already knew
included Boas, who presided in the absence of AAA President Putnam, and
MacCurdy, the AAA Secretary, and immediately following the Ithaca meeting,
he traveled to New York and New Haven to consult with them about his Costa
Rican research. MacCurdy (1906) also reported at the Ithaca meeting on the plans
for AAA participation at the 16th ICA in Quebec. The AAA provided European
anthropologists in Quebec with voluminous information about “Recent Progress
in American Anthropology” (Anonymous, 1906).
Hartman served on the American Anthropological Association’s Committee for
the Preservation of American Antiquities, one of the committees that ultimately
lobbied successfully for enactment of the Antiquities Act of 1906. He published
two articles in American Anthropologist (1907Z?, 1907c), one on his Costa Rican
research and the other on engraved shell artifacts from Tennessee in the Carnegie
Museum collections.
Museums
Hartman’s anthropological career was centered in museums, not in universities.
He was employed in museums; his two monographs on Costa Rican archaeology
were museum publications; field methods he used in Costa Rica were learned
from a museum anthropologist; and his professional colleagues were anthropol-
ogists associated primarily with museums (including university museums). The
Hartman Archives reveal that he performed the full gamut of museum duties
relating to exhibition and collections while at Carnegie Museum. However, his
2002 Watters and Fonseca Zamora — C. V. Hartman’s Anthropological Career 279
five=year (1903-1908) term of employment is more appropriately perceived as a
time span intervening between periods of employment at the Naturhistoriska Riks-
museet (Royal Museum of Natural History) in Sweden.
Ethnographical Department of the Royal Museum of Natural History of Sweden
Hjalmar Stolpe was the key person responsible for mentoring Hartman in mu-
seum anthropology and for involving him with the Swedish Museum of Natural
History. Both were trained in the natural sciences, Stolpe in zoology and Hartman
in botany, yet each later embraced anthropology as his life’s work and ultimately
ended his career as a museum administrator, rightfully having achieved recogni-
tion as an archaeologist as well as an ethnologist (Brunius, 1984; Culin, 1906;
Lindblom, 1941). Hartman’s association with Stolpe began at least as far back as
1894, when both participated in the 10th ICA in Stockholm.
Soon after the 10th ICA, Stolpe introduced Hartman to Ake Sjogren and it is
clear that Sjogren’s interest in Costa Rica’s archaeology was the determining fac-
tor in Hartman’s decision to conduct research in that country (Stolpe, 1905).
Undoubtedly even more compelling was Sjogren’s willingness to fund the ex-
panded (in anthropological and geographical scope) Central American expedition
of 1896-1899. Hartman modeled his field methods for excavating Costa Rican
burial grounds on Stolpe’s meticulous excavation and mapping of Swedish Iron
Age cemeteries at Bjorko and Vendel (Alvarsson and Brunius, 1994:43-45; Brun-
ius, 1984; Rowe, 1959). He acknowledged this debt: “Having received thorough
instructions from Dr. Hj. Stolpe in the investigation of graves, according to his
well-known method for similar research in Sweden, I started for Costa Rica in
the spring of 1896” (Hartman, \90\a:\). After the Central American expedition
concluded, Hartman traveled for six months in the United States visiting the major
museums in order to prepare himself for museum work.
Hartman’s return to Sweden in 1899, following the three-year Central American
expedition, was auspicious because Stolpe was achieving his long-term goal of
creating a separate Ethnographical Department (sometimes termed Ethnographical
Museum) within the Royal Museum of Natural History. The department came
into being in 1900 and Stolpe was appointed its Director (Brunius, 1990). Stolpe
hired Hartman as his assistant, a position affording him opportunities to perform
curatorial duties, develop exhibits, catalog collections, and write up his research.
This appointment provided Hartman with his first true museum experience, al-
though he had been involved earlier in exhibit development at the World’s Co-
lumbian Exposition.
Hartman acknowledged his indebtedness for Stolpe’s training:
In Stockholm I undertook new training in the various duties of a mu-
seum worker under the immediate supervision of Professor Hjalmar
Stolpe. I classified, arranged and labelled anew my collections and wrote
descriptive catalogues. At the same time I had the opportunity day after
day of following in detail the work Dr. Stolpe was doing in the different
departments,. at that time especially valuable and instructive, on account
of the complete reorganization of the institution then going on, accord-
ing to thorough scientific and systematic plan. Although the experience
thus gained during the three years of daily intercourse with Professor
Stolpe does not seem to cover a very long time I consider the same
really valuable, because my instructor is undoubtedly one of the fore-
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VOL. 71
most museum men of Europe, having worked some 30 years inside the
walls of the National Museum in Stockholm . . . and having studied
nearly all the museums of Europe, North & South America, and Asia,
(Hartman Archives, letter, February 20, 1903, C. V. Hartman to W. J.
Holland).
Within a single decade, the 1890s, Hartman had transformed himself from a
botanist on the Lumholtz expedition to a professional anthropologist employed at
a major natural history museum in Europe. Within a year of his museum em-
ployment in Sweden, he mounted a major exhibit of the Central American ar-
chaeological and ethnographic materials that his benefactor, Ake Sjogren, by then
had donated to the Ethnographical Department (Anonymous, 1900). By 1902, he
(1901«, 1901Z7, 1902) had produced a monograph in English and two articles in
Swedish about his Costa Rican archaeological and Salvadoran ethnographic re-
search and had been honored with the Due de Loubat award.
His research had been focused almost exclusively in the New World, yet his
museum employment remained firmly rooted in the Old World until Carnegie
Museum hired him in 1903. Five years later he returned to Sweden to take over
the position of Director of the Ethnographical Department, vacant since the death
of his mentor, Hjalmar Stolpe in 1905. Thereafter, he became a full-time museum
administrator whose duties precluded further productivity in the realms of research
and publication. While it is clear that he planned to publish more about his re-
search in Costa Rican archaeology and Central American ethnology, these plans
failed to come to fruition for the most part. He took a medical leave of absence
from the Ethnographical Department in 1923, from which he entered retirement
in 1928; he died in 1941 (Lindblom, 1941).
Carnegie Museum of Carnegie Institute
Hartman attended the 13th ICA with the intent of seeking a position at a mu-
seum in the United States. During the succeeding three months, he used his net-
work of contacts with American anthropologists to explore employment oppor-
tunities at museums in New York, Washington, and Philadelphia before he ap-
proached Carnegie Museum. He was seeking a museum position at the same time
Holland (Fig. 5) sought an anthropology curator, yet neither was the first choice
of the other. Holland had been unsuccessful in his effort to woo Stewart Culin
from the Museum of the University of Pennsylvania in Philadelphia; instead,
Culin went to the Brooklyn Institute of Arts and Sciences, now the Brooklyn
Museum, as its first Curator of Ethnology (Fane, 1991). Culin declined Holland’s
offer during the first week of February 1903, thus opening the door for his serious
consideration of Hartman’s application letter of January 28. Thereafter, negotia-
tions progressed rapidly and Hartman was employed exactly one month later
(Watters and Fonseca Zamora, 200 IZ?).
Holland’s assessment of Hartman’s qualifications for becoming Curator of the
Section of Ethnology and Archaeology was quite favorable. In his Monthly Report
to the Museum Committee, delivered February 28, 1903, Holland wrote:
Dr. C. V. Hartmann [sic], whose acquaintance I formed at the recent
Congress of Americanists, is in the building, and 1 shall take pleasure
in presenting him to you at sometime this evening. He has had expe-
rience in Museum work, having been associated with Dr. Stolpe, the
celebrated Swedish ethnologist, whose assistant he was at Stockholm.
2002 Watters and Fonseca Zamora — C. V. Hartman’s Anthropological Career 28
Fig. 5. — William Jacob Holland was Director of Carnegie Museum during Hartman’s tenure (1903
1908). (Carnegie Museum of Natural History Archives).
He was for a long time with Dr. Lumholtz in Mexico and has written
extensively upon the ethnology of Costa Rica. It may be that he is the
man, for whom we are looking. He produces excellent testimonials.
(Holland Archives, Monthly Reports of the Director to the Museum
Committee, volume 1, March 1, 1903).
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VOL. 71
Among those “excellent testimonials” were letters from Boas at the American
Museum of Natural History, McGee at the Bureau of American Ethnology, Smith-
sonian Institution, and German anthropologists Eduard Seler and A. B. Meyer.
Holland offered and Hartman (who had traveled to Pittsburgh) accepted employ-
ment on Eebruary 28, the same day the Committee approved Holland’s recom-
mendation to hire.
In his speech at the 13th ICA in 1902, Holland (1905<2) alluded to Carnegie
Museum, then in existence for six years, being in its “infancy” (Watters, 2002a).
Thus, acquiring collections was a priority for Holland. Hartman, being aware of
the Director’s desire, emphasized during their negotiations his personal knowledge
of Costa Rican collections available for purchase (Eig. 6). He likewise was aware
of the plans afoot for the new Carnegie Museum, having heard Holland expound
upon his vision at the 13th ICA and afterward in Pittsburgh with the visiting
delegation. He contributed to the realization of that vision when the new facility
was inaugurated in 1907. His Costa Rican artifacts, displayed prominently in the
new Gallery of Archeology, were one of only two archaeology collections men-
tioned in the booklet (Pig. 7) prepared for that dedication (Carnegie Institute,
1907). In his letter to Mellor soon after the dedication, Hartman estimated about
2,000 Costa Rican artifacts were on exhibit in 1907.
Holland and Hartman experienced differences of opinion within a few months
of the hiring, and the difficulties resurfaced at various times during Hartman’s
employment. The fundamental issue lay in how each conceived of the curatorial
position. In Holland’s view, acquisition of collections and preparation of exhibits
were the priorities. Hartman consistently emphasized research, from the resump-
tion of his fieldwork in Costa Rica to conducting comparative studies at other
museums. The situation escalated on May 27, 1907 when Hartman wrote a twen-
ty-page letter stating his case for a salary increase to C. C. Mellor, Chairman of
the Museum Committee. The Committee referred the letter to Holland for action;
Holland’s response to the Committee was less than enthusiastic. Although a year
was to pass before Hartman resigned, he began to lay the groundwork for that
move soon after his request for a salary increase. In July 1907, he used his
accumulated vacation to visit Sweden for three months, after which he returned
to Pittsburgh for about six months, resigned from Carnegie Museum on May 1,
1908, and soon thereafter moved back permanently to Sweden, where he assumed
the directorship of the Ethnographical Department.
Hartman took advantage of the professional enhancements afforded by Car-
negie Museum, including his trips to the Louisiana Purchase Exposition in St.
Louis, the 15th ICA in Quebec, and the AAA meeting in Ithaca. He interacted
with anthropologists in the United States through a series of trips authorized by
Holland to facilitate access to library and collection resources not available at
Carnegie Museum. He reported on one trip:
Nearly whole the month of January I have spent in New York at the
private libraries of Professors M. H. Saville and A. E Bandelier of the
American Museum of Natural History and at the Lennox library, in
order to look up references regarding Spanish-American archaeology
and history. I have also consulted the collections from said countries in
the Museums of New York, Cambridge, New Haven, and Philadelphia.
(Hartman Archives, Monthly Report of the Section of Ethnology, Jan-
uary 1905, C. V. Hartman to W. J. Holland).
2002 Watters and Fonseca Zamora — C. V. Hartman’s Anthropological Career 283
SiBiS
0 I t 3 4 5
Fig. 6. — An elaborately decorated “jaguar metate” obtained by Hartman in Costa Rica in 1903 (Ac-
cession #2793/2076).
Other museum trips include visits to New York and Philadelphia in March 1904;
to the Bureau of American Ethnology, Washington, D.C. in August 1905; to New
York, New Haven, Washington, D,C., and Philadelphia from December 1905
through March 1906; and to New York in September 1906 after the 15th ICA in
Quebec.
The tension between Holland and Hartman surfaced during the latter’s lengthy
absence, between late December 1905 and March 1906, when he conducted re-
search at the American Museum of Natural History and Yale Peabody Museum
in consultation respectively with Boas and MacCurdy. Hartman requested and
Holland approved, albeit reluctantly, an extension to this trip. However, his pro-
284
Annals of Carnegie Museum
VOL. 71
M itime to arrange all its collections in the new buikliiifr-
but enough will be seen of the 1,300,000 obiects to give
the visitor some idea of the present scope and possibilities
rf,the Mu-seum - v/h;i-“ endea'^Diing tc inteiest and edu-
fe cate the masses in as many directions as possible, it aims
, g . also to thoroughly illustrate the surrounding Appalachian
p region, in its fauna and flora, its geology and mineral
resources; and to further scientific knowledge and re-
search, by its “Annals” and “Memoirs,” its library and
I?] systematic collections.
The hurried lay visitor may be aided by having his
p attention called to the following items:
; The Cabinet of Coins arid Gems contains a valuable
y collection of timepieces, some of them having very inter-
U esting historical associations. In the Hall of Geology and
y Mineralogy, the large and accurate Relief Map of Pitts-
burgh, and the collection of Minerals. Further on are
the gigantic skeleton of the Diplodocus Carnegiei; the
remarkable skull of the Triceratops; fossil fishes and
ii; reptiles; and types of recently discovered extinct mam-
tc- mals, many of them the results of special expeditions
p sent out by the Museum.
fe. The Gallery of Ornithology affords artistically and
S truthfully mounted groups of birds, both native and for-
eign, which constitute part of 27,000 specimens.
The rare specimen of White Rhinoceros, the life-like
groups of fur seals and sea lions, are seen in the Gallerv
ot Mammals; while butterflies, beetles and shells in end-
g:, less variety of color and form adorn the Gallery of Recent
Invertebrates.
g Botany is represented by 169,000 recent and fossil
P specimens, and by illustrations of plants and their uses.
U In the Gallery of Archeology will be found Etruscan
> and Costa Rican collections, and a boat from Egypt
l.j which has escaped the wreckage of several thousand
if
f; Habits and customs of Indians of the Plains of Alaska
jfo' ar>-. depicted in the Gallery of Ethnology. The evolution
of transportation and of various industries; collections of
coins, pottery, etc., can be studied in the .section of Me-
|:;i chanic and Useful Arts.
!■' Zbc Carneflfe ITecbnical Schools
• : founded by Andrew Carnegie, are located adjacent to the
® Carnegie Library and Institute. The City of Pittsburgh
M haa provided for them a site of 32 acres adjoining Schenley
CARNEGIE
INSTITUTE
A PITTSBURGH S
Fig. 7. — The cover of the 1907 dedication booklet and the page mentioning the Costa Rican collection
exhibited in the Gallery of Archeology.
tracted stay increasingly annoyed Holland, to the point that in early March he
ordered him to return to Pittsburgh. Between that trip and his resignation in 1908,
Hartman never again traveled under Carnegie Museum auspices to any museum
except for the brief visit to New York in September 1906 after attending the 15th
ICA. Ironically, besides conducting collection research, Hartman actually was
complying with Holland’s mandate to acquire collections. While visiting Clark
Wissler at the American Museum of Natural History, he made anangements for
a major exchange of artifacts between the two museums.
Carnegie Museum published Hartman’s (1907a) second Costa Rican mono-
graph in its Memoirs series and it is clear that Holland expected to publish more
monographs. When departing Carnegie Museum, Hartman promised he would
submit from Sweden the manuscript for his second Memoirs monograph dealing
with ceramic artifacts from Costa Rica, a text that was well along and possibly
even nearly completed. However, he never sent the manuscript and no draft ver-
sions are present in the Hartman Archives. A third Memoirs monograph was
planned or at least under consideration before he left Carnegie Museum.
His presence at Carnegie Museum afforded him the opportunity to expand his
2002 Watters and Fonseca Zamora — C. V. Hartman’s Anthropological Career 285
contacts within the field of museology and this network gained importance when
he was appointed Director of the Ethnographical Department in Sweden, He was
in Pittsburgh when the newly founded American Association of Museums (AAM)
met there in June 1907 to celebrate the dedication of the new Carnegie Institute
and in May 1908 he attended the AAM annual meeting in Chicago.
National Museum of Costa Rica
Hartman interacted with personnel at the Museo Nacional de Costa Rica
(MNCR) during the Swedish expedition (1896-1897) and Carnegie expedition
(1903). Anastasio Alfaro, known to Hartman through the World’s Columbian Ex=
position and the 10th ICA in Stockholm, headed the MNCR during the first visit.
Juan Fernandez Ferraz, whom he had known since the Swedish expedition, suc-
ceeded Alfaro in 1898 and headed the MNCR until 1904, according to Kandler
(1987:28). Henri Pittier de Fabrega, Swiss by birth, was Director of Costa Rica’s
Instituto Fisico-Geografico (IFG) and had been associated with the MNCR since
1888, being one of the first persons appointed to serve on its Board of Directors.
Fernandez Ferraz and Pittier de Fabrega represented Costa Rica at the 13th ICA
in New York, thereafter visited Pittsburgh with Hartman during the post-Congress
excursion, and were the recipients of the letters of introduction that Holland wrote
on Hartman’s behalf for the Carnegie Museum expedition (Watters, 2Q02a). Fer-
nandez Ferraz, a Spaniard, and Pittier de Fabrega, Swiss, exemplify the many
foreigners who moved to Costa Rica for career advancement (Gonzalez Flores,
1921).
A shared interest in and involvement with scientific research allied the MNCR
and IFG. Although normally independent entities, they were on occasions con-
solidated during various reorganization schemes (Anonymous, 1904Z?; Conejo
Guevara, 1975:26, 149-150; Stone, 1956:17). One such reorganization scheme
had taken place not long before the Carnegie expedition began:
A great advantage for me just now is, that one of my most intimate
friends. Professor H. Pittier de Fabrega, the head of the “Instituto Phys-
ico-Geographico” [sic] in San Jose has recently, when Dr. Juan Ferraz
was discharged, got the arch, museum affiliated with his institution. At
the Congress in N.Y. he urged me ardently to try to go down to C.R.
again. He has no time for arch, and he wants me during the rainy season
to arrange and classify the somewhat heterogenous [sic] collections in
the museum. (Hartman Archives, letter, January 28, 1903, C. V. Hart-
man to W. J. Holland).
Hartman’s statement discloses that the IFG under Pittier de Fabrega “affiliated”
the MNCR in 1903, a point conflicting with Kandler’s (1987:28) contention that
Fernandez Ferraz headed the MNCR until 1904. His statement would indicate
that it was Pittier de Fabrega, not Fernandez Ferraz, who arranged access to the
MNCR archaeological collections studied and photographed by Hartman in the
midst of the Carnegie expedition (Fig. 8). He used images of some MNCR arti-
facts for comparative purposes in his 1907 monograph. He also used his acquain-
tance with MNCR scientists (e.g., entomologist P Biolley) to obtain botanical and
zoological specimens that Holland desired for Carnegie Museum.
Pittier de Fabrega links with Hartman and with Costa Rican anthropology in
other ways. He wrote reports on languages and customs of Costa Rica’s indige-
nous peoples (Conejo Guevara, 1975:157) and was involved with mapping Costa
286
Annals of Carnegie Museum
VOL. 71
Fig. 8. — Artifacts photographed by Hartman in 1903 in the collection of the Museo Nacional de Costa
Rica. (Section of Anthropology glass-plate negative G484).
Rica, a project in the course of which he explored the entire country and routinely
encountered archaeological sites. Hartman attests to the close ties he maintained
with Pittier de Fabrega:
In case I can continue the investigations even during the following dry
season Professor P [Pittier de Fabrega] places all his knowledge about
ancient remains, rockcarvings, idols, etc. at my disposition. While en-
gaged since 20 years ago on the work on the map of C. R. he has
travelled almost all over the Republic. All arch, work that can be done
in C.R. ought to be carried out soon before the rush of laborers to
Panama [for canal construction] begins and wages rise. (Hartman Ar-
chives, letter, January 28, 1903, C. V. Hartman to W. J. Holland).
Hartman’s mounting of the expedition in 1903 was fortuitous because Pittier de
Fabrega soon thereafter left Costa Rica, having lived there for seventeen years,
and moved to the United States, where he worked for the U. S. Department of
2002 Watters and Fonseca Zamora — C. V. Hartman’s Anthropological Career 287
Agriculture starting in 1905 (Chase, 1950; Standley, 1950). Pittier de Fabrega is
accredited as a founding member of the American Anthropological Association
(Anonymous, 1903:191; recorded incorrectly as Fabrega, H. R de).
Though not connected to the MNCR, other foreigners residing in Costa Rica
were a valuable link for Hartman. He explained the ways they assisted his ex-
pedition:
My success in the excavations during this short period [the Carnegie
Museum expedition], I attribute in the first hand to my own local ex-
perience in the field; second, to the valuable friendships, previously
made both among the foreigners, Scandinavians, Americans and Ger-
mans, as well as among the natives in various places. Through their
generous help and disinterested assistance I was enable to obtain per-
mission from the owners of the land to excavate in various localities;
and owning to the hospitality and courtesy of various of these gentle-
men, my expenses were considerably lowered. (Hartman Archives, let-
ter, May 27, 1907, C. V. Hartman to C. C. Mellor).
Minor C. Keith, the American entrepreneur who built railroads in Costa Rica,
was an avid collector of antiquities and assembled a large collection while living
there (Stewart, 1964:160-168). Keith obtained antiquities from various sources
but especially at the site of Las Mercedes, situated on land he owned on the
Atlantic coastal plain (Mason, 1945; Spinden, 1915). With Keith’s authorization,
Hartman (1901s %■■.'■;■ f.* iii!:^ilr‘^,^ ', " ’■ "
* ■ ■ ^ - i -■. -ujt/u'tfT* ' '' ,5 '•ft*
'.' ' ■;/, , 'C
j. .. ' • ^' 1 . :.-vi ,n/
'-'f ' * " .V ^ •■ ; ■' :
INDEX TO VOLUME 71
CONTENTS
ARTICLES
New specimens of picromomyids (Plesiadapiformes, Primates) with description of a new species
of Alveojunctus Mary T. Silcox, Kenneth D. Rose, and Stephen L. Walsh 1
Review of the Neotropical genus Bredinia (Trichoptera: Hydroptilidae: Stactobiini) . . .
Steven C. Harris, Ralph W. Holzenthal, and Oliver S. Flint, Jr. 13
New species of microcaddisflies (Trichoptera: Hydroptilidae) from northern Florida .......
Steven C. Harris 47
A Miocene bone bed from Agate Fossil Beds National Monument
W. Orr Goehring and Mary R. Dawson 59
Comparing pottery from the Proto-historic McKees Rocks Village and Eisiminger sites of south-
western Pennsylvania Richard L. George 63
Fleas (Siphonaptera) from Ancash Department, Peru with the description of a new species,
Ectinorus alejoi (Rhopalopsyllidae), and the description of the male of Plocopsylla pallas
(Rothschild, 1914) (Stephanocircidae)
. . Michael W. Hastriter, Michael D. Zyzak, Ruler Soto, Roberto Fernandez, Nelson Solorzano,
and Michael F Whiting 87
Review of Leptomeryx (Artiodactyla, Leptomeryicidae) from the Orellan (Oligocene) of Ne-
braska . William W. Korth and Margaret E. Diamond 107
W. J. Holland’s speech at the International Congress of Americanists, 13th session, in 1902
David R. Watters 1 3 1
Revision of the New World abariform genera Neotalus n.gen. and Abaris Dejean (Coleoptera:
Carabidae: Pterostichini (Auctorum)) Kipling W. Will 143
W. J. Holland’s roles in the 1902 meetings of the American Association for the Advancement
of Science and the American Anthropological Association David R. Watters 215
A Late Miocene-Early Pliocene population of Trachemys (Testudines: Emydidae) from East
Tennessee . . . Paul W. Parmalee, Walter E. Klippel, Peter A. Meylan, and J. Alan Holman 233
Redescription of Eopelobates grandis, a Late Eocene anuran from the Chadron Eormation of
South Dakota Amy C. Henrici 241
Expeditions, Expositions, Associations, and Museums in the Anthropological Career of C. V.
Hartman David R. Watters and Oscar Eonseca Zamora 261
NEW TAXA
NEW GENERA AND SPECIES
Abaris aquilonaria, new species 164
301
302 Annals of Carnegie Museum vol. 71
Abaris bicolor, new species 173
Abaris convexa, new species 179
Abaris erwini, new species 163
Abaris franiai, new species . 176
Abaris impunctata, new species . 166
Abaris inaeqiialoides, new species 181
Abaris inflata, new species 177
Abaris metaUica, new species 171
Abaris mina, new species 169
Abaris napoensis, new species 159
Abaris nigra, new species 179
Abaris nitida, new species 175
Abaris nobilis, new species 173
Abaris opaca, new species 182
Abaris tachypoides, revised combination 178
Abaris wardi, new species 185
fAlveojunctus bowni, new species 3
Bredinia alza, new species 35
Bredinia davenporti, new species 24
Bredinia emarginata, new species 37
Bredinia espinosa, new species 20
Bredinia guanacasteca, new species 17
Bredinia manabiensis, new species 27
Bredinia mexicana, new species 35
Bredinia pilcopata, new species 32
Bredinia selva, new species 19
Bredinia spangleri, new species 34
Bredinia sucrensis, new species 37
Bredinia venezuelensis, new species 29
Bredinia zulia, new species 39
Ectinorus (Ectinoriis) alejoi, new species 92
Hydroptila bribriae, new species . 50
Hydroptila eglinensis, new species 49
Hydroptila hamiltoni, new species 54
Hydroptila okaloosa, new species 53
Hydroptila sarahae, new species 52
Hydroptila sykorai, new species 56
'fLeptomeryx elissae, new species 114
Neotalus, new genus .................................................. 154
Neotalus portal, new combination . 155
2002
Index to Volume 71
303
Oxyethira chrysocara, new species 47
t Fossil taxa
AUTHOR INDEX
Dawson, Mary R. ....... 59
Diamond, Margaret E 107
Fernandez, Roberto 87
Flint, Oliver S. . . . 13
Fonseca Zamora, Oscar 261
George, Richard L. 63
Goehring, W. Orr . 59
Harris, Steven C. 13, 47
Hastriter, Michael W. . 87
Henrici, Amy C. . 241
Holman, J. Alan 233
Holzenthal, Ralph W. 13
Klippel, Walter E. . . 233
Korth, William W. 107
Meyian, Peter A. 233
Parmalee, Paul W. 233
Rose, Kenneth D. . 1
Silcox, Mary T. 1
Solorzano, Nelson 87
Soto, Ruler 87
Walsh, Stephen L. 1
Watters, David R. . . 215, 261
Whiting, Michael F. 87
Will, Kipling W. 143
Zyzak, Michael D. 87
INSTRUCTIONS FOR AUTHORS
ANNALS OF CARNEGIE MUSEUM consists of
contributions to the earth sciences (including paleontol-
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format (127 by 195 mm or 5 by 7% inches). Submit all
manuscripts to the Office of Scientific Publications. Au-
thors should give particular attention to scientific con-
tent, format, and general style for the ANNALS. Manu-
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NALS will be returned to the author immediately.
Every manuscript will be subjected to external peer re-
view. Authors should submit the names, addresses,
phone and fax numbers, and e-mail addresses of at least
four qualified potential reviewers for each manuscript.
Authors will be asked to subsidize, if funds are available,
any or all costs of publication (approximately $ 1 00/page
printed). A detailed set of instructions to authors may be
requested from the Office of Scientific Publications.
Manuscript Style. — Articles should include the fol-
lowing items in this order: title page, abstract, text (with
desired headings), acknowledgments, literature cited, ta-
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manuscripts must be typed double-spaced on standard
814- by 1 1 -inch white bond paper, with at least one-inch
margins all around, and submitted in quadruplicate — an
original for the editors, and three review copies. All
pages should be numbered, including tables, literature
cited, and the list of figure captions. Only correspon-
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(e.g.. Courier) is required. A disk in WordPerfect x.x or
ASCII is helpful.
Title Page. The title should be brief, include the ani-
mal or plant group involved, be placed on a separate
page, and appear two to three inches below the top mar-
gin. Include the author’s name(s) and the affiliations of
non-CMNH authors. In the case of multiple authorship,
indicate the address to which proofs should be sent.
Abstracts. Abstracts should be short, but substantive,
and included at the head of the first page of text.
Text. Do not right justify text or break (hyphenate) a
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Tables and Figure Legends. The list of figure legends
and each table should be typed consecutively on indi-
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Literature Cited. All references to literature in text
must appear in the Literature Cited section. The data
(author, date, and page) in both citations must agree. Do
not abbreviate the titles of periodicals or serials. The
following style, double-spaced, should be used in Lit-
erature Cited:
1) Two authors in a journal series:
Soltis, D. E., and P. S. Soltis. 1992. The distri-
bution of selling rates in homosporous ferns.
American Journal of Botany, 79:97-100.
2) Same authors repeated — use three-em dash:
. 1923. The fauna of the Ardyn Obo For-
mation. American Museum of Natural History
Novitates, 98:1-5.
3) Same authors plus a third author — repeat all authors:
Knutson, L. V, R. E. Orth, and W. L. Murphy.
1986. Catalog of Sciomyzidae (Diptera) of
America north of Mexico. Entomography, 4:
1-53.
4) Chapter in an edited volume:
Rausch, R. L. 1963. A review of the distribution
of Holarctic mammals. Pp. 29^3, in Pacific
Basin Biogeography (J. L. Gressitt, ed.).
Bishop Museum Press, Honolulu, Hawaii.
5) Unpublished dissertation:
Smith, J. P. 1976. Review of Eocene Mammals.
Unpublished Ph.D. Dissert., University of
California, Berkeley, California.
6) Book:
White, M. J. D. 1961. The Chromosomes. Meth-
euen and Co., Ltd., London, United Kingdom.
7) Journal articles with usual volume and issue number:
Anderson, W. I. 1969. Lower Mississippian con-
odonts from northern Iowa. Journal of Pale-
ontology, 43:916-928.
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