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e integripennis species group of Geocharidius Jeannel, 1963... 61
The integripennis species group of Geocharidius Jeannel,
1963 (Carabidae, Bembidiini, Anillina) from Nuclear
Central America: a taxonomic review with notes
about biogeography and speciation
Igor M. Sokolov1, David H. Kavanaugh1
1 Department of Entomology, California Academy of Sciences, Golden Gate Park, 55 Music Concourse Drive,
San Francisco, CA 94118, USA
Corresponding author: David H. Kavanaugh (dkavanaugh@calacademy.org)
Academic editor: Terry Erwin|Received 13 May 2014|Accepted 14 August 2014|Published 30 September 2014
http://zoobank.org/E3384139-6A6E-426C-840D-85BC32A12E78
Citation: Sokolov IM, Kavanaugh DH (2014) e integripennis species group of Geocharidius Jeannel, 1963 (Carabidae,
Bembidiini, Anillina) from Nuclear Central America: a taxonomic review with notes about biogeography and speciation.
ZooKeys 443: 61–118. doi: 10.3897/zookeys.443.7880
Abstract
Our review recognizes 15 species of the integripennis species group of Geocharidius from Nuclear Central
America, include three species previously described (G. gimlii Erwin, G. integripennis (Bates) and G. zul-
linii Vigna Taglianti) and 12 described here as new. ey are: G. andersoni sp. n. (type locality: Chiapas,
Chiapas Highlands, Cerro Huitepec) and G. vignatagliantii sp. n. (type locality: Chiapas, Motozintla,
Sierra Madre de Chiapas, Benito Juárez) from Mexico; G. antigua sp. n. (type locality: Sacatepéquez, 5 km
SE of Antigua), G. balini sp. n. (type locality: Suchitepéquez, 4 km S of Volcan Atitlán), G. erwini sp. n.
(type locality: Quiché Department, 7 km NE of Los Encuentros), G. jalapensis sp. n. (type locality: Jalapa
Department, 4 km E of Mataquescuintla), G. longinoi, sp. n. (type locality: El Progreso Department,
Cerro Pinalón), and G. minimus sp. n. (type locality: Sacatepéquez Department, 5 km SE of Antigua)
from Guatemala; and G. celaquensis sp. n. (type locality: Lempira Department, Celaque National Park),
G. comayaguanus sp. n. (type locality: Comayagua Department, 18 km ENE of Comayagua), G. disjunc-
tus sp. n. (type locality: Francisco Morazán, La Tigra National Park), and G. lencanus sp. n. (type locality:
Lempira Department, Celaque National Park) from Honduras. For all members of the group, adult struc-
tural characters, including male and female genitalia, are described, and a taxonomic key for all members
of the integripennis species group is presented based on these characters. Behavioral and biogeographical
aspects of speciation in the group are discussed, based on the morphological analysis. In all cases of sym-
patry, pairs of closely related species show greater dierences in sizes than pairs of more remotely related
ZooKeys 443: 61–118 (2014)
doi: 10.3897/zookeys.443.7880
http://zookeys.pensoft.net
Copyright I.M. Sokolov, D.H. Kavanaugh. This is an open access article distributed under the terms of the Creative Commons Attribution License
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RESEARCH ARTICLE
Launched to accelerate biodiversity research
A peer-reviewed open-access journal
Igor M. Sokolov & David H. Kavanaugh / ZooKeys 443: 61–118 (2014)
62
species. Integripennis group species occupy six dierent montane areas at elevations above 1300m, with
no species shared among them. Major faunal barriers in the region limiting present species distributions
include the Motagua Fault Zone and a gap between the Guatemalan Cordillera volcanic chain and the
Honduran Interior Highlands no higher than 900m in elevation. Highest species diversity is in the Gua-
tematan Cordillera (six species), second highest in the Honduran Interior Highlands area (four species).
Keywords
Coleoptera, Adephaga, Carabidae, Bembidiini, Anillina, Geocharidius, new species, Nuclear Central
America, Motagua Fault Zone, biogeography, sympatric speciation
Introduction
Geocharidius Jeannel, 1963 was established for a Guatemalan species, G. integripennis
(Bates), described in “Biologia Centralia-Americana” (Bates 1882). Jeannel’s descrip-
tion of Geocharidius omitted or misinterpreted several important morphological de-
tails, leading Vigna Taglianti (1973) to re-describe the genus on the basis of the two
species, G. integripennis and G. zullinii Vigna Taglianti, known to him at that time.
Providing new evidence, Vigna Taglianti (l.c.) proposed a new phyletic lineage for Geo-
charidius, which had been placed by Jeannel (1963) in a lineage with the Mediterra-
nean Geocharis Ehlers and Rhegmatobius Jeannel. e new lineage integrated the New
World anillines of Jeannel’s “scotodipnienne” stock of genera (i.e. those taxa, members
of which have a mental tooth along with the umbilicate series of elytral pores of type
B (Jeannel 1937), where pores 7 and 8 and 8 and 9 are separated from each other by
equal distances. According to Vigna Taglianti (1973) this lineage included also Mexa-
nillus Vigna Taglianti, described in the same paper, and perhaps also Mystroceridius
Reichardt (1970) from the Galapagos Islands. Since then, several new genera of the
“scotodipnienne” stock of anilline genera have been described from the New World
(Zaballos 1997; Mateu and Etonti 2002). At present, Geocharidius includes 5 species
(Lorenz 2005; Zaballos 2004), four of which are limited in their distribution to Gua-
temala (Erwin 1982). Ecologically, representatives of Geocharidius are adapted for life
in forest litter, and these beetles are comparatively easy to collect using litter sifting
techniques.
From 2008 to 2011, the “Leaf Litter Arthropods of Mesoamerica” (LLAMA)
project (http://llama.evergreen.edu/) generated the rst signicant samples of the leaf
litter invertebrate fauna of Mesoamerica (including southern Mexico). is project
focused on sampling ants and weevils from the litter layer of the tropical forest oor,
but it also sampled many dierent non-target taxa, including many litter anillines. By
2012, the second author (DHK) had assembled on loan most available material repre-
senting Mesoamerican Anillina at the California Academy of Sciences, San Francisco.
Several hundred specimens of the subtribe were borrowed from the collections of the
six institutions noted below. is material served as the basis of and inspiration for the
current report. In this paper, we present the results of a taxonomic study of one intra-
generic group of species of Geocharidius, the integripennis species group.
e integripennis species group of Geocharidius Jeannel, 1963... 63
Materials and methods
is study is based on examination of 455 specimens belonging to the integripennis
group of species of Geocharidius, which includes 15 species. Material was borrowed
from and/or is deposited in the following institutions, identied in the text by the fol-
lowing associated codens:
CAS California Academy of Sciences, 55 Music Concourse Drive, San Francisco,
California 94118 (D. H. Kavanaugh, Curator)
CMNC Canadian Museum of Nature, Entomology, P.O. Box 3443, Station D,
Ottawa, Ontario, Canada, K1P 6P4 (R. S. Anderson, Curator)
CMNH Carnegie Museum of Natural History, Pittsburgh, Pennsylvania, U.S.A.
15213 (R. L. Davidson, Collections Manager)
KUNHM University of Kansas Natural History Museum, 1345 Jayhawk Blvd.,
Lawrence, Kansas, 66045-7593USA (Z. Falin, Collection Manager)
MNHN Muséum national d’Histoire naturelle de Paris, 57 Rue Cuvier, Paris,
75005, France (T. Deuve and A. Taghavian)
NMNH Department of Entomology, United States National Museum of Natural
History, Smithsonian Institution, Washington, D. C., U.S.A. 20013-
7012 (T. L. Erwin, Curator)
Verbatim label data are given for type specimens of all newly described taxa, with
label breaks indicated by a slash (“\”). For a series of KUNHM specimens with the
same geographical labels but diering in various barcode numbers only, these numbers
were replaced in the text by periods of ellipsis (“…”).
Measurements. All specimens were measured electronically using a Leica M420
macroscope equipped with a Syncroscopy AutoMontage photomicroscopy system
(SYNCROSCOPY, Synoptics Ltd.). Measurements for various body parts are encoded
as follows: LH = length of head, measured along midline from anterior margin of la-
brum to a virtual line connecting posterior supraorbital setae; WH = width of head, at
level of anterior supraorbital setae; WPm = maximum width across pronotum; WPa
= width across anterior angles of pronotum; WPp = width across posterior angles of
pronotum; LP = length of pronotum from base to apex along midline; WE = width of
elytra, at level of 4th umbilicate setae; LE = length of the elytra, from apex of scutellum
to apex of left elytron; SBL = standardized body length, a sum of LH, LP and LE. Meas-
urements of SBL are given in millimeters; others are presented as eight ratios: mean
widths-WH/WPm and WPm/WE and body parts-WPa/WPp, WPm/WPp, WPm/LP,
WE/LE, LE/SBL and WE/SBL. All values are given as mean ± standard deviation.
Illustrations. Digital photographs of the dorsal habitus of new species were taken
with the AutoMontage system using a Leica M420 macroscope. Line drawings of se-
lected body parts were made using grids on a Labomed Lx400 compound microscope.
Scanning electron micrographs were made with coating on a LEO 1450VP SEM.
Diagrams were prepared using Statistica 6.0. (StatSoft Inc.).
Igor M. Sokolov & David H. Kavanaugh / ZooKeys 443: 61–118 (2014)
64
Dissections. Dissections were made using standard techniques. Genitalia were dis-
sected from abdomens of specimens previously softened in boiling water for 20–30
minutes. Contents of the abdomen were cleared using boiling 10% KOH for 2-3
minutes to remove internal tissues, and then washed in hot water before examination.
After examination, genitalia were mounted on plastic transparent boards in dimethyl-
hydantoin formaldehyde resin (DMHF) and pinned beneath the specimen. In some
species, investigation of body parts was undertaken in the following way. e whole
specimen was cleared using boiling 10% KOH for ~5 minutes, then washed and dis-
sected. Disassembled body parts from a single specimen were placed on plastic trans-
parent cardboard, properly oriented, mounted in DMHF and pinned together with
the specimen labels.
Type material. e authors had no opportunity to investigate type material of the
Mexican species of Anillina described by A. Vigna Taglianti. e identication of Geo-
charidius zullinii was made only on the basis of the original description of the species
(Vigna Taglianti 1973). Types of the Guatemalan species of Geocharidius described
by T. L. Erwin in his revision of Central American Bembidiini (Erwin 1982) were
examined. All paratypes listed in the treatments for new species have been labeled with
appropriate yellow paratypes labels, which have not been included in the verbatim
label data provided for each specimen.
Terms. Terms used in the paper are largely of general use and follow the literature
(Ball and Bousquet 2000; Ball and Shpeley 2005, 2009; Erwin 1974; Jeannel 1963),
except those for ventral surface structures, terms for which follow the Handbook of
Zoology (Lawrence et al. 2010). We use the term “dorsal sclerites” (eg. Fig. 9A, D, E
and H) to refer to a complex of more or less sclerotized plates and/or agellum-like
pieces in the dorsobasal region of the retracted internal sac of the male median lobe.
ese sclerotized elements are highly varied in their size, shape and number and/or
degree of fusion among males of dierent species of this species group, and we have
not yet distinguished individual homologies among the varied elements. However, we
distinguish this complex of sclerites as a group from the “ventral sclerites” complex
found in males of many species of Anillinus Casey (1918) along with the dorsal sclerite
complex (eg., see Sokolov 2014, g.6K, vsc). We dened Nuclear Central America
as the region between the Isthmus of Tehuantepec and the Nicaraguan Depression
(Schuchert, 1935).
Species ranking. Species recognition is in accordance with our previous approach
(Sokolov et al. 2004), except for cases explained in the text.
Arrangement of taxa in the text. Taxonomic treatments of species are arranged
separately by country for the region (i.e. Mexico, Guatemala and Honduras) consistent
with the geographical distinctions made in our key to species. For each country, species
treatments are arranged in alphabetical order..
Descriptions. e scheme of description generally follows that of Ball and Shpeley
(2005, 2009).
Map. e map (Fig.22) was downloaded from the web-site: http://www.maps-
for-free.com/ and adjusted using Adobe Photoshop® software.
e integripennis species group of Geocharidius Jeannel, 1963... 65
Taxonomy
Geocharidius Jeannel
Geocharidius Jeannel, 1963: 107 (type species Anillus integripennis Bates, 1882, by
original designation)
Recognition. e members of this genus are distinguished from those of the other
North and Central American Anillina by the following combination of characters:
frontal area of head with small median tubercle; maxillary palps with palpomere 4
shorter than 1/4 that of palpomere 3; labial ligula without paraglossae, mentum and
submentum separated by mental-submental suture; pronotum convex, with short ves-
titure throughout, including the areas forward of the lateral setae; elytra without xed
discal setae and with the 7th and the 8th and the 8th and the 9th pores of the umbilicate
series separated by equal distances; metendoventrite linear without lateral arms; and
intercoxal process between the hind legs widely triangular (Sokolov 2013).
Included taxa. e species of Geocharidius, as treated at present (Lorenz 2005),
are arranged in two groups, based on body form: those with a subdepressed form and
those with a globose habitus (Erwin 1982). Species with members subdepressed in
habitus (Fig.2A–C) correspond to the type species of the genus and are treated be-
low as the integripennis species group. Members of the genus with a globose habitus
(Fig.2D), like Geocharidius phineus Erwin, Geocharidius romeoi Erwin and similar
undescribed species, are not treated in this report.
e integripennis species group
Recognition. Members of this group are distinguished from the other representatives
of the genus by the following combination of external characters: head totally covered
with microlines, microsculpture mesh pattern isodiametric (Figs 1A–C) and elytra
only moderately convex (subdepressed) (Fig.2A–C). Most species also have members
with the elytra totally covered with microlines in form of isodiametric mesh pattern
(Figs 1G–H), males with long copulatory sclerites of the internal sac (Figs9, 13, 19)
and females with simple, not bilobate, spermatheca (Figs11, 17, 21).
Description. Size. SBL range 1.15-1.61 mm.
Habitus. Body form slightly to moderately convex, subparallel or slightly ovoid.
Color. Body monocolorous, brunneorufous or rufotestaceous, appendages testaceous.
Microsculpture. Dorsal surface with polygonal sculpticells present on head in all
species, also on elytra except in G. andersoni members with smooth elytra (Fig.1I).
Development of microsculpture on pronotum and proepisternum (pes) varied among
dierent species (Figs 1D–F, Figs 3A–C).
Luster. Body surface shiny.
Macrosculpture. Body surface sparsely and nely punctate.
Igor M. Sokolov & David H. Kavanaugh / ZooKeys 443: 61–118 (2014)
66
Figures 1. SEM illustrations of structural features of body parts of Geocharidius species. A–C head,
dorsal aspect: A G. balini B G. zullinii C G. andersoni. D–F right half of pronotum, dorsal aspect: D G.
balini E G. zullinii F G. andersoni. G–I basal part of right elytron: G G. balini H G. zullinii I G. andersoni
Legend: bm – basal margin; cl – clypeus; ft – frontal tubercle; lb – labrum; mp3 – maxillary palpomere 3;
mp4 – maxillary palpomere 4. Scale = 0.05mm.
Vestiture. Body surface covered with sparse yellow setae of moderate and more or
less equal length throughout.
Fixed setae. Primary head setae include one pair of clypeal, one pair of frontal and
two pairs of supraorbital setae. Mentum (Fig.5) with two pairs of long primary (para-
medial [pms] and lateral [lms]) setae. Submentum (Fig.5) with three groups of setae:
two (Fig.5D) to three (Fig.5C) in medial row (mss), two (Fig.5F) to three (Fig.5C)
e integripennis species group of Geocharidius Jeannel, 1963... 67
Figures 2. SEM illustrations of structural features and shape of elytra of Geocharidius species, left lat-
eral aspect. A G. erwini B G. jalapensis C G. comayaguanus D G. phineus. Legend: ed2 – scutellar seta;
ed8–apical seta; eo1-9 – setae 1-9 from the umbilicate series. Scale = 0.2mm.
Igor M. Sokolov & David H. Kavanaugh / ZooKeys 443: 61–118 (2014)
68
in lateral rows (lss) and one pair of primary basal setae (prss). Elytra without discal
setae (Fig.2), but with scutellar (ed2) and apical setae (ed8) present. Last three (7th, 8th
and 9th) pores of umbilicate series (eo7, eo8 and eo9) in line and equally spread apart
(Fig.2B). Fifth abdominal ventrite (Fig. 3G–I) of male with one pair (Fig.3H) and of
female with two pairs (Fig.3I) of abdominal setae along the posterior margin.
Head (Fig.1A–C). Anterior margin of clypeus straight. Frontal area with small
tubercle medially near frontoclypeal suture. Fronto-lateral carinae distinct and long.
Eyes. Absent.
Antennae. Submoniliform, with 11antennomeres, extended to about posterior
margin of pronotum. Antennomeres 1 and 2 elongate, of approximately equal length
and 1.4-1.5 times longer than antennomere 3, which is only slightly elongate and 1.1-
1.2 times longer than antennomere 4. Antennomere 4 the shortest and 1.1.-1.2 times
shorter than antennomere 5. Antennomeres 5 to 10 globose, antennomere 11 conical
and 1.6-1.7 times longer than antennomere 10.
Labrum (Fig.1A–C). Transverse with straight, entire anterior margin, with six
setae apically, increasing in size from central pair outward.
Mandibles (Fig.4). Right mandible with distinct anterior (art) and posterior (prt)
retinacular, terebral (tt), and molar (mt) teeth. Left mandible with terebral and molar
teeth only. Premolar teeth absent from both mandibles.
Maxillae. Cardo trianguloid, stipes with dorsal and ventral lobes, galea dimer-
ous, lacinia standard for bembidiines. Palpus (Fig.1A–B) with short 4th palpomere,
0.2–0.25 length of palpomere 3.
Labium (Fig.5). Mental tooth present; mentum (m) and submentum (sm) divided
by mental-submental suture. Glossal sclerite (gsc) bisetose, without distinct paraglossae.
Prothorax. Pronotum cordiform, of moderate length (LP/SBL varied from 0.23
to 0.24 among species, LP/LE varied from 0.38 to 0.42 among species), moderately
convex, not sinuate posteriorly. Basal margin of pronotum with slightly protruding
medial portion. Anterior angles indistinct, broadly rounded. Posterior angles dentic-
ulate, with two or three small denticles anterior to angles. Prosternum (Fig.3A–C,
ps) slightly protruding at the anterior margin medially, there with a group of longer
setae relative to other prosternal vestiture, also with a pair of long ambulatory sen-
sor setae (pas) at the middle of sclerite. Prosternal intercoxal process unmargined,
slightly dilated apically and obtusely truncate at apex, with a row of sparse setulae
along midline.
Scutellum. Externally visible, triangular, with rounded apex.
Elytra. Moderate in length (LE/SBL varied from 0.57 to 0.60 among species),
without visible interneurs. Basal margination varied (long in some species, extended
halfway between humeral angle and scutellar pore (Fig.1H–I, bm), very short in oth-
ers, length about equal to diameter of basal setiferous pore socket (Fig.1G)) but dis-
tinct in all species. Lateral elytral margin without subapical sinuation in apical half.
Hind wings. Absent.
Pterothoracic venter (Fig.3D–F). Metaventrite (mtv) short, distance between meso-
and metacoxae (mtcx) slightly less than the diameter of mesocoxa (mscx). Metanepisternum
e integripennis species group of Geocharidius Jeannel, 1963... 69
Figures 3. SEM illustrations of structural features of body parts of Geocharidius species. A–C prothorax,
ventral aspect: A G. zullinii B G. erwini C G. comayaguanus. D–F pterothorax, ventral aspect: DG.jalapensis
E G.minimus F G. comayaguanus. G–I abdominal ventrites 3-5: G G. minimus, male HG. jalapensis, male
I G. jalapensis female. Legend: asts – abdominal sternal terminal seta; ipa –intercoxal process of abdomi-
nal ventrite 2; mscx – mesocoxa; msp – mesosternal process; mtcx – metacoxa; mtp – metasternal process;
mtv–metaventrite; pas – prosternal ambulatory seta; pep – proepipleuron; pes – proepisternum; prcx– pro-
coxa; ps – prosternum; psp – prosternal process; v3-v5 – abdominal ventrites 3-5. Scale = 0.05mm.
Igor M. Sokolov & David H. Kavanaugh / ZooKeys 443: 61–118 (2014)
70
Figures 4. SEM illustrations of structural features of mandibles of Geocharidius species. A–B G. zullinii:
dorsal aspect of left and right mandibles, respectively C–D G. jalapensis: dorsal aspect of left and right
mandibles, respectively E–F G. zullinii:–ventral aspect of left and right mandibles, respectively G–H G.
jalapensis: ventral aspect of left and right mandibles, respectively (right mandible with Λ-shaped crack
apically from retinacular ridge). Legend: art – anterior retinacular tooth; bemr – molar ridge, basal exten-
sion; it – incisor tooth; mr – molar ridge; mt – molar tooth; pog – posterior occlusal grove; prt – poste-
rior retinacular tooth; rr – retinacular ridge; tr – terebral ridge; tt – terebral tooth; vg – ventral groove;
vm–microtrichia. Scale = 0.05mm.
e integripennis species group of Geocharidius Jeannel, 1963... 71
Figures 5. SEM illustrations of structural features of labial complex of Geocharidius species. A G. erwini
BG. balini C G. lencanus D G. zullinii E G. longinoi F G. minimus. Legend: gsc – glossal sclerite; lms
– lateral mental seta; lss – lateral submental seta; m – mentum; mss – medial submental seta; pms – para-
medial mental seta; prss – primary basal submental seta; sm – submentum. Scale = 0.05mm.
short, subquadrate, with anterior and outer margins of equal length. Metendoventrite linear
without lateral arms.
Legs. Moderate in length, not elongate. Prothoracic legs of males with 1st protar-
somere not dilated, but with varied setal pattern. In some species (Fig.6A) tarsomeres
1-4 of males with one to three pairs of slightly dilated adhesive setae (Stork 1980),
which are absent from protarsi of females (Fig.6B); in males of other species, these
setae on tarsomeres 2-4 only (Fig.6C); and in other species, males have adhesive vesti-
ture similar to females (Fig.6D–E). Protibiae (Fig.6F–I) with antenna cleaner of type
Igor M. Sokolov & David H. Kavanaugh / ZooKeys 443: 61–118 (2014)
72
Figures 6. SEM illustrations of structural features of front legs of Geocharidius species. A–E protarsi, ven-
tral aspect: A right protarsus of G. jalapensis, male B left protarsus of G. jalapensis, female C right protarsus
of G. erwini, male D right protarsus of G. minimus, female E left protarsus of G. minimus, male F–I proti-
bia: F right protibia of G. andersoni, dorso-lateral aspect G left protibia of G. lencanus, lateral aspect H left
protibia of G. comayaguanus, ventral aspect I right protibia of G. erwini, ventral aspect. Legend: ac – antenna
cleaner; as – adhesive seta; asp – anterior spur; asr – anterior setal row; cls – clip seta; psp – posterior spur;
psr – posterior setal row; sb – setal band; ta1-ta4 – tarsomeres 1-4. Scale = 0.05mm.
B (Hlavac 1971), with both anterior (asr) and posterior (psr) apical setal rows and con-
cave apico-lateral notch (tbn). Profemora moderately swollen. Mesotibiae (Fig.7A–D)
with two terminal spurs (mss), tibial brush (msb) and a row of modied setae posterio-
laterally (msms). Metafemora unmodied, metatibiae (Fig.7E–H) with two terminal
spurs (mts), tibial brush (mtb) and one modied seta (mtms) in posteriolateral setal
row. Tarsi pentamerous, last and 1st tarsomeres are the longest, 2-4 tarsomeres of equal
length on the tarsi of all legs, 1st tarsomere shorter than combined length of 2-4 tar-
someres. Tarsal claws simple, untoothed.
e integripennis species group of Geocharidius Jeannel, 1963... 73
Figures 7. SEM illustrations of structural features of meso- and metatibia of Geocharidius species, vari-
ous aspects. A–D mesotibia: A right mesotibia of G. jalapensis, ventral aspect B left mesotibia of G. mini-
mus, medial aspect C left mesotibia of G. longinoi, ventral aspect D left mesotibia of G. comayaguanus,
medial aspect E–H metatibia: E right metatibia of G. zullini, medial aspect F left metatibia of G. balini,
medial aspect G left metatibia of G. longinoi, ventral aspect H right metatibia of G. comayaguanus, ventral
aspect. Legend: msb – mesotibial brush; msms – mesotibial modied seta; mss – mesotibial spur; msta1
– mesotarsus 1; mtb – metatibial brush; mtms – metatibial modied seta; mts – metatibial spur; mtta1 –
metatarsus 1. Scale = 0.02mm.
Abdominal ventrites. Five visible abdominal ventrites: 2nd ventrite longest, more
than 3 times longer than 3rd or 4th, 3rd and 4th equal in length; the last, 5th, approximately
1.5 times longer than 4th. Intercoxal process (ipa) of 2nd ventrite broad (Fig.3D–F),
widely triangular.
Male genitalia. Median lobe of aedeagus anopic, elongate, arcuate and twisted.
Internal sac with dorsal copulatory sclerites only, which are long, longer than half of
length of the median lobe, except in G. comayaguanus male in which they are rather
short (Fig. 19K–L, O–P). Sclerites in form of a long plate, rounded or pointed at basal
end, and typically tapered into a long agellum in apical half, in a few species tapered
Igor M. Sokolov & David H. Kavanaugh / ZooKeys 443: 61–118 (2014)
74
as a short blade-like structure. Additional spines of internal sac absent. Parameres bi-
setose. Left paramere large and relatively narrow, mostly with long and narrow apical
constriction. Right paramere small and narrow. Ring sclerite (Figs 10, 16, 20) ovate or
triangular-ovate with an elongate handle-like extension of varied shape.
Ovipositor. Gonocoxite 1 asetose. Gonocoxite 2 triangular, 1.8–2.0 times longer
than its basal width, slightly to moderately curved, with lateral and medial ensiform
and two apical nematiform setae. Ensiform setae of similar length and shape. Later-
otergite with 5-7 setae.
Female internal genitalia. Spermatheca simple (Figs11, 17, 21), not bilobate, ei-
ther unsclerotized fusiform or sclerotized of varied shape, typically fusiform with a
bulb-like enlargement apically, straight or bent rectangularly. Parts of spermatheca
mostly undierentiated and named in relation to point of attachment of the spermate-
cal gland: cornu from point of gland attachment to the apex, and nodulus from point
of gland attachment to point of duct attachment (Fig.11). e ramus, the protruding
area of attachment of the spermathecal gland (Maddison 1993), is at and not devel-
oped in the species under consideration here.
Included taxa. e integripennis species group includes three previously described spe-
cies: G. integripennis (Bates), G. zullinii Vigna Taglianti, G. gimlii Erwin, and twelve new
species, described below: G. andersoni, sp. n., G. vignatagliantii, sp. n., G. erwini, sp. n.,
G.minimus, sp. n., G. jalapensis, sp. n., G. balini, sp. n., G. longinoi, sp. n., G. antigua, sp.n.,
G. lencanus, sp. n., G. celaquensis, sp. n., G. comayaguanus, sp. n. and G. disjunctus, sp. n.
Geographical distribution. e species of this group now are known from moun-
tain ranges of southern Mexico (state of Chiapas), Guatemala and Honduras (Fig.22).
Way of life. According to label information, specimens of this group were collected
from leaf litter within the 2050–2950 m elevation range in the mountains of southern
Mexico, within the 1600–3200 m range in the mountains of Guatemala, and within the
1300–2500 m range in the mountains of Honduras. Beetles were extracted from litter
in oak, pine, pine-oak, mixed hardwood (without oaks), cloud and lower and upper
montane forests. Months of collection include May through September and November.
Relationships. e position of the integripennis group species within the genus is
unclear at present and awaits further morphological study of the globose representa-
tives of Geocharidius and a molecular phylogenetic analysis of all species.
A key for identication of adults of the integripennis species group of Geocharidius
Jeannel
e following key includes all known members of the integripennis species group. e
key makes use of distributional information because each of the three countries men-
tioned has its own Geocharidius assemblage, the ranges of which are non-overlapping
with those of neighboring countries. Our current information on species distributions
may prove to be incomplete with additional sampling, so dissection and examination
of genitalia should be used for conrmation wherever possible.
e integripennis species group of Geocharidius Jeannel, 1963... 75
1 Body form slightly to moderately convex (Fig. 2A–C) and EITHER with
head (Fig. 1A–C) and elytra (Fig. 1G–H) totally covered with microsculpture
throughout OR, if only head covered with microsculpture, then elytra sub-
parallel with prominent rounded humeri (Fig.8C) .....................................2
– Body forms globose, with markedly convex elytra (Fig.2D) and EITHER
frontal area of head without microsculpture OR, if frontal area with mi-
crosculpture, then elytra ovoid with widely rounded humeri (g.78, p. 76,
Sokolov 2013) .............................................. other groups of Geocharidius
2 Specimen from Mexico ............................................................................... 3
– Specimen from Guatemala..........................................................................5
– Specimen from Honduras ......................................................................... 12
3 Specimen larger (SBL range 1.40–1.61 mm). Habitus as in Fig.8C. Elytra
almost subparallel in basal two-thirds with rectangularly rounded humeri.
Microsculpture developed only on head; pronotum and elytra smooth, with-
out evident microscuplture ............................................ G. andersoni sp. n.
– Specimen smaller (SBL range 1.18–1.40 mm). Habitus as in Fig. 8A–B.
Elytra subparallel only in middle part, humeri broadly rounded. Microsculp-
ture present on head and elytra, only pronotum smooth ............................. 4
4 Specimen from the Chiapas Highlands. Male with shaft of median lobe of
male aedeagus (Fig.9A, D) widened apically and with dorsal sclerites of inter-
nal sac compact. Female with spermatheca unsclerotized, long and fusiform
(Fig.11A) ..........................................................G. zullinii Vigna Taglianti
– Specimen from the Sierra Madre de Chiapas. Male with shaft of median lobe
of aedeagus (Fig.9E) subparallel and with dorsal sclerites of internal sac parti-
tioned. Female with spermatheca sclerotized, short and bean-like (Fig.11B) ..
................................................................................ G. vignatagliantii sp. n.
5 Elongate (Fig.12H), smaller on average (SBL range 1.11–1.24 mm). Prono-
tum with basal margin narrower (WPa/WPp 1.06±0.024). Pronotum and
proepisternum smooth. Male with dorsal sclerites of internal sac in form of
a agellum with basal part slightly widened and bent laterally (Fig.13G, H).
Female with spermatheca as in Fig.17C ..........................G. minimus sp. n.
– Elongate ovoid (Fig.12A–G), larger on average (SBL range 1.16–1.57 mm).
Pronotum with basal margin wider (WPa/WPp<1.03). If pronotum smooth,
then proepisternum with microsculpture. If male with dorsal sclerites of in-
ternal sac formed as a long agellum (Fig.19A–B), then their base not bent
laterally. Spermatheca of female, if fusiform, then with distinct apical bulb-
like enlargement (Fig.17B, D, E) ...............................................................6
6 Pronotum wider basally, width between posterior angles greater than between
anterior angles (Wm/Wp<0.97) .................................................................. 7
– Pronotum with narrower basal margin, width between posterior angles equal
to that of anterior angles (0.97<Wm/Wp<1.03) .........................................8
7 Specimen smaller (SBL range 1.26–1.29 mm). Habitus as in Fig.12G. Male
with dorsal sclerites of internal sac formed as a agellum-like structure apical-
Igor M. Sokolov & David H. Kavanaugh / ZooKeys 443: 61–118 (2014)
76
ly, abruptly and markedly widened towards rounded basal part (Fig.19A–B).
Female with spermatheca short (Fig.21A), with swollen nodulus. Specimen
from the volcanic chain of the Guatemalan Cordillera .......G. antigua sp. n.
– Specimen larger (SBL range 1.34–1.51 mm). Habitus as in Fig.12C. Male
with dorsal sclerites of internal sac formed as a wavy, ribbon-like structure
(Fig.13R), of approximately equal width along its entire length. Female with
spermatheca elongate, of similar breadth along entire length (Fig.17F). Spec-
imen from the interior: the Sierra de las Minas range ........G. longinoi sp. n.
8 Pronotum smooth (as in Fig.1E). Male with dorsal sclerites of internal sac
markedly extended basally through basal orice (Fig. 13A, D) as a long and
narrow plate, widened and rounded basally. Female with spermatheca, if fusi-
form, then also curved (Fig.17B) ...............................................................9
– Pronotum covered with microsculpture (Fig.1D). Male with dorsal sclerites
of internal sac slightly extended basally through basal orice, EITHER elon-
gate and pointed basally (Figs 13K–L) OR formed as a wide plate (Fig.13O).
Female with spermatheca fusiform and straight, not bent (Fig.17D–E) ...11
9. Specimen smaller on average (SBL range 1.16–1.31 mm). Habitus as in
Fig.12E. Male with dorsal sclerites of internal sac gradually dilated basally to
straight basal part (Fig.13D). Spermatheca of female fusiform (Fig.13B) .....
............................................................................................G. erwini sp. n.
– Specimen larger on average (SBL range 1.33–1.43 mm). Habitus as in
Fig.12A–B. Male with dorsal sclerites of internal sac abruptly dilated basally
with enlargement curved ventrally (Figs13A, 15). Female with spermatheca
bean-shaped (Fig.17A) .............................................................................10
10. Body form broad, ovoid (Fig.12A). Pronotum markedly transverse (Wm/
Le=1.32) and distinctly constricted posteriorly (Wm/Wp= 1.38). Specimen
from the Sierra de los Cuchumatanes (Huehuetenango Department) ...........
........................................................................................... G. gimlii Erwin
– Body form narrow, elongate ovoid (Fig.12B). Pronotum less transverse (Wm/
Le=1.25±0.009) and less constricted toward base (Wm/Wp= 1.33±0.002).
Specimen from the Cerro Maria Tecún (Totonicapán Department) .............
.............................................................................. G. integripennis (Bates)
11. Specimen smaller on average (SBL range 1.22–1.34 mm). Habitus as in
Fig.12F. Male with dorsal sclerites of internal sac formed as a wide plate wid-
ened basally (Fig.13O). Female with spermathecal gland short (Fig.17E) ....
........................................................................................... G. balini, sp. n.
– Specimen larger on average (SBL range 1.33–1.57 mm). Habitus as in
Fig.12D. Male with dorsal sclerites of internal sac formed as an elongate
plate, tapered and pointed basally (Figs 13K–L). Female with spermathecal
gland long, longer than spermatheca (Fig.17D) ............ G. jalapensis sp. n.
12. Pronotum and proepisternum covered with microsculpture. Male with dor-
sal sclerites of internal sac long, only slightly widened basally (Fig. 19E, H).
e integripennis species group of Geocharidius Jeannel, 1963... 77
Female with spermatheca slightly dilated in apical fourth and with point of
spermathecal gland attachment closer to the apex (Fig.21B–C) ................13
– Pronotum and proepisternum smooth, without evident microsculpture.
Male with dorsal sclerites of internal sac EITHER short (Fig. 19K–L, O–P)
OR, if long, then these abruptly and markedly dilated basally (Fig. 19Q, T).
Female with point of spermathecal gland attachment to spermatheca closer to
the basal end and EITHER tapered apically OR of similar breadth over apical
half (Fig. 21D–E) .....................................................................................14
13. Specimen larger (SBL range 1.30–1.47 mm). Habitus as in Fig.18B. Male
with apical part of median lobe of aedeagus attenuated (Fig.19H). Female
with spermatheca curved apically (Fig.21C) .................... G. lencanus sp. n.
– Specimen smaller (SBL range 1.15–1.20 mm). Habitus as in Fig.18A. Male
with apical part of median lobe of aedeagus of average shape (Fig.19E). Fe-
male with spermatheca straight (Fig.21B) ................... G. celaquensis sp. n.
14. Male with dorsal sclerites of internal sac short (Figs 19K–L, O–P). Female
with spermatheca tapered apically (Fig.21D) ........ G. comayaguanus sp. n.
– Male with dorsal sclerites of internal sac long (Figs 19Q, T). Spermatheca of fe-
male of similar breadth throughout apical half (Fig.21E) .....G. disjunctus sp. n.
Species from Mexico
Geocharidius andersoni sp. n.
http://zoobank.org/97465882-51B8-473D-899A-6A0300DDAB82
Figs 1C, F, I, 6F, 8C, 9H–J, 10C, 11C, 22, 23
Type material. HOLOTYPE, a male, in CMNH, point-mounted, dissected, labeled:
\ MEXICO: Chiapas, Cerro Huitepec (Pico), ca. 5km W San Cristobal, 2750m, 15
IX 1991, R. Anderson,#91-101, ex: cloud forest litter \ CMNH \ HOLOTYPE Geo-
charidius andersoni Sokolov and Kavanaugh 2014 [red label] \. PARATYPES: A total
of 4 specimens (1 male and 1 female were dissected), deposited in CAS and CMNH,
labeled same as holotype except for one female, which has an additional label \ Anil-
linus sp. det. D. Shpeley 1997 \.
Type locality. Mexico, Chiapas, Chiapas Highlands, Cerro Huitepec.
Etymology. e specic epithet is a Latinized eponym in the genitive case, and is
based on the surname of Robert S. Anderson, Curator of Entomology at the Canadian
Museum of Nature, Ottawa, Canada, the collector of the type series of this species.
Recognition. Adults of this new species are distinguished from those of other
members of the integripennis species group by the following combination of external
characters: size large, elytra wide and smooth (without microsculpture); and males are
further distinguished by the size and structure of the median lobe (Figs 9H–J, 10) and
the form of the ring sclerite (Fig.10C).
Igor M. Sokolov & David H. Kavanaugh / ZooKeys 443: 61–118 (2014)
78
Description. Size. Large for genus (SBL range 1.40–1.61 mm, mean 1.51±0.077 mm,
n=5).
Habitus. Body form (Fig.8C) moderately convex, elongate ovoid, general propor-
tions (WE/SBL 0.42±0.015) wide, head narrow relative to pronotum (WH/WPm
0.69±0.017), pronotum narrow in comparison with elytra (WPm/WE 0.73±0.028).
Color. Body rufotestaceous, appendages testaceous.
Microsculpture. Mesh pattern of irregularly isodiametric sculpticells distinctly pre-
sent over dorsal surface of head only. Pronotum, elytra and proepisternum smooth
(without evident microsculpture).
Head, dorsal aspect (Fig.1C).
Prothorax. Pronotum (Fig. 1F) moderately transverse (WPm/LP 1.29±0.007),
with lateral margins slightly constricted posteriorly (WPm/WPp 1.29±0.020). Pos-
terior angles slightly obtuse (100–110°). Width between posterior angles greater than
between anterior angles (WPa/WPp 0.94±0.019).
Elytra (Fig.8C). Moderately convex, slightly depressed along suture, widest in
this species group (WE/LE 0.70±0.025), without traces of striae. Humeri distinct,
rounded, in outline forming right angle with longitudinal axis of body. Lateral margins
subparallel, slightly divergent at basal fth, evenly rounded to apex in apical third.
Legs. Protibia (Fig.6F).
Male genitalia. Median lobe (Fig.9H) with very long subparallel shaft, and
moderately enlarged apex, broadly rounded at tip. Ventral margin slightly convex
Figures 8. Digital images of habitus, dorsal aspect, of Mexican Geocharidius species. A G. zullinii
(MEXICO, Chiapas, Reserva Huitepec) B G. vignatagliantii (MEXICO, Chiapas, Motozintla), paratype
C G. andersoni (MEXICO, Chiapas, Cerro Huitepec), holotype. Scale = 0.5mm.
e integripennis species group of Geocharidius Jeannel, 1963... 79
medially. Dorsal sclerite of internal sac in form of a long plate, apically tapered into
a short agellum, and gradually widened basally with basal margin bent ventrally.
Membranous eld near ostium ag with numerous small scales. Right paramere
with long and narrow apical constriction (Fig.9J). Left paramere with very long and
narrow apical constriction (Fig.9I). Ring sclerite with handle triangular, rounded
apically (Fig.10C).
Female internal genitalia. Spermatheca unsclerotized, fusiform, arcuate, with cor-
nu long and subparallel and nodulus short and tapered basally (Fig.11C). Length of
spermathecal gland less than length of spermatheca. Spermathecal duct loosely wavy,
but not coiled.
Geographical distribution. is species is known only from the type locality in
the mountains of the Cerro Huitepec, part of the Chiapas Highlands, State of Chiapas,
Mexico (Fig.22, white diamond).
Way of life. Specimens were extracted from cloud forest litter at an elevation of
2750 m.
Figures 9. Line drawings of aedeagus of Mexican Geocharidius species. G. zullinii (MEXICO, Chiapas,
Guadalupe Shankala): A median lobe, right lateral aspect B left paramere, left lateral aspect Cright
paramere, right lateral aspect. G. zullinii (MEXICO, Chiapas, Reserva Huitepec): Dpart of median
lobe, right lateral aspect. G. vignatagliantii (MEXICO, Chiapas, Motozintla), paratype: E median lobe,
right lateral aspect F left paramere, left lateral aspect G right paramere, right lateral aspect. G. andersoni
(MEXICO, Chiapas, Cerro Huitepec), holotype: H median lobe, right lateral aspect I left paramere, left
lateral aspect J right paramere, right lateral aspect. Legend: ds – dorsal sclerites. Scale = 0.05mm.
Igor M. Sokolov & David H. Kavanaugh / ZooKeys 443: 61–118 (2014)
80
Relationships. e shape of the spermatheca of females (Fig.11C) suggests that
this species is closely related to the sympatric G. zullinii Fig.11A). Dorsal sclerites in the
internal sac of G. andersoni males (Fig.9H) resemble a shortened version of the dorsal
sclerites of males of G. gimlii (Fig.13A), which can be considered as a remote relative.
Geocharidius vignatagliantii sp. n.
http://zoobank.org/B1AB6E98-4312-4F87-8B7D-48EE76DB2C82
Figs 8B, 9E–G, 10B, 11B, 22
Type material. HOLOTYPE, a male, in CMNC, point-mounted, labeled: \ MEXI-
CO: Chiapas: Mpio: Motozintla, Benito Juarez, 2050m, 15°22.1'00"N, 92°19'07"W,
28.VII.2005, R. Anderson, oak/pine forest litter 2005-013C \ CMNC \ HOLOTYPE
Geocharidius vignatagliantii Sokolov and Kavanaugh 2014 [red label] \. PARATYPES:
A total of 14 specimens (2 males and 4 females were dissected), deposited in CAS,
CMNC and KUNHM; 3 specimens labeled same as holotype; 4 specimens labeled:
\ MEXICO: Chiapas: Mpio: Motozintla, Benito Juarez, 2050m, 15°22.1'00"N,
92°19'07"W, 28.VII.2005, R. Anderson, oak/pine forest litter 2005-013A \ CMNC\;
7 specimens labeled: \ MEXICO: Chiapas: Mpio, Motozintla, Benito Juarez
Figures 10. Line drawings of ring sclerite of Mexican Geocharidius species, male genitalia, dorsal aspect.
AG. zullinii (MEXICO, Chiapas, Reserva Huitepec) B G. vignatagliantii (MEXICO, Chiapas, Motozintla),
paratype C G. andersoni (MEXICO, Chiapas, Cerro Huitepec), holotype. Legend: hd – handle of ring scle-
rite. Scale = 0.1mm.
e integripennis species group of Geocharidius Jeannel, 1963... 81
15°22.017'N, 92°19.117'W, 2050m, 28.VII.2005, R. Anderson, oak/pine forest litter
MEX 1A05-013 \ SM0711461 KUNHM-ENT \.
Type locality. Mexico, Chiapas, Motozintla, Sierra Madre de Chiapas, Benito Juárez.
Etymology. e specic epithet is a Latinized eponym in the genitive case, and is
based on the surname of Prof. Augusto Vigna Taglianti, Director of the Museum of
Zoology at the Sapienza University of Rome, Roma, Italia, the rst reviser of the spe-
cies of Geocharidius.
Recognition. Adults of this new species are practically indistinguishable externally
from those of G. zullinii but can be distinguished from the latter and from the other
members of the integripennis species group by the structure of the median lobe of males
and the shape of spermatheca of females.
Description. Size. Medium for genus (SBL range 1.27–1.40 mm, mean
1.32±0.038 mm, n=5).
Figures 11. Line drawings of spermatheca of Mexican Geocharidius species. A G. zullinii (MEXICO,
Chiapas, Reserva Huitepec) B G. vignatagliantii (MEXICO, Chiapas, Motozintla), paratype C G. ander-
soni (MEXICO, Chiapas, Cerro Huitepec), paratype. Legend: c – cornu; n – nodulus; sd – spermathecal
duct; sg – spermathecal gland; sp – spermatheca. Scale = 0.05mm.
Igor M. Sokolov & David H. Kavanaugh / ZooKeys 443: 61–118 (2014)
82
Habitus. Body form (Fig.8B) moderately convex, elongate ovoid, general pro-
portions (WE/SBL 0.40±0.008), proportions of head (WH/WPm 0.73±0.014) and
pronotum (WPm/WE 0.75±0.016) average for group.
Color. Body brunneorufous, appendages testaceous.
Microsculpture. Present over all dorsal surfaces of head and elytra. Pronotum and
proepisternum smooth.
Prothorax. Pronotum moderately transverse (WPm/LP 1.26±0.021), with lateral
margins moderately constricted posteriorly (WPm/WPp 1.32±0.020). Posterior an-
gles obtuse (110–120°). Widths between anterior and posterior angles of equal length
(WPa/WPp 0.99±0.020).
Elytra. Moderately convex, slightly depressed along suture, moderately wide (WE/
LE 0.68±0.016), without traces of striae. Humeri broadly rounded, in outline forming
right angle with longitudinal axis of body. Lateral margins convex, evenly divergent at
basal third, evenly rounded to apex in apical third.
Male genitalia. Median lobe of aedeagus (Fig.9E) with long subparallel shaft, and
small rounded apex. Ventral margin almost straight. Dorsal sclerites of internal sac in
form of a long plate, apically tapered into a rather long agellum, and abruptly widened
basally as a semicircular end plate near basal orice. Right paramere with long and nar-
row apical constriction (Fig.9G). Left paramere with long and narrow apical constric-
tion (Fig.9F). Ring sclerite with handle triangular, widely rounded apically (Fig.10B).
Female internal genitalia. Spermatheca sclerotized, bean-shaped, arcuate, with
cornu short and nodulus long (Fig.11B). Length of spermathecal gland greater than
length of spermatheca. Spermathecal duct not coiled.
Geographical distribution. is species is known only from the type locality in
the mountains of the Sierra Madre de Chiapas, located in the municipality of Motoz-
intla, State of Chiapas, Mexico (Fig.22, white squares).
Way of life. Specimens were collected by sifting oak/pine forest litter at an eleva-
tion of 2050 m.
Relationships. Adults of this species closely resemble those of G. zullinii from
the Chiapas Highlands externally and in the shape of dorsal sclerites of the internal
sac (Fig.9E; cf. Fig.9A). e shape of the spermatheca of females (Fig.11B) suggests
a relationship with the Guatemalan G. integripennis (Fig.17A) from the Sierra de los
Cuchumatanes of the Guatemalan Cordillera.
Geocharidius zullinii Vigna Taglianti
Figs 1B, E, H, 3A, 4A–B, E–F, 5D, 7E, 8A, 9A–D, 10A, 11A, 22, 23
Geocharidius zullinii Vigna Taglianti, 1973: 314.
Holotype. A male, dissected, deposited in A. Vigna Taglianti’s private collection [not
examined]. Type locality: Mexico, Chiapas, Chiapas Highlands, Comitàn, S of Ago-
stin (Fig.22, white circle with dot).
e integripennis species group of Geocharidius Jeannel, 1963... 83
Recognition. Adults of this species (Fig.8A) are practically indistinguishable from
the adults of G. vignatagliantii, described below, and are distinguished from the lat-
ter and from the other members of the integripennis species group by structure of the
median lobe and shape of spermatheca.
Description. e original description provides a thorough accounting of external
features of this species and is absolutely sucient for species characterization. Below,
we add references to illustrations of structural features presented here and descriptions
of genitalia, which, for females, has not been done previously.
Head, dorsal aspect (Fig.1B).
Mouthparts. Mandibles (Figs 4A–B, and 4E–F). Maxillae and labium ( Fig.5D).
Prothorax. Ventral aspect (Fig.3A). Pronotum, lateral margin ( Fig.1E).
Elytra. Lateral margin (Fig.1H).
Legs. Metatibia (Fig.7E).
Male genitalia. Median lobe (Fig.9A) with shaft long, slightly widened apically,
apex small and rounded. Ventral margin almost straight. Dorsal sclerites of internal
sac in form of a long plate, tapered apically to a rather long blade, and abruptly
widen basally as a semicircular end plate near basal orice. Specimens from the
northern part of the geographical range demonstrate slightly dierent shape of dor-
sal sclerites (Fig.9D). Right paramere with long and narrow apical constriction
(Fig.9C). Left paramere with moderately long and gradually tapered apical con-
striction (Fig.9B). Ring sclerite with handle triangular, narrowly rounded apically
(Fig.10A).
Female internal genitalia. Spermatheca unsclerotized, fusiform, arcuate, with cor-
nu long and subparallel and nodulus short and basally tapered (Fig.11A). Length of
spermathecal gland less than length of spermatheca. Spermathecal duct loosely wavy,
but not coiled.
Geographical distribution. is species is widely distributed across the Chiapas
Highlands, State of Chiapas, Mexico (Fig.22, white circles). We have examined a
total of 15 specimens (6 males and 3 females dissected) from the following locali-
ties: 4 specimens labeled: MEXICO: Chiapas: Mpio, Huixtán, Guadalupe Shankala,
16°38'N, 92° 25'W, 2350m, 25.VII.2005, R. Anderson, mixed hardwood (no oaks)
forest litter MEX 1A05-007 \ SM0701883 KUNHM-ENT \; 2 specimens labeled:
MEXICO: Chiapas: Reserva Huitepec, 16°44.686'N, 92°41.312'W, 2600m, 9-VII-
2007 M.G.Branstetter ex. winkler, cloud forest leaf litter LLAMA07 MGB629 \
SM0781461 KUNHM-ENT \; 3 specimens labeled: \ MEXICO: Chiapas: Reserva
Huitepec, 16°44.686'N, 92°41.312'W, 2600m, 11-VII-2007 J.Longino ex. win-
kler, under pines, cloud forest edge, leaitter LLAMA07 JTL6037-s \ SM0786461
KUNHM-ENT \; 3 specimens labeled: \ MEXICO: Chiapas: Reserva Huitepec,
16°44.686'N, 92°41.312'W, 2600m, 11-VII-2007 J.Longino ex. winkler, cloud for-
est, leaitter LLAMA07 JTL6036-s \ SM0786461 KUNHM-ENT \; 1 specimen
labeled: \ MEXICO: Chiapas, Mpio, San Cristobal de las Casas, Reserva Huitepec,
2450m, 16°45.84'N, 92°40.70'W, 2600m, 11-VII-2003, R. Anderson, cloud forest
Igor M. Sokolov & David H. Kavanaugh / ZooKeys 443: 61–118 (2014)
84
lit., MEX1A03 108 \ SM0477446 KUNHM-ENT \; 1 specimen labeled: \ MEX-
ICO: Chiapas: 15km E San Cristobal 16.74689°N, 92.48985°W, 2500m, 29-V-
2008, sifted leaf litter, cloud forest LLAMA08 Wm-A-05-1 \ SM0836667 KUNHM-
ENT \; 1 specimen labeled: \ MEXICO: Chiapas: Mpio, Huixtán, Bazóm, 2450m,
16°44'19.0N 92°29', 18.3W, 9-VII-2003, R. Anderson, oak forest litter, MEX1A03
107 \ SMO517781 KUNHM-ENT \.
Way of life. Specimens were sifted from litter in a wide range of dierent forest types
(hardwood without oaks, oak, pine and cloud forests) at elevations of 2350–2600 m.
Relationships. e shape of spermatheca (Fig.11A) of females suggests that this
species is closely related to the sympatric G. andersoni (Fig.11C), described above.
Species from Guatemala
Geocharidius antigua sp. n.
http://zoobank.org/2A03829D-0D34-4608-A5A6-9D8909EAEDC2
Figs 12G, 19A–D, 20A, 21A, 22, 23
Type material. HOLOTYPE, a male, in KUNHM, point-mounted, dissected, la-
beled: \ GUATEMALA: Sacatepéquez: 5km SE Antigua, 14.52779 -90.68971±200m,
2350m, 10-VI-2009, ex. sifted leaf litter, oak forest LLAMA09 Wm-B-08-2-all \ KUN-
HM \ HOLOTYPE Geocharidius antigua Sokolov and Kavanaugh 2014 [red label] \.
PARATYPES: 1 female, dissected, labeled same as holotype (deposited in KUNHM).
Type locality. Guatemala, Sacatepéquez, 5 km SE of Antigua.
Etymology. e specic epithet is a noun in apposition and refers to the city in
the vicinity of which the type series was collected.
Recognition. Adults of this new species are practically indistinguishable in body
shape from those of other Guatemalan species of Geocharidius with small body size;
but the smooth pronotum and presence of microsculpture on the proepisternum form
a basis for distinguishing adults of G. antigua from those of sympatric G. minimus and
allopatric G. balini, described below. Males and females of G. antigua are distinguished
from those of the other members of the integripennis species group by the structure of
the median lobe and the shape of spermatheca, respectively.
Description. Size. Medium for genus (SBL range 1.26–1.29 mm, mean
1.28±0.019 mm, n=2).
Habitus. Body form (Fig.12G) moderately convex, elongate ovoid, general pro-
portions (WE/SBL 0.39±0.009) and proportions of head (WH/WPm 0.71±0.002)
and pronotum (WPm/WE 0.78±0.020) average for group.
Color. Body brunneorufous, appendages testaceous.
Microsculpture. Mesh pattern of irregularly isodiametric sculpticells present over
all dorsal surfaces of head and elytra. Pronotum smooth. Proepisternum with evident
microsculpture.
e integripennis species group of Geocharidius Jeannel, 1963... 85
Prothorax. Pronotum moderately wide (WPm/LP 1.28±0.011), with lateral mar-
gins slightly constricted posteriorly (WPm/WPp 1.29±0.004). Posterior angles slightly
obtuse (100–110°). Width between posterior angles greater than between anterior angles
(WPa/WPp 0.95±0.022).
Elytra. Moderately convex, slightly depressed along suture, moderately wide (WE/
LE 0.67±0.015), without traces of striae. Humeri broadly rounded, in outline forming
right angle with longitudinal axis of body. Lateral margins convex, evenly divergent at
basal third, evenly rounded to apex in apical third.
Male genitalia. Median lobe of aedeagus (Fig.19A) with long subparallel shaft,
and small rounded apex. Ventral margin almost straight. Dorsal sclerites of internal
Figures 12. Digital images of habitus of Guatemalan Geocharidius species. A G. gimlii (GUATEMALA,
Huehuetenango, San Juan Ixcoy), holotype B G. integripennis (GUATEMALA, Totonicapán, “Totoni-
capam”), lectotype C G. longinoi (GUATEMALA, El Progreso, Cerro Pinalón), paratype D G. jalapensis
(GUATEMALA, Jalapa, Mataquescuintla), paratype E G. erwini (GUATEMALA, Quiché, Los Encuen-
tros), paratype F G. balini (GUATEMALA, Suchitepéquez, Volcano Atitlán), paratype G G. antigua
(GUATEMALA, Sacatepéquez, Antigua), paratype H G. minimus (GUATEMALA, Sacatepéquez, Anti-
gua), paratype. Scale = 0.5mm.
Igor M. Sokolov & David H. Kavanaugh / ZooKeys 443: 61–118 (2014)
86
sac in form of a long plate, tapered apically as a long agellum, and abruptly widened
basally as a nearly circular complex of structures near basal orice (Figs 19A–B). Right
paramere with short and narrow apical constriction (Fig.19D). Left paramere with
long and narrow apical constriction (Fig.19C). Ring sclerite with handle triangular,
slightly asymmetrical, pointed apically (Fig.20A).
Female internal genitalia. Spermatheca sclerotized, bulb-shaped, straight, and very
wide, with cornu short and nodulus swollen (Fig.21A). Length of spermathecal gland
greater than length of spermatheca. Spermathecal duct loosely coiled.
Geographical distribution. is species is known only from the type locality,
situated on the northern slopes of volcano Agua in the volcanic chain of the Guatema-
lan Cordillera (Fig.22, yellow quadrangle).
Way of life. Specimens were collected by sifting oak forest litter at an elevation of
2350 m.
Relationships. e shape of dorsal sclerites of the internal sac (Fig. 19A–B) sug-
gests that this species is closely related to the Honduran G. disjunctus (Fig. 19Q, T),
described below.
Geocharidius balini sp. n.
http://zoobank.org/4AC8143D-8FDF-49B2-AC46-0AEBC7FE0D32
Figs 1A, D, G, 5B, 7F, 12F, 13O–Q, 16E, 17E, 22, 23
Type material. HOLOTYPE, a male, in KUNHM, point-mounted, labeled: \ GUA-
TEMALA: Suchitepéquez: 4km S Vol. Atitlán, 14.54915- 91.19055 ±200m, 1625m,
15-VI-2009 ex sifted leaf litter, cloud forest, LLAMA09 Wa-B-09-1-all \ KUNHM \
HOLOTYPE Geocharidius balini Sokolov and Kavanaugh 2014 [red label] \. PARA-
TYPES: A total of 117 specimens (3 males and 2 females were dissected), deposited
in CAS and KUNHM; 99 specimens labeled same as holotype; 10 specimens labeled:
\ GUATEMALA: Suchitepéquez: 4km S Vol. Atitlán, 14.55103- 91.19350 ±306m,
1750m, 15-VI-2009 ex sifted leaf litter, cloud forest, LLAMA09 Wm-B-09-2-01 \
KUNHM \; 7 specimens labeled: \ GUATEMALA: Suchitepéquez: 4km S Vol. Ati-
tlán, 14.55311- 91.19337 ±35m, 1750m, 15-VI-2009 ex sifted leaf litter, cloud for-
est, LLAMA09 Wm-B-09-2-02 \ KUNHM \; 1 specimen labeled: \ GUATEMALA:
Jalapa: 4km E Mataquescuintla, 14.53257 -90.15253 ±200m, 2400m, 1-VI-2009,
ex. sifted leaf litter, cloud forest, LLAMA09 Wa-B-07-2-all \ KUNHM \.
Type locality. Guatemala, Suchitepéquez, 4 km S of Volcan Atitlán.
Etymology. e specic epithet is a Latinized eponym in the genitive case, and is
based on the given name of the dwarf Balin, a refounder of the underground kingdom of
Moria, one of orin Oakenshield’s Company of Dwarves who had accompanied Bilbo
Baggins on the Quest of Erebor in the book “e Hobbit, or ere and Back Again” by
J.R.R.Tolkien.
Recognition. Adults of this new species are distinguished from those of other
members of the integripennis species group externally by their small size and the pres-
e integripennis species group of Geocharidius Jeannel, 1963... 87
Figures 13. Line drawings of aedeagus of Guatemalan Geocharidius species. A–C G. gimlii (GUATE-
MALA, Huehuetenango, San Juan Ixcoy), holotype: A median lobe with internal sac and dorsal sclerites,
right lateral aspect B left paramere, left lateral aspect C right paramere, right lateral aspect D–FG. erwini
(GUATEMALA, Quiché, Los Encuentros), paratype: D median lobe with internal sac and dorsal sclerites,
right lateral aspect E left paramere, left lateral aspect F right paramere, right lateral aspect G–JG.minimus
(GUATEMALA, Sacatepéquez, Antigua), paratype: G median lobe with internal sac and dorsal sclerites,
right lateral aspect H dorsal sclerite of median lobe, dorsal aspect Ileft paramere, left lateral aspect J right
paramere, right lateral aspect K–N G. jalapensis (GUATEMALA, Jalapa, Mataquescuintla), paratype:
Kmedian lobe with internal sac and dorsal sclerites, right lateral aspect L dorsal sclerite of median lobe,
dorsal aspect M left paramere, left lateral aspect Nright paramere, right lateral aspect O–Q G. balini
(GUATEMALA, Suchitepéquez, Volcano Atitlán), paratype: O median lobe with internal sac and dor-
sal sclerites, right lateral aspect P left paramere, left lateral aspect Q right paramere, right lateral aspect
R–TG. longinoi (GUATEMALA, El Progreso, Cerro Pinalón), paratype: R median lobe with internal sac
and dorsal sclerites, right lateral aspect S left paramere, left lateral aspect T right paramere, right lateral
aspect. Scale = 0.05mm.
Igor M. Sokolov & David H. Kavanaugh / ZooKeys 443: 61–118 (2014)
88
ence of microsculpture on the pronotum and internally by the structure of the median
lobe of males and the shape of spermatheca of females.
Description. Size. Small to medium for genus (SBL range 1.22–1.34 mm, mean
1.27±0.040 mm, n=26).
Habitus. Body form (Fig.12F) moderately convex, elongate ovoid, general propor-
tions (WE/SBL 0.38±0.008) and proportions of head (WH/WPm 0.72±0.013) and
pronotum (WPm/WE 0.78±0.017) average for group.
Color. Body brunneorufous, appendages testaceous.
Microsculpture. Mesh pattern of irregularly isodiametric sculpticells present over all
dorsal surfaces of head, pronotum and elytra. Proepisternum with evident microsculpture.
Head (Fig.1A).
Mouthparts. Maxillae and labium (Fig.5B)
Prothorax. Pronotum (Fig.1D) moderately transverse (WPm/LP 1.26±0.022),
with lateral margins moderately constricted posteriorly (WPm/WPp 1.32±0.021).
Posterior angles obtuse (110–120°). Widths between anterior and posterior angles of
equal length (WPa/WPp 0.99±0.022).
Elytra (Fig.1G). Moderately convex, slightly depressed along suture, moderately
narrow (WE/LE 0.64±0.018), without traces of striae. Humeri rounded, in outline
forming right angle with longitudinal axis of body. Lateral margins convex, evenly
divergent at basal third, evenly rounded to apex in apical third.
Legs. Metatibia (Fig.7F)
Male genitalia. Median lobe (Fig.13O) with shaft short and broad and apex of
moderate size and rounded. Ventral margin greatly enlarged and convex, with nu-
merous poriferous canals. Dorsal sclerites of internal sac of peculiar shape, in form of
anastomosing short plates, connected in apical and basal thirds, pointed apically as a
short blade. Right paramere long and narrow (Fig.13Q). Left paramere with long and
narrow apical constriction (Fig.13P). Ring sclerite with handle almost rectangularly
rounded, slightly asymmetrical (Fig.16E).
Female internal genitalia. Spermatheca sclerotized, fusiform with apical bulb en-
largement, straight, with cornu long and nodulus short (Fig.17E). Lengths of sper-
mathecal gland and spermatheca equal. Spermathecal duct not coiled.
Geographical distribution. is species is known only from two localities remote
from each other: one situated on the southern slopes of volcano Agua in the Suchite-
péquez Department, and the other situated on the northern slopes of a former twinned
volcano, remains of which form the caldera of Laguna de Ayarza, in Jalapa Depart-
ment. Both localities are in the volcanic chain of the Guatemalan Cordillera (Fig.22,
green diamonds).
Way of life. Specimens were collected by sifting cloud forest litter at middle
(1600–1750 m) to high elevations (2400 m).
Relationships. e shapes of handle of ring sclerite (Fig.16E) and of the sper-
matheca (Fig.17E) suggest that this species is closely related to G. jalapensis (Figs 16D,
17D, described below.
e integripennis species group of Geocharidius Jeannel, 1963... 89
Geocharidius erwini sp. n.
http://zoobank.org/6B696BFC-D7AE-4893-BCBC-80579DFAA5B7
Figs 2A, 3B, 5A, 6C, I, 12E, 13D–F, 16B, 17B, 22
Geocharidius integripennis, Sokolov, 2013:53
Type material. HOLOTYPE, a male, in NMNH, glued on cardboard, labeled: \ Guate-
mala: QUICHÉ, 7km NE Los Encuentros, 2400m, 18.XI.1991 leg. R.Baranowski \ sift-
ing litter under bushes at roadside pine forest \ Loan from USNMNH 2051867 \ HOL-
OTYPE Geocharidius erwini Sokolov and Kavanaugh 2014 [red label] \. PARATYPES: A
total of 45 specimens (4 males and 4 females were dissected), deposited in CAS, CMNC,
NMNH; 23 specimens labeled same as holotype; 10 specimens labeled: \ Guatemala:
QUICHÉ, 7km NE Los Encuentros, 2400m, 14.XI.1991 leg. R.Baranowski \ sifting
litter under bushes at roadside pine forest \ Loan from USNMNH 2051867 \; 8 speci-
mens labeled: \ Guatemala QUICHÉ, 6km S Chichicastenango, 2140m. 16.XI.1991 leg.
R.Baranowski \ sifting litter, pine-oak forest \ Loan from USNMNH 2051867 \; 4 speci-
mens labeled: \ Guatemala: QUICHÉ, 5km S Chichicastenango, 2000m. 18.XI.1991
leg. R.Baranowski \ sifting litter, pine-oak forest \ Loan from USNMNH 2051867 \; 1
specimen labeled: \ Guat.: QUEZALTEN. 12km S.E. Zunil, Fuentes Georginas, 2460m,
21.VI.1993, R. Anderson, cloud for. litter 93-10A\ CMNC \; 1 specimen labeled: \ Guat.:
QUEZALTEN. 12km S.E. Zunil, Fuentes Georginas, 2460m, 21.VI.1993, R. Ander-
son, cloud for. litter 93-10E\ CMNC \; 1 specimen labeled: \ Guat.: QUEZALTEN.
12km S.E. Zunil, Fuentes Georginas, 2450m, 19.VI.1993, R. Anderson, cloud for. lit-
ter 93-5CC \ CMNC \; 1 specimen labeled: \ Guat.: QUEZALTEN. 12km S.E. Zunil,
N.W.face Cerro Zunil, 2700m, 20.VI.1993, R. Anderson, hardwood for. litter 93-8F \
CMNC \; 1 specimenslabeled: \ Guat.: QUEZALTENANGO: 12km SE Zunil, NW face
Cerro Zunil, hardwd.for.litter, 2700–2760m, R. Anderson 91-30 28.V.1991. \ CMNC \.
Type locality. Guatemala, Quiché Department, 7 km NE of Los Encuentros.
Etymology. e specic epithet is a Latinized eponym in the genitive case and is
based on the surname of Terry L. Erwin, Curator of Entomology at the Smithsonian
Institution, United States National Museum of Natural History, Washington, D. C.,
U.S.A., the rst reviser of the Guatemalan Anillina.
Recognition. Adults of this new species are practically indistinguishable from
those of other the Guatemalan species of Geocharidius with small body size. Males
and females of G. erwini are distinguished from those of the other members of the
integripennis species group by the structure of the median lobe of males and the shape
of spermatheca of females, respectively.
Description. Size. Small to medium for genus (SBL range 1.16–1.31 mm, mean
1.23±0.057 mm, n=30).
Habitus. Body form (Fig.12E) moderately convex, elongate ovoid, general pro-
portions (WE/SBL 0.38±0.009) and proportions of head (WH/WPm 0.72±0.012)
and pronotum (WPm/WE 0.79±0.015) average for group.
Igor M. Sokolov & David H. Kavanaugh / ZooKeys 443: 61–118 (2014)
90
Color. Body rufotestaceous, appendages testaceous.
Microsculpture. Mesh pattern of irregularly isodiametric sculpticells present over
all dorsal surfaces of head and elytra. Pronotum smooth. Proepisternum with evident
microsculpture.
Mouthparts. Labium (Fig.5A).
Prothorax. Pronotum moderately transverse (WPm/LP 1.26±0.019), with lateral
margins slightly constricted posteriorly (WPm/WPp 1.31±0.025). Posterior angles
slightly obtuse (100–110°). Widths between anterior and posterior angles of equal
length (WPa/WPp 1.01±0.025). Ventral aspect (Fig.3B).
Elytra (Fig.2A). Moderately convex, slightly depressed along suture, moderate-
ly wide (WE/LE 0.67±0.017), without traces of striae. Humeri rounded, in outline
forming right angle with longitudinal axis of body. Lateral margins convex, evenly
divergent at basal third, evenly rounded to apex in apical third.
Legs. Protibia (Fig.6I). Protarsus (Fig.6C).
Male genitalia. Median lobe (Fig.13D) with shaft moderately long, slightly wid-
ened apically, and apex small and rounded. Ventral margin almost straight. Dorsal
Figures 14. Photographs of labels for type specimens of Geocharidius species. A G. gimlii, holotype
BG.integripennis, lectotype C G. integripennis, paralectotype.
e integripennis species group of Geocharidius Jeannel, 1963... 91
sclerites of internal sac (Fig.13D) in form of a long plate, tapered apically as a long a-
gellum, and gradually widen towards semicircular basal end extended basally through
the basal orice. Right paramere with short and wide apical consriction (Fig.13F). Left
paramere with long and moderately narrow apical constriction (Fig.13E). Ring sclerite
with almost rectangularly rounded, subparallel, handle (Fig.16B).
Female internal genitalia. Spermatheca sclerotized, fusiform with bulb enlarge-
ment apically, twice bent rectangularly in opposite directions, with cornu long and
nodulus short (Fig.17B). Length of spermathecal gland less than length of spermathe-
ca. Spermathecal duct coiled.
Geographical distribution. is species is known from a few scattered localities
in the Quiché and Quetzaltenango Departments of Guatemala (Fig.22, green circles).
Way of life. Specimens were collected by sifting litter from dierent habitats:
cloud, hardwood, pine and pine-oak forests at elevations of 2140-2760 m.
Relationships. e shape of dorsal sclerites of the internal sac in males suggests a
remote relationship with G. minimus (Fig.13G), described below.
Geocharidius gimlii Erwin
Figs 12A, 13A–C, 14A, 15B, 16A, 22
Geocharidius gimlii Erwin, 1982: 488.
Holotype. A male, deposited in NMNH, point-mounted, dissected, labeled (Fig.14A):
\ ♂ –wing \ ADP 26556 \ GUATEMALA: Hue. 7.7km S SanJuan Ixcoy, 2780 MS 15
35'N, 091 27'W 11 August 1974 \ G. E. Ball, H. Frania, D.R. Whitehead Colls. leaf lit-
ter \. Type locality. Guatemala, Huehuetenango Department, 7.7 km S of San Juan Ixcoy.
Recognition. Males of this species are distinguished from those of other members
of the integripennis species group by the following combination of characters: prono-
tum small, transverse, elytra comparatively wide and structure of median lobe of male
as in Fig.13A.
Description. Size. Medium for genus (SBL 1.42 mm).
Habitus. Body form (Fig.12A) moderately convex, broadly ovoid; general propor-
tions (WE/SBL 0.41) rather wide; proportions of head (WH/WPm 0.72) average for
group; pronotum narrow (WPm/WE 0.74) relative to elytra.
Color. Body rufotestaceous, appendages testaceous.
Microsculpture. Mesh pattern of irregularly isodiametric sculpticells present over
all dorsal surfaces of head and elytra. Pronotum smooth (without evident microsculp-
ture). Proepisternum with microsculpture.
Prothorax. Pronotum transverse (WPm/LP 1.32), with lateral margins markedly
constricted posteriorly (WPm/WPp 1.38). Posterior angles obtuse (112°). Widths be-
tween anterior and posterior angles equal (WPa/WPp 1.01).
Elytra. Moderately convex, slightly depressed along suture, markedly wide (WE/
LE 0.70), without traces of striae. Humeri broadly rounded, in outline forming slightly
Igor M. Sokolov & David H. Kavanaugh / ZooKeys 443: 61–118 (2014)
92
obtuse angle with longitudinal axis of body. Lateral margins convex, evenly divergent
at basal half, evenly rounded to apex in apical half.
Male genitalia. Median lobe of aedeagus (Fig.13A, 15B) with shaft long, widened
apically, and apex small and acutely rounded. Ventral margin straight. Dorsal sclerites
of internal sac in form of very long plate, protruding from basal orice, and tapered
apically in rather long agellum, abruptly widened basally as a semicircular dilation,
bent ventrally and surrounded by complex of semisclerotized sheaths of peculiar shape.
Right paramere with long, narrow apical constriction (Fig.13C). Left paramere with
long, narrow and curved apical constriction (Fig.13B). Ring sclerite of triangular
shape, with sinuations on both sides of long basal handle (Fig.16A).
Female internal genitalia. Females unknown.
Geographical distribution. is species is known only from the type locality in
the mountains of the Sierra de los Cuchumatanes, located in the Huehuetenango De-
partment of Guatemala (Fig.22, white triangle).
Way of life. e unique type specimen was sifted from leaf litter in Lower Mon-
tane Wet Forest (Erwin 1982) at an elevation of 2780 m.
Figures 15. Schematic line drawings of median lobe and dorsal sclerites of internal sac of two previously
described Guatemalan Geocharidius species. A G. integripennis (GUATEMALA, Totonicapán, “Totoni-
capam”), holotype B G. gimlii (GUATEMALA, Huehuetenango, San Juan Ixcoy), holotype, both left
lateral aspect.
e integripennis species group of Geocharidius Jeannel, 1963... 93
Relationships. e shape of the median lobe in the holotype of G. gimlii (Fig.15B) is
almost identical to that of the male holotype of G. integripennis (Fig.15A); hence, at least
for now, the latter can be considered as its closest relative. e general shape of the dorsal
sclerites of the internal sac (namely the apically tapered plate, widened and ventrally bent
at the basal end) is also similar to that in G andersoni (Fig.9H) males described above.
Figures 16. Line drawings of ring sclerite of Guatemalan Geocharidius species, male genitalia, dorsal
aspect. A G. gimlii (GUATEMALA, Huehuetenango, San Juan Ixcoy), holotype B G. erwini (GUA-
TEMALA, Quiché, Los Encuentros), paratype C G. minimus (GUATEMALA, Sacatepéquez, Antigua),
paratype D G. jalapensis (GUATEMALA, Jalapa, Mataquescuintla), paratype E G. balini (GUATE-
MALA, Suchitepéquez, Volcano Atitlán), paratype F G. longinoi (GUATEMALA, El Progreso, Cerro
Pinalón), paratype. Scale = 0.1mm.
Igor M. Sokolov & David H. Kavanaugh / ZooKeys 443: 61–118 (2014)
94
Geocharidius integripennis (Bates)
Figs 12B, 14B, 14C, 15A, 17A, 22
Anillus integripennis Bates, 1882: 145.
Lectotype. A male, deposited in MNHN, glued on cardboard, dissected, labeled
(Fig.14B): \ Totonicapam, 85- 10,500 ft. Champion \ integripennis Bates \ LEC
TOTYPUS ♂ det. A. Vigna 1972 \ LECTOTYPUS ♂ Geocharidius integripennis
Bates Desig. Vigna- Taglianti 1972 Det. J.P. Zaballos 2004\. Paralectotype female,
also in MNHN, glued on cardboard, dissected, labeled (Fig.14C): \ Totonicapam, 85-
10,500 ft. Champion \ Anillus integripennis Bates \ TYPE \ ♀ \ PARALECTOTYPUS
♀ Geocharidius integripennis Bates Det. J.P.Zaballos 2004 \. Type locality: Guatemala,
Totonicapán Department, Totonicapam [=Totonicapán].
Geocharidius tagliantii Erwin, 1982: 494; synonymized by Zaballos (2004).
Recognition. Males and females of this species are distinguished from those of oth-
er members of the integripennis species group (except G. gimlii, see Relationships above)
by the structure of the median lobe of males and the spermatheca of females. Adults of
G. gimlii have proportionately much wider elytra than those of G. integripennis.
Description. Size. Medium for genus (SBL range 1.33–1.43 mm, mean
1.38±0.070 mm, n=2).).
Habitus. Body form (Fig.12B) moderately convex, elongate ovoid, general pro-
portions (WE/SBL 0.40±0.002), proportions of head (WH/WPm 0.74±0.022) and
pronotum (WPm/WE 0.77±0.026) average for group.
Color. Body rufotestaceous, appendages testaceous.
Microsculpture. Mesh pattern of irregularly isodiametric sculpticells present
over all dorsal surfaces of head and elytra. Pronotum smooth. Proepisternum with
microsculpture.
Prothorax. Pronotum slightly transverse (WPm/LP 1.25±0.009), with lateral
margins moderately constricted posteriorly (WPm/WPp 1.33±0.002). Posterior an-
gles slightly obtuse (100-110°). Widths between anterior and posterior angles of equal
length (WPa/WPp 0.99±0.018).
Elytra. Moderately convex, slightly depressed along suture, moderately wide (WE/
LE 0.67±0.005), without traces of striae. Humeri rounded, in outline forming right
angle with longitudinal axis of body. Lateral margins convex, evenly divergent at basal
fourth, evenly rounded to apex in apical third.
Male genitalia. Male genitalia of the lectotype are mounted in old gum, covered
now with a network of numerous cracks, making the objects inside hard to see. Hence,
we could examine only general shape of the median lobe and could not discern details of
the inner sac or of the parameres or the round sclerite. Based on what we could see, the
median lobe of the aedeagus (Fig.15A) is very similar to that of the G. gimlii holotype.
Female internal genitalia. Spermatheca of the paralectotype sclerotized, bean-shaped,
with apical constriction, almost straight, cornu short and nodulus long (Fig.17A). Length
of spermathecal gland greater than length of spermatheca. Spermathecal duct loosely coiled.
e integripennis species group of Geocharidius Jeannel, 1963... 95
Geographical distribution. Precise locality at which the type series of this species
was collected is unknown. Presumably, the material that was collected by Champion
and served as the basis for the Bates’ description came from the Cerro Maria Tecún
Figures 17. Line drawings of spermatheca of Guatemalan Geocharidius species. A G. integripennis
(GUATEMALA, Totonicapán, “Totonicapam”), holotype B G. erwini (GUATEMALA, Quiché, Los
Encuentros), paratype C G. minimus (GUATEMALA, Sacatepéquez, Antigua), paratype D G. jalapensis
(GUATEMALA, Jalapa, Mataquescuintla), paratype E G. balini (GUATEMALA, Suchitepéquez, Volcano
Atitlán), paratype F G. longinoi (GUATEMALA, El Progreso, Cerro Pinalón), paratype. Scale = 0.05mm.
Igor M. Sokolov & David H. Kavanaugh / ZooKeys 443: 61–118 (2014)
96
mountains in the Totonicapán Department of Guatemala (Fig.22, black and white
triangle), as was shown by Ball and Roughley (1982) for Pterostichus (Percolaus) cham-
pioni (Bates), the locality label of which is identical with that of the G. integripennis type
specimens.
Way of life. e type specimens were collected at an elevation of “10,500 ft.”
(=3200 m).
Relationships. Without doubt, the closest relative of G. integripennis is G. gimlii.
In view of the similarity in the shape of the median lobes (Fig.15B; cf. Fig.15A) of their
males and the range of variation of the median lobes seen among other species of the
group, it may seem reasonable to consider these taxa as two subspecies of a single spe-
cies. However, in the absence of sucient material for more thorough investigation of
variation of the external features and structure of the genitalia, we prefer to preserve the
“status quo” and consider G. gimlii and G. integripennis as close, but separate species.
Geocharidius jalapensis sp. n.
http://zoobank.org/BF792073-053E-4FA8-BA99-759634A24AC7
Figs 2B, 3D, H–I, 4C–D, G–H, 6A–B, 7A, 12D, 13K–N, 16D, 17D, 22, 23
Type material. HOLOTYPE, a male, in KUNHM, point-mounted, labeled: \ GUA-
TEMALA: Jalapa: 4km E Mataquescuintla, 14.52943 -90.14775 ±105m, 2620m,
2-VI-2009, ex. sifted leaf litter, cloud forest, LLAMA09 Wm-B-07-1-06 \ KUNHM
\ HOLOTYPE Geocharidius jalapensis Sokolov and Kavanaugh 2014 [red label] \.
PARATYPES: A total of 78 specimens (4 males and 4 females were dissected), depos-
ited in CAS and KUNHM; 10 specimens labeled same as holotype; 2 specimens la-
beled: \ GUATEMALA: Jalapa: 4km E Mataquescuintla, 14.53409 -90.15290 ±28m,
2325m, 3-VI-2009, ex. sifted leaf litter, cloud forest, LLAMA09 Wm-B-07-1-10 \
KUNHM \; 1 specimen labeled: \ GUATEMALA: Jalapa: 4km E Mataquescuintla,
14.52950 -90.14802 ±254m, 2600m, 1-VI-2009, ex. sifted leaf litter, cloud forest,
LLAMA09 Wm-B-07-1-01 \ KUNHM \; 20 specimens labeled: \ GUATEMALA:
Jalapa: 4km E Mataquescuintla, 14.53257 -90.15253 ±200m, 2400m, 1-VI-2009, ex.
sifted leaf litter, cloud forest, LLAMA09 Wa-B-07-2-all \ KUNHM \; 23 specimens la-
beled: \ GUATEMALA: Jalapa: 4km E Mataquescuintla, 14.52780 -90.14671 ±105m,
2655m, 2-VI-2009, ex. sifted leaf litter, cloud forest, LLAMA09 Wm-B-07-1-04 \
KUNHM \; 16 specimens labeled: \ GUATEMALA: Jalapa: 4km E Mataquescuintla,
14.52987 -90.14908 ±200m, 2600m, 1-VI-2009, ex. sifted leaf litter, cloud forest on
ridge top, LLAMA09 Wa-B-07-1-all \ KUNHM \; 6 specimens labeled: \ GUATE-
MALA: Jalapa: 4km E Mataquescuintla, 14.52705 -90.14671 ±105m, 2660m, 2-VI-
2009, ex. sifted leaf litter, cloud forest, LLAMA09 Wm-B-07-1-05 \ KUNHM \.
Type locality. Guatemala, Jalapa Department, 4 km E of Mataquescuintla.
Etymology. e specic epithet is a Latinized adjective in the masculine form based
on the name Jalapa, the Department of Guatemala in which the type series was collected.
e integripennis species group of Geocharidius Jeannel, 1963... 97
Recognition. Adults of this new species are distinguished from those of other
members of the integripennis species group by the following combination of external
characters: size large and pronotum transverse and fully covered with microsculpture.
Males and female of G. jalapensis are distinguished from those of other members of
the integripennis species group by the structure of the median lobe and the shape of
spermatheca, respectively.
Description. Size. Medium to large for genus (SBL range 1.33–1.57 mm, mean
1.46±0.081mm, n=25).
Habitus. Body form (Fig.12D) moderately convex, elongate ovoid, general pro-
portions (WE/SBL 0.39±0.007), proportions of head (WH/WPm 0.71±0.019) and
pronotum (WPm/WE 0.76±0.021) average for group.
Color. Body brunneorufous, appendages testaceous.
Microsculpture. Mesh pattern of irregularly isodiametric sculpticells present over all
dorsal surfaces of head, pronotum and elytra. Proepisternum with evident microsculpture.
Mouthparts. Mandibles (Figs 4C–D, G–H).
Prothorax. Pronotum markedly transverse (WPm/LP 1.30±0.026), with lateral
margins moderately constricted posteriorly (WPm/WPp 1.33±0.029). Posterior an-
gles obtuse (110–120°). Widths between anterior and posterior angles of equal length
(WPa/WPp 0.99±0.023).
Pterothorax (Fig.3D).
Elytra (Fig.2B). Moderately convex, slightly depressed along suture, moderate-
ly wide (WE/LE 0.65±0.009), without traces of striae. Humeri rounded, in outline
forming right angle with longitudinal axis of body. Lateral margins convex, evenly
divergent at basal third, evenly rounded to apex in apical third.
Legs. Mesotibia (Fig.7A). Protarsus (Fig. 6A–B).
Abdomen. Ventrites 3-5 (Fig. 3H–I).
Male genitalia. Median lobe (Fig.13K) with shaft long and apex slightly enlarged
and rounded. Ventral margin almost straight. Dorsal sclerites of internal sac in form of
a long plate, tapered apically and basally in short extensions (Fig.13K–L). Right para-
mere with short and rather wide apical constriction (Fig.13N). Left paramere with
long and narrow apical constriction (Fig.13M). Ring sclerite with almost rectangularly
rounded, slightly asymmetrical, handle (Fig.16D).
Female internal genitalia. Spermatheca sclerotized, fusiform with apical bulb en-
largement, straight, with long cornu and short nodulus (Fig.17D). Length of sper-
mathecal gland greater than length of spermatheca. Spermathecal duct coiled.
Geographical distribution. is species is known only from type locality, situ-
ated on the northern slopes of the former twinned volcano, remains of which form the
caldera now lled with the waters of Laguna de Ayarza (Jalapa Department). Physi-
ographically, the region is part of the volcanic chain of the Guatemalan Cordillera
(Fig.22, green triangle).
Way of life. Specimens were collected by sifting cloud forest litter at elevations of
2325–2620 m.
Igor M. Sokolov & David H. Kavanaugh / ZooKeys 443: 61–118 (2014)
98
Relationships. e shapes of handle of the ring sclerite (Fig.16D) and of the sper-
matheca (Fig.17D) suggest that this species is closely related to G. balini (Figs 13O
and 17E), described above.
Geocharidius longinoi sp. n.
http://zoobank.org/2037D9B1-5260-4836-8ED4-0D0AE950111F
Figs 5E, 7C, G, 12C, 13R–T, 16F, 17F, 22
Type material. HOLOTYPE, a male, in KUNHM, point-mounted, labeled: \ GUA-
TEMALA: El Progreso : Cerro Pinalón, 15.08392-89.93013 ±55m, 2750m, 1-V-2009
ex. sifted leaf litter, cloud forest, LLAMA09 Wm-B-01-1-04 \ KUNHM \ HOLOTYPE
Geocharidius longinoi Sokolov and Kavanaugh 2014 [red label] \. PARATYPES: A total
of 13 specimens (2 males and 1 female were dissected), deposited in CAS, CMNC
and KUNHM; 5 specimens labeled same as holotype; 7 specimens labeled: \ GUATE-
MALA: El Progreso : Cerro Pinalón, 15.08411-89.93239 ±57m, 2715m, 1-V-2009
ex. sifted leaf litter, cloud forest, LLAMA09 Wm-B-01-1-05 \ KUNHM \; 1 specimen
labeled: \ GUAT.: EL PROGRESO : 19.6km.N.Estancia de la Virgen, 2000m, Finca
la Illuciones, 24.VI.1993, R.Anderson, cloud for. litter, 93-13C \ CMNC \.
Type locality. Guatemala, El Progreso Department, Cerro Pinalón.
Etymology. e specic epithet is a Latinized eponym in the genitive case, and is
based on the surname of John T. (Jack) Longino, Professor of the Biology Department
of the University of Utah, and one of Co-PI’s of the LLAMA project, which provided
the material on which the description of this species is based.
Recognition. Adults of this new species are distinguished from those of other members
of the integripennis species group by the large size, distinctive shape of the pronotum with
very wide basal margin, and the proepisternum with evident microsculpture. Males and
females of G. longinoi are distinguished from those of other members of the integripennis
species group by the structure of the median lobe and the shape of spermatheca, respectively.
Description. Size. Medium to large for genus (SBL range 1.34–1.51 mm, mean
1.41±0.071mm, n=12).
Habitus. Body form (Fig.12C) moderately convex, elongate ovoid, general propor-
tions (WE/SBL 0.38±0.008) and proportions of head (WH/WPm 0.71±0.012) aver-
age for group, pronotum markedly wide compared to elytra (WPm/WE 0.80±0.013).
Color. Body brunneorufous, appendages testaceous.
Microsculpture. Mesh pattern of irregularly isodiametric sculpticells present over all
dorsal surfaces of head and elytra. Pronotum smooth (without evident microsculpture).
Proepisternum with evident microsculpture.
Mouthparts. Maxillae and labium (Fig.5E).
Prothorax. Pronotum slightly transverse (WPm/LP 1.25±0.019), with lateral mar-
gins slightly constricted posteriorly (WPm/WPp 1.29±0.018). Posterior angles slightly
obtuse (100–110°). Width between posterior angles greater than between anterior an-
gles (WPa/WPp 0.94±0.020).
e integripennis species group of Geocharidius Jeannel, 1963... 99
Elytra. Moderately convex, slightly depressed along suture, moderately wide (WE/
LE 0.66±0.020), without traces of striae. Humeri rounded, in outline forming right
angle with longitudinal axis of body. Lateral margins convex, evenly divergent at basal
forth, evenly rounded to apex in apical third.
Legs. Mesotibia (Fig.7C). Metatibia (Fig.7G).
Male genitalia. Median lobe (Fig.13R) with shaft long, apically slightly widened,
apex small and narrowly rounded. Ventral margin straight. Dorsal sclerites of internal
sac (Fig.13R) in form of long waved ribbon, tapered apically and slightly dilated ba-
sally, where sclerite forms small hook-like extension. Right paramere with short apical
constriction (Fig.13T). Left paramere with rather short and narrow apical constriction
(Fig.13S). Ring sclerite with triangularly rounded handle (Fig.16F).
Female internal genitalia. Spermatheca sclerotized, elongate, subparallel, almost
straight, with long cornu and short nodulus (Fig.17F). Length of spermathecal gland
less than length of spermatheca. Spermathecal duct not coiled.
Geographical distribution. is species is known only from Cerro Pinalón, part
of the Sierra de las Minas range of Guatemala (Fig.22, green squares).
Way of life. Specimens were extracted from cloud forest litter at elevations of
2000–2750 m.
Relationships. e shape of handle of the ring sclerite (Fig.16F) and the structure
of dorsal sclerites of the internal sac (Fig.13R) suggest a relationship with the Hondu-
ran G. celaquensis (Figs 20B and 19E), described below.
Geocharidius minimus sp. n.
http://zoobank.org/38CA85F3-81F4-4B3F-B23B-EB29892CEDAD
Figs 3E, G, 5F, 6D–E, 7B, 12H, 13G–J, 16C, 17C, 22, 23
Type material. HOLOTYPE, a male, in KUNHM, glued on cardboard, labeled:
\ GUATEMALA: Sacatepéquez: 5km SE Antigua, 14.53577 -90.69428±200m,
2150m, 10-VI-2009, ex. sifted leaf litter, hardwood forest LLAMA09 Wa-B-08-1-
all \ KUNHM \ HOLOTYPE Geocharidius minimus Sokolov and Kavanaugh 2014
[red label] \. PARATYPES: A total of 121 specimens (6 males and 4 females were
dissected), deposited in CAS and KUNHM; 53 specimens labeled same as holotype;
11 specimens labeled: \ GUATEMALA: Sacatepéquez: 5km SE Antigua, 14.53439
-90.69340±36m, 2175m, 11-VI-2009, ex. sifted leaf litter, hardwood forest LLA-
MA09 Wm-B-08-2-08 \ KUNHM \; 1 specimen labeled: \ GUATEMALA: Sacate-
péquez: 5km SE Antigua, 14.53666 -90.69491±255m, 2140m, 10-VI-2009, ex.
sifted leaf litter, hardwood forest LLAMA09 Wm-B-08-1-07 \ SEMC0896573 KUN-
HM-ENT \; 48 specimens labeled: \ GUATEMALA: Sacatepéquez: 5km SE Antigua,
14.53482-90.69398±33m, 2175m, 10-VI-2009, ex. sifted leaf litter, hardwood forest
LLAMA09 Wm-B-08-1-04 \ SEMC0888829 KUNHM-ENT \; 4 specimens labeled:
\ GUATEMALA: Suchitepéquez: 4km S Vol. Atitlán, 14.55311- 91.19337 ±35m,
1750m, 15-VI-2009 ex sifted leaf litter, cloud forest, LLAMA09 Wm-B-09-2-02 1 \
Igor M. Sokolov & David H. Kavanaugh / ZooKeys 443: 61–118 (2014)
100
KUNHM \; 2 specimens labeled: GUATEMALA: Suchitepéquez: 4km S Vol. Atitlán,
14.54915- 91.19055 ±200m, 1625m, 15-VI-2009 ex sifted leaf litter, cloud forest,
LLAMA09 Wa-B-09-1-all \ SEMC0889856 KUNHM-ENT \; 2 specimens labeled:
\ GUATEMALA: Suchitepéquez: 4km S Vol. Atitlán, 14.55972- 91.18951 ±27m,
2164m, 17-VI-2009 ex sifted leaf litter, cloud forest, LLAMA09 Wm-B-09-2-06 \
SEMC0896573 KUNHM-ENT \.
Type locality. Guatemala, Sacatepéquez Department, 5 km SE of Antigua.
Etymology. e specic epithet is a Latin adjective, minimus (superlative of par-
vus), in the masculine form, meaning “smallest”.
Recognition. Adults of this new species are distinguished from those of other
members of the integripennis species group by the combination of small size, elongate
habitus and smooth proepisternum. Males and females of G. minimus are distinguished
from those of other members of the integripennis species group by the structure of the
median lobe and the shape of spermatheca, respectively.
Description. Size. Small for genus (SBL range 1.11–1.24 mm, mean 1.18±0.041
mm, n=26).
Habitus. Body form (Fig.12H) moderately convex, elongate, general proportions
narrow (WE/SBL 0.37±0.009), proportions of head (WH/WPm 0.75±0.016) and
pronotum (WPm/WE 0.80±0.015) wide for group.
Color. Body rufotestaceous, appendages testaceous.
Microsculpture. Mesh pattern of irregularly isodiametric sculpticells present over
all dorsal surfaces of head and elytra. Pronotum and proepisternum smooth (without
evident microscupture).
Mouthparts. Maxillae and labium (Fig.5F).
Prothorax. Pronotum moderately narrow (WPm/LP 1.24±0.027), with lateral
margins markedly constricted posteriorly (WPm/WPp 1.35±0.022). Posterior angles
obtuse (110–120°). Width between anterior angles greater than between posterior an-
gles (WPa/WPp 1.06±0.024).
Pterothorax (Fig.3E).
Elytra. Moderately convex, slightly depressed along suture, moderately narrow
(WE/LE 0.64±0.017), without traces of striae. Humeri rounded, in outline forming
right angle with longitudinal axis of body. Lateral margins subparallel, evenly diver-
gent at basal fth, evenly rounded to apex in apical fourth.
Legs. Mesotibia (Fig, 7B). Protarsus (Figs 6D–E).
Abdomen. Ventrites 3-5 (Fig.3G).
Male genitalia. Median lobe (Fig.13G) with shaft moderately long, subparallel,
apex small and narrowly rounded. Ventral margin almost straight. Dorsal sclerites of
internal sac in form of a long plate, tapered apically as a long agellum, and gradually
widen towards narrow and rounded basal end extended through basal orice and bent
laterally (Fig.13G–H). Right paramere with short apical consriction (Fig.13J). Left
paramere moderately long, with rather short apical constriction (Fig.13I). Ring scler-
ite with handle triangular, slightly asymmetrical, pointed apically (Fig.16C).
e integripennis species group of Geocharidius Jeannel, 1963... 101
Female internal genitalia. Spermatheca sclerotized, fusiform, only slightly dilated
apically, straight, with cornu and nodulus of approximately equal length (Fig.17C).
Lengths of spermathecal gland and spermatheca equal. Spermathecal duct not coiled.
Geographical distribution. is species is known from the slopes of two volca-
nos, Agua and Atitlán, in Sacatepéquez and Suchitepéquez Departments of Guate-
mala, respectively (Fig.22, green owers).
Way of life. Specimens were collected by sifting litter in hardwood and cloud
forests at middle and high elevations of 1600 and 2200 m, respectively.
Relationships. e shape of dorsal sclerites of the internal sac (Fig.13G) of males
suggests a distant relationship with G. erwini (Fig 13D), described above, whereas the
shape of the handle of the ring sclerite (Fig.16C) of males suggests relationships with
the Guatemalan G. antigua (Fig.20A), described above, and the Honduran G. disjunc-
tus (Fig.20E), described below.
Species from Honduras
Geocharidius celaquensis sp. n.
http://zoobank.org/2AE3AACE-E66B-4F68-B8D8-AF5974E29BC6
Figs 18A, 19E–G, 20B, 21B, 22, 23
Type material. HOLOTYPE, a male, in CMNC, point-mounted, dissected, labeled:
\ HONDURAS: Lempira Dept., P.N. Celaque, nr. Gracias, Campamiento Naranjo,
2500m, N14°32.7', W88°39.7', 12–13.V.2002, cloud forest litter R. Anderson, 2002-
020C \ CMNC \ HOLOTYPE Geocharidius celaquensis Sokolov and Kavanaugh 2014
[red label] \. PARATYPES: A total of 2 females (both were dissected), deposited in
CAS and KUNHM; labeled same as holotype, except label of the holder: SEMC0…
KUNHM-ENT \.
Type locality. Honduras, Lempira Department, Celaque National Park.
Etymology. e specic epithet is a Latinized adjective in the masculine form
based on the name of Celaque National Park, from which the new species is described.
Recognition. Adults of this new species are distinguished from those of other mem-
bers of the integripennis species group by their small size, fully microsculptured dorsal
body surface and pronotum with wide basal margin. Males and females of G. celaquensis
are distinguished from those of the other members of the integripennis species group by
the structure of the median lobe and the shape of spermatheca, respectively.
Description. Size. Small for genus (SBL range 1.15–1.20 mm, mean
1.18±0.023mm, n=3).
Habitus. Body form (Fig.18A) moderately convex, ovoid, general proportions
(WE/SBL 0.40±0.005), proportions of head (WH/WPm 0.73±0.016) and pronotum
(WPm/WE 0.78±0.015) moderately wide.
Color. Body rufotestaceous, appendages testaceous.
Igor M. Sokolov & David H. Kavanaugh / ZooKeys 443: 61–118 (2014)
102
Microsculpture. Mesh pattern of irregularly isodiametric sculpticells present over
all dorsal surfaces of head, pronotum and elytra. Proepisternum also with evident
microsculpture.
Prothorax. Pronotum markedly transverse (WPm/LP 1.32±0.025), with lateral
margins markedly constricted posteriorly (WPm/WPp 1.35±0.002). Posterior angles
obtuse (110–120°). Width between posterior angles slightly greater than between an-
terior angles (WPa/WPp 1.04±0.004).
Elytra. Moderately convex, slightly depressed along suture, moderately wide (WE/
LE 0.68±0.015), without traces of striae. Humeri rounded, in outline forming right
angle with longitudinal axis of body. Lateral margins convex, evenly divergent at basal
third, evenly rounded to apex in apical third.
Male genitalia. Median lobe (Fig.19E) with shaft subparallel, apex small and nar-
rowly rounded. Ventral margin almost straight. Dorsal sclerites of internal sac in form
of long, waved ribbon, tapered apically and slightly dilated and narrowly rounded
basally. Right paramere with short apical constriction (Fig.19G). Left paramere with
long and narrow apical constriction (Fig.19F). Ring sclerite with handle triangular,
widely rounded at apex (Fig.20B).
Female internal genitalia. Spermatheca sclerotized, fusiform, slightly dilated api-
cally, straight, with short cornu and long nodulus (Fig.21B). Length of spermathecal
gland less than length of spermatheca. Spermathecal duct not coiled.
Geographical distribution. is species is known only from Celaque National
Park, part of the Cerro las Minas range of Honduras (Fig.22, yellow ower).
Way of life. Specimens were extracted from cloud forest litter at an elevation of 2500 m.
Relationships. e shape of dorsal sclerites of the internal sac (Fig.19E) in males
and the point of the attachment of the spermathecal gland (Fig.21B) in females suggest
Figures 18. Digital images of habitus of Honduran Geocharidius species. A G. celaquensis (HONDURAS,
Lempira, Celaque National Park), paratype B G. lencanus (HONDURAS, Lempira, Celaque National
Park), paratype C G. comayaguanus (HONDURAS, Comayagua, Comayagua), paratype D G. disjunctus
(HONDURAS, Francisco Morazán, La Tigra National Park), holotype. Scale = 0.5mm.
e integripennis species group of Geocharidius Jeannel, 1963... 103
that this species is closely related to G. lencanus (Figs 19H and 21C), described below,
and perhaps also, but more remotely, to the Guatemalan G. longinoi (Figs 13R and
17F), described above.
Figures 19. Line drawings of aedeagus of Guatemalan and Honduran Geocharidius species. A–D G.an-
tigua (GUATEMALA, Sacatepéquez, Antigua), holotype: A median lobe with internal sac and dorsal scle-
rites, right lateral aspect B dorsal sclerite of median lobe, dorsal aspect C left paramere, left lateral aspect
D right paramere, right lateral aspect E–G G. celaquensis (HONDURAS, Lempira, Celaque National
Park), holotype: E median lobe with internal sac and dorsal sclerites, right lateral aspect F left paramere,
left lateral aspect G right paramere, right lateral aspect H–J G. lencanus (HONDURAS, Lempira, Cel-
aque National Park), paratype: H median lobe with internal sac and dorsal sclerites, right lateral aspect
Ileft paramere, left lateral aspect J right paramere, right lateral aspect K–N G. comayaguanus (HON-
DURAS, Comayagua, Comayagua), paratype: K median lobe with internal sac and dorsal sclerites, right
lateral aspect L variation in a shape of dorsal sclerite of internal sac, right lateral aspect M left paramere,
left lateral aspect N right paramere, right lateral aspect O–P G. comayaguanus (HONDURAS, La Paz,
Guajicuiro): variations in a shape of dorsal sclerite of internal sac, right lateral aspect Q–S G. disjunctus
(HONDURAS, Francisco Morazán, La Tigra National Park), holotype: Q median lobe with internal sac
and dorsal sclerites, right lateral aspect R left paramere, left lateral aspect S right paramere, right lateral
aspect. TG.disjunctus (HONDURAS, Yoro, Pico Pijol National Park): shape of dorsal sclerite of median
lobe, right lateral aspect. Scale = 0.05mm.
Igor M. Sokolov & David H. Kavanaugh / ZooKeys 443: 61–118 (2014)
104
Geocharidius comayaguanus sp. n.
http://zoobank.org/CC2857E4-7428-4FC0-A2BC-CFA5A571E8C3
Figs 2C, 3C, 3F, 6H, 7D, H, 18C, 19K–P, 20D, 21D, 22, 23
Type material. HOLOTYPE, a male, in CMNC, point-mounted, dissected, labeled:
\ HONDURAS: Comayagua, 18km ENE Comayagua, 1950m, 20.VIII.1994, S. & J.
Peck, wet oak-pine forest litter, S&JPeck 1994-52 \ CMNC \ HOLOTYPE Geocha-
ridius comayaguanus Sokolov and Kavanaugh 2014 [red label] \. PARATYPES: A total
of 23 specimens (6 males and 2 females were dissected), deposited in CAS, CMNC
and KUNHM; 4 specimens labeled same as holotype; 5 specimens labeled: \ HON-
DURAS: Comayagua, Comayagua (18km E.N.E.), 1950m, 20.VIII.1994, S. Peck wet
oak-pine forest litter, SBP 94-52 \ CMNC \; 8 specimens labeled: HONDURAS:
La Paz Dept. Tutule, Res. Biol. Guajiquiro, 14°10'N, 87°50'W, 2130m, 7-V-2002,
R.Anderson, cloud forest litter, RSA2002-010 \ SM0… KUNHM-ENT \; 1 specimen
labeled: HONDURAS: LA PAZ: Tutule, Res. Biol. Guajiquiro, N14°10', W87°50',
2130m, 7.V.2002, R.Anderson, cloud forest litter, 2002-010H \ CMNC \; 1 specimen
labeled: \ HONDURAS: LA PAZ: Tutule, Res. Biol. Guajiquiro, N14°10', W87°50',
2130m, 7.V.2002, R.Anderson, cloud forest litter, 2002-010D \ CMNC \; 1 specimen
labeled: \ HONDURAS: LA PAZ: Tutule, Res. Biol. Guajiquiro, N14°10', W87°50',
2130m, 7.V.2002, R.Anderson, cloud forest litter, 2002-010E \ CMNC \; 2 specimens
labeled: \ HONDURAS: LA PAZ: Tutule, Res. Biol. Guajiquiro, N14°10', W87°50',
2130m, 7.V.2002, R.Anderson, cloud forest litter, 2002-010I \ CMNC \; 1 specimen
labeled: \ HONDURAS: Yoro Dept., P.N. Pico Pijol, 1300m, N15°09.4', W87°37.6',
11.V.2002, R. Anderson, upper montane forest litter, 2002-017A\ CMNC.
Type locality. Honduras, Comayagua Department, 18 km ENE of Comayagua.
Etymology. e specic epithet is a Latinized adjective in the masculine form
based on the name of the city of Comayagua, from the vicinity of which the new spe-
cies is described.
Recognition. Adults of this species are practically indistinguishable externally
from those of G. disjunctus, described below, and are distinguished from the latter, as
from those of the other members of the integripennis species group, by the structure of
the male median lobe and the shape of spermatheca in females.
Description. Size. Small to medium for genus (SBL range 1.19–1.34 mm, mean
1.28±0.072mm, n=20).
Habitus. Body form (Fig.18C) moderately convex, ovoid, general proportions
(WE/SBL 0.40±0.011), proportions of head (WH/WPm 0.74±0.017) and pronotum
(WPm/WE 0.78±0.018) moderately wide.
Color. Body brunneorufous, appendages testaceous.
Microsculpture. Mesh pattern of irregularly isodiametric sculpticells present over
all dorsal surfaces of head and elytra. Pronotum and proepisternum smooth (without
evident microsculpture).
Prothorax. Pronotum moderately transverse (WPm/LP 1.29±0.024), with lateral
margins markedly constricted posteriorly (WPm/WPp 1.35±0.027). Posterior angles
e integripennis species group of Geocharidius Jeannel, 1963... 105
obtuse (110–120°). Width between posterior angles equal to the width between ante-
rior angles (WPa/WPp 1.02±0.026). Ventral aspect (Fig.3C).
Pterothorax (Fig.3F).
Elytra (Fig.2C). Moderately convex, slightly depressed along suture, moderate-
ly wide (WE/LE 0.68±0.022), without traces of striae. Humeri rounded, in outline
forming right angle with longitudinal axis of body. Lateral margins convex, evenly
divergent at basal third, evenly rounded to apex in apical third.
Legs. Protibia (Fig.6H). Mesotibia (Fig.7D). Metatibia (Fig.7H).
Male genitalia. Median lobe (Fig.19K) with shaft long and dorsally convex, apex
small and narrowly rounded. Ventral margin straight. Dorsal sclerites of internal sac
small, in form of a short hook-like plate, slightly varied among dierent populations
Figures 20. Line drawings of ring sclerite of Guatemalan and Honduran Geocharidius species, male
genitalia, dorsal aspect. A G. antigua (GUATEMALA, Sacatepéquez, Antigua), holotype B G. celaquen-
sis (HONDURAS, Lempira, Celaque National Park), holotype C G. lencanus (HONDURAS, Lempira,
Celaque National Park), paratype D G. comayaguanus (HONDURAS, Comayagua, Comayagua), paratype
E G. disjunctus (HONDURAS, Francisco Morazán, La Tigra National Park), holotype. Scale = 0.1mm.
Igor M. Sokolov & David H. Kavanaugh / ZooKeys 443: 61–118 (2014)
106
(Fig.19K–L, O–P). Right paramere with long apical constriction (Fig.19N). Left
paramere with long and narrow apical constriction (Fig.19M). Ring sclerite with han-
dle triangular, pointed at apex (Fig.20D).
Female internal genitalia. Spermatheca sclerotized, fusiform, slightly tapered api-
cally, straight, with cornu and nodulus of equal length (Fig.21D). Length of sper-
mathecal gland less than length of spermatheca. Spermathecal duct not coiled.
Geographical distribution. is species is known from La Paz, Comayagua and
Yoro Departments, thus having a range that crosses nearly the entire Honduran Inte-
rior Highlands from the Pacic to the Antlantic slope (Fig.22, yellow circles).
Way of life. Specimens were collected in litter samples from cloud, upper montane
and wet oak-pine forests at middle and high elevations of 1300 and 2130 m, respectively.
Relationships. is species is unique within the integripennis species group in the
shape of dorsal sclerites of the internal sac (Fig.19K) of males and of the spermatheca
(Fig.21D) of females. Hence, G. comayaguanus appears to be only remotely related to
the other members of the species group.
Geocharidius disjunctus sp. n.
http://zoobank.org/A0E6C548-3EC5-40B6-933F-414125D61068
Figs 18D, 19Q–T, 20E, 21E, 22, 23
Type material. HOLOTYPE, a male, in CMNC, point-mounted, dissected, labeled: \
HONDURAS: FRANC. MOR: P.N. La Tigra, 23.2km N Tegucigalpa, 15.VIII.1994-
201A, 2100m, R.Anderson, cloud forest litter berlese \ CMNC \ HOLOTYPE Geo-
charidius disjunctus Sokolov and Kavanaugh 2014 [red label] \. PARATYPES: A total
of 2 specimens (both were dissected), deposited in CAS and CMNC ; 1 male la-
beled: \ HONDURAS: Yoro Dept., P.N. Pico Pijol, 1400m, N15°09.4'W87°37.6',
11.V.2002, R. Anderson, upper montane forest litter, 2002-016C\ CMNC \; 1 female
labeled: \ HONDURAS: Yoro Dept., P.N. Pico Pijol, 1300m, N15°09.4'W87°37.6',
11.V.2002, R. Anderson, upper montane forest litter, 2002-017A \ CMNC \.
Type locality. Honduras, Francisco Morazán, La Tigra National Park.
Etymology. e specic epithet is a Latin adjective, disjunctus, in the masculine
form, meaning “separated”, and refers to the species distinctness from the sympatric G.
comayaguanus, described above.
Recognition. Adults of this new species are practically indistinguishable from
those of the sympatric G. comayaguanus in body shape. Males and females of G. dis-
junctus are distinguished from those of the other members of the integripennis species
group by the structure of the median lobe and the shape of spermatheca, respectively.
Description. Size. Small to medium for genus (SBL range 1.17–1.36 mm, mean
1.28±0.101 mm, n=3).
Habitus. Body form (Fig.18D) moderately convex, elongate ovoid, general pro-
portions (WE/SBL 0.38±0.005) and proportions of head (WH/WPm 0.74±0.020)
and pronotum (WPm/WE 0.78±0.018) average for group.
e integripennis species group of Geocharidius Jeannel, 1963... 107
Color. Body brunneorufous, appendages testaceous.
Microsculpture. Mesh pattern of irregularly isodiametric sculpticells present over
all dorsal surfaces of head and elytra. Pronotum and proepisternum smooth (without
evident microsculpture).
Prothorax. Pronotum moderately transverse (WPm/LP 1.28±0.010), with lateral
margins moderately constricted posteriorly (WPm/WPp 1.33±0.004). Posterior angles
slightly obtuse (100–110°). Width between posterior angles equal to width between
anterior angles (WPa/WPp 1.00±0.022).
Figures 21. Line drawings of spermatheca of Guatemalan and Honduran Geocharidius species. A G. an-
tigua (GUATEMALA, Sacatepéquez, Antigua), paratype B G. celaquensis (HONDURAS, Lempira, Cel-
aque National Park), paratype C G. lencanus (HONDURAS, Lempira, Celaque National Park), paratype
D G. comayaguanus (HONDURAS, Comayagua, Comayagua), paratype E G. disjunctus (HONDURAS,
Yoro, Pico Pijol National Park). Scale = 0.05mm.
Igor M. Sokolov & David H. Kavanaugh / ZooKeys 443: 61–118 (2014)
108
Elytra. Moderately convex, slightly depressed along suture, moderately wide (WE/
LE 0.65±0.015), without traces of striae. Humeri broadly rounded, in outline forming
right angle with longitudinal axis of body. Lateral margins convex, evenly divergent at
basal third, evenly rounded to apex in apical third.
Male genitalia. Median lobe of aedeagus (Fig.19Q) with shaft long and subpar-
allel, apex small and rounded. Ventral margin straight. Dorsal sclerites of internal
sac in form of a long plate, tapered apically into a long agellum, and abruptly wid-
ened basally, with a ventral appendix and pointed semicircular enlargement near
basal orice (Fig.19Q–T). Right paramere with long and narrow apical constric-
tion (Fig.19S). Left paramere with long and narrow apical constriction (Fig.19R).
Ring sclerite with handle triangular, slightly asymmetrical and pointed apically
(Fig.20E).
Female internal genitalia. Spermatheca sclerotized, fusiform, almost straight, ta-
pered basally, with cornu and nodulus of approximately equal length (Fig.21E). Length
of spermathecal gland greater than length of spermatheca. Spermathecal duct coiled.
Geographical distribution. is species is known only from two remote localities
in the Honduran Interior Highlands, situated in Yoro and Francisco Morazán Depart-
ments.(Fig.22, yellow triangles).
Way of life. Specimens were collected by sifting cloud and upper montane forest
litter at middle to high elevations (1300–2100 m).
Relationships. e shape of dorsal sclerites of the internal sac (Fig.19Q) of males
suggests that this species is closely related to the Guatemalan G. antigua (Fig.19A),
described above
Geocharidius lencanus sp. n.
http://zoobank.org/B41C1418-E0FB-4896-BE0F-827FE08A84D1
Figs 5C, 6G, 18B, 19H–J, 20C, 21C, 22, 23
Type material. HOLOTYPE, a male, in KUNHM, point-mounted, dissected,
labeled: \ HONDURAS: Lempira Dept., P.N. Celaque, nr. Gracias, Campami-
ento Naranjo, 14°32.7'N, 88°39.7'W, 2500m, 12-13-V-2002, cloud forest litter R.
Anderson, RSA2002-020 \ SM0… KUNHM-ENT \ HOLOTYPE Geocharidius
lencanus Sokolov and Kavanaugh 2014 [red label] \. PARATYPES: A total of 6
specimens (3 males and 2 females were dissected), deposited in CAS, CMNC and
KUNHM; 4 specimens labeled same as holotype; 2 specimens labeled: \ HONDU-
RAS: Lempira Dept., P.N. Celaque, nr. Gracias, Campamiento Naranjo, 2500m,
N14°32.7', W88°39.7', 12–13.V.2002, cloud forest litter R. Anderson, 2002-020E
\ CMNC \.
Type locality. Honduras, Lempira Department, Celaque National Park.
Etymology. e specic epithet is a Latinized adjective in the masculine form based on
the name of the indigenous people, the Lenca, living in the territory of Celaque National
Park during historic times.
e integripennis species group of Geocharidius Jeannel, 1963... 109
Figure 22. Map of southern Mexico, Guatemala and adjacent part of Honduras, showing positions of locality records for the species of Geocharidius: white diamond,
G. andersoni; white circles (black point in a circle shows “terra typica” for the species), G. zullinii; white squares, G. vignatagliantii; white triangle, G. gimlii; black and
white triangle, G. integripennis; green squares, G. longinoi; green circles, G. erwini; green owers, G. minimus; green diamonds, G. balini; green triangle, G.jalapensis;
yellow quadrangle, G. antigua; yellow diamond, G. lencanus; yellow ower, G. celaquensis; yellow triangles, G. disjunctus; yellow circles, G.comayaguanus. Physiographic
features: CH, Chiapas Highlands; GC, Guatemalan Cordillera; HIH, Honduran Interior Highlands; MTZ, Motagua Fault Zone; SC, Sierra de los Cuchumatanes;
SMC, Sierra Madre de Chiapas. Elevation scale bar is given in meters.
Igor M. Sokolov & David H. Kavanaugh / ZooKeys 443: 61–118 (2014)
110
Recognition. Externally, members of this species represent a larger version of G.
celaquensis adults, described above. Adults of G. lencanus are distinguished from those
of other members of the integripennis species group by the following combination of
external characters: medium to large size, rather wide habitus and fully microsculp-
tured dorsal body surface. Males and females of G. lencanus are distinguished from
those of the other members of the integripennis species group by the structure of the
median lobe and the shape of spermatheca, respectively.
Description. Size. Medium to large for genus (SBL range 1.30–1.47 mm, mean
1.39±0.091mm, n=4).
Habitus. Body form (Fig.18B) moderately convex, ovoid, general proportions
(WE/SBL 0.40±0.006), proportions of head (WH/WPm 0.73±0.008) and pronotum
(WPm/WE 0.77±0.010) average for group.
Color. Body rufotestaceous, appendages testaceous.
Microsculpture. Mesh pattern of irregularly isodiametric sculpticells present over all dor-
sal surfaces of head, pronotum and elytra. Proepisternum also with evident microsculpture.
Mouthparts. Maxillae and labium (Fig.5C).
Prothorax. Pronotum moderately transverse (WPm/LP 1.28±0.009), with lateral
margins moderately constricted posteriorly (WPm/WPp 1.34±0.019). Posterior angles
slightly obtuse (100–110°). Width between posterior angles nearly equal to the width
between anterior angles (WPa/WPp 1.01±0.012).
Legs. Protibia (Fig.6G).
Elytra. Moderately convex, slightly depressed along suture, markedly wide (WE/
LE 0.69±0.013), without traces of striae. Humeri rounded, in outline forming right
angle with longitudinal axis of body. Lateral margins convex, evenly divergent in basal
fourth, evenly rounded to apex in apical third.
Male genitalia. Median lobe (Fig.19H) with shaft subparallel with a long attenu-
ated preapical part, apex small and narrowly rounded. Ventral margin almost straight.
Dorsal sclerites of internal sac in form of a long narrow plate, agellum-like in apical
two-thirds, and slightly dilated and curved dorsally in basal third. Right paramere with
long and narrow apical constriction (Fig.19J). Left paramere with long and narrow api-
cal constriction (Fig.19I). Ring sclerite with handle rectangularly rounded (Fig.20C).
Female internal genitalia. Spermatheca sclerotized, fusiform, slightly dilated and
rectangularly bent apically, with short cornu and long nodulus (Fig.21C). Length of
spermathecal gland less than length of spermatheca. Spermathecal duct not coiled.
Geographical distribution. is species is known only from Celaque National
Park, in the Cerro las Minas range of Honduras (Fig.22, yellow diamond).
Way of life. Specimens were collected by sifting cloud forest litter at an elevation
of 2500 m.
Relationships. e rectangular shape of the handle of the ring sclerite (Fig.20C)
in males and the shape of spermatheca (Fig.21C) in females suggest a close relation-
ship with G. celaquensis (Figs 20B, 21B), described above.
e integripennis species group of Geocharidius Jeannel, 1963... 111
Figures 23. Diagrams illustrating size variation for sympatric pairs of closely and remotely related species
of the integripennis group at dierent geographical localities of Nuclear Central America. Closely related
pairs (for shared characters see subchapters on Relationships in the text for corresponding species): A Chi-
apas Highlands, Mexico B Mataquescuintla, Guatemala C Celaque National Park, Honduras. Remotely
related species: D Volcano Agua, Guatemala E Volcano Atitlan, Guatemala F Pico Pijol National Park,
Honduras. Legend: black dot – median; box – 25–75% range of values; whiskers – non-outlier range of
values; b – coecient of regression.
Discussion
A comprehensive phylogenetic and biogeographic analysis of Geocharidius is post-
poned until a thorough revision of all species of the genus has been completed. Below
we discuss only a few biogeographical and evolutionary issues, raised during our mor-
phological and distributional studies of the integripennis group species.
Igor M. Sokolov & David H. Kavanaugh / ZooKeys 443: 61–118 (2014)
112
Biogeographical notes
Distributional records of the integripennis species group to date are represented in Table1
and can be summarized as follows: e group includes mostly high elevation species: all
15 species live at elevations greater than 2000m, and only four of these also inhabit the
1300-2000m mid-elevation range. Physiographically (Fig.22), species of the group in-
habit the interior mountain ranges of the Chiapas (CH), Guatemalan (SC+MTZ) and
Honduran Highlands (HIH) along with the coastal Sierra Madre de Chiapas (SMC) and
its continuation as the Guatemalan Cordillera (GC). Geologically, these territories are
part of the Maya Block of the North American and the Chortis Block of the Carribean
Tectonic Plates, divided by the Motagua (or Motagua-Polochic) Fault Zone (Marschall
2007). e Motagua Fault Zone (Fig.22, MTZ) has been identied as the most impor-
tant physiographic barrier in Nuclear Central America, corresponding to many phylo-
geographic breaks in the distributions of dierent vertebrate taxa (Perdices et al. 2005,
Conchero Pérez et al. 2006; Castoe et al. 2009, Daza et al. 2010, Hardy et al. 2013). For
Geocharidius species, this zone separates montane areas with higher species diversity (to
the south and east) from those with lower diversity (to the north and west).
Each of the six montane areas has its own unique assemblage of Geocharidius spe-
cies, ranging in number from one to six species; and none of these species are shared
between montane areas. Consequently, any faunal connections between them are
through sister species rather than through conspecic populations; and this pattern
has implications for the timing of past dispersal and vicariance events (i.e., suggesting
somewhat greater antiquity for such events). As mentioned above, the Motagua Fault
Zone is a major physiographic barrier limiting the present distributions of integripennis
group species. A second evidently strong barrier is one between the faunas of the Gua-
temalan Cordillera and the Honduran Interior Highlands, separating the ranges of six
and four species endemic to each of these regions, respectively. e headwater valleys
of the Rio Paz to the south and Rio Motagua/Rio Shutague to the north, respectively,
are linked by gaps in the intervening uplands that do not exceed 900m in elevation,
creating a continuous break across these highlands that is 400m lower than the lowest
elevations at which any intergripennis species in the region has been recorded.
Among integripennis group members, six species have quite wide ranges within
their own montane area, while the other nine species are known from only one locality
or from two very close localities (G. longinoi) within their area. Within-group diversity
varies markedly between dierent parts of the region. Four of the six montane areas
are inhabited by one or two species, the Honduran Interior Highlands by four species,
and the Guatemalan Cordillera volcanic chain (Fig.22, GC) by six species. Within
these areas, such diversity is not based solely on a high number of locally restricted
forms. For example, three of the six species of the Guatemalan Cordillera have rather
wide ranges for Geocharidius species. is distributional pattern results in three cases
of sympatry among the species within the Cordillera. MacVean and Schuster (1981)
recorded similarly wide ranges for passalid beetle species and sympatry among them on
volcanoes of the Guatemalan Cordillera.
e integripennis species group of Geocharidius Jeannel, 1963... 113
Within the range of the integripennis species group, the Guatemalan Cordillera
occupies a special place and can be characterized by the highest number of the species
in total, the highest number of species with wide ranges and the highest number of
localities in which sympatry has been recorded. is combination of parameters may
indicate that, historically, this region played an important role as a staging area for
immigrants and as an intersection of dispersal routes of integripennis group species
dispersing between dierent areas.
Two cases of evident similarities in morphology of male and female genitalia be-
tween species inhabiting the Cordillera and their relatives outside the region seem to
support the above mentioned assertion. e similarity in median lobe structure be-
tween the eastern Guatemalan G. antigua (Fig.19A) and the Honduran G disjunctus
(Fig.19Q) is unequivocal; and presumably homologous structures in the median lobe
of males of G. integripennis and G. gimlii (Fig.15) are virtually identical. is leads
us to consider these pairs as sister species. ese examples connect the Cordilleran
fauna (Fig.22, GC) with faunas of the Honduran Highlands (HIH) and Sierra de los
Cuchumatanes (SC), respectively, thus, supporting our evaluation of the role of the
Guatemalan Cordillera as important in the dispersal history of the integripennis group.
Further, certain morphological similarities can be found between the west Gua-
temalan pair of species, G. gimlii, and G. integripennis, and among the Mexican trio
of species, G. andersoni, G. zullinii andG. vignatagliantii. Males of all three Mexican
species share a similar shape of the dorsal sclerites of the median lobe (Fig.9), and a
Table 1. Montane areas in Nuclear Central America occupied by the species of the Geocharidius integ-
ripennis species group.
Species Montane areas Elevation range
(in meters)
Number of
localities
CH SMC SC MTZ GC HIH
G. andersoni X 2750 1
G. zullinii X 2350–2600 6
G. vignataglianti X 2050 1
G. gimlii X 2780 1
G. longinoi X 2000–2750 2
G. integripennis X 3200 1
G. balini X 1625–2400 2
G. jalapensis X 2325–2660 1
G. erwini X 2140–2760 4
G. minimus X 1625–2175 2
G. antigua X 2350 1
G. disjunctus X 1300–2100 2
G. celaquensis X 2500 1
G. lencanus X 2500 1
G. comayaguanus X 1300–2130 3
TOTAL SPP. 2 1 1 1 6 4
Legend: CH, Chiapas Highlands; GC, Guatemalan Cordillera; HIH, Honduran Interior Highlands;
MTZ, Motagua Fault Zone; SC, Sierra de los Cuchumatanes; SMC, Sierra Madre de Chiapas.
Igor M. Sokolov & David H. Kavanaugh / ZooKeys 443: 61–118 (2014)
114
triangular handle of the ring sclerite (Fig.10). At the same time, females of G. vigna-
tagliantii can be connected with those of the west Guatemalan G. integripennis by the
short, sclerotized, and basally swollen basally spermatheca (Figs 11B, 17A), while the
dorsal sclerites of the median lobe of G. andersoni males (Fig.9H) are somewhat simi-
lar to the shortened variant of the dorsal sclerites of G. gimlii males (Fig.13A).
ese examples suggest that the Sierra de los Cuchumatanes (Fig.22, SC) may
have served as an important dispersal route from the Pacic coastal Guatemalan Cor-
dillera northward to the Chiapas Highlands (CH). Given its proximity to the Guate-
malan Cordillera, the Sierra Madre de Chiapas (Fig.22, SMC) would appear to have
been a more likely dispersal route northward, but we have no evidence that this route
has been used. It is worth noting that, based on morphology, all species of the group
living to the north of the Motagua Fault Zone appear to be rather closely related to
each other, whereas the species living to the south and to the east of the Motagua Fault
Zone appear to represent several morphologically dierent lineages. Perhaps the Mexi-
can representatives of the group are descendants of a comparatively recent dispersal
event involving one of the southern lineages.
Searching for concordant taxon-area relationships in other taxa reveals other car-
abids with similar distribution patterns. e distribution pattern for species of the
pterostichine subgenus Percolaus Bates, as described by Ball and Roughley (1982), is
identical to the distribution pattern of the Mexican-west Guatemalan set of Geocha-
ridius’ species and encompasses the Chiapas Highlands, the Sierra Madre de Chiapas,
Cerro Maria Tecún and Sierra de los Cuchumatanes. Interestingly, these authors sug-
gested that Pterostichus (Percolaus) championi Bates, from the Cerro Maria Tecún has
its closest known relative in the Sierra de los Cuchumatanes, the same relationship we
see between G. integripennis, presumably collected in the Cerro Maria Tecún, and G.
gimlii from the Sierra de los Cuchumatanes. Also, the distribution patterns of three
Mexican and one western Guatemalan Geocharidius (namely G. andersoni, G. zullinii,
G. vignatagliantii and G. gimlii ) correspond perfectly to the distribution pattern of the
species of Platynus jaegeri group (namely P. dilatipes Liebherr, P. robustus (Chaudoir)
and P. strictinotum Liebherr (Liebherr 1988).
Sympatric speciation
One common evolutionary trend among Anillina is syntopic miniaturization, a type
of sympatric speciation that produces a number of related species diering in size and
descendant from a common ancestor (Sokolov 2013). So, comparing average sizes of
adults of integripennis group species in localities where sympatry has been recorded
presented an interesting test of this idea. As noted above, we recognized six cases of
sympatry (Fig.22) involving the following species pairs (pairs marked by star are syn-
topic cases): G. andersoni – G. zullinii (Chiapas, Mexico), G. antigua – G. minimus
(Volcano Agua, Guatemala), G. minimus – G. balini* (Volcano Atitlan, Guatemala),
G. balini – G. jalapensis* (Mataquescuintla, Guatemala), G. lencanus – G. celaquensis*
e integripennis species group of Geocharidius Jeannel, 1963... 115
(Celaque National Park, Honduras), and G. comayaguanus – G. disjunctus* (Pico Pi-
jol National Park, Honduras). ese pairs of species can be grouped by the number
of shared morphological characters into two categories: (1) a group of more closely
related species that share two characters of male or female genitalia, namely the shape
of the male ring sclerite and the shape of the female spermatheca; and (2) a group of
more remotely related species that share only one character from either male or female
genitalia. Data on size dierences between species in all pairs are presented graphically
as box-and-whiskers plots with regression lines (Fig.23). For all pairs, we recorded
the dierences in averages of standardized body length between species. Our data sup-
port previous observations that the co-occurrence of taxonomically related anilline
species in the same habitat is often accompanied by dierentiation in their size (body
length) (Pérez-Gonzáles and Zaballos 2013, Sokolov 2013). Perhaps the persistent
(simultaneous) coexistence of two forms (a “larger” and a “smaller” form) in the litter
reects specic adaptations for living in only grossly overlapping microniches, which
dier in some unknown parameters of substrate interspaces and thereby harbor dif-
ferent microbiotas. Hypothetically, slight divergence in niche preferences might result
in divergence in target food preferences and decrease the number of contacts between
representatives of “larger” and “smaller” forms. is, in turn, which could reduce com-
petition between them and allow each to exploit resources more eectively.
At least in some cases, sympatry among anillines is a result of the dispersal of for-
merly allopatric taxa, typically in response to historical geological events and/or climate
change. Interestingly, dierence in sizes between a “larger” and a “smaller” species is
evidently greater in pairs of more closely related species than in the pairs of more re-
motely related species. is dierence can be seen visually in the slopes of regression
lines and the means of the regression coecients of these lines (Fig.23, b). Unfortu-
nately, the low number of observations does not allow us to analyze our data statisti-
cally and thereby evaluate how signicant the observed dierences between groups may
be. We can only speculate about the origins and signicance of dierences between the
two groups. For the present, we interpret our ndings as reecting dierences in his-
torical time at which each case of sympatry developed and, accordingly, by the length
of time during which disruptive selection was occurring. We presume that, in the cases
of the closely related species, we are dealing with intraspecic divergence, which was
continuing for much longer times than in the cases of remotely related species, sympa-
try among which we consider a result of postspeciation dispersal, and thus of compara-
tively recent origin. In the latter case, interspecic divergence occurred over a much
shorter time period, resulting in lesser dierences in size between co-occuring species.
Acknowledgements
Important loans were received from the University of Kansas Natural History Museum,
Canadian Museum National Collection, Carnegie Museum of Natural History and
the U.S. National Museum of Natural History, through the help of Zachary H. Falin
Igor M. Sokolov & David H. Kavanaugh / ZooKeys 443: 61–118 (2014)
116
(Collections Manager), Robert S. Anderson (Curator), Robert L. Davidson (Collections
Manager), and Terry L. Erwin (Curator) at those institutions, respectively. We thank
Terry Erwin (NMHH), ierry Deuve and Azadekh Taghavian (MNHN) for the loan
of types and other specimens under their care. is paper is based in part on material ob-
tained through sampling supported by the National Science Foundation through grant
DEB-0640015 (J.T. Longino, R.S. Anderson, P.S. Ward, Principal Investigators). e
rst author acknowledges the Evert I. Schlinger Foundation for the Schlinger Postdoc-
toral Fellowship at the California Academy of Sciences in 2012–2014.
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