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MEMOIRS
7 | OF THE
AMERICAN ENTOMOLOGICAL SOCIETY
ne : a:
=—_— NUMBER 36
BIOSYSTEMATICS OF THE GENUS
DICROTENDIPES KIEFFER, 1913 (DIPTERA:
-CHIRONOMIDAE: CHIRONOMINAE) OF
| - THE WORLD _
By
JOHN H. EPLER
THIS PUBL ATION IS ENT 10
ESTABLISH DATE OF ISSUE. PLEA
STAMP DATE OF RECtI
PUBLISHED BY THE AMERICAN ENTOMOLOGICAL SOCIETY
AT THE ACADEMY OF NATURAL SCIENCES
PHILADELPHIA
1988
MEMOIRS
OF THE
AMERICAN ENTOMOLOGICAL SOCIETY
NUMBER 36
BIOSYSTEMATICS OF THE GENUS
DICROTENDIPES KIEFFER, 1913 (DIPTERA:
CHIRONOMIDAE: CHIRONOMINAE) OF
THE WORLD
By
JOHN H. EPLER
—
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PUBLISHED BY THE AMERICAN ENTOMOLOGICAL SOCIETY
AT THE ACADEMY OF NATURAL SCIENCES
PHILADELPHIA
1988
SELWYN S. ROBACK
EDITOR
(Issued July 29, 1988)
PRINTED IN THE UNITED STATES OF AMERICA
TABLE OF CONTENTS
JNIBISSIEI SVAN COG et re gee re Eig gene lB er ar eg 1
CHAPTER I. INTRODUCTION AND NOTES ON THE HOLARCTIC
IDI GROMEN DUPE o5 apne re 6 8 ote Bee Siesta fae 2
INDNOLCKONOERY, Ts oes Gee eds, oa ote ob Aho Oke io oben Sos Men cernete eter 3
Terminology and Morphology .................----+-+-- 4
BAKO) OPEAY, | Saeis Oho MetON eC ERIGRO: OPER ORE mS eo ca eather ero et CLs ut ris nee 4
TED NO VOVOIONY 43:5 srauanaees nec ona a cmncons oie encmoW anor or tao tea ieeorer cemnecined 6
Notes on the Holarctic Dicrotendipes .................-.---- 12
Key to Adult Males of Holarctic Dicrotendipes ............... 13
Key to Known Pupae of Holarctic Dicrotendipes ............. 15
Key to Known 4th Instar Larvae of Holarctic Dicrotendipes...... 16
CHAPTER II. A REVIEW OF THE AFROTROPICAL
DICROTENDIPES, WITH DESCRIPTIONS OF THE
TIN TIN ACI SSAVAIGE Sree iia sates acum aeeie eeeue re 32
Key to Adult Males of Afrotropical Dicrotendipes ............ 33
CHAPTER III. THE NEOTROPICAL DICROTENDIPES ....... 58
Key to Adult Males of Neotropical Dicrotendipes ............. 58
CHAPTER IV. THE DICROTENDIPES OF THE ORIENTAL-
AUSTRATASTAN REGION 2 234 csc 6 ee Ps ee 110
Key to Adult Males of Oriental-Australasian Dicrotendipes ....... 111
Key to Known Pupae of Oriental-Australasian Dicrotendipes ...... 112
Key to Known Larvae of Oriental-Australasian Dicrotendipes ..... 113
CHAPTER Vo ZOOGEOGRAPEIY: _. cis 280 5.oe eae cehs bs: Ie 178
CHAPTER VAt PEIVISOGEINNG ony tern e ee ogo eire a sist Gs es aces oe acne 194
ANCKNOWEEDGE MENUS v5 ee. cakoncrit tke cols uo tings i Gocusereebentucne 2 204
PEPE RAROREICIMED re ails. a eee eee aoe pr eee 205
APPENDIX 1-List of recognized species names in the genus Dicrotendipes,
andithein distri Duttomiesy< eb ceneiees sc westua a aeter sid &, oe nee: 211
APPENDIX 2-List of recent name changes and current name as recognized
MMNCMISHPAPSH ce teasers owes tne eet ree cha Auta he cece te Goa e amen 212
HINT D) EDC Pe ee ee nee geen Seen nent res: ceanomene owe 213
MEM. AMER. ENT. SOC., 36
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MEMOIRS
OF THE
AMERICAN ENTOMOLOGICAL SOCIETY
NUMBER 36
BIOSYSTEMATICS OF THE GENUS DICROTENDIPES
KIEFFER, 1913 (DIPTERA: CHIRONOMIDAE:
CHIRONOMINAE) OF THE WORLD
BY
JOHN H. EPLER!
ABSTRACT — The taxonomy, zoogeography and phylogeny of the genus Dicrotendipes Kief-
fer, 1913 (Diptera: Chironomidae) are reviewed on a world-wide basis. Data from a previous study
of the Nearctic fauna (Epler 1983, 1987a) are supplemented by studies of the Holarctic, Neotrop-
ical, Afrotropical, Oriental, Australian and Oceanian faunas. Seventy-two species names are rec-
ognized; the type-species of the genus is Chironomus septemmaculatus Becker, 1908.
Notes on the Holarctic fauna are provided; 25 species are recognized from the area. Dicroten-
dipes incurvus (Sublette, 1964) is considered a junior synonym of D. tritomus (Kieffer, 1916). Keys
for the adult males, pupae and larvae of the Dicrotendipes of the Holarctic region are included.
The taxonomy of 14 species recognized from the Afrotropical region is reviewed; these are: D.
bredoi (Goetghebuer, 1936), D. chambiensis (Goetghebuer, 1936), D. collarti (Goetghebuer, 1936),
D. cordatus Kieffer, 1922, D. ealae (Freeman, 1957), D. freemani [nom. nov. for D. binotatus
(Kieffer, 1911)], D. fusconotatus (Kieffer, 1922), D. kribiicola (Kieffer, 1923), D. leucolabis Kieffer,
1922, D. nigrolineatus (Freeman, 1957), D. peringueyanus Kieffer, 1924, D. schoutedeni (Goet-
ghebuer, 1936), D. septemmaculatus (Becker, 1908) and D. sudanicus (Freeman, 1957). The larvae
and pupae of D. fusconotatus, D. kribiicola, D. septemmaculatus and D. sudanicus, and the pupa
of D. cordatus are described. New junior synonyms for D. septemmaculatus are: D. formosanus
Kieffer, 1916, D. frontalis Kieffer, 1916, Chironomus hirtitarsis Johannsen, 1932, and D. rajas-
thani Singh & Kulshrestha, 1977; Ch. punctatipennis Kieffer, 1910 is considered a probable junior
synonym. A key to adult males is provided.
Nineteen species are recorded from the Neotropical region. Previously described species dis-
cussed include: D. aethiops (Townes, 1945), D. alsinensis (Paggi, 1975), D. californicus (Jo-
hannsen, 1905), D. crypticus Epler, 1987, D. embalsensis Paggi, 1987, D. nestori Paggi, 1978, D.
1 Entomology, Florida A&M University, Tallahassee, FL 32307.
1
MEM. AMER. ENT. SOC., 36
D; GENUS DICROTENDIPES
obrienorum Epler, 1987, D. pellegriniensis Paggi, 1987 and D. sinoposus Epler, 1987. The adult
males of 10 new species are described: D. amazonicus, D. dasylabidus, D. demissus, D. fittkaui,
D. palearivillosus, D. paradasylabidus, D. paterjohni, D. radinovskyi, D. reissi and D. soccus;
adult females are described for D. amazonicus and D. demissus; and pupae are described for D.
fittkaui and D. reissi. A key is provided to identify Neotropical adult males.
A total of 19 species are recognized from the combined Oriental, Australian and Oceanian re-
gions. Three species previously considered to be Dicrotendipes are removed: Ch. blandellus Kieffer,
1906, D. paxillus Guha, Chaudhuri & Nandi, 1982, and D. socionotus Guha, Chaudhuri & Nandi,
1982. New species and stages described are: D. balciunasi (adult male), D. cumberlandensis (adult
male and female, pupa, larva), D. jobetus (adult male), D. jonmartini (adult male, pupa, larva),
D. lindae (adult male), D. pseudoconjunctus (adult male, pupa, larva) and D. sarinae (adult male
and female, pupa, larva). The adult males and females are redescribed, and pupae and larvae
described, for: D. candidibasis (Edwards, 1924), D. conjunctus (Walker, 1856) and D. pelochloris
(Kieffer, 1912); adult males are redescribed for D. bilobatus Kieffer, 1917 and D. tenuiforceps
(Kieffer, 1913). The pupa and larva of D. flexus (Johannsen, 1932) are redescribed, and the tax-
onomy of D. /eei (Freeman, 1961) and D. taylori (Freeman, 1961) is reviewed; D. semiviridis is
considered a species inquirenda. New synonyms include: Ch. melanocnemis Edwards, 1928 (junior
synonym of D. candidibasis); Limnochironomus niveicauda Kieffer, 1921, Ch. inferior Johannsen,
1932, Ch. (D.) wirthi Freeman, 1961, Xenochironomus loripes Guha & Chaudhuri, 1981, Kimius
hoonsooi Ree, 1981 Gunior synonyms of D. pelochloris) and Ch. (D.) innisfailensis Freeman, 1961
Gunior synonym of D. tenuiforceps); D. bilobatus is removed from synonymy with D. conjunctus.
Keys are provided for adult males, pupae and larvae.
Distributions of selected species are discussed and distribution maps provided for D. aethiops,
D. californicus complex, D. flexus, D. lobiger, D. modestus, D. nervosus, D. pelochlo-
ris, D. septemmaculatus, D. sinoposus, D. tenuiforceps and D. tritomus.
The phylogeny of Dicrotendipes species known in all 3 life stages is analyzed cladistically. Results
indicate that 3 major lineages occur within the genus, with at least 9 species groups.
CHAPTER I.
INTRODUCTION AND NOTES ON THE
HOLARCTIC DICROTENDIPES
The genus Dicrotendipes was recently revised for the Nearctic region (Epler
1987a). This monograph is basically a continuation of that revision, and includes
general notes on the genus and the Holarctic species (Chapter I), a review of the
Afrotropical species (Chapter II), a revision of the Neotropical species (Chapter
III), and a revision of the species found in the combined Oriental, Australasian and
Oceanian region (termed the Oriental-Australasian region in this paper) (Chapter
IV). The zoogeography of selected species in the genus is discussed in Chapter V.
A phylogenetic analysis is included in Chapter VI. Appendix 1 is a list of the rec-
ognized species (72) in the genus; Appendix 2 presents a list of the most recent syn-
onymies in the genus.
JOHN H. EPLER 3
METHODOLOGY
Methods used for specimen preparation and measurement are similar to those
explained in Epler (1983, 1987a, 1987b). Most measurement methods are illus-
trated in Figs. 1-5. All measurements are in micrometers (um) (unless stated oth-
erwise) and consist of the range, mean, and, in parentheses, the number of speci-
mens utilized if different from the number (n) cited at the beginning of the
description.
All illustrations were drawn by the author. The hypopygial figures are drawn to
show a ventral and internal view (of apodemes) on the left, and a dorsal view to
the right. Lateral aspects of hypopygia are drawn with the gonocoxae and
gonostyli removed. The dorsal hypopygial figures are drawn with the sensilla chae-
tica of the superior volsella as they may appear when viewed with a compound mi-
croscope, i.e., dorsal. In actuality the sensilla chaetica of the superior volsella are
generally ventral (Fig. 3A).
Specimens examined in this study were borrowed from various individuals
and/or institutions. The following abbreviations are used to denote collections
from which material was borrowed or in which type material will be placed:
AN — Australian National Insect Collection, CSIRO, Canberra, Australia
(D.H. Colless).
BM — British Museum (Natural History), London, England, U.K. (P.S.
Cranston).
CU — Clemson University, Clemson, SC, U.S.A. (M.W. Heyn).
FS — Florida State Collection of Arthropods, Florida A & M University, Tal-
lahassee, FL, U.S.A.
HH — H. Hashimoto, Shizuoka University, Shizuoka, Japan.
IL — Instituto de Limnologia, Universidad Nacional de la Plata, Berisso, Ar-
gentina (A.C. Paggi).
IN — Instituto Nacional de Pesquisas da Amazonia, Manaus, Brazil.
IP — Institut ftir Pflanzenschutzforschung Kleinmachnow, Eberswalde-Fi-
now, D.D.R (H.J. Miiller).
JB — J. Balciunas, Fort Lauderdale, FL, U.S.A.
JE — J.H. Epler, Florida A & M University, Tallahassee, FL, U.S.A.
JM — J. Martin, University of Melbourne, Melbourne, Australia.
KM — Koninklijk Museum voor Midden-Afrika, Tervuren, Belgium (E.
DeConinck).
LH — L. Hare, Université du Quebec, Quebec, Canada.
NM — Naturhistorisches Museum, Vienna, Austria (R. Contreras-
Lichtenberg).
SP — School of Public Health and Tropical Medicine, University of Sydney,
Sydney, Australia (M.L. Debenham).
MEM. AMER. ENT. SOC., 36
4 GENUS DICROTENDIPES
UB — University of Burdwan, Burdwan, India (P.K. Chaudhuri).
US — United States National Museum, Washington, D.C., U.S.A. (B.V.
Peterson).
ZH — Zoologisches Museum, Humboldt University, Berlin, FR.G (through
P.S. Cranston).
ZM — Z.Moubayed, Université Paul Sabatier, Toulouse, France.
ZS — Zoologische Staatssammlung, Munich, F.R.G (E.J. Fittkau & FE. Reiss).
TERMINOLOGY AND MORPHOLOGY
Terminology and abbreviations for body parts and ratios follow Saether (1980)
and Epler (1987a, 1987b), and are illustrated in Figs. 1-7. LWR refers to the
length/width ratio of the superior volsella. See Epler (1987a) for a more detailed
discussion of the morphology of Dicrotendipes. Some new terms are introduced
here.
If setae present on the dorsum of the hypopygium (adult male) could not be dis-
tinguished as medial and/or dorsal basal setae, the combined term dorsomedial
setae was used.
Freeman (1957:366) noted the presence of ‘‘5-6 black spines ventrally’ on ab-
dominal sternite VI of D. cordatus Kieffer. Epler (1987a) also noted the presence
of ‘‘darker, flattened, bluntly tipped setae, which are easily lost’’ on most Nearctic
species. Epler (1987b) referred to these setae as ““flattened setae on S VI.”
Throughout the present monograph these setae, which are an apparent autapo-
morphy for Dicrotendipes, are termed ventral accessory setae. Although they are
most often found onS VI, ventral accessory setae may also occur onS V andS VII,
and are present on males and females of most species in the genus. They are absent
from members of the D. californicus complex and the septemmaculatus group.
Ventral accessory setae may appear as darker, more robust setae, as slightly flat-
tened, darker and more robust setae, or as darker, stout spine-like setae (Fig. 1F).
I was mistaken in my Nearctic revision (Epler 1987a) when I stated that the large
subquadrate to oval areas on the frontoclypeal apotome of D. /obiger (Kieffer) and
the frontal apotome of D. /eucoscelis (Townes) were dorsal (on the apotome). These
areas are ventral (on the apotome) or internal. Similar areas are found on the larval
frontal apotomes of D. kribiicola (Kieffer), D. notatus (Meigen), D. conjunctus
(Walker) and its close relatives, and D. pelochloris (Kieffer).
BIOLOGY
Adults of Dicrotendipes have been considered as pests due to large emergences
(Frommer & Rauch 1971; Ali & Mulla 1980), and have been implicated in allergic
reactions in humans in Africa (Cranston et al. 1983).
JOHN H. EPLER 5
The immature stages are found in both lentic and lotic habitats, but are generally
more prevalent in lentic situations. The larvae in general are not ‘‘burrowers”’ as
listed in Coffman & Ferrington (1984), but would be better classified as ‘‘clingers’’
and/or “‘sprawlers.’’ Although often reported in the literature as ‘‘bottom-dwell-
ing’’ (Roback 1957) or “‘epibenthic, unattached’’ (Beck 1977), the majority of Di-
crotendipes larvae are found on the surface of aquatic vegetation, and among or
on vegetation or Aufwuchs on rocks, logs, or similar substrata. The larvae con-
struct silken tubes which are attached to the substrate. The pupal stage is also spent
in the tube. I have rarely collected Dicrotendipes larvae from bottom mud, al-
though larvae do occur commonly in algal mats on the bottom. Darby (1962) re-
ported finding D. californicus (Johannsen) larvae on the surface of bottom mud
in late May and early June in California, but found that as submerged vegetation
increased, the vegetation became the favored habitat. Lenz (1954-1962) reported
Dicrotendipes \arvae (as Limnochironomus) as predominantly littoral and living
onrocks, plant stems, and among aquatic vegetation, but also noted their existence
in benthic detritus in water 10 m or more in depth, as well as among mosses and
other vegetation on rocks in flowing water. I have found D. fumidus (Johannsen)
on Myriophyllum and algae covered rocks, D. lobus (Beck) associated with Najas,
Juncus and Spartina in salt marshes and D. modestus (Say) on Myriophyllum and
Typha; and I have examined reared D. candidibasis (Edwards) specimens collected
from streamside vegetation in Fijiand D. septemmaculatus larvae collected on Hy-
drilla from Burma, Indonesia and Malaysia. Edward (1964) found the larvae of
D. conjunctus inhabiting ‘‘algal mats and slime.’’ I have collected D. crypticus
Epler larvae from vegetation on rocks in the swiftly flowing portions of a New
Mexican creek; I have scraped D. neomodestus (Malloch) larvae from algae in
sheets of water flowing over a dam in Pennsylvania, and have collected the ap-
parent larva of D. adnilus Epler from periphyton from midstream rocks in the
mountains of southeastern Arizona (Epler 1987a).
Hudson (1987:table 1) listed chironomid genera that have species with
unusual larval habitats or life histories. Among the 9 unusual habitats were
listed 2 in which Dicrotendipes larvae may be found. The first of these
categories is “‘symbiotic.’’ Disney (1975) found D. peringueyanus Kieffer
living phoretically on the African river crabs Potomonautes africanus (A.
Milne-Edwards) and P. pobeguini (Rathbun) in Cameroon. The second un-
usual larval habitat is ‘‘water held in plants.’’ Epler (1987a) examined a D.
leucoscelis larva which had been collected and reared from the water in a
bromeliad from Florida. At least 4 species (D. inouei Hashimoto, D. lobus
(Beck), D. modestus (Say) & D. pallidicornis (Goetghebuer)) occur in brack-
ish water; 2 of which, D. inouei and D. lobus, apparently occur exclusively
in brackish water. The larvae of D. flexus and D. pelochloris (as inferior)
have been reported from hot springs (40°) from Sumatra (Lenz 1937).
MEM. AMER. ENT. SOC., 36
6 GENUS DICROTENDIPES
Dicrotendipes larvae feed on algae, detritus, or the microorganisms as-
sociated with it. Lenz (1954-1962) reported that he observed larvae spinning
small ‘‘catch-nets’’ within their tubes. By undulating their bodies, the larvae
produced a current which forced water through the catch-net. After a period
of time, the catch-net and its catch of detritus and microorganisms were
consumed by the larvae. I have not observed feeding in Dicrotendipes, but
many larval guts I’ve examined were packed with algae and detrital material.
The larval undulations probably serve a respiratory function, introducing
fresh water in the larval tube. I have also observed pupae undulating in their
tubes.
TAXONOMY
Kieffer (1913b:23) established the genus Dicrotendipes from African ma-
terial, stating (my translation) ‘‘this genus differs from all others in that
the inferior appendages of the male forceps are bifurcated.’’ Only one spe-
cies, D. pictipennis, was included, which by monotypy (International Code
of Zoological Nomenclature (ICZN), Art. 68(d)) was the type-species for
the genus. When Freeman (1957) relegated Dicrotendipes to subgeneric sta-
tus within Chironomus, the epithet pictipennis became a junior secondary
homonym of Ch. pictipennis Philippi, 1865. The next available name for
this species was quatuordecimpunctatum Goetghebuer, originally described
in Polypedilum by Goetghebuer (1936). Freeman (1957) considered qua-
tuordecimpunctatum a subspecies of D. pilosimanus Kieffer, 1914; thus, D.
pilosimanus became the type-species of Dicrotendipes. Although Dicroten-
dipes was later returned to full generic status (Hamilton et al. 1969), the
name D. pictipennis was invalid under ICZN Art. 59(b), because it was a
junior secondary homonym replaced before 1961. Cranston and Armitage
(1988) rediscovered the holotype of Chironomus septemmaculatus Becker,
1908, and considered it to be a senior synonym of D. pilosimanus Kieffer,
1914. The correct name for the type-species of the genus is now Ch. sep-
temmaculatus Becker.
The genus Limnochironomus was established from the Palaearctic by
Kieffer (1920:166), who described the apex of the inferior volsella (appen-
dage) as ‘‘sometimes simple, sometimes imperfectly bi- or trilobed.’’ Several
species were included. The type-species, by original designation, is Tendipes
falciformis Kieffer, 1912. This species is a junior synonym of Chironomus
nervosus Staeger, 1839.
Kieffer (1922) later described 3 more species in Dicrotendipes: trilabis,
cordatus, and leucolabis. In his description of the inferior volsella of D.
cordatus, Kieffer (1922:65) stated ‘‘the apex strongly broadened like a heart,
JOHN H. EPLER 7
the 2 lobes not as long as wide, divided by a curved indentation.”’ By in-
cluding cordatus in Dicrotendipes, Kieffer expanded the concept of the ge-
nus. In the other species of Dicrotendipes he described, the apex of the
inferior volsella is deeply bifid. However, the cordiform apex of the inferior
volsella of D. cordatus would also fit the description of the same structure
in Limnochironomus, as can be seen from Kieffer’s figure (Kieffer 1922:fig.
66). My examination of the pupa of D. cordatus (see Chapter IJ) indicates
that Kieffer was correct in including the species in Dicrotendipes.
Calochironomus was first mentioned by Kieffer (1921a) in a key to genera.
No species were mentioned. In a later paper on African chironomids (Kieffer
1922:66), he included 6 species in the genus and designated the type-species
as fusconotatum, which was described from a female.
Kieffer (1921b:590) established the genus Carteria based on material from
the Philippines and Formosa, naming Chironomus longilobus (Kieffer, 1916)
(originally described in Tendipes), as the type-species. However, Carteria
was preoccupied by Carteria Diesing, 1866 (a protozoan), and was renamed
Carteronica by Strand (1928).
Edwards’ (1929) concept of Chironomus was much broader than that of
the continental European workers (e.g. Thienemann) at that time. He did
not consider Limnochironomus distinctive enough to rate subgeneric rank,
but relegated it to his Group C of Chironomus.
Goetghebuer (1936, 1937-1954) also defined Chironomus in a broad sense.
He considered Carteria, Calochironomus, Dicrotendipes and Limnochiron-
omus to be subgenera of Chironomus. He apparently was unaware of
Strand’s 1928 paper renaming Carteria as Carteronica.
Aristovskaya (1935:114) established the genus Sernowia in a footnote,
stating (loosely translated): ‘‘name at present for form previously described
by N.N. Lipina as Chironominae genuinae No. 6.’’ However, no type-spe-
cies was designated. Pankratova’s subsequent listing (in Chernovskii 1949)
of Chironominae genuinae No. 6 and Sernowia (as Sernovia) as synonyms
of Limnochironomus ex gr. nervosus Staeger does not satisfy the require-
ments for designation of a type-species (ICZN Art. 69). The name Sernowia
thus becomes a nomen nudum and is not available (ICZN Art, 13(b)). Ser-
nowia is also misspelled as ‘‘Sernorwia’’ in a table (Aristovskaya 1935:118).
Lenz (1937:6) described the larva and pupa of Chironomus (Limnochi-
ronomus) flexus Johannsen, 1932, and established it as a new genus, Lim-
notendipes. By monotypy, the type-species was flexus.
Cladotendipes was established by Lenz (1937) for a single species, Chi-
ronomus inferior Johannsen, 1932. Sublette and Sublette (1973) placed Ch.
inferior in Dicrotendipes, but made no mention of Cladotendipes. See also
Ashe (1983).
MEM. AMER. ENT. SOC., 36
8 GENUS DICROTENDIPES
Townes (1945), in his comprehensive monograph of the Nearctic Chi-
ronomini (as Tendipedini), kept Limnochironomus as a subgenus of Ten-
dipes (=Chironomus), and considered Limnotendipes a junior synonym.
Freeman (1955a) synonymized Calochironomus and Limnochironomus
with Dicrotendipes. The type-species of Calochironomus, now known as D.
fusconotatus, was described from a female. The male, apparently associated
with the female by the pattern of wing spots and banded legs, has an inferior
volsella with a deeply bifid apex, and is a Dicrotendipes in the sense of
Kieffer’s (1913b) original description of the genus. My examination of the
immature stages (Chapter II) confirms this. Freeman also noted that the
only species of Calochironomus in which the male was known to Kieffer,
C. oxylabis Kieffer, 1922, was quite unlike the other species in the genus
and was a Chironomus (Einfeldia). This species is now considered a junior
synonym of Chironomus formosipennis Kieffer, 1908 (Freeman 1957; Free-
man and Cranston 1980). Freeman (1955a) considered Limnochironomus a
synonym of Dicrotendipes mainly because when Kieffer (1922) expanded
the concept of Dicrotendipes by including D. cordatus, a species with a
codiform apex to its inferior volsella, there was no longer a difference be-
tween the 2 genera. By Kieffer’s (1920:106) own definition the inferior vol-
sella of Limnochironomus could be imperfectly bi- or trilobed; a
cordiform apex (as in D. cordatus) is imperfectly bilobed. As has been shown
in Contreras-Lichtenberg (1986) and Epler (1987a), on the basis of the im-
mature life stages Limnochironomus is a junior synonym of Dicrotendipes.
Freeman (1957) redefined Chironomus in much broader terms and included Di-
crotendipes as a subgenus. He also synonymized Carteria and Carteronica with
Chironomus (Dicrotendipes). The gonostylus of Carteronica species was very short
and wide. Goetghebuer (1936:465) had described a species from Africa with a sim-
ilar gonostylus, Chironomus (Carteria) regalis, and Freeman (1957) included this
species and 2 new species with similar gonostyli, penicillatus and multispinosus, in
his Chironomus (Dicrotendipes). He did not use structural similarities in the males
to synonymize these 2 genera, but relied on similar thoracic color patterns of the
females.
Hamilton et al. (1969) used the genus Chironomus in a stricter sense. Instead of
a large genus with many subgenera, they preferred to use several smaller genera,
in accordance with current European workers. Dicrotendipes was elevated back to
the genus level.
Calochironomus and Limnochironomus are definitely junior synonyms of Di-
crotendipes. However, Carteronica is not. This is based on examination of reared
material of Chironomus (Carteria) regalis Goetghebuer and Chironomus longi-
lobus (Kieffer) (type-species of Carteronica) from the Orient and Africa made
available to me by Dr. L. Hare. Carteronica (replacement name for Carteria Kief-
JOHN H. EPLER 9
fer, 1921) was removed from synonymy with Dicrotendipes (Epler 1987a). It should
also be noted that the Afrotropical species which resemble Carteronica placed in
Dicrotendipes by Freeman & Cranston (1980) (crispi Freeman, multispinosus
Freeman, penicillatus Freeman and regalis Goetghebuer), are not Carteronica, but
represent another, new, genus (manuscript in preparation).
Kimius, a monotypic genus established by Ree (1981) for the species hoonsooi,
was included as a synonym of D. niveicaudus (Kieffer) (= D. pelochloris) by Sasa
& Hasegawa (1983). I have seen specimens of this species and agree with the
synonymy.
Generic synonyms are listed below. These are followed by an emended diagnosis
for the genus incorporating new information discovered during the present study.
A list of recognized species names (72) is given in Appendix 1. Appendix 2 is a list
of recent specific synonymies in Dicrotendipes.
Genus DICROTENDIPES Kieffer
Dicrotendipes Kieffer 1913:23. Type-species: Dicrotendipes pictipennis Kieffer, 1913 (junior
homonym, preoccupied by pictipennis, Philippi, 1865; = Polypedilum quatuordecimpuncta-
tum Goetghebuer, 1936 = Dicrotendipes pilosimanus Kieffer, 1914 = Chironomus septem-
maculatus Becker, 1908), by monotypy.
Limnochironomus Kieffer 1920:166. Type-species: Tendipes falciformis Kieffer, 1912, (=nervosus
Staeger), by original designation.
nec Carteria Kieffer 1921b:590. Type-species: Chironomus longilobus (Kieffer, 1916), by original des-
ignation (junior homonym preoccupied by Carteria Diesing 1866); Freeman & Cranston 1980:190;
Ashe 1983:15,21.
Calochironomus Kieffer 1921a:274. Type-species: Calochironomus fusconotatum Kieffer, 1922, by
designation of Kieffer (1922:66).
nec Carteronica Strand 1928:48 (replacement name for Carteria Kieffer 1921); Freeman & Cranston
1980:190; Ashe 1983:15,21.
Chironomus (Limnochironomus), Goetghebuer 1928:50, Goetghebuer 1936:464, Goetghebuer 1937-
1954:19.
Chironomus (Chironomus) Group C, Edwards 1929:386.
Sernowia Aristovskaya 1935:114. Nomen nudum. (No type-species designated).
nec Chironomus (Carteria), Goetghebuer 1936:465.
Chironomus (Dicrotendipes), Goetghebuer 1936:466; Goetghebuer 1937-1954:31.
Chironomus (Calochironomus), Goetghebuer 1936:467.
Chironomus (sensu stricto), Goetghebuer 1936:470 (partim).
Limnotendipes Lenz 1937:6. Type-species: Chironomus (Limnochironomus) flexus Johannsen, 1932,
by monotypy.
Cladotendipes Lenz 1937:7. Type-species: Chironomus inferior Johannsen, 1932. by monotypy.
Tendipes (Limnochronomus), Townes 1945:102; Hauber & Morrissey 1945:287; Roback 1957:109.
Dicranotendipes Kruseman 1949:254 (misspelling).
Kimius Ree 1981:217. Type-species: Kimius hoonsooi Ree, 1981 (synonym of D. pelochloris (Kieffer,
1912)) by original designation; Sasa & Hasegawa 1983:321.
MEM. AMER. ENT. SOC., 36
10 GENUS DICROTENDIPES
DESCRIPTION: Adult male. Medium sized chironomids, light yellow-green to dark green or light
brown to dark red-brown. Legs sometimes banded, wings sometimes spotted or banded.
Eyes bare. Temporal setae in 1-3 rows beginning mesad to dorsomesal extension of eye, ending be-
hind eye. Frontal tubercles present, small (2 um) to medium (28 zm), very rarely absent. Antennal fla-
gellum with 11 flagellomeres. Maxillary palp 5-segmented, basal segment weakly sclerotized and bear-
ing one large lateral seta; segment 3 with specialized sensillae near apex. Clypeus subquadrate, setose.
Cibarial setae present.
Antepronotum bare, narrowed and weakly notched dorsomesally. Thoracic scar well developed; hu-
meral pit usually present dorsocaudally to thoracic scar. Scutal tubercle well to poorly developed, or
absent. Acrostichal setae when present in double row (absent or reduced in some species), anteriorly
beginning close to antepronotum and running posteriorly to anterior base of scutal tubercle (if present)
or approximately mid-scutum. Dorsocentral setae in 1-3, usually 2, rows/side. Scutellar setae in 1-3
rows. Supraalar seta 1, rarely 2, /side. Prealar setae 2-7/side. Wing membrane without setae; squama
with setal fringe. Brachiolum with 1-5 setae and 2 groups of campaniform sensilla; R, R; and R4+5 with
setae; costa ends at R4+5; FCu proximal, below, or distal to RM.
Metatarsal beard present or absent on foreleg, almost always present on hind leg. Foretibia with in-
ner apical rounded scale which projects slightly beyond similar scale on outer tibial apex. Middle and
hind tibiae with 2 combs each, barely separated, each comb bearing one spine which projects beyond
others. Sensilla chaetica present on metatarsus of middle leg, usually confined to apical 1/5, occasion-
ally running almost entire length of tarsomere; also sometimes present on hind metatarsus. Pulvilli 2
entire lobes; empodium thin, with sparse ventral fringe.
Abdominal sternite VI (and sometimes V or VII) with or without a medial group of ventral accessory
setae, which are easily lost.
Gonostylus usually evenly curved on inner and outer margins, bearing one short, stout seta apically
and several to many longer setae on inner preapical margin. Superior volsella well developed; digiti-
form, deltoid, pediform, elongate-cylindrical or bifid with sclerotized superior portion and mem-
branous-lamelliform inferior portion; usually bare dorsally, bare or with microtrichia ventrally; often
membranous apically; bearing several to many large sensilla chaetica on mesal and/or ventral surface.
Inferior volsella well developed, usually strongly bowed dorsoventrally, usually with an expanded
clubbed or slightly to deeply bifid or trifid apex bearing several to many long, strong sensilla chaetica.
Rarely, a membranous median volsella present (2 species). Anal point of hypopygium pyriform to elon-
gate-spatulate, occasionally with shelf-like basal lateral extension, usually slightly deflexed (strongly
deflexed in some species).
Female. Generally similar to male; abdomen and wings stouter, and overall generally more setose
than male. Genitalia with well developed dorsomesal and ventrolateral lobes; a well developed apo-
deme lobe present, usually covered with fine microtrichia (reduced in some species, very well developed
in some species). Labia without microtrichia. Seminal capsules spherical to ovoid, with weak to mod-
erate neck, spermathecal ducts without loops or bends.
Pupa. Light green to green or brown in life; exuviae almost colorless to dark brown. Cephalic
tubercles present (essentially absent in D. flexus), poorly to well developed, broadly to narrowly
conical, each with a short preapical frontal seta. Dorsum of thorax weakly to strongly granulose
or pebbled, usually with a circular humeral callus lateroventral to base of thoracic horn. A
scutal tubercle sometimes present. Thoracic horn with more than 50 branches emanating from
2 main trunks. Base of thoracic horn usually somewhat dumbbell-shaped, tracheal bundles
usually separate (but joined in several species). Precorneal setae 2-3, dorsocentral setae 4,
rarely 5.
Abdominal segment I without lateral setae, segments II-IV with 3 lateral hairlike setae, V-
VII with 4 lateral lamellar setae, VIII with 4 or 5 lateral lamellar setae. Anal lobe with an
anterior pair of dorsal setae, easily lost; a pair of dorsal caudolateral lamellar setae; and 30+
JOHN H. EPLER 11
ventral lateral lamellar setae on each lobe, usually uniserial, at times partially biserial. An
uninterrupted row of caudal hooklets on T II. Intersegmental conjunctiva of T III/IV-T V/
VI usually with fine spinules. Sternites I-III with or without 1-2 transverse rows of spines.
Caudolateral corners of VIII with one to many, weakly to strongly developed, straight to
strongly sinuate spurs. Shagreen occasionally present on T I, present as fine longitudinal band(s)
or generally spread spinules on S I. Ventral shagreen on S II and III present as 2-4 longitudinal
bands connected by a transverse anterior band; weak to absent on S IV-VI (strongly developed
in 1 species). Dorsal shagreen on T II-VI subquadrate, hourglass-shaped, or triangular in
outline; often with small, separate elliptical to round anterolateral shagreen areas which may
merge with median shagreen area. Segment VII often with rounded anterolateral shagreen
areas, better developed dorsally. Dorsal shagreen of VIII usually U-shaped, or 2 longitudinal
bands, or a pair of anterior and caudal oval to round areas, or almost completely covering
dorsal area (D. flexus); ventral shagreen of VIII at most a weak copy of dorsal pattern, usually
more reduced. Small caudolateral spine groups present on T V-VII. Pedes spurii A well de-
veloped on S IV. Pedes spurii B present on II. Segments II-VIII with one dorsal and one
ventral pair O-setae. Segment I with 2-3 ventral, 2-4 dorsal pairs of setae; II with 3-4 ventral,
3-5 dorsal pairs; III with 3-4 ventral, 5 dorsal pairs; IV-VII with 4 ventral, 5 dorsal pairs;
VIII with 1 ventral and 1 dorsal pair of setae. T VIII usually with small posterolateral dorsal
lobes mediad of caudolateral spurs. In species with larvae possessing ventral tubules, weak
ventral tubules should be present caudolaterally on S VIII (I have not yet observed such tubules
on Dicrotendipes pupae).
Larva. Body pale green to green suffused with cream and/or red in life. Head capsule pale
yellow to red-brown, often with a dark middorsal stripe over frontal apotome; postmentum
often darker than remainder of head capsule. Mentum and mandibular teeth dark red-brown
to black. Three pairs of eyespots, the ventral 2 pairs often joined, giving the appearance of
only 2 pairs of eyespots. Dorsal eyespot largest, roughly oval to triangular in outline.
Antenna with 5 segments, segment 1 2-4X longer than second, segment 4 greater or ap-
proximately equal to 3. Antennal blade arises from apex of segment 1, reaching to 4th or 5th
segment. Lauterborn organs and well developed style present at apex of segment 2.
Frontal apotome usually concave and roughly tuberculate along frontal suture, sometimes
fused with 1st labral sclerite (forming a frontoclypeal apotome); usually with a small to large
ventral anteromedian frontal pit and/or frontal process; or a larger ventral suboval to subquad-
rate area. Labrum with setae I-IVA + B present; S I moderately plumose; S II large, unfringed;
S III hairlike; S IVA small. 2-segmented, S IVB subequal to S IVA, simple. Laterad to S II
is a group of 3 fringed labral chaetae, subequal to S II, and 3-4 smaller chaetae. Labral lamella
with fringe of 20-75 teeth. Pecten epipharyngis usually with 3-9, rarely 10-13, usually rounded,
ventral lobes. 5-8 chaetulae laterales. Premandible distally bifid, the inner blade subequal to
2-3X wider than outer blade; 1-3 inner medial teeth usually present; a medial premandibular
brush present.
Mandible with apical tooth, 1-2 dorsal preapical teeth, and 3 inner teeth, the proximal inner
tooth sometimes modified. Pecten mandibularis composed of 6-18 strong setae. Seta subden-
talis widest at middle, 4-7 times longer than wide; sometimes with accessory tooth. Seta interna
with 4 main branches, united basally.
Mentum with 11-13 teeth, the median tooth and Ist laterals usually subequal, median tooth
often notched mesolaterally, 2nd lateral tooth often fused or partially fused with 1st lateral;
5th and 6th lateral teeth sometimes rounded and fused. Ventromental plates 1.4X-2.2X wider
than long, deeply striate, with smooth or crenulated anterior margin. Setae submenti usually
simple, sometimes distally divided in many species.
Triangulum occipitale very narrow, scarcely visible in ventral view.
MEM. AMER. ENT. SOC., 36
12 GENUS DICROTENDIPES
Ventral tubules usually absent, occasionally 1 pair present on 8th abdominal segment. Pro-
cercus wider than long or as wide as long, with 6-8 apical setae; 2 anterolateral preapical setae
located on lightly sclerotized preapical plate. A pair of well developed supraanal setae present.
Two pairs of anal tubules, usually somewhat conical, sometimes rounded and reduced (in
brackish water species); ventral pair usually larger than dorsal pair.
NOTES ON THE HOLARCTIC DICROTENDIPES
Epler (1987a) revised the Nearctic Dicrotendipes. The western Palaearctic
species have recently been revised (Contreras-Lichtenberg 1986). A consid-
erable amount of Palaearctic material, mostly of species with western
Holarctic distributions, was available and is included in this study. In ad-
dition, eastern Palaearctic material of D. pelochloris (see Chap. IV) and D.
septemmaculatus (see Chap. II) was available. Two additional species have
recently been described from the eastern Palaearctic: D. inouei Hashimoto,
1984 and D. tamaviridis Sasa, 1981. Because the descriptions of these species
are incomplete and no material was made available to me, D. inouei and
D. tamaviridis are not included in this study.
Epler (1987a:51) noted the similarities between the immature stages of D.
incurvus (Sublette, 1964) and D. tritomus (Kieffer, 1916), but had insuffi-
cient comparative material to make a more definite statement. Observing
the illustration of D. tritomus (as Limnochironomus) in Pinder (1978:Fig.
158B), I erroneously stated that the apex of the superior volsella of D tri-
tomus was turned out, not in as in D. incurvus. Recent examination of
additional reared Palaearctic material of D. tritomus (through the kindness
of Drs. P.H. Langton and F. Reiss) and Contreras-Lichtenberg’s (1986:Fig.
14) recent redescription and illustration of this species revealed that the apex
of the superior volsella of D. tritomus is directed mesad. The adult was
originally described (Thienemann & Kieffer 1916) as having a trifid apex on
the inferior volsella, or at least 3 apicodorsal rows of sensilla chaetica (Con-
treras-Lichtenberg 1986). Contreras-Lichtenberg (1986:671) used the num-
ber of dorsal rows of sensilla chaetica and the bifid or trifid nature of the
apex of the inferior volsella as a secondary character in her key to separate
D. tritomus from D. modestus (Say). However, many specimens, particu-
larly from the Nearctic (D. incurvus), possess only a bifid apex, or 2 rows
of apicodorsal sensilla chaetica (although I have also seen many Nearctic
specimens with 3 dorsal rows), and Goetghebuer (1937-1954:19) stated in
his key that the apex could be bifid or trifid. I have examined many spec-
imens of D. modestus which have 3 or more rows of apicodorsal sensilla
chaetica on the inferior volsella (see Epler 1987a:71; Fig. 92).
The immature stages of D. incurvus and D. tritomus are also inseparable
to me and I conclude that the 2 species are synonymous. The holotype of
JOHN H. EPLER 13
D. tritomus, as many of Kieffer’s species, is apparently lost; I have not
located one and Contreras-Lichtenberg did not examine it. I am using D.
tritomus here in the sense of Lenz (1954-1962), Langton (1984), Pinder (1978)
and Contreras-Lichtenberg (1986). Thus, D. tritomus is the fourth species
of Dicrotendipes with a Holarctic distribution (see Chap. V).
The character, the presence of a second dorsal mandibular tooth, that
Contreras-Lichtenberg (1986:670) uses for separating the larvae of D. mo-
destus and D. tritomus is not reliable. I have seen a specimen of D. tritomus
from Great Britain without a second tooth, and have seen many specimens
of D. modestus with a second dorsal tooth or at least a definite large notch
in the dorsal tooth. The second dorsal tooth is often difficult to observe
unless the mandible is positioned correctly. The larvae of these 2 species are
difficult to separate; usually the count of the ventromental plate strial ridges
and differences in postmental coloration will separate them (see key below),
but specimens must be reared to confirm species identity. Likewise, Con-
treras-Lichtenberg’s (1986) pupal key will fail to separate these 2 species.
The pupa of D. peringueyanus as described by Contreras-Lichtenberg
(1986) is not separable from D. fusconotatus or D. pallidicornis (see also
Chap. II). In her pupal key, Contreras-Lichtenberg (1986) separates D. per-
ingueyanus from D. pallidicornis based on the absence of conjunctival
spinules on D. pallidicornis. However, all specimens of D. pallidicornis which
I have examined do possess these spinules.
The following keys are offered for the identification of all known life
stages (excepting the adult females) of the Holarctic Dicrotendipes. The
reader is advised to consult the illustrations, descriptions and diagnoses in
Contreras-Lichtenberg (1986) and Epler (1987a).
KEY TO ADULT MALES OF HOLARCTIC DICROTENDIPES
(D. inouei, D. tamaviridis not included)
1. Inferior volsella deeply bifid apically; Palaearctic species ................ 2
Inferior volsella with simple apex, at most notched, appearing shallowly bifid or trifid;
EVOAaTCiHCRSPECleSe tn cae eee ee oe Sty Sees Ge, Garey ne tke on asa 5
2. _ Wings with spots or bands, or clouds along veins .................... 3
Wiingssimmadculates) 44. 2.6 14660 0e os eae D. pallidicornis (Goetghebuer)
3. Small, membranous, triangular flap-like appendages present near base of anal point
J coast S uO cee aga c Seite ee RE Ier Bee eer eee areca D. fusconotatus (Kieffer)
Base of anal point without appendages................++.-++--e00. 4
4. Wing with 6-7 well-defined spots, with 1 spot usually present in cell m3,4.......
Sep IRR uM REO Gok So vin Toki we ae cae Oe D. septemmaculatus (Becker)
Wing with weakly defined spots, none present in cell m3,4 .D. peringueyanus Kieffer
5. Inferior volsella with membranous dorsal extension; Palaearctic .D. notatus (Meigen)
Inferior volsella without membranous dorsal extension; Holarctic........... 6
MEM. AMER. ENT. SOC., 36
14
10.
11.
12.
13.
14.
15.
16.
17.
18.
19.
20.
GENUS DICROTENDIPES
Anal point sharply reflexed ventrad, usually not visible in dorsal view (Fig. 36); eastern
Paldéarctic:s% 0a te Se a eee D. pelochloris (Kieffer)
Anal point not sharply reflexed ventrad, visible in dorsal view; Holarctic....... 7
Superior volsella strongly pediform, apex directed outward; or triangular....... 8
Superior volsella digitiform, long and slender, or long with weakly expanded membra-
nous apex; if somewhat pediform, then apex directed inward .......... 14
Superior volsella triangular, or if weakly pediform, sensilla chaetica restricted to pos-
terior margin of superior volsella; coastal, brackish water species of SE U.S.A.. .
ee rd Cnc Cte an hoster eco okale cn. a DhosG D. lobus (Beck)
Superior volsella pediform; sensilla chaetica not restricted to posterior margin of
VOlSell as, se: cat se Se Pat ec Elesh ROS as te ee as GLEE CE se 9
Legs strongly banded; Nearactic species... . . D. californicus complex .... . 10
Legs not banded, at most distal portions of some leg segments darker; Holarctic
SPECIES 55.0%. Ad: SOR AS Red cee remetleatencs Soro nce TESA: GutcRE TOM SRR EMI mean ieee 11
AR 2.13-2.36, mean 2.29; SV, 4.15-4.41, mean 4.29; SV; 2.71-2.89, mean 2.80; known
from central and eastern New Mexico and Imperial Dam vicinity, California (pos-
Sibly Kansas)) 3s... cca x. 6 Saas Se ee ee eee D. crypticus Epler
AR 2.29-2.69, mean 2.47, SV> 3.81-4.42, mean 4.07; SV; 2.61-2.84, mean 2.71; wide-
spread in western U.S. and Mexico........... D. californicus (Johannsen)
Anal point with raised truncate base; known only from Chiricahua Mountains of SE
ATIZONA) he AS oS oe ees «oe eee D. adnilus Epler
Anal point not as above; widespread. ..............2..+.+++2202- 12
Dorsum of tergite [IX with many long setae laterad of anal point; SE U.S.A. .....
Re are re eet oc NGen te ere ei eer el Ricks Geototsroson D. thanatogratus Epler
DorsumvotatergitesPxenotrasia Ove enna nen cn ne nen nnn nen nner 13
Gonostylus inflated medially, narrowed proximally and preapically; general coloration
browns Neéarctich. cc nee eH wet £ Shae, ee a egos D. neomodestus (Malloch)
Gonostylus not as above; general coloration green to red-brown; Holarctic.......
ee ere ee ten cle: foucka: ceo eet cc! oS uloromont! GAB 6 D. modestus (Say)
Superior volsella short, digitiform .....................2-+-++---: 15
Superior volsella long and slender or long with weakly expanded membranous apex
PN ECE Poe oS atte c rel cf ord eon ase ta oh oED Si oto lured ole Go aS" S-0 oO 18
Anal point with wide shelf-like base, tapering gradually to apex; superior volsella cy-
lindrical, with slightly out-turned apex; Nearctic..... D. fumidus (Johannsen)
Anal point spatulate or narrowed at base .....................-.-. 16
Superior volsella with thin membranous preapical extension; metatarsi usually with wide
basal white band (lacking in most Florida specimens); Nearctic ...........
OR Tar crceoronc oretc a once arb boo D. leucoscelis (Townes)
Superior volsella not as above; metatarsi without wide basal white band ...... 17
Anal point long, narrowly spatulate; superior volsella with sclerotized area at apex;
Holarctichcc. 2c. Ae cs Foe ees ee ee a ye D. lobiger (Kieffer)
Anal point and superior volsella not as above...............-.-.--+-- 18
Superior volsella long, slender, recurved, with an acute apex . . D. botaurus (Townes)
Superior volsella not as: above <5 o> Kecscicusiy os) Ciao: Cece ce) ee ce 19
Apex of superior volsella turned in. .................+e-e05+++e0s 20
Apex of superior volsella turned out; or straight, semiglobose ............ 21
Gonostylus long, thin, and strongly curved; superior volsella usually without microtri-
chia; apex at most semimembranous; Nearctic......... D. milleri (Townes)
Gonostylus not as above; at least basal half of superior volsella with microtrichia, apex
NASM ONEMOUGS IBIOEMEANG 5560005 0c cb ooo oD Hb OOe D. tritomus (Kieffer)
21.
DD
10.
JOHN H. EPLER 15
Superior volsella cylindrical, curving outward; apex bare, not expanded (Fig. 13); thorax
with well developed scutal tubercle; SW U.S.A., Mexico . . D. aethiops (Townes)
Superior volsella not as above, apex expanded or inflated; thorax with or without scutal
tubercles widespread’ :.ees iar. Riese te | NS eels Seneca oc, ahs 22
Sensilla chaetica of superior volsella in a line often reaching middle of appendage tip,
not directed exclusively inward; length of superior volsella 2-3.25X maximum width;
wing with more than 35 setae on R & Rj; Holaretic.... . D. nervosus (Staeger)
Sensilla chaetica of superior volsella distributed on inner surface of appendage, the
majority directed inward; length of superior volsella 4.25-5.25X maximum width;
wing with 35 or fewer setae on R & Rj; Nearctic....................
. . . . D. lucifer complex (D. lucifer (Johannsen), D. simpsoni Epler). (D. inouei
Hashimoto & D. tamaviridis Sasa, both eastern Palaearctic species, may key out
here.)
KEY TO KNOWN PUPAE OF HOLARCTIC DICROTENDIPES
Ventral spine row(s) presenton SII ..................0002200005 8
WVentrallspine:row(S)iabsent. (3 4 20) 4. cis eh Ge os ee ee He eR el Sree 2
4ilateralilamellansetaeon’h VII. 2.2 he ae ee ee eee ee 3
Srlateraltlamellarisetaeronly VAIN, ee FS eo SI a eS I hs 5
Shagreen spinules on T II-V largest posteriorly, tips often rounded; exuviae light yellow-
brown to dark yellow-brown; Nearctic................... D. fumidus
Shagreen spinules on T II-V more or less equal; exuviae clear or light to dark brown;
FVOLAT CLC AM tke yo eee ancl eiec eects DE me RSS SUL RMS Be ae 4
Median shagreen spinules more or less equal on T VI; exuviae light to dark brown;
strongly reticulate cuticular pattern on T VI-VIII, especially on T VI; coastal, brack-
ishawaterspeciesvOfuSE WES AN 2 ais cn es ee ee le ee D. lobus
Median shagreen area on T VI with longest spinules in middle of area; exuviae clear
with yellowish borders; reticulate cuticular pattern on T VI-VIII at most moderately
developed: Holarcticss is wie ew ale Se oe ee RR ee es D. nervosus group
(D. lucifer, D. nervosus, D. simpsoni). (D. tamaviridis may key here.)
Anal lobe with dorsal shagreen; Palaearctic......... D. septemmaculatus group
(D. fusconotatus, D. pallidicornis, D. peringueyanus, D. septemmaculatus)
Anal lobe without dorsal shagreen ..............-----2-2-2-+-+----- 6
Intersegmental spines present between T V and T VI; Nearctic and western Palaearctic
SDECIESHSE eaten ete ek eoniie Go Se Uae oe CORO ee ee ete 7]
T V and T VI without intersegmental spines (Fig. 41J); eastern Palaearctic.......
iat. Sr tancrtri et een: bee eS Gea iain eee GE eT ee D. pelochloris
INCANCliCYSPECIES! ie Gk os Re ee ee new Se me D. leucoscelis
Ralacarcticispeciesiytes Eis AAT ee ee es ge I D. notatus
Silateralisetacionle WIM). .5. sion ee Ste we ee Se See D. lobiger
Aulateraltsetacions DLs VLIIMy Sih is ee Be SR OE 33 9
Caudolateral spur on T VIII double or triple, spurs well separated ......... 10
Caudolateral spur on T VIII single or closely appressed double ........... 11
Cephalic tubercles long, thin, sharply acute, 33-40, mean 37, anal fin setae; New York,
Indiana, Michigan, Minnesota, Oregon, Ontario and Manitoba.... . D. milleri
Cephalic tubercles shorter, wider; 33-55, mean 45, anal fin setae; Holarctic .......
AT Su, Ways tas tas ras guste mae crcelie Taree Soe at tI ec ea emote ta Teme, Naive tare D. tritomus
MEM. AMER. ENT. SOC., 36
16
Ihe
12.
13.
14.
GENUS DICROTENDIPES
Median shagreen area on T VI with larger anterior and/or posterior spinules; wide-
Spread... 2... - eects oR a BS ses ee eee ee ee 72
Median shagreen spinules on T VI largest anteriorly only or those of posterior portion
only slightly larger than middle; west and southwest U.S.A..............
Ais ee eee ae D. californicus complex (D. californicus, D. crypticus)
Dorsum of anal disc with thinly spined anteromesal shagreen areas; known only from
Floridas..55) 2. coe ee Pa A ee eee D. thanatogratus
Dorsum of anal disc without shagreen.................22-2-2-.-00- 13
Anterior and posterior spinules of median shagreen area on T VI more strongly devel-
oped than those of center; caudolateral spur on T VIII usually strongly recurved or
SINUS serps Macias 2,5. 24 tes SRA ae eee Se ee ee 14
Posterior spinules of median shagreen area on T VI larger, more rounded or longer than
those of anterior portion of shagreen area; caudolateral spur on T VIII not strongly
Sinuate Ole Cunvedssal Most siiall tases nee nena nnn nee D. tritomus
Anal lobe with 31-50, mean 38, lateral setae; Nearctic......... D. neomodestus
Anal lobe with 30-64, mean 50, lateral setae (1 specimen seen from Florida with only
30-33 anal lobe setae); Holarctic..................... D. modestus
KEY TO KNOWN 4TH INSTAR LARVAE OF HOLARCTIC DICROTENDIPES
Frontal apotome with a large anteromesal oval or subquadrate area; frontal pit or pro-
CeSsvabsent:.ccsiers eed ed Bae ee A Ve ee eee ee eee 2,
Frontal apotome without such an area; a small anteromedian frontal pit or frontal
processiusually:preséntine Sine He Ree ae ee PR ee eee 5
A frontoclypeal apotome present; sixth lateral tooth of mentum well developed; 30-41,
mean 35, ventromental strial ridges ..................... D. lobiger
A frontal apotome present; sixth lateral tooth of mentum reduced or closely appressed
1d | ee henna in een ORS sir ob Sir 4 co. Gha-6, ols, o ores On8 3
Palaearctic specieSin..iaiets, wkere.e) Ae Ss As FR. 2 ie eR SE ee Clore oe 4
Nearctic species; 37-60, mean 52 ventromental strial ridges. ....... D. leucoscelis
Eastern Palaearctic species; 30-34, mean 32 ventromental strial ridges (Fig. 41N) . . .
ee ee Ee ies oh oe RS Te eae oe RY Showa ateee Bre € D. pelochloris
Western Palaearctic species; about 40 ventromental strial ridges ...... D. notatus
Second lateral tooth of mentum almost completely fused with or closely appressed to
Ist so that Ist lateral tooth appears notched (Fig. 13)................ 6
Second lateral tooth at most only partially fused to Ist lateral at base, Ist lateral tooth
not appearinp notched”.s . see oo ais Swe Oe «SENS ROS Seen 9
Head capsule integument appears coarsely granular at 400X; Nearctic species . . . 16
Head capsule integument at most appears slightly granular at 400X; Nearctic or Pa-
laGarcticSPeCleS ss. spe wie ee cre a RUS AEE: eeu at ROO re tear a,
A long, ventral frontal process present on anteromedian margin of frontal apotome;
Palaearctic.ionly. 2 = sure 2 es eee ee D. septemmaculatus
Long frontal process absent, but a frontal pit usually present; Nearctic only. .... 8
Anal tubules reduced; anterior margin of ventromental plate mostly smooth; coastal,
brackishtwatemspecies/ of SEW: SAG ie eieiea- ie aie neni nena D. lobus
Anal tubules normal; at least anterior outer margin of ventromental plate usually cren-
ulated= widespreadsins Nearcticus) iene Cena nner D. neomodestus
60-70 ventromental strial ridges; Palaearctic only............ D. pallidicornis
Less than 50 ventromental strial ridges; Nearctic or Palaearctic............ 10
10.
11.
12.
13.
14.
15.
16.
17.
18.
19.
20.
JOHN H. EPLER 17
Ventromental plate with 22 or fewer strial ridges; anterior margin deeply scalloped . .
eee tee ees Coat, sia © ve. ahem enn wb oh. ech G6) atin eee RU ROMI Ua cele, tak cyt nes ets 11
Ventromental plate with more than 22 strial ridges; anterior margin smooth or with
shallow tojmoderatecrenulations= 52555 4---.-2-e400005¢ 4400+" 12
Nearctic species (known only from Florida)............... D. thanatogratus
alacanchiGysPecleSmp:, esis ys cuts te. Seven ste see Ge Gees ne ure CMTE D. fusconotatus
Proximal tooth of mandible saddle-shaped or with ? points; or with inner surface of
mandible adjacent to proximal tooth with deep semicircular incision; Nearctic. 13
Proximal tooth of mandible mostly triangular in outline, not as above; Holarctic . 14
Sixth lateral tooth of mentum rounded and closely appressed or fused to Sth lateral
tooth; inner surface of mandible adjacent to proximal tooth with deep semicircular
LEAGISI OM GWE gai Pe eee Pees ee cu re. suk Sa at Bue Boos eae et eet cele D. simpsoni
Sixth lateral tooth of mentum pointed; inner surface of mandible adjacent to proximal
tooth without deep semicircular incision. .................. D. lucifer
Sixth lateral tooth of mentum rounded and closely appressed or fused to Sth lateral
TOOL eee peace crates Nae eee nis Ge eo er eth cae wee oe re D. nervosus
Sixthglateralitoothypomtedsseparatere seis - oe cncnneen carieencnenencncnencne 15
Head capsule integument appears coarsely granular at 400X; head capsule color yellow-
brown to yellow-red-brown; Nearctic species.................... 16
Head capsule integument at most appears slightly granular at 400X; head capsule color
light brown to brown or pale yellow; Holarctic species .............. 18
Postmentum or posterior portion of head capsule usually not darkened; Ist lateral teeth
of mentum often turned outward; occurs throughout the Nearctic. . . D. fumidus
Postmentum or posterior portion of head capsule much darker than rest of head capsule;
1st lateral teeth of mentum rarely turned outward; western U.S.A. ....... 17
Ventromental plate with crenulated anterior margin; 23-28, mean 25, ventromental strial
ridges; known from central and eastern New Mexico and Imperial Dam vicinity,
Ealifornial(possibly,Kansas)s ns 3 2 4 so oe eee D. crypticus
Ventromental plate with smooth anterior margin; 34-42, mean 37, ventromental strial
ridges; widespread in western U.S.A. ...............-.-. D. californicus
Meadkcapsuleipaleyellows*) 4 aa 25 2 ase Seeks cee eee ee ate ee 19
lnlgag! Gaps Welt lOKONAN TO IOOWN, 5 oo oo poop DoD odo o ooo Oooo OOO 20
Ventromental strial ridges 28-36, mean 32; postmentum usually darkened .......
Mire eo isan re See cries Sree 12s ey GOR EM epics CG rad fate es D. modestus
Ventromental strial ridges 23-29, mean 25; postmentum rarely darkened . D. tritomus
Anterior margin of ventromental plates mostly smooth; known only from Chiricahua
Mountainsrof-SEvArizonale) aes cies os cea er et ees eee D. adnilus
Anterior margin of ventromental plates with shallow to moderate crenulations; wide-
SpreadbinulTolanctich yay ct sk eee a cee ete eaest ce) co ou Lenco tommy D. modestus
MEM. AMER. ENT. SOC., 36
18 GENUS DICROTENDIPES
Fic. 1. Adult morphology. A) Structures of head, frontal view. B) An-
tennal flagellum and pedicel; AR measurement method. C) Cibarial pump
and associated structures. D) Method of abdominal length measurement.
E) Thoracic structures. F) Ventral accessory setae of (left to right): D. neo-
modestus, D. leucoscelis, D. cordatus. (A, abdominal length; Ac, acrosti-
chal setae; Cls, clypeal setae; CP, cibarial pump; CS, cibarial setae; Dc,
dorsocentral setae; HP, humeral pit; LL, labial lonchus; Pa, prealar setae;
Ped, pedicel; Pm,_,, maxillary palpomeres 1-5; PPS, postpronotal suture;
PS, parapsidal suture; PSS, prescutoscutal suture; Scp, scape; Sct, scutellar
setae; Scut, scutellar tubercle; Sp, spiracle; Su, supraalar seta; T, thorax
length; Te, tentorium; Tem, temporal setae; THS, thoracic scar; To, torma.)
MEM. AMER. ENT. SOC., 36
JOHN H. EPLER
20 GENUS DICROTENDIPES
Fic. 2. Adult morphology. A) Method of measuring all femora & the
fore tibia. B) Method of measuring fore metatarsus. C) Method of mea-
suring tibiae & tarsomeres of mid & hind legs. D) Wing (see Saether 1980
for abbreviations). E) Methods of wing measurement. (WL, wing length;
WW, wing width.)
JOHN H. EPLER
a
eal ee
earnest
tars
Bo Ar C Se RM Ry Rog
\ . - R45
One SSS> ——— SS = a
21
MEM. AMER. ENT. SOC., 36
22 GENUS DICROTENDIPES
Fic. 3. Adult male morphology. A) Lateral view of hypopygium (nearest
gonostylus, inferior & superior volsellae removed). B) Dorsal view of right
side of hypopygium. C) Ventral view of right side of hypopygium, showing
internal apodemes. D) Ventral view, superior volsella. (AnP, anal point;
ATB, anal tergal band; Ca, coxapodeme; DBS, dorsal basal setae; Gc, gon-
ocoxite; Gs, gonostylus; I Vo, inferior volsella; L, length of superior volsella;
LBS, lateral basal setae; MS, median setae; Pha, phallapodeme; SCh, sen-
silla chaetica; SVo, superior volsella; TSa, transverse sternapodeme; VAS,
ventral apical seta; W, width of superior volsella.)
MEM. AMER. ENT. SOC., 36
JOHN H. EPLER
23
24 GENUS DICROTENDIPES
Fic. 4. Adult female morphology: A) Ventral view of terminalia. B) DmL
and VIL. C) ApL. (ApL, apodeme lobe; Ce, cercus; Csa, coxosternapo-
deme; DmL, dorsomesal lobe; Ge IX, gonocoxite IX; SCa, seminal capsule;
S VIII, sternite VIII; VIL, ventrolateral lobe; X, segment X.)
JOHN H. EPLER 25
el 86 3 TS 2 ge ts
x} Kee en eg Rife
nM ih bh S 2b ia Degas tape y at
Te eg ~ a fess wd
UY NN Nice eee arate Re
di AYA Ai x \ Y ¢ /
|, se Yip eee ENG We te -. Ace
h Ay /; S - = BR) er
4 7 7 y} WF, Wy 7. 1 é -. Be
\y Win Cy) Y LL \ v =, PS
\ BS i tik a
py > Dy eG
} ON. eK
= o
1
Q ee
a3 4
ie
Ap ES
AE ey
ht 7:
MEM. AMER. ENT. SOC., 36
26 GENUS DICROTENDIPES
Fic. 5. Pupal morphology. A) Ventral view of spread cephalothorax. B)
Lateral view of cephalothorax. C) Dorsal view of anal lobe, depicting method
of calculating disc ratio, DR. DR is the total number of ventral anal lobe
setae (V) divided by the number of ventral setae anterior to and including
dorsal seta position (D); DR = V/D. D) Dorsal view abdomen. E) Ventral
view of abdomen. (A, abdomen length; ADS, anterodorsal seta; AS, an-
terolateral shagreen area; C, cephalothoracic length; CS, caudolateral spur;
CT, cephalic tubercle; D, number of ventral setae from dorsal seta position
to anterior margin of anal lobe; Dc, dorsocentral setae; DS, dorsal seta; H,
hooklets; HC, humeral callus; LLS, lateral lamellar setae; MS, median sha-
green area; Pc, precorneal setae; PSA, pedes spurii A; PSB, pedes spurii
B; ScuT, scutal tubercle; THB, thoracic horn base; V, total number of ven-
tral anal lobe setae; VS, ventral setae of anal lobe; VSR, ventral spine rows.)
JOHN H. EPLER
Ween,
ven
fl
\
\ \ 4
“CHW VAN
MEM. AMER. ENT. SOC., 36
27
28 GENUS DICROTENDIPES
Fic. 6. Larval morphology: A) Ventral view of head capsule. B) Dorsal
view of head capsule. C) Mentum. D) Ventromental plate. E) Premandible.
F) Mandible. G) Antenna.(ABI, accessory blade; Bl, antennal blade; FA,
frontal apotome; L, length; LO, lauterborn organ; M, mentum width; Man,
mandible; Max, maxilla; MT, width of median teeth; PM, postmentum
length; PMa, pecten mandibularis; Pm, premandible; PmB, premandibular
brush; Po, postocciput; RO, ring organ; S, style; Si, seta interna; SI 1,2,
labral sclerites 1,2; SR, strial ridges; SSd, seta subdentalis; TO, triangulum
occipitale; Vm, ventromental plate; W, width.)
JOHN H. EPLER 29
30 GENUS DICROTENDIPES
Fic. 7. Larval morphology: A) Palatal surface of labrum. B) Ventral view
of maxilla. C) Dorsal view of maxilla. D) Anterior portion of frontal apo-
tome and related labral sclerites 1-5 of D. fusconotatus. (Ch, chaetae; ChL,
chaetulae laterales; ChP, chaetulae of palpiger; FA, frontal apotome; FP,
frontal process; G, galea; La, lacinia; LL, labral lamella; MP, maxillary
palp; P, palpiger; PE, pecten epipharyngis; SM, >, setae maxillaris; SI-SIVA,
SIVB, labral setae; 1-5, labral sclerites 1-5.)
JOHN H. EPLER
31
m
SSS Peal -
—— ra
a
= |
y
=
=
C QS
») Zz
Z
\
MEM. AMER. ENT. SOC., 36
a2 GENUS DICROTENDIPES
CHAPTER II.
A REVIEW OF THE AFROTROPICAL DICROTENDIPES WITH
DESCRIPTIONS OF THE IMMATURE STAGES
This chapter presents a review of the 14 species of Afrotropical Dicro-
tendipes and describes or redescribes the immature stages of 5 species (only
the pupal stage is described for D. cordatus).
The genus Dicrotendipes was originally described from the Afrotropical
region (Kieffer 1913b). Freeman (1957) provided a first revision of the Af-
rotropical species; he included 19 species in Chironomus (Dicrotendipes), 6
of which he described as new. Freeman & Cranston (1980) provided an up-
dated checklist. These works concentrated only on the adult stages. The
immature stages had never been fully described until, in a recent revision
of the western Palaearctic Dicrotendipes, Contreras-Lichtenberg (1986) de-
scribed the immature stages of some species which are also found in the
Afrotropical region; D. fusconotatus (larva and pupa), D. peringueyanus
(pupa) and D. septemmaculatus (larva and pupa, as OD.
quatuordecimpunctatus).
A key to identify adults was provided by Freeman (1957), but one should
be aware that Freeman’s key included several species no longer considered
to be members of Dicrotendipes. A new key to the adult males of Afro-
tropical Dicrotendipes is provided below. The species chloronotus Kieffer
is now considered a Kiefferulus (Freeman and Cranston 1980). Freeman
(1957) and Freeman and Cranston (1980) erroneously synonymized Carter-
onica Strand (= Carteria Kieffer) with Dicrotendipes (cf. also Ashe 1983).
As noted by Epler (1987a), these genera are not synonymous. The Afro-
tropical species crispi Freeman, regalis Goetghebuer, multispinosus Free-
man and penicillatus Freeman, formerly included in Dicrotendipes, some-
what resemble Carteronica in the adult stage (the species regalis was
originally described as a Carteria). Although these species resemble Car-
teronica, based upon examination of the immature stages of reared mul-
tispinosus and immature Chironomus longilobus (Kieffer) (type-species for
Carteronica), these Afrotropical species must be placed in a new genus.
At least one other described Afrotropical species, Xenochironomus tri-
setosus (Kieffer, 1922) may be a Dicrotendipes (see description in Freeman
1957:381). It possesses an inferior volsella similar to that of D. bredoi. Until
reared immature stages are available, I believe it best to retain frisetosus in
Xenochironomus.
The following key will replace that found in Freeman (1957:359-360).
Descriptions of adults and illustrations of wings and male terminalia can
be found in Freeman (1957).
JOHN H. EPLER 33
KEY TO ADULT MALES OF AFROTROPICAL DICROTENDIPES
1. Wings with spots or bands, or clouds along veins .................6.. 2
WARNES HTT ETSH CMOS eel mel ce PRT Bel aWNBL ee Snes oop o alate: co laied dicot EU Pe eter 8
2. Inferior volsella simple, at most apex deeply notched .................. 3
Inferior volsella with apex deeply bifid..................2.-200085 5
3. Wing with dark distal band covering or touching RM.................. 4
Wing with dark distal band not touching RM........... D. leucolabis Kieffer
4. Superior volsella preapically expanded, appearing somewhat globose
PE TE aig orc aa gece rss ater 126 no okay Sec Se aa See ce D. cordatus Kieffer
Superior volsella not preapically expanded........... D. collarti (Goetghebuer)
5. Wing pattern weak, consisting of clouds along major veins . .D. sudanicus (Freeman)
Wing pattern consisting of spots (which may be weak in | species) .......... 6
6. Small, membranous, triangular flap-like appendages present laterad of anal point. . .
5 tans Gg CoP OLA OLE ORTOP ELE OPEL eS ae ee D. fusconotatus (Kieffer)
Membranous appendages near anal point absent. .................... fl
7. Wing with 6-7 well defined spots, with 1 spot usually present in cell m3, 4
Sado ON Sat Cha tavick OLA cack te Deemer cs D. septemmaculatus (Becker)
Wing with weakly defined spots, none present in m3,4 . . . D. peringueyanus Kieffer
8. Base of anal point with lateral projections; apex of inferior volsella narrow, not
expanded tk SRN coe Ski aes 9 Ee ae D. bredoi (Goetghebuer)
Base of anal point without lateral projections; apex of inferior volsella expanded . . 9
9. Anal point broad and strongly reflexed ventrad..................... 10
AN AlapointenOtrassa DOVE. oy cc, 5 hat oe Sulasast Ger for St sce Sone sa ahead Gita Sold, A cop aged wae eee are 6 11
10. Superior volsella somewhat cylindrical with globose to pediform apex; apex of superior
volsella with mediad membranous area bearing sensilla chaetica; foretarsi
WithoutibeardWigss anes Siciete chest taaeaers 2.4 D. kribiicola (Kieffer)
Superior volsella not cylindrical with globose apex, but rather leaf shaped with attenuate
apex; apex of superior volsella without membranous area bearing sensilla
chaeticas| foretansi bearded’) 7-7-5. 2252550 -5-6 D. ealae (Freeman)
11. Superior volsella short, globose, with medially directed apical point
MSE SUM Rete ta ae oan an et ae re D. nigrolineatus (Freeman)
Superior volsella somewhat cylindrical or with elongate pedicel............ 12
12. Inferior volsella with large swollen apex; superior volsella with small mesally directed
apicalipoints oy.0s ses ae Seeley ede D. schoutedeni (Goetghebuer)
Inferior volsella with normal, moderately expanded apex; superior volsella without small
mesalivadinectedsaplcalgnoltememeiememeE en cnenenen Carne nine ran 13
13. Anal point long and thin, with 2-4 dorsal basal setae . D. freemani Epler, nom. nov.
Anal point shorter, wider, with more than 6 dorsal basal setae
Li Mee ears Lie cetera Bins AL accra SoM cerca Reig te D. chambiensis (Goetghebuer)
Dicrotendipes bredoi (Goetghebuer)
Chironomus Bredoi Goetghebuer, 1936:473.
Chironomus (Dicrotendipes) bredoi Goetghebuer: Freeman 1957:369.
Dicrotendipes bredoi (Goetghebuer): Freeman and Cranston 1980:190; Hare & Carter 1987:70.
See adult description in Goetghebuer (1936:473) and Freeman (1957:369);
the immature stages are unknown.
MEM. AMER. ENT. SOC., 36
34 GENUS DICROTENDIPES
This is quite an unusual species for a Dicrotendipes, and may not belong
here but in another, possibly new, genus. The superior and inferior volsellae
are not typical for Dicrotendipes. Examination of the immature stages is
needed before this species can be placed with certainty. There are no ventral
accessory setae apparent on S VI. The specimen from Nigeria represents the
first record for this species from that country.
MATERIAL EXAMINED: NIGERIA: Lake Opi nr Nsukka, light trap, 29-30-XII-1979,
leg. L. Hare, 1 male (LH). [ZAIRE]: ‘Belgian Congo,”’ Eala, X-1929, H.J. Bredo, 1 male
(holotype) (KM).
Dicrotendipes chambiensis (Goetghebuer) (Fig. 8)
Chironomus (Limnochironomus) chambiensis Goetghebuer, 1936:464.
Chihronomus (Dicrotendipes) chambiensis Goetghebuer: Freeman 1957:368.
Dicrotendipes chambiensis (Goetghebuer): Freeman and Cranston 1980:190.
See adult description in Goetghebuer (1936:464) and Freeman (1957:368);
the immature stages are unknown.
The superior volsella of the holotype of D. chambiensis (Fig. 8C) is some-
what stouter than that of D. freemani (Fig. 8A), and is apparently distorted
from preparation procedures. Specimens from Zaire and South Africa in
the BM determined as chambiensis have superior volsellae (Fig. 8D) which
are more similar to the Nearctic D. /ucifer complex (Epler 1987a: Figs. 111,
112), and possess numerous dorsal basal or dorsomesal setae on T IX. D.
freemani specimens have 2-4 T IX dorsal basal setae and superior volsellae
more similar to those of D. nervosus (Epler 1987a: Figs. 105-109). I could
not discern any dorsal setae on T IX of the D. chambiensis holotype spec-
imen. The Zaire and South Africa specimens might represent a new species,
but without reared associations I cannot justify “‘splitting’’ the species. The
differences between the anal points and inferior volsellae of D. freemani
(as binotatus) and D. chambiensis cited by Freeman (1957:368) and utilized
in the key above could be attributed to individual variation, as can the dif-
ferences in the superior volsellae. It should also be noted that superior vol-
sellae with menbranous apices, such as those possessed by D. freemani and
D. chambiensis, are very sensitive to pressure induced variations caused by
mounting procedures.
The hypopygium of the D. chambiensis holotype was mounted between
2 pieces of celluloid on the pin with the rest of the specimen and was in
very poor condition. The mount had split open, the mounting medium had
dried and broken the hypopygium. I remounted the hypopygium in balsam
on a microscope slide and was able to salvage the gonocoxae, one superior
JOHN H. EPLER 35
volsella and T IX with the anal point; the inferior volsellae and the gonostyli
are lost.
MATERIAL EXAMINED: [SOUTH AFRICA): Transvaal: Blaaubank River, IV-1957, A.D.
Harrison & B.R. Allanson, 1 male (BM); Blaaubank River nr Sterkfontein, IV-1957, A.D.
Harrison & B.R. Allanson, 1 male (BM). [ZAIRE]: Elizabethville, 17, 24-XII-1938, H.J. Bredo,
2 males (BM); Escpm. Kabasha: Chambi, X-1933, Dr. De Wulf, 1 male (holotype) (KM).
Dicrotendipes collarti (Goetghebuer)
Chironomus (Dicrotendipes) collarti Goetghebuer, 1936:466; Freeman 1957:366.
Dicrotendipes collarti (Goetghebuer): Freeman and Cranston 1980: 190.
See Goetghebuer (1936:466) and Freeman (1957:366) for the adult de-
scription; the immature stages are unknown. I have seen one male with 9
heavy spine-like setae on S VI.
MATERIAL EXAMINED: KENYA: Aberdare Range, Chania Falls, 4,000 feet, B.M.E. Afr.
Exp., X-1934, F.W. Edwards, 1 male (BM). [ZAIRE]: ‘Belgian Congo’’: Ituri, Alokoko, 11-
II-1930, A. Collart, 1 male (holotype) (KM); Katanga, Kafubu Mission, IX-1931, Miss A.
Mackie, 1 male (BM).
Dicrotendipes cordatus Kieffer (Figs. 1, 9)
Dicrotendipes cordatus Kieffer, 1922:64; Freeman 1955a:22; Freeman and Cranston 1980:190;
Hare & Carter 1987:70.
Paratendipes pictus Goetghebuer, 1934:199.
Chironomus (Paratendipes) pictus (Goetghebuer): Freeman 1954a: 443.
Chironomus (Dicrotendipes) cordatus (Kieffer): Freeman 1957:365; Dejoux 1968:56.
See adult description in Kieffer (1922:64) and Freeman (1957:365). There
are 4-6 heavy spine-like ventral accessory setae on S VI (Fig. 1F). The holo-
type of D. cordatus is apparently lost (Freeman 1957:366).
This species is probably closely related to the Afrotropical D. kribiicola,
the Nearctic D. leucoscelis and the Palaearctic D. notatus.
PUPA: (n=3)
COLOR. Light brown, with darker areas along lateral margins of tergites.
LENGTH. Total 4.38-5.03, 4.64 mm. Cephalothorax 1.11-1.16, 1.13 mm. Abdomen 3.27-
3.92, 3.52 mm.
CEPHALOTHORAX. Cephalic tubercles well developed (Fig. 9A). Dorsum moderately to
well pebbled. Dc2 closer to Dc3. Thoracic horn base (Fig. 9B) with tracheal bundles narrowly
joined or fused medially.
ABDOMEN (Fig. 9C). Sternites I-III with fine lateral shagreen bands; S I with posterior
band; S II occasionally with anterior band. Tergite I without shagreen; T II with median quad-
MEM. AMER. ENT. SOC., 36
36 GENUS DICROTENDIPES
rilateral shagreen area; T IIJ-V with larger median quadrilateral shagreen area, areas extended
laterally on anterior portion; T VI with Y or T shaped shagreen area; T VII with anterior band
of fine shagreen and a pair of posterolateral ovoid shagreen areas; T VIII with an anterior
pair of ovoid fine shagreen areas and a smaller posterior pair of ovoid shagreen areas; shagreen
areas on T II-VI with spines larger in middle of area. Tergites III-V with posterior conjunctival
band of fine spinules; T V posteriorly with 2 groups of intersegmental spines, 2-6 spines in
each group; T VI-VIII with weak to moderate reticulate cuticular pattern posterolaterally.
Posterior margin of T II with transverse row of 64-101, 84 hooklets. T VIII with 5 lateral
setae. Caudolateral spurs on T VIII (Figs. 2D, E) single or multiple, small. Anal lobes with
45-54, 49 setae. DR 2.76-3.16, 2.92.
MATERIAL EXAMINED: NIGERIA: Lake Opi nr Nsukka, 14-27-I-1979, leg. Landis Hare,
1 male, 1 female/Pex, 2 Pex (LH). Kafanchan, 6-V-1979, J.C. Deeming, 1 male (BM). SUDAN:
Khartoum, at light, XI-1951, D.J. Lewis, 3 males (BM). Liednum nr Wau, 13-IV-1955, E.T.M.
Reid, 1 male (BM). UGANDA: Lake Nabugabo, 13-XI-1934, RW. Edwards, 1 male (BM).
Dicrotendipes ealae (Freeman)
Chironomus (Dicrotendipes) ealae Freeman,-1957:369; Dejoux 1968:57.
Dicrotendipes ealae (Freeman): Freeman and Cranston 1980:190.
See adult description in Freeman (1957:369); the immature stages are un-
known. There are no ventral accessory setae apparent on S VI.
MATERIAL EXAMINED: S[OUTH] A[FRICA].: Natal: Howick Falls, 6-IV-1953, G.H.
Satchell, 2 males (holotype, paratype) (BM):
Dicrotendipes freemani nom. nov. (Fig. 8)
Chironomus binotatus Kieffer, 1911b:354 (junior homonym of Ch. binotatus Wiedemann,
1817).
Chironomus seychelleanus Kieffer, 1911b:356 (part) (junior synonym of Chironomus calli-
chirus Kieffer, 1911b); (part) = Ch. binotatus Kieffer, 1911b.
Chironomus (Dicrotendipes) binotatus Kieffer: Freeman 1957:367.
Dicrotendipes binotatus (Kieffer): Freeman and Cranston 1980:190.
See Freeman (1957) for the description of the adult; the immature stages
are unknown.
Chironomus seychelleanus was described from a male and a female (Kief-
fer 1911b). Freeman (1957:341, 367) found that the male was a Ch. calli-
chirus Kieffer and the female a D. binotatus. I have examined Kieffer’s
female specimen; it is a Dicrotendipes and has at least 2 ventral accessory
setae on S VI. It matches the holotype male of Ch. binotatus, which I also
examined, in coloration, and probably is the female of this species. Freeman
(1957:341) fixed the male specimen of Ch. seychelleanus as the lectotype;
by page priority he considered it a junior synonym of Ch. callichirus. He
JOHN H. EPLER 37
(Freeman 1957:368) listed the female as a ‘‘cotype’’; it automatically became
a paralectotype of Ch. seychelleanus (ICZN, Art. 73(b)(ii)). Because Ch.
binotatus Kieffer, 1911 is a junior homonym of Ch. binotatus Wiedemann,
1817 (now considered the type-species of Krenopelopia Fittkau; see Fittkau
1962:262), and Ch. seychelleanus Kieffer is not available for this species, a
new name must be given this species. I am happy to name this species for
Dr. Paul Freeman, in recognition for the work he has done on the African
chironomid fauna.
Dicrotendipes freemani is very similar to D. chambiensis (Goetghebuer).
These may be the same species, but without a comparison of the immature
stages of the 2 species I cannot consider synonymizing them.
The superior volsella of this species (Figs. 8A, B) also resembles that of
the Holarctic D. nervosus (Staeger) and/or the Indo-Pacific D. flexus (Jo-
hannsen). Again, reared specimens are needed to confirm species relation-
ships. See also remarks under D. chambiensis.
MATERIAL EXAMINED: ETHIOPIA: Bahar Dar, Tana-See, 26-I-1977, 1 male (ZS).
[SEYCHELLES ISLANDS]: La Reunion: St. Pierre Villa, 5-I-1956, J. Hamon, 1 male (BM);
Mahé, ’08-9, Seychelles Exp., 1 male, 1 female (holotype male of binotatus and paralectotype
female of seychelleanus) (BM) (Kieffer (1911b) lists the locality data for both specimens as:
““Seychellen, Mahé: marshes on coastal plain at Anse aux Pins and Anse Royal, 19-21-I-1909.’’)
Dicrotendipes fusconotatus (Kieffer) (Fig. 10)
[?] Dicrotendipes trilabis Kieffer, 1922:63.
Calochironomus fusconotatum Kieffer, 1922:68.
Calochironomus griseonotatus Kieffer, 1922:69.
Calochironomus griseosparsus Kieffer, 1922:69.
Dicrotendipes forficula Kieffer, 1925:298.
Dicrotendipes nilicola Kieffer. 1925:300.
Polypedilum quatuorpunctatum Goetghebuer, 1936:489.
Dicrotendipes fusconotatus (Kieffer): Freeman 1955a:22; Dejoux 1977:293; Freeman and Cran-
ston 1980:190; Reiss 1986:159; Contreras-Lichtenberg 1986:717.
Chironomus (Dicrotendipes) fusconotatus (Kieffer): Freeman 1957:362; Dejoux 1968:56.
See adult description in Freeman (1957:362) and Contreras-Lichtenberg
(1986:718). There are no ventral accessory setae apparent on S VI. Con-
treras-Lichtenberg (1986) also described the pupa and larva of D. fusco-
notatus. I cannot reliably distinguish the pupa of D. fusconotatus from D.
pallidicornis Goetghebuer, D. peringueyanus (as described by Contreras-
Lichtenberg 1986) or D. sudanicus. Characters given in couplets 7 and 8 of
Contreras-Lichtenberg’s (1986) pupal key will not separate these species (the
number of T II hooklets given for D. fusconotatus is very low; see descrip-
tion below); see also remarks under D. peringueyanus below.
MEM. AMER. ENT. SOC., 36
38 GENUS DICROTENDIPES
I follow Freeman (1957) in listing D. trilabis as a probable synonym; the
type is apparently lost and the description of the species appears to be that
of a teneral D. fusconotatus. Kieffer described fusconotatus as 4 different
species in one paper (Kieffer 1922) and as 2 more species in a later paper
(Kieffer 1925).
PUPA: (n=4)
COLOR. Very light brown, with light brown along lateral margins of tergites.
LENGTH. Total 4.08-4.90 mm (2).Cephalothorax 0.95-1.20 mm (2). Abdomen 3.13-3.69
mm (2).
CEPHALOTHORAX. Cephalic tubercles well developed (Fig. 10A). Dorsum moderately
to well pebbled. Dc, closer to Dc3. Thoracic horn base (Fig. 10B) with tracheal bundles separate.
ABDOMEN (Fig. 10C). Sternites II-III with fine lateral shagreen bands; also fine spinules
scattered over S II-III; T I with posterolateral weak to well-developed reticulate pattern; T II
with median quadrilateral shagreen area; T III-V with larger median quadrilateral shagreen
area, areas somewhat extended laterally on anterior and/or posterior portion; T VI with V
shaped shagreen area; T VII with an anterior pair of suboval shagreen areas; T VIII with a
pair of longitudinal bands of fine shagreen; shagreen areas of T II-VI with spines larger in
middle and posterior portion of area. Weak shagreen present dorsally on anal lobe. Tergites
III-V with posterior conjunctival band of fine spinules; T VI-VIII with well developed reticulate
cuticular pattern posterolaterally. Posterior margin of T II with transverse row of 59-68, 58
hooklets (40 in Contreras-Lichtenberg (1986)). T VIII with 5 lateral setae. Caudolateral spurs
on T VIII (Figs. 10D, E) single or multiple, very small. Anal lobes with 61-78 setae (3), partially
biserial. DR 2.35-2.64 (3).
FOURTH INSTAR LARVA: (n=7)
COLOR. Head capsule light brown to brown, mentum and mandibular apodemes darker,
sometimes with darker area anterior to postocciput; postocciput dark brown-black.
HEAD. Postmentum length 215-253, 234 (6). Mandible (Figs. 10F, G) length 203-223, 212
(6), with second and third lateral teeth partially fused, a shallow to deep incision proximal to
these, followed by a fourth lateral tooth or ‘‘hump’’; 2 well developed dorsal teeth present.
Pecten mandibularis composed of 11-15, 12 (6) setae. Mentum (Fig. 10H) with 13 teeth, Sth
and 6th teeth fused or closely appressed; width 133-143, 139 (6); MR 2.51-2.64, 2.59 (6).
Ventromental plate with crenulate anterior margin; width 99-106, 103 (6); length 40-50, 45.
VPR 2.10-2.50, 2.30 (6); IPD 72-79, 75 (6); PSR 1.27-1.44, 1.37 (6); 20-22, 21 strial ridges.
Length of antennal segments: 64-79, 69 (5); 16-19, 18 (4); 11-12, 11 (5S); 12-14, 13 (5); 6-7, 6
(5). AR 1.28-1.49, 1.40 (4) (Fig. 101). Inner blade of premandible (Fig. 10J) greater than outer
blade. Pecten epipharyngis (Fig. 10K) with 3-5, 5 (6) lobes. Anterior margin of frontal apotome
(Fig. 7D) with long, thin ventral frontal process; labral sclerite 1 smooth. S I with 6-9, 7 fringes
(Fig. 10L).
BODY. Ventral tubuli absent.
MATERIAL EXAMINED: ‘‘CONGO BELGE’: Ishango bae, 20-II-1954, J. Verbeke—KEA,
1 male (BM); P.N.A. Vitshumbi (S. Lac. Ed.), 12-14-I-1953, J. Verbeke-KEA, 4 males (BM).
EGYPT: Lake Nasser, El Madin, 7-XI-1971, M. Gillies, 1 male (BM); Lake Nasser, Nubia,
9-11-I-1981, P.S. Cranston, 2 males (BM). KENYA: Lake Beringo, Yellow Kay, XII-1977,
Waddesdon School Exp., 1 female (BM). SUDAN: Abu Hamed, March 1948, 1 male (US).
Blue Nile, Ummbenane nr Singa, light trap, 19-IV-1981, P. Mellor, 1 male (BM). Khartoum,
24-I-1953, D.J. Lewis, 2 males (BM). Wad Medani, Feb. 1952, D.J. Lewis, 1 male (BM). White
Nile, Kalakla, 20 km S Khartoum, to light, I-1980, P.S. Cranston, 4 males, 1 female (BM).
White Nile, Jebel Aulia, I-1980, P.S. Cranston, 1 male/Pex/Lex, 1 pharate male pupa/Lex,
2 males, 1 female/Pex 1 pharate female pupa, 5 larvae (BM).
JOHN H. EPLER 39
Dicrotendipes kribiicola (Kieffer) (Fig. 11)
Phytochironomus kribiicola Kieffer, 1923:152.
Chironomus (Dicrotendipes) kribiicola (Kieffer): Freeman 1957:368; Dejoux 1968:57.
Dicrotendipes kribiicola (Kieffer): Freeman and Cranston 1980:190; Hare & Carter 1987:70.
See adult description in Freeman (1957:368); the type is apparently lost.
There are 4-5 ventral accessory setae on S VI.
Freeman and Cranston (1980:190) erroneously listed Polypedilum as the
genus in which this species was originally placed. The record from Nigeria
is the first record of this species from that country.
I also examined a male specimen from Lake Kainji in Nigeria (in the BM
collection) which superficially resembles D. kribiicola, differing mainly in
the shape of the superior volsella. This specimen may represent a new spe-
cies, but is in too poor condition to allow sufficient description.
PUPA: (n=7)
COLOR. Light brown, with darker areas along lateral margins of tergites.
LENGTH. Total 3.92 mm (1). Cephalothorax 0.94—0.96 mm (2). Abdomen 2.70-3.17, 2.96
mm (5).
CEPHALOTHORAX. Cephalic tubercles well developed (Fig. 5A), 40-53, 49 (3) high, 60-
105, 88 (3) wide. Dorsum moderately pebbled. Dc, closer to Dc3. Thoracic horn base (Fig. 5B)
with tracheal bundles fused medially or very closely appressed.
ABDOMEN (Fig. 11C). Sternites I-III with fine shagreen. Tergite I sometimes with weak
lateral shagreen areas; T II-V with median quadrilateral shagreen area; T VI with a pair of
median triangular shagreen areas, areas sometimes narrowly joined; T VII with anterior band
of fine shagreen or a pair of anterolateral ovoid shagreen areas; T VIII with an anterior pair
of ovoid fine shagreen areas and a smaller posterior pair of ovoid shagreen areas of these two
areas joined in a pair of lateral bands; shagreen areas on T II-VI with spines larger in middle
of area. Tergites III-V or IV-V with posterior conjunctival band of fine spinules; T V posteriorly
with 2 groups of intersegmental spines, 2-4 spines in each group. Posterior margin of T II
with transverse row of 52-68, 63 hooklets. T VIII with 5 lateral setae. Caudolateral spurs on
T VIII (Fig. 11D) single or multiple, small to thorn-shaped. Anal lobes with 28-39, 36 setae.
DR 1.50-2.41, 1.83.
FOURTH INSTAR LARVA: (n=4)
COLOR. Head capsule light yellow-brown, postmentum darker brown; postocciput, men-
tum and mandibular apodemes dark red-brown.
HEAD. Postmentum length 180-195, 190 (3). Mandible (Fig. 11E) length 157-165, 160 with
3 lateral teeth; 2 well-developed dorsal teeth present. Pecten mandibularis composed of 9-10,
9 setae. Mentum (Fig. 11F) with apparently only 11 teeth, 5th and 6th teeth fused, 4th closely
appressed; width 105-115, 110 (3); MR 2.10-2.20, 2.16 (3). Ventromental plate with smooth
anterior margin; width 78-85, 83; length 38-43, 42. VPR 1.81-2.24, 1.99; IPD 41-45, 43 (3);
PSR 1.81-2.07, 1.91 (3); 24-28, 25 strial ridges. Length of antennal segments: 54-61, 57; 18-
20, 19; 9-10, 9 (3); 12-14, 13 (3); 4-5, 5 (3). AR 1.17-1.29, 1.23 (3) (Fig. 11G). Inner blade
of premandible (Fig. 11H) subequal to outer blade. Pecten epipharyngis (Fig. 111) with 5 lobes.
Frontal apotome (Fig. 11J) with large ventral mesal ovoid pit; labral sclerite 1 smooth, reduced,
much broader than long. S I with 6-9 (2) fringes.
BODY. Ventral tubuli absent.
MEM. AMER. ENT. SOC., 36
40 GENUS DICROTENDIPES
MATERIAL EXAMINED: GHANA: Accra, J.D. Thomas, 12 males (BM). NIGERIA: Lake
Opi nr Nsukka, mass rearing, 16-22-X-1977, leg. L. Hare, 1 male/Pex, 1 Pex; same locality,
light trap, 31-XII-1978-1-I-1979, leg. L. Hare, 1 male (LH); same locality, mass rearing, 14-
25-I-1979, leg. L. Hare, 2 males/Pex (LH); same locality, 28-I-3-II-1979, leg. L. Hare, 1 female/
Pex/Lex (LH); same locality, 10-11-III-1979, leg. L. Hare, 1 female/Pex/Lex (LH); same lo-
cality, 22-I1V-1979, leg. L. Hare, 3 larvae (LH); same locality, emergence trap, 24-25-XI-1979,
leg. L. Hare, 1 male/Pex (LH). [ZAIRE]: “‘Congo Belge,’’ Eala, I-1935, J. Ghesquiere, 1 male
(BM).
Dicrotendipes leucolabis Kieffer
Dicrotendipes leucolabis Kieffer, 1922:65; Freeman and Cranston 1980:190.
Polypedilum (?) aequatoris Goetghebuer, 1936:482.
Chironomus (Dicrotendipes) leucolabis (Kieffer): Freeman 1957:367.
See adult descriptions in Kieffer (1922:65), Goetghebuer (1936:482) and
Freeman (1957:367); immature stages are unknown.
This species may be a senior synonym of D. collarti. However, the type
of leucolabis is probably lost and Kieffer’s figure (1922: Fig. 67) of the
hypopygium is insufficiently clear. Freeman (1957) synonymized Polype-
dilum aequatoris, known only as a female, with this species on the basis of
wing maculation and body coloration. Kieffer’s (1922) description of the
wing of D. leucolabis does seem to match that illustrated for the female
aequatoris specimen (Goetghebuer 1936:Fig. 35). The wing pattern of Jeu-
colabis differs from that of collarti: the transverse band is adjacent to RM
and FCu in collarti and more distal in /eucolabis; also the quadrate spot
over the anal vein is smaller in collarti. These differences may be variations
within one species; reared associations would have to be examined to de-
termine whether 1 or 2 species are present.
Goetghebuer (1936) was apparently unsure of the generic position of this
species, which he denoted with a parenthetical question mark following the
genus name; Freeman examined the type and considered it a Dicrotendipes
on the basis of tibial spurs. I have not seen any specimens of D. leucolabis.
Dicrotendipes nigrolineatus (Freeman)
Chironomus (Dicrotendipes) nigrolineatus Freeman, 1957:370.
Dicrotendipes nigrolineatus (Freeman): Freeman and Cranston 1980:190.
See adult description in Freeman (1957:370); immature stages are un-
known. This species may be an Einfeldia; examination of reared immature
stages is necessary for correct placement. There are no ventral accessory
setae apparent on S VI.
JOHN H. EPLER 4]
MATERIAL EXAMINED: [ZAIRE]: ‘‘Belgian Congo’’: Elisabethville, 17-31-XII-1932, C.
Seydel, 3 females (paratypes) (BM); same locality & collector, II-1933, 1 female (paratype)
(BM); same locality & collector, 1-12-I1I-1933, 1 male (holotype) (BM).
Dicrotendipes peringueyanus Kieffer
Dicrotendipes peringueyanus Kieffer, 1924:257; Freeman 1955b:372; Disney 1975; Reiss
1977b:91, 93; Freeman and Cranston 1980:190; Prat 1981:58; Contreras-Lichtenberg
1986:706.
Polypedilum griseovittatum Goetghebuer, 1936:485.
Chironomus (Dicrotendipes) peringueyanus (Kieffer): Freeman 1957:364; Dejoux 1968:57.
See adult description in Freeman (1957:364) and Contreras-Lichtenberg
(1986:707); the pupa has been described by Contreras-Lichtenberg
(1986:706); the larva is unknown. There were no ventral accessory setae
present on the males I examined.
This species may be a variant of D. fusconotatus. The genitalia are similar
and the 2 species differ only slightly in wing maculation (see Freeman 1957).
Without associated larval stages a decision regarding the status of the 2
species cannot be made.
Disney (1975) reported the phoretic association of the immature stages
of D. peringueyanus with the African river crabs Potomanautes africanus
(A. Milne-Edwards) and P. pobeguini (Rathbun) in Cameroon. The chi-
ronomid identifications were based on larvae reared to adults. Disney placed
this material in the BM (Disney, pers. comm.). However, this material could
not be located recently (Cranston, pers. comm.).
The pupa of D.peringueyanus as described by Contreras-Lichtenberg
(1986) is not separable from D. fusconotatus, D. pallidicornis or D. su-
danicus. In her pupal key, Contreras-Lichtenberg (1986) separates D. per-
igueyanus from D. pallidicornis based on the absence of conjunctival spi-
nules on D. pallidicornis. However, all specimens of D. pallidicornis which
I have examined do possess these spinules.
MATERIAL EXAMINED: [CAMEROON]: Kumba, Mambonjese, [on] crab P. africanus,
13-V-1969, [Disney], 1 male hypopygium & leg, det. A.D. Harrison (BM). [SOUTH AFRICA]:
Cape Prov: Upington, XI-1950, P. Brinck, 2 males (BM); Transvaal: Nelspruit, XI-1956-II-
1957, G.H. Frank, 1 male (BM).
Dicrotendipes schoutedeni (Goetghebuer)
Chironomus (Limnochironomus) Schoutedeni Goetghebuer, 1936:465.
Chironomus (Dicrotendipes) schoutedeni Goetghebuer: Freeman 1957:370; Freeman 1961b:247.
Dicrotendipes schoutedeni (Goetghebuer): Freeman and Cranston 1980:190; Hare & Carter
1987:70.
MEM. AMER. ENT. SOC., 36
42 GENUS DICROTENDIPES
See adult descriptions in Goetghebuer (1936:465) and Freeman (1957:370);
the immature stages are unknown. The anal tergal bands are unusual for a
Dicrotendipes; there are no apparent ventral accessory setae on S VI. I have
not examined the holotype.
MATERIAL EXAMINED: [BENIN] ‘“‘DAHOMEY”’: cercle de Porto Novo, GBE home,
7-12-54, J. Hamon, 1 male (BM). NIGERIA: Lake Kainji, A. Bidwell, 1 male (BM); Lake
Opi nr Nsukka, light trap, 13-14-I-1979, leg. L. Hare, 1 male (LH).
Dicrotendipes septemmaculatus (Becker) (Figs. 12, 49)
Chironomus septemmaculatus Becker, 1908:77.
[?] Chironomus (Prochironomus) punctatipennis Kieffer, 1910:234. NEW SYNONYMY.
Dicrotendipes pictipennis Kieffer, 1913b:23; Freeman 1955a:22.
[?] Tendipes punctatipennis (Kieffer): Kieffer 1913a:138.
Dicrotendipes pilosimanus Kieffer, 1914:262; Freeman 1955b:372; Sublette and Sublette
1973:404; Reiss 1977b:93; Reiss 1978:75; Freeman and Cranston 1980:190; Reiss 1986:159;
Contreras-Lichtenberg 1986:716; Chaudhuri & Guha 1987:27.
Dicrotendipes formosanus Kieffer, 1916:115; Sublette and Sublette 1973:403; Hashimoto et al.
1981:12; Sasa and Hasegawa 1983:320. NEW SYNONYMY.
Dicrotendipes formosanus var frontalis Kieffer, 1916:116.
Dicrotendipes speciosus Kieffer, 1924:256; Kieffer 1925:299.
Stictochironomus sexnotatus Goetghebuer, 1930:95.
Chironomus hirtitarsis Johannsen, 1932:534; Sublette and Sublette 1973:402. NEW
SYNONYMY.
Polypedilum quatuordecimpunctatum Goetghebuer, 1936:48.
Dicranotendipes speciosus Kieffer: Kruseman 1949:254 (misspelling).
Chironomus (Dicrotendipes) pilosimanus (Kieffer): Freeman 1954b:19; Freeman 1957:360;
Freeman 1961a:247; Freeman 1961b:694; Dejoux 1968:56 (misspelled as pilosinamus).
Chironomus (Dicrotendipes) pilosimanus subsp. quatuordecimpunctatus (Goetghebuer): Free-
man 1957:361.
Dicrotendipes frontalis Kieffer: Sublette and Sublette 1973:403. NEW SYNONYMY.
[?] Dicrotendipes punctatipennis (Kieffer): Sublette and Sublette 1973:404; Chaudhuri & Guha
1987:27 (misspelled as punctipennis).
Dicrotendipes rajasthani Singh and Kulshrestha, 1977:233. NEW SYNONYMY.
Dicrotendipes hiritarsis (Johannsen): Guha et al. 1982:30; Chaudhuri & Guha 1987:27
(misspelling).
Dicrotendipes quatuordecimpunctatus (Goetghebuer): Contreras-Lichtenberg 1986:710. NEW
SYNONYMY.
Dicrotendipes septemmaculatus (Becker): Cranston and Armitage 1988.
See adult descriptions in Freeman (1957:360; 1961b:694; as pilosimanus)
and Contreras-Lichtenberg (1986:711, 716, as pilosimanus and quatuor-
decimpunctatus). There are no ventral accessory setae apparent on S VI.
Dicrotendipes septemmaculatus is one of the most widely distributed
members of this genus in the world (Fig. 49). I have examined reared ma-
terial from the Afrotropical, southern Palaearctic (Lebanon) and Australian
JOHN H. EPLER 43
regions. Afrotropical pupae possess long, well developed cephalic tubercles;
in the one reared Australian pupa examined, the cephalic tubercles are not
as well developed. Cephalic tubercle length often varies greatly within spe-
cies in other species of Dicrotendipes. Pupae of D. septemmaculatus are
difficult to separate from D. fusconotatus, D. pallidicornis, D. peringuey-
anus and D. sudanicus. Shagreen patterns in all 5 species are similar, and
T II hooklet counts and anal lobe setal numbers overlap. Dicrotendipes
septemmaculatus possesses a relatively well developed humeral comb or
ridge; however, the development of this structure varies intraspecifically. It
is also present, although more weakly developed, in D. fusconotatus and
D. sudanicus. In D. septemmaculatus and D. sudanicus, Dc, is closer to De,
than to Dc,; however, D. septemmaculatus generally is larger, possesses long,
well developed cephalic tubercles (shorter, squatter in D. suwdanicus) and
generally has more anal lobe setae. I would consider specific identification
of isolated pupal exuviae of these 5 species to be risky.
Wing spots are variable in D. septemmaculatus. They may be absent in
teneral specimens, and the pair of spots in cell r,,,; is sometimes combined
into one spot.
Based on Afrotropical material, Freeman (1957) considered D. pilosi-
manus (=D. septemmaculatus) to consist of 2 geographical subspecies, D.
p. pilosimanus and D. p. quatuordecimpunctatus (Goetghebuer). These dif-
fer only in the presence (D. p. pilosimanus) or absence (D. p. quatuorde-
cimpunctatus) of a fore metatarsal beard, and, as Freeman (1957) noted,
intermediates occur. In all reared material made available to me for this
study, the adults possessed a sparse to moderately developed beard on fore
ta, and ta,. The moderately developed beard on many specimens was at least
as well developed as that on specimens in the British Museum collection
determined as D. p. pilosimanus. Although some workers believe 2 species
may be present, until more reared material of both “‘subspecies’’ is made
available, I see little need to continue the usage of quatuordecimpunctatus.
Fortunately, the discovery of the Becker type of septemmaculatus no longer
necessitates the usage of the name pilosimanus and its subspecies (Cranston
& Armitage 1988). I have examined the Becker female holotype specimen.
Cranston & Armitage (1988) also rediscovered the type of Stictochiron-
omus sexnotatus Goetghebuer and considered the species to be a junior
synonym of D. septemmaculatus. I have also examined the male lectotype
specimen and concur.
I have listed Ch. punctatipennis Kieffer as a probable synonym of sep-
temmaculatus. Kieffer’s (1910) original description of punctatipennis lacked
illustrations and for the most part described wing maculation. In a later
redescription based on additional specimens, Kieffer (1913a) provided an
MEM. AMER. ENT. SOC., 36
44 GENUS DICROTENDIPES
illustration of the hypopygium and more morphological details. In the sec-
ond description (plate XI, Fig. 10, mislabeled as Tendipes punctatissimus),
Kieffer stated (and illustrated) that the male of punctatipennis possessed 3
volsellae. Kieffer may have been mistaking the proximal arm of the deeply
bifid inferior volsella (as in septemmaculatus) for a third volsella. However,
without examining a male from this series, or the type series (if one exists),
I can not be positive that Kieffer may have made such a mistake.
Some of the material redescribed as Ch. punctatipennis by Kieffer (1913a)
came from the collection of E. Brunetti; Brunetti’s collection is now with
the British Museum. I examined 3 female specimens from Bosundhur (Bang-
ladesh), leg. J.T. Jenkins, from that collection. Two are labeled ‘‘ 7endipes
punctatipennis’’ in what appears to be Kieffer’s hand-writing; the third is
labeled ‘‘Chiron. punctatipennis Kieffer.’’ These females are to me in-
separable from D. septemmaculatus.
The type or type series (if one exists) of D. punctatipennis and perhaps
D. semiviridis (Kieffer, 1911) may be with the Zoological Survey of India,
Calcutta. According to Dr. M. Datta, Zoological Survey of India (pers.
comm., 12 Sept. 1986), specimens of punctatipennis (and semiviridis) are
present in the collection, but ‘‘are in extremely miserable condition and are
suggestive of not being mailed to anybody so as to save from further de-
terioration.’’ Obviously, an on-site examination of these specimens by a
competent chironomid worker will be necessary to clear up the taxonomy
of D. punctatipennis.
I consider D. formosanus Kieffer to be a junior synonym of D. septem-
maculatus; only minor differences in coloration had separated the 2 species,
which I do not consider to be sufficient reason to maintain the separation.
The type of formosanus was apparently destroyed in the 1956 fire at the
Hungarian National Museum.
Hashimoto et al. (1981:12) designated a holotype and paratypes for for-
mosanus from Thailand material. These designations are invalid (ICZN,
Chapter XVI); holotypes and paratypes can not be designated by any other
than the original author. Lectotypes and paralectotypes can not be desig-
nated from Thailand material either, because such types can only be des-
ignated from the original series. Perhaps a neotype could be designated, but
because Hashimoto et al. (1981) is not a revisory work, such a designation
could also be considered invalid.
PUPA: (n=6)
COLOR. Light yellow-brown, with light brown along lateral margins of tergites.
LENGTH. Total 5.90-6.14, 5.98 mm (3). Cephalothorax 1.30-1.50, 1.41 mm (3). Abdomen
4.23-4.71, 4.49 mm (5).
CEPHALOTHORAX. Cephalic tubercles well developed (Fig. 6A), 210-250, 228 (3) high,
125-162, 139 (3) wide. Dorsum and frontal apotome moderately to well pebbled. Dc, closer
JOHN H. EPLER 45
to Dc,. Thoracic horn base (Fig. 6B) with tracheal bundles separate. Dorsal portion of humeral
callus usually with moderately developed ‘‘comb’’ or ridge.
ABDOMEN. Similar to D. fusconotatus (Fig. 10C). Sternite I sometimes with very fine
median shagreen spinules; sternites II-III with fine lateral shagreen bands; also fine spinules
scattered over S II-III; Tergite I with posterolateral weak reticulate pattern; T II with median
quadrilateral shagreen area; T III-V with larger median quadrilateral shagreen area, areas
somewhat extended laterally on anterior and/or posterior portion; T VI with broadly V shaped
shagreen area; T VII with an anterior pair of suboval shagreen areas, sometimes joined me-
dially; T VIII with a pair of longitudinal bands of fine shagreen; shagreen areas on T II-VI
with spines larger in middle and posterior portion of area. Well developed shagreen area present
dorsally on anal lobe. Tergites III-V with posterior conjunctival band of fine spinules; T V-
VIII with reticulate cuticular pattern posterolaterally, well developed on T VI-VIII. Posterior
margin of T II with transverse row of 66-93, 79 hooklets. T VIII with 5 lateral setae. Cau-
dolateral spurs on T VIII single or multiple, small (similar to D. fusconotatus, Figs. 10D, E).
Anal lobes with 47-67, 57 setae, partially biserial. DR 2.25-3.03, 2.63.
FOURTH INSTAR LARVA: (n=4)
COLOR. Head capsule light brown to brown, mentum and postocciput red-brown, with
slight caudal darkening of postmentum.
HEAD. Postmentum length 240-253, 245. Mandible (Fig. 12C) length 220-240, 230, with
three lateral teeth; 2 well developed dorsal teeth present. Pecten mandibularis composed of
10-12, 11 setae, apical seta very broad. Mentum (Fig. 12D) with 13 teeth, Ist and 2nd teeth
fused or closely appressed; width 150-162, 153; MR 2.58-2.88, 2.71. Ventromental plate with
mostly smooth anterior margin, sometimes with extremely shallow crenulations; width 100-
109, 104; length 52-65, 55. VPR 1.72-2.04, 1.88; IPD 63-72, 67; PSR 1.47-1.63, 1.55; 27-
29, 28 strial ridges. Length of antennal segments: 79-87, 84; 25-27, 26; 12; 15-18, 17; 6. AR
1.32-1.43, 1.39 (Fig. 12E). Inner blade of premandible (Fig. 12F) greater than outer blade.
Pecten epipharyngis (Fig. 12G) with 5 lobes. Anterior margin of frontal apotome (Fig. 12H)
with long, thin ventral frontal process; labral sclerite 1 smooth. S I with 8-10 (2) fringes.
BODY. Ventral tubuli absent.
MATERIAL EXAMINED: ALGERIA: Oued Bechar, 26-III-1955, leg. E.J. Fittkau, 1 male/
Pex/Lex, 3 Pex (ZS). AUSTRALIA: New South Wales: Deniliquin, from egg mass #6, 6-I-
1969, 1 male (JM). Northern Territory: Fogg Dam, 15 km NE Humpty Doo, at UV light, 5-
X-1982, leg. J.K. Balciunas, J. Gillett, 5 males, 1 female (JB). Queensland: Charleville, light
trap, 17-II-1969, leg. A.L. Dyce, 1 male (JM); Sandgate, 14-V-1974, leg. J. Martin, 1 male/
Pex (JM); Somerset Dam edge, 80 mi N Brisbane, 8-VI-1968, leg. J. Martin, 1 male (JM).
[BANGLADESH]: Khulna, Bosondhur, Ganges delta, on launch, 29-VII-1909, J.T. Jenkins,
3 females (BM) (punctatipennis specimens). BURMA: Shan State, Lnle Marsh, % mi N of
Shwevanpye, on Hydrilla, 26-V1-1982, leg. J.K. Balciunas, 2 larvae (JB). CANARY ISLANDS:
Tenerife, Puerto Orotava, ‘‘51307.8,’’ 1 female (holotype of Ch. septemmaculatus Becker)
(ZH). EGYPT: Luxor, 12-I-1981, leg. P.S. Cranston, 1 male (BM). INDIA: Karnataka: Ban-
galore, 20 May 1982, leg. J.K. Balciunas, M.C. Minno, 1 pharate female pupa (JB); Dasap-
paddodi Pond, 35.5 km WSW Bangalore, at UV light, 24-V-1982, leg. J.K. Balciunas, M.C.
Minno, 1 female (JB); Medahalli Well, 15 km E of Bangalore, at UV light, 21-V-1982, leg.
J.K. Balciunas, M.C. Minno, 3 males (JB); Nandi Hills, el. 1200 m, 5 Oct 1985, leg. C.W. &
L.B. O’Brien, 1 male (FS). Kerala: Chalakuby, 32 km S Trichur, 10-X-1985, leg. C.W. & L.B.
O’Brien, 1 male (FS). Mahar: Rahuri, Mula River, Oct. 17, 1985, leg. C.W. & L.B. O’Brien,
1 male (FS). West Bengal: Burdwan, 7-VII-1978, leg. S.K. Nandi, 1 male (BM); Malda, 4-IV-
1975, 1 female (BM). INDONESIA: Java: Central Java, Jombor Lake, 10 km S Klaten, on
Hydrilla, 28 Aug 1981, leg. J.K. Balciunas, 1 larva (JB). South Sulawesi: Lake Lampulung,
MEM. AMER. ENT. SOC., 36
46 GENUS DICROTENDIPES
5 km NE Seng Kang, on Hydrilla, 3-1X-1982, leg. J.K. Balciunas, 1 larva (JB); same data,
except at black light, 65 males (JB). Sumatra: Bay of Meat, Lake Toba, 10-IV-1929, 1 male
(minus wing, hypopygium) (paratype Ch. hirtitarsis) (BM). ISRAEL: Beth Netufa, Sept 1968,
leg. J. Kugler, 1 male (JM). JAPAN: Hamamatsu City, Sept-Nov. 1984, leg. H. Hashimoto,
3 males, 5 females (HH). LEBANON: Ammik, el. 850 m, 16-V-1982, leg. Z. Moubayed, 4
males, 2 females, 3 Pex (ZM); Baalbek, el. 1150 m, 18-X-1982, leg. Z. Moubayed, 1 male, 1
pharate male pupa, 1 pharate female pupa, 13 Pex (ZM); River Litani at Jib-Jennine, 1 female,
1 pharate female pupa, 8 Pex, 4 larvae (ZM). MALAGASY: Pr. Tanan, Ankeniheny River
(28°C) 4 km S Manjakatompo Forest Sta., 1-XI-71, G.F. & C.H. Edmunds, F. Emmanuel, 2
males (FS). MALAYSIA: Penang State: irrigation canal in Balik Pulau Village, on Hydrilla,
Sept 7 1983, leg. J.K. Balciunas, 1 larva (JB). [NAMIBIA]: ‘‘S.W. AFRICA”: Swakopmund,
26-30-I-1972, Southern African Exp., 2 males (BM). NIGERIA: Jos., 24-XI-1970, J.C. Deem-
ing, 1 male, 2 females (BM). [SOUTH AFRICA): Transvaal: Olifantsvlei, nr Johannesburg,
22-1X-54, A.D. Harrison, 1 male (BM); Jukskei River, nr Johannesburg, VII-1956, A.D. Har-
rison & B.R. Allansen, 1 male (BM). SPAIN: Algeciras, 1 male (lectotype of Stictochironomus
sexnotatus Goetghebuer) (NM). SRI LANKA: 1975, leg. F. Schiemer, 1 male (ZS). [SUDAN]:
W. Darfur: N. Jebel Murra, Kurra, 5600 ft, 4-VII-1932, M. Steele, 1 male (BM). UGANDA:
L. Victoria, 21-VI-1950, W.W. Macdonald, 1 male (BM). YEMEN: San/’a, ca. 7900 ft., at
night, 10-15-X-1937, C. Rathjens, 1 male (BM). [ZIMBABWE] ‘‘N. RHODESIA”’: Salisbury,
V-1956, E.T.M. Reid, 1 male (BM). [ZAIRE]: Elizabethville, XI-1933, C. Seydel, 1 male (BM);
Elizabethville, 17-XII-1938, H.J. Bredo, 1 female (BM).
Dicrotendipes sudanicus (Freeman) (Fig. 12)
Chironomus (Dicrotendipes) sudanicus Freeman, 1957:365; Dejoux 1968:56.
Dicrotendipes sudanicus (Freeman): Dejoux 1977:294; Freeman and Cranston 1980:190; De-
joux 1984:161; Hare & Carter 1987:70.
See adult description in Freeman (1957:365). There are no ventral acces-
sory setae apparent on S VI.
This species appears to be a diminutive D. fusconotatus. The hypopygia
of the 2 species are basically identical. The adults of the species differ only
in relative size and in wing maculation; the wing markings of swdanicus are
“‘clouds’’ along and over the veins while fusconotatus possesses spots in the
cells. The pupae differ slightly. The pupa of fusconotatus is larger, has more
anal lobe setae (61-78) and Dc, is closer to Dc,. The pupa of sudanicus is
smaller, has fewer anal lobe setae (35-47) and Dc, is closer to Dc,. The larval
mentum of D. sudanicus is similar to D. septemmaculatus and has the 2nd
lateral tooth fused/appressed to the 1st lateral tooth; in D. fusconotatus
the 6th lateral tooth is fused/appressed to the Sth, and the Ist and 2nd lateral
teeth are separate. Larvae also differ in size, D. fusconotatus being much
larger, and in ventromental strial ridge counts (20-22 in fusconotatus, 27-
29 in septemmaculatus, 22-26 in sudanicus). The pupal and larval data are
based on a very small sample.
JOHN H. EPLER 47
PUPA: (n=5)
COLOR. Light yellow-brown, with light brown along lateral margins of tergites.
LENGTH. Total 3.21 mm (1). Cephalothorax 0.83 mm (1). Abdomen 2.38-3.10, 2.77 mm
(3).
CEPHALOTHORAX. Cephalic tubercles well developed (Fig. 12J), 45-47 (2) high, 65-88
(2) wide. Dorsum moderately pebbled. Dc, closer to Dc;. Thoracic horn base (Fig. 12K) with
tracheal bundles separate.
ABDOMEN. Similar to D. fusconotatus (Fig. 10C). Sternite I with very fine median shagreen
spinules; sternites II-III with fine lateral shagreen bands; also fine spinules scattered over S
II-III; Tergite I with anteromedian weak reticulate pattern; T II with median quadrilateral
shagreen area; T III-V with larger median quadrilateral shagreen area, areas somewhat ex-
tended laterally on anterior and/or posterior portion; T VI with broadly V shaped shagreen
area; T VII with an anterior pair of suboval shagreen areas; T VIII with a pair of longitudinal
bands of fine shagreen, or an anterior and posterior ovoid to reniform pair of fine shagreen
areas; shagreen areas on T II-VI with spines larger in middle and posterior portion of area.
Well developed shagreen area present dorsally on anal lobe. Tergites III-V with posterior con-
junctival band of fine spinules; T VI-VIII with reticulate cuticular pattern posterolaterally well
developed on T VI-VII. Posterior margin of T II with transverse row of 56-70, 63 hooklets.
T VIII with 5 lateral setae. Caudolateral spurs on T VIII single or multiple, small (similar to
D. fusconotatus, Figs. 10D, E). Anal lobes with 35-47, 40 setae. DR 2.50-2.69, 2.57.
FOURTH INSTAR LARVA: (n=5)
COLOR. Head capsule light brown, mentum and postocciput red-brown; postmentum not
darkened.
HEAD. Postmentum length 175-193, 182. Mandible length 163-165, 164 (3) with three lateral
teeth; 2 dorsal teeth present. Pecten mandibularis composed of 8-10, 9 (4) setae. Mentum (Fig.
12L) with 13 teeth, Ist and 2nd teeth fused or closely appressed; width 100-110, 105 (4); MR
2.38-2.64, 2.51 (3). Ventromental plate with mostly smooth anterior margin, sometimes with
extremely shallow crenulations; width 81-87, 83 (4); length 40-43, 41 (4). VPR 1.91-2.18, 2.03
(4); IPD 46-51, 49 (4); PSR 1.61-1.89, 1.69 (4); 22-26, 24 strial ridges. Length of antennal
segments: 52-61, 57 (4); 14-16, 15 (4); 8-9, 9 (3); 13 (2); 6 (3). AR 1.21-1.39 (2) (Fig. 12M).
Inner blade of premandible greater than outer blade. Pecten epipharyngis with 4-5, 5 lobes.
Anterior margin of frontal apotome with long, thin ventral frontal process; labral sclerite 1
smooth. S I with 5-7, 6 fringes.
BODY. Ventral tubuli absent.
MATERIAL EXAMINED: MALAGASY: Pr. Tanan, Antanifotsy R. (20° C), at forest
station, 31-X-71, G.E. & C.H. Edmunds, F. Emmanuel, 1 male (FS). NIGERIA: Kaduna, 19-
X-1956, B. McMillan, 2 males (BM); Lake Opi nr Nsukka, light trap, 26-26-XI-1978, leg L.
Hare, 1 female (CH). SUDAN: Adok, 21-XI-1953, 1 male (holotype) (BM); Leidnum nr Wau,
III-IV-1955, E.T.M. Reid, 1 male (paratype) (BM); Melut, 17-XI-1953, E.T.M. Reid, 1 female
(paratype) (BM). TANZANIA: U. Pangani Rv, Ngumba Ya Mungu Reservoir, emergence trap,
15-VIII-1973, R.G. Bailey, 1 male (BM). [ZIMBABWE?]: Lake Karibe, 1966, leg. A.J.
McLachlan, 1 male, 6 Pex, 6 Lex (BM).
MEM. AMER. ENT. SOC., 36
48 GENUS DICROTENDIPES
Fic. 8. Superior volsellae of some Afrotropical Dicrotendipes. A, B) D.
freemani. C, D) D. chambiensis.
MEM. AMER. ENT. SOC., 36
JOHN H. EPLER
49
50 GENUS DICROTENDIPES
Fic. 9. D. cordatus, pupa. A) cephalic tubercles. B) Thoracic horn bases
from 2 specimens. C) Abdomen, dorsal. D, E) Caudolateral spurs on T
VIII.
51
JOHN H. EPLER
aft
MEM. AMER. ENT. SOC., 36
3 GENUS DICROTENDIPES
Fic. 10. D. fusconotatus, pupa (A-E) and larva (F-L). A) Cephalic tu-
bercle. B) Thoracic horn base. C) Abdomen, dorsal. D, E) Caudolateral
spurs on T VIII. F) Mandible, ventral. G) Mandible, dorsal. H) Mentum
and ventromental plate. 1) Antenna. J) Premandible. K) Pecten epiphar-
yngis. L) S I.
5)3)
JOHN H. EPLER
MEM. AMER. ENT. SOC., 36
54 GENUS DICROTENDIPES
Fic. 11. D. kribiicola, pupa (A-D) and larva (E-J). A) Cephalic tubercle.
B) Thoracic horn base. C) Abdomen, dorsal. D) Caudolateral spur on T
VIII. E) Mandible, ventral. F) Mentum and ventromental plate. G) An-
tenna. H) Premandible. I) Pecten epipharyngis. J) Frontal apotome and
labral sclerites. K) S I.
JOHN H. EPLER 55
Nf
SS
MEM. AMER. ENT. SOC., 36
56 GENUS DICROTENDIPES
Fic. 12. D. septemmaculatus, pupa (A, B) and larva (C-I). A) Cephalic
tubercle. B) Thoracic horn base. C) Mandible, ventral. D) Mentum and
ventromental plate. E) Antenna. F) Premandible. G) Pecten epipharyngis.
H) Frontal apotome and labral sclerites. I) S I. D. sudanicus, pupa (J, K)
and larva (L, M). J) Cephalic tubercle. K) Thoracic horn base. L) Mentum
and ventromental plate. M) Antenna.
37)
JOHN H. EPLER
58 GENUS DICROTENDIPES
CHAPTER III.
THE NEOTROPICAL DICROTENDIPES
With only a few exceptions (Brundin 1966; Edwards 1931; Reiss 1972,
1974), the Chironomidae of South America have not been studied in great
detail. Reiss (1977a, 1982) has summarized the majority of papers dealing
with the taxonomy of the family from South America.
The Dicrotendipes of the Neotropical region are poorly known. Rempel
(1939:213) described a species from Brazil, Chironomus (Calochironomus)
atripennis, which may be a Dicrotendipes. However, I have not seen any
material of this species, and the description and figures are not sufficient
to place this species in Dicrotendipes. Fittkau & Reiss (1979) listed one de-
scribed and 10 undescribed species of Dicrotendipes from tropical and sub-
tropical South America. To date, only 6 species have been described from
the region (Paggi 1975, 1978, 1987; Epler 1987b). In this chapter, I review
the known species, and supply descriptions for 10 new species, 9 of which
are from the Amazon area.
It is unfortunate that the immature stages, especially the larvae, remain
unknown for the majority of the species described here. As a result, the
majority of the Neotropical species could not be used in the phylogenetic
analysis (Chap. VI).
Many of the species from the Amazon area are quite bizarre, e.g., D.
soccus, when compared with the Holarctic or Afrotropical fauna. All species
from South America with deeply bifid inferior volsellae known to me pos-
sess palmate sensilla chaetica on the hind metatarsus. Such sensilla chaetica
are lacking in Afrotropical forms with deeply bifid inferior volsellae (but
are present in 2 species with simple inferior volsellae from the Oriental-
Australasian region, and in one species from the Neotropics, D. alsinensis).
I have little doubt that more species of South American Dicrotendipes
remain to be discovered and described. The following key must be consid-
ered as incomplete in this respect. Because the immature stages of Neo-
tropical Dicrotendipes remain largely unknown, no keys are offered for their
identification. Paggi (1987) offered a key for all stages of Argentinian Di-
crotendipes known to her.
KEY TO ADULT MALES OF NEOTROPICAL DICROTENDIPES
1. Inferior volsella simple or at most deeply notched apically (Figs. 13-16); palmate sensilla
chaetica absent on male hind metatarsus (except in D. alsinensis; see couplet 5) . 2
Inferior volsella deeply bifid, each lobe well separated (Figs. 17, 19, 20, 24-27) or inferior
volsella stout and extremely setose (Figs. 18, 23); palmate sensilla chaetica present
on maleshind\metatarsuss 75 oc a ce eee ee eee 7
10.
11.
12.
13.
14.
15.
16.
JOHN H. EPLER 59
Base of anal point raised, truncate, with more than 10 dorsal basal setae (Fig. 16) . .
OF Ar diors: Chola nen cn ace ee BERG Rainer tac, "GA awr in Ampeouc esd D. sinoposus Epler
Base of anal point not truncate; 7 or fewer dorsal basal setae ............. 3
Superior volsella with short ventral, basal membranous flap (Fig. 22)..........
6. hal Be Ger OL OOs ee eta acho Sou ORR one ean ite ont D. palearivillosus sp. nov.
Superior volsella without membranous flap..................+.2-.4- 4
Superior volsella cylindrical-digitiform with membranous apex............. 5
Superior volsellaypediformim-cces suscan a as sucech ) neice ee cae ao Gees aes 6
Foretarsal beard well developed; gonostylus moderately heavy; superior volsella stout
(Paggi 1975: Figs. 5, 8); with palmate sensilla chaetica on hind metatarsus.... .
SGOT OEE CL RRC ONCE COCO een Shc aE ene er Teer D. alsinensis (Paggi)
Foretarsal beard absent; gonostylus thin; superior volsella thin (Fig. 13); without pal-
mate sensilla chaetica on hind metatarsus........... D. aethiops (Townes)
Superior volsella with elongate thin basal portion, ‘‘foot’’ narrow; inferior volsella deeply
notched apically, lobes almost completely separated (Paggi 1978: Figs. 3,5)... .
B18 “6, CEO VO "a: Gee SLO Oe DMC non ee COCR ORISTIONG cme ECTS ROP ania nan D. nestori Paggi
Superior volsella with thicker basal portion, ‘‘foot’’ robust; inferior volsella shallowly
notchedvorsimplel(Eigssl3=16)eeesseeeneien ene eiienenicnen iene enna rrr en aneie 7
Mid leg metatarsus very short, SV, > 5.00; inferior volsella simple; wings mostly clear
(RageilO8i Bigs: 2a. (Ga). 2. 2 elo ws Be eee D. pellegriniensis Paggi
Metatarsus normal, SV, < 5.00; inferior volsella simple or slightly notched apically
(Figs. 14-15); wings with faint markings at base of r4,5, at FCu and along M;, 4 and
Cu, PS OGCOTOOECORDOSODSGOODCHGCADFTOCOODOOD OOOO ODDO OOM OD 8
General coloration yellow-brown to brown; legs stramineous-brown, fore femora with
darkened apices... .. . D. californicus (Johannsen) complex (D. californicus,
bee hee eee A ee es eee D. crypticus Epler, D. embalsensis Paggi
General coloration red-brown; legs dark brown, fore femora solid dark brown ....
Pe ese ig aceasta fon ees ey USL ce cate pean eee ee D. obrienorum Epler
Inferior volsella stout, covered with long, dense setae ................. 10
Inferior volsella thin to moderate, without long, dense setae ............. 11
Ventral portion of bifid superior volsella with sensilla chaetica (Fig. 18) (ventral portion
of volsella is often difficult to discern)............ D. dasylabidus sp. nov.
Ventral portion of bifid superior volsella without sensilla chaetica (Fig. 23) (ventral
portion of volsella is often difficult to discern). . . . D. paradasylabidus sp. nov.
Base of anal point with ventral bulbous swelling (Figs. 17, 20, 24, 25)........ 14
Base of anal point without ventral bulbous swelling (Figs. 19, 26, 27)........ 12
Disto-ventral portion of distal arm of inferior volsella with long sensilla chaetica; su-
perior volsella slipper-shaped; posterior margin of T IX strongly cordiform-emar-
ginate at base of anal point (Fig. 27)................ D. soccus sp. nov.
INOtaSHADOV ER errr rete terse er ee ee ele Geee Gr te es ce May ah a ore Reea arate fa fos ay te 13
Superior volsella deltoid-pediform in dorsal view (Fig. 26) ...... D. reissi sp. nov.
Superior volsella elongate-cylindrical with pointed apex and dorsal basal lobe from which
volsella is suspended (Fig. 19).................. D. demissus sp. nov.
Superior volsella bifid (Fig. 25).................. D. radinovskyi sp. nov.
Superior volsellainot bifid! 15.25 ae wo oe oe a Be ee oe ee Se te 15
Superior volsella with acute apex directed caudad (Fig. 17) . .D. amazonicus sp. nov.
Superior volsella deltoid-pediform or pediform (Figs. 21, 24)............. 16
Superior volsella pediform, apex directed laterad (Fig. 24) . . .D. paterjohni sp. nov.
MEM. AMER. ENT. SOC., 36
60 GENUS DICROTENDIPES
Superior volsella deltoid-pediform, apex membranous and ventrally produced (may be
directed mediad or laterad) (Fig. 20)............... D. fittkaui sp. nov.
PREVIOUSLY DESCRIBED SPECIES
Dicrotendipes aethiops (Townes) (Figs. 13, 48)
Tendipes (Limnochironomus) aethiops Townes, 1945:107.
Tendipes (Limnochironomus) figueroai Vargas, 1952:48.
Dicrotendipes aethiops (Townes): Epler 1987a:30.
See Epler (1987a) for complete synonymy and description of adult male,
and Epler (1987b) for additional distribution records. This species occurs
in the southwestern United States and Mexico, and may occur farther south.
Dicrotendipes alsinensis (Paggi)
Chironomus (Dicrotendipes) alsinensis Paggi, 1975:149.
Dicrotendipes alsinensis (Paggi): Paggi 1978:235.
See Paggi (1975:149) for description of adult male, pupa and larva. To
Paggi’s (1975) description I can add the following data: Adult male: 12
cibarial setae; mid leg metatarsus with 10 palmate sensilla chaetica, hind
metatarsus with 3 palmate sensilla chaetica; wing setae: R 18, R, 7, Ry,; 45
no ventral accessory setae on S VI. Larva: 23 ventromental plate strial ridges,
anterior margin of ventromental plate crenulated.
On the basis of superior volsella morphology (digitiform-cylindrical with
a membranous apex), D. alsinensis would appear to be a member of the D.
nervosus group as defined by Epler (1987a). However, Paggi’s (1975:Fig.
12) pupal illustration does not indicate that the median shagreen area on T
VI is V-shaped and the one pupal specimen I examined is damaged; T VI
is not observable. Because the frontal apotome and labral sclerite 1 are not
clearly visible on the larval exuviae associated with this pupal specimen (and
they are not described by Paggi 1975), this species is difficult to place. It
may be related to the modestus group, the nervosus group or D. fumidus.
However, the one male of this species that I examined possesses palmate
sensilla chaetica on the hind metatarsus; these sensilla chaetica are not pres-
ent on any of the members of the aforementioned groups.
MATERIAL EXAMINED: ARGENTINA: Prov. Buenas Aires, Laguna Alsina, 27-III-1975,
1 male (IL); same locality, 27-IV-1975, leg. Paggi, 1 pharate male pupa/Lex (IL).
JOHN H. EPLER 61
Dicrotendipes californicus (Johannsen) (Figs. 14, 47)
Chironomus californicus Johannsen, 1905:217.
See Epler (1987a) for full synonymies and descriptions of adult male, pupa
and larva.
This species is widespread from California, southern Idaho and South
Dakota in the U.S. south to Chile. Four other apparently closely related
species have been described: D. crypticus Epler from New Mexico and Cal-
ifornia (Epler 1987a), D. obrienorum Epler from Mexico (Epler 1987b), and
D. embalsensis Paggi and D. pellegriniensis Paggi from Argentina (Paggi
1987). These 4 species may only represent variants or ecotypes of D. cali-
fornicus. More data are needed to clarify their status. See also discussions
under each species below.
I have examined a single male specimen from the NM (collected at Finca
Richter nr Bogota, Colombia, by Lichtenberg) which may represent a new
species. However, it is quite possible that this specimen is a D. californicus
(or D. obrienorum) with deformed superior volsellae. I would have to see
more specimens of this type before I could justify describing it as a new
species.
There is minor variation in wing maculation over this species’ range. A
specimen from Peru has wing markings which are similar, but darker and
more pronounced, than the typical D. californicus. See also comments under
D. pellegriniensis below.
In addition to the material listed below, material from Costa Rica and
Mexico has been examined (Epler 1987a).
MATERIAL EXAMINED: CHILE: Coquimbo, Punta Teatinos, 16-IX-1952, leg. P.G. Kus-
chel, 1 male (US); Santa Maria Fuadu [?], 28-I-1943, E. Melland, 1 male (BM). COLOMBIA:
Palmira, Lichtfang light trapj, 25-I-1975, leg. Lichtenberg, 1 male (NM); Palmira, Finca Aus-
tria, b. Schwimmbecken, 26-I-1975, leg. Lichtenberg, 1 male (NM). PANAMA: Chiriqui Prov.,
Presa Fortuna, holding pond above Aoki camp, 3900’, light trap, 25-V-1985, leg. R.W. Flow-
ers, 1 male (JE). PERU: Laguna de Medio Mundo, vegetation, brackish water of northern
Peruvian coast, 17-[V-1975, leg. Gloria S. Minaya Gomez, 1 male, 1 larva (ZS).
Dicrotendipes crypticus Epler? (Fig. 47)
Dicrotendipes crypticus Epler, 1987a:39.
I have seen one larval specimen from South America which fits my con-
cept of this species. The specimen possesses a head capsule with grainy in-
tegument, 27 ventromental plate strial ridges and a crenulate anterior margin
MEM. AMER. ENT. SOC., 36
62 GENUS DICROTENDIPES
on the ventromental plate. This specimen may belong with D. embalsensis.
The larvae of these 2 species appear identical, and the 2 species may prove
to be synonyms. The date of publication of both Epler 1987a and Paggi
1987 is March 1987. Although D. crypticus was originally described in my
thesis (Epler 1983), theses are not considered publications by the Interna-
tional Code of Zoological Nomenclature, Article 9 (1985). However, until
more specimens of both species are examined, no decision can be made
concerning their status.
MATERIAL EXAMINED: PARAGUAY: Rio Pilcomayo, 31-III-1974, leg. H. Sioli, 1 larva
(ZS).
Dicrotendipes embalsensis Paggi
Dicrotendipes embalsensis Paggi, 1987:703.
See Paggi (1987:703) for description of adult male, pupa and larva. To
her description I can add the following data: Adult male: 8 cibarial setae;
mid leg metatarsus with 13-14 palmate sensilla chaetica, 0 on hind meta-
tarsus; wing setae; squama 18-21, R 18, R, 4, R,,; 12; no ventral accessory
setae apparent on S VI. Larva: 29-30 ventromental plate strial ridges, an-
terior margin of plate weakly and shallowly crenulated.
The key character of adult male frontal tubercle length used by Paggi
(1987:697) will probably not separate D. embalsensis from D. pellegrinien-
sis. The one specimen of D. embalsensis I examined lacked frontal tubercles.
The length and presence or absence of frontal tubercles is extremely variable
in all species of Dicrotendipes 1 have examined when sufficient material
(usually more than 10 specimens) was available. The same would probably
hold true for squamal setal counts (the one specimen I examined had a lower
squamal setal count, 18, than recorded in the description or used in the
keys). As adults, the 2 species can be separated by the short metatarsus and
apparently clear wings of D. pellegriniensis. The characters used to separate
the larvae of these 2 species in Paggi (1987) will probably also fail to separate
them when more material is examined. Postmental darkening may be en-
vironmentally influenced (it apparently is in many Nearctic species; see Epler
1987a); and there probably is not a significant difference between a pecten
epipharyngis with 5 teeth (D. embalsensis) and one with 6-7 teeth (D. pel-
legriniensis). More specimens of all species of the D. californicus complex
must be examined in order to determine which characters, if any, will sep-
arate the species. It is possible that all of these species are nothing more
than variants or ecophenotypes of D. californicus.
JOHN H. EPLER 63
MATERIAL EXAMINED: ARGENTINA: Prov. Neuquén, Lago Ramos Mexia, 4-X-1983,
leg. Kaisin, 1 male/Pex/Lex (IL).
Dicrotendipes nervosus (Staeger) group (Fig. 45)
I have seen several larval specimens from Brazil which conform to my
definition of the D. nervosus group (cf. Epler 1987a). All of these specimens
possess a pair of ventral tubuli on the eighth abdominal segment. It is pos-
sible that these larvae may belong with one or several of the species described
as new below. Pupae are known for only 2 of these species, D. fittkauii and
D. soccus. None of the pupal specimens examined possessed ventral tubule
remnants on S VIII; these remnants should have been present had the larvae
possessed ventral tubules. The pupae of both of these species possess a V-
shaped median shagreen area on T VI, which would place them in the D.
nervosus group. However, without associated larvae, such placement must
be considered uncertain.
MATERIAL EXAMINED: BRAZIL: Amazonas: Lago do Calado, lower Rio SolimGes, nr
village Manacapuru, experimental box filled with wood wool as artificial substrate, exposed
free floating at border of marginal floating meadows, Autumn 1968, leg. W. Junk, 1 larva
(ZS). Para: Lago Salgado, Cabaciera do boi, zw. Tabacorana-Wurzeln, 13-IV-1948, leg. Braun,
1 larva (ZS); same locality & collector, Aufwuchs auf Canaranca, 16-IV-1948, 1 larva (ZS).
Dicrotendipes nestori Paggi
Dicrotendipes nestori Paggi, 1978:235.
See Paggi (1978:235) for description of adult male, pupa and larva. To
her description I can add the followng data: Adult male: 9 cibarial setae;
mid leg metatarsus with 7 palmate sensilla chaetica, 0 on hind metatarsus;
wing setae: R 19, R, 7, R,,; 12; no ventral accessory setae apparent on S
VI.
I have not examined any larvae or pupae of this species. Because Paggi
(1978) did not describe the immature stages completely, this species can not
be placed phylogenetically.
MATERIAL EXAMINED: ARGENTINA: Prov. Buenas Aires, Laguna Alsina, 27-III-1975,
1 male (paratype) (IL).
Dicrotendipes obrienorum Epler (Figs. 15, 47)
Dicrotendipes obrienorum Epler, 1987b.
MEM. AMER. ENT. SOC., 36
64 GENUS DICROTENDIPES
See description in Epler (1987b). Known only from Mexico, this species
may be a lower latitude but higher elevation variant of D. californicus.
Dicrotendipes pellegriniensis Paggi
Dicrotendipes pellegriniensis Paggi, 1987:695.
See Paggi (1987:695) for description of adult male, pupa and larva. To
her description I can add the following data: 10 cibarial setae; at least 14
palmate sensilla chaetica on mid leg metatarsus (not clearly discernible), 0
palmate sensilla chaetica on hind metatarsus; wing setae: R 16, R, 6, R,,;
9; no ventral accessory setae apparent on S VI. Larva: ventromental plate
with 34 strial ridges, anterior margin of plate smooth.
The larvae and pupae of this species are morphologically inseparable from
D. californicus. The adult male differs from D. californicus in the very short
mid leg metatarsus which results in a very high SV, (6.46 in the one specimen
made available to me), and perhaps in wing maculation. Paggi (1987) de-
scribed the wings of this species as transparent, without spots. Unfortu-
nately, in the one specimen of this species available to me, the mountant
under the cover slip over the wings is unclear and the wings can not be
viewed clearly. However, there appears to be a spot in the apex of cell r,,;.
I have found the amount or intensity of wing maculation to be variable in
Nearctic specimens of D. californicus; thus, this character may not be of
use in species separation within the D. californicus complex. The inferior
volsella of D. pellegriniensis differs from that of typical D. californicus in
that there is no trace of an apical indentation. This character is unreliable,
however, for in many species the amount of apical indentation of the inferior
volsella is variable. It is quite possible that D. pellegriniensis is but an eco-
phenotype of D. californicus. Paggi (1987) does not supply any physico-
chemical data for the collection sites of this species; perhaps an unknown
pollutant could cause morphological changes which may produce a popu-
lation of aberrant individuals, which may then be described as a ‘‘new spe-
cies.’’ More research is needed to solve the complexities of this problem.
MATERIAL EXAMINED: ARGENTINA: Prov. Rio Negro, Lago Pellegrini, 25, 26-III-
1979, leg. Paggi, 1 male, 1 pupa/Lex (IL).
Dicrotendipes sinoposus Epler (Figs. 16, 48)
Dicrotendipes sinoposus Epler, 1987b.
JOHN H. EPLER 65
See description in Epler (1987b). Originally described from Mexico, I have
since examined more material from the Neotropics. These additional spec-
imens enable me to add the following data to the original description: AR
2.22-2.70; 7-9 palmate sensilla chaetica on metatarsus of middle leg; 0-4
ventral accessory setae on S VI.
MATERIAL EXAMINED: BRAZIL: [Parad]: Rio Tocantins, nr village Jatobal, light trap,
5-XI-1960, leg. E.J. Fittkau, 1 male (ZS); Amazonas: Lago Cabaliana, lower Rio SolimGes,
drift, 16-VI-1971, leg. F. Reiss, 1 male (ZS). COLOMBIA: Arbeldez, light trap, 26-XI-1974,
leg. Lichtenberg, 1 male; same locality & collector, 30-XI-1974, 1 male (NM). DOMINICA:
Pont Casse, 1.5 mi. N, rain forest, 12 Feb 1965, leg. W.W. Wirth, Bredin-Archbold-Smith-
sonian Biol. Survey Dominica, 2 males (US).
NEw SPECIES DESCRIPTIONS
Dicrotendipes amazonicus sp. nov. (Fig. 17)
TYPE LOCALITY: Rio Tupani, lower Rio Madeira, Amazonas, Brazil.
TYPE MATERIAL: Holotype: male, BRAZIL: Amazonas: Rio Tupani, lower Rio Madeira,
light trap, 14-15-IX-1960, leg. E.J. Fittkau (ZS). Paratypes (2572): same data as holotype, 2553
males, 13 females (ZS); lower Rio Madeira, Parana Madeirinha, light trap, 12-IX-1960, leg.
E.J. Fittkau, 4 males (ZS); Rio Tupani (wide, lake-like), light trap, 21-I[X-1960, leg. E.J. Fitt-
kau, 3 males (ZS); Upper Rio Tapajos, slightly downstream from mouth of Rio Juruena &
Rio Sao Manuel, light trap at village Barra, 13-I-1961, leg. E.J. Fittkau, 1 male (ZS). Holotype
to be deposited in IN; paratypes in ZS, JE.
DIAGNOSIS: The immaculate wings, deeply bifid inferior volsella, dis-
tinctive superior volsella and bulbous swelling beneath the anal point will
distinguish this species.
ETYMOLOGY. This species is named for the Amazon region.
MALE IMAGO (n=6)
COLOR (slide mounted specimens). Head, body and legs light brown. Wing immaculate,
slightly dusky brown, with light brown veins.
LENGTH. Total 3.20-3.65, 3.33 mm. Thorax 0.72-0.82, 0.78 mm. Abdomen 2.43-2.85,
2.56 mm.
HEAD. Setae: temporal 22-32, 27; clypeal 10-14, 12; cibarial 7-8, 8 (4). Palpomere lengths:
27-40, 35; 35-40, 38; 70-88, 79; 106-113, 111; 148-180, 170 (4). Frontal tubercles 7-20, 12
long, 6-7, 7 wide (5). AR 2.03-2.15, 2.09 (3).
THORAX. Scutal tubercle well developed; humeral pit with 5-7 small tubercles. Acrostichals
6-9, 7; dorsocentrals 13-17, 15 (5); scutellars 7-8, 7; prealars 6-8, 7 (5).
WING. Length 1.28-1.40, 1.36 mm (5); width 0.39-0.41, 0.40 mm (5). FCu distal to RM.
VR 0.85-0.88, 0.86 (5). Setae: brachiolum 2; squama 1-2, 2; R 12-16, 14; R; 5-8, 7; R34,5 9-
14, 12.
LEGS. Foretarsal beard absent. Palmate sensilla chaetica: 5-9, 7 on middle metatarsus, 2-
4, 3 on hind metatarsus. Lengths and proportions of legs:
MEM. AMER. ENT. SOC., 36
66 GENUS DICROTENDIPES
P, P, P;
fe 745-850, 620-700, 680-790,
789 655 727
ti 455-530, 540-610, 700-800
486 577 743
ta, 1050-1170, 310-365, 460-530,
1109 (5) 338 497
ta) 490-550, 140-170, 225-265,
519 (5) 160 247
ta; 420-490, 100-120, 180-210,
456 (5) 112 197
tay 305-420, 45-50, 90-105,
388 (5) 49 (5) 98
tas 150-175, 50-55, 65-80,
168 (5) 51 (5) 76 (5)
LR DIDDLG, 0.57-0.61, 0.66-0.69,
2.26 (5) 0.59 0.67
BV 1.51-1.62, 4.09-4.41, 3.14-3.29,
1.57 (5) 4.26 (5) 3.19 (5)
SV 1.14-1.18, 3.55-3.76, 2.88-3.00,
1.16 (5) 3.64 2.96
ABDOMEN. 1-4 ventral accessory setae on S VI; 1 specimen seen with 1 ventral accessory
seta on S V.
HYPOPYGIUM (Figs. 17A, B) with 2-5, 4 medial setae. Gonostylus normal, curved me-
dially, with 4-7, 6 preapical setae. Superior volsella (Figs. 17C, D) length 50-72, 61 (4); width
27-30, 29 (4); LWR 1.9-2.6, 2.2 (4); slipper-shaped in dorsal view; with 4-S, 5 sensilla chaetica
and fine setae on medial surface. Inferior volsella length 105-130, 117 (4); deeply bifid, with
3-5 sensilla chaetica in single row on proximal lobe, 4-5 sensilla chaetica in single row on
distal lobe; distal lobe with 1 well developed ventral preapical seta. Anal point bare dorsally,
with basal peduncle and bulbous ventral extension, weakly deflexed; with 1-2 dorsal basal
setae and 6-13, 10 lateral basal setae.
FEMALE IMAGO (n = 3)
COLOR (slide mounted specimens). Similar to male.
LENGTH. Total 2.23-3.04, 2.56 mm. Thorax 0.74-0.81, 0.78 mm. Abdomen 1.42-2.30,
1.77 mm.
HEAD. Setae: temporal 22-23, 23; clypeal 11-14, 12; cibarial 7-10, 9. Palpomere lengths:
30-35, 33; 40-42, 41; 80-85, 83; 112-115, 115; 165-175, 170. Frontal tubercles 9-18, 12 long,
5-10, 7 wide. AR 0.47-0.55, 0.50.
THORAX. Scutal tubercle well developed; humeral pit with about 4 small tubercles. Ac-
rostichals 9-10 (2); dorsocentrals 22-29, 25; scutellars 7-9, 7; prealars 6-8, 7.
WING. Length 1.49-1.55, 1.53 mm; width 0.47-0.51, 0.50 mm. FCu distal to RM. VR 0.82-
0.89, 0.87. Setae: brachiolum 2; squama 1-4, 2; R 16-18, 17; R, 11-16, 13; R4,5 27-32, 29.
LEGS. Foretarsal beard absent. Palmate sensilla chaetica: 15-20, 17 on middle metatarsus,
5-7, 6 on hind metatarsus. Lengths and proportions of legs:
JOHN H. EPLER 67
P, P> P;
fe 790-800, 660-695, 745-790,
793 675 762
ti 500-520, 595-620, 745-790,
510 606 778
ta, 1125-1240 355-370, 470-520,
(2) 362 493
tay 530-570 150-160, 225-250,
(2) 157 233
ta; 450-510 105-110 180-195,
(2) 107 188
ta, 430-560 45-50, 90
(2) 48
tas 190 50 70-75,
(2) 72
LR 2.25-2.43 0.60 0.62-0.65,
(2) 0.63
BV 1.47-1.51 4.44-4.64, 3.43-3.58,
(2) 4.54 3.49
SV 1.05-1.15 3.51-3.56, 3.06-3.19,
(2) 3.54 3.12
ABDOMEN. 2-4 ventral accessory setae on S VI. Notum 148-157, 153; cerci 72-90, 82. S
VIII with 11-15 setae/side; X with 4-6, 5 setae; Gc IX with 1-2, 2 setae/side. DmL, VIL and
ApL as in Fig. 17E.
Dicrotendipes dasylabidus sp. nov. (Fig. 18)
TYPE LOCALITY: Brazil, Amazonas, Upper Rio Solim6es, Florianopolis.
TYPE MATERIAL: Holotype: male, BRAZIL: Amazonas: Upper Rio Solimoes, Floriano-
polis, light trap, 15-VIII-1961, leg. E.J. Fittkau (ZS). Holotype to be deposited in IN.
DIAGNOSIS: The immaculate wings, stout, deeply bifid and densely se-
tose inferior volsella, distinctive bifid superior volsella and cordiform-emar-
ginate base of the anal point will distinguish this species.
ETYMOLOGY. From the Greek dasys, hairy and /abidos, forceps; refers
to the densely setose inferior volsella.
MALE IMAGO (n= 1)
COLOR (slide mounted specimens). Head, body and legs light brown. Wing immaculate,
slightly dusky brown, with light brown veins.
LENGTH. Total 3.64 mm. Thorax 0.91 mm. Abdomen 2.73 mm.
HEAD. Setae: temporal 33; clypeal 11; cibarial 9. Palpomere lengths: 41, 47; 93; 115; 190.
Frontal tubercles 6 long, 6 wide. AR 2.08.
MEM. AMER. ENT. SOC., 36
68 GENUS DICROTENDIPES
THORAX. Scutal tubercle well developed; humeral pit indiscernible. Acrostichals 9; dor-
socentrals 14; scutellars 10; prealars 8.
WING. Length 1.58 mm; width 0.47 mm. FCu distal to RM. VR 0.83. Setae: brachiolum
2; squama 2; R 15; Ry; 6; R4,5 11.
LEGS. Foretarsal beard absent. Palmate sensilla chaetica: 8 on middle metatarsus, 5 on hind
metatarsus. Lengths and proportions of legs:
P, P, P;
fe 855 710 830
ti 550 655 860
ta 1130 390 550
tay 530 190 280
ta; 450 135 220
tay 400 50 100
tas 190 55 70
LR 2.05 0.60 0.64
BV 1.61 4.08 3.34
SV 1.24 3.50 3.07
ABDOMEN. Ventral accessory setae on S VI not apparent.
HYPOPYGIUM (Fig. 18A) with 4 medial setae. Gonostylus normal, curved medially, with
4 preapical setae. Superior volsella (Fig. 18B) bifid; length of dorsal portion 54, ventral portion
90; width of dorsal portion 11, ventral portion 30; dorsal portion small, digitiform, sclerotized,
with 2 sensilla chaetica; ventral portion thin, lamellar, densely setose, with 2-4 sensilla chaetica.
Inferior volsella length 135; deeply bifid, stout, with 4 sensilla chaetica each on proximal and
distal lobes, densely setose on inner margin, with 1 well developed ventral preapical seta. Anal
point cordiform-emarginate basally, bare dorsally; with 14 dorsal basal setae and 9 lateral basal
setae.
Dicrotendipes demissus sp. nov. (Fig. 19)
TYPE LOCALITY: Brazil, Amazonas, Lago do Rei.
TYPE MATERIAL: Holotype: male, BRAZIL: Amazonas: Lago do Rei, Abends an bord
bei Licht, 28-[X-1959, leg. Sioli-Settler (ZS). Paratypes (152): 132 males, 20 females, same data
as holotype (ZS). Holotype to be deposited in IN; paratypes in ZS, JE.
DIAGNOSIS: The immaculate wings, deeply bifid inferior volsella and
distinctive superior volsella will distinguish the male of this species. The
immature stages are unknown.
ETYMOLOGY. From the Latin demissus, hanging down, drooping; re-
fers to the superior volsella when viewed laterally.
MALE IMAGO (n=7)
COLOR (slide mounted specimens). Head, body and legs light brown. Wing immaculate,
slightly dusky brown, with yellow-brown veins.
LENGTH. Total 3.73-4.10, 3.85 (4) mm. Thorax 0.97-1.01, 0.99 (4) mm. Abdomen 2.73-
3.15, 2.90 (6) mm.
JOHN H. EPLER 69
HEAD. Setae: temporal 31-37, 36 (4); clypeal 15-21, 18 (6); cibarial 7-10, 9 (4). Palpomere
lengths: 33-40, 38; 40-52, 46; 85-108, 97; 100-118, 111; 180-198, 189 (6). Frontal tubercles
20-28, 24 long, 8-10, 9 wide (5). AR 1.87-2.18, 1.96 (6).
THORAX. Scutal tubercle well developed; humeral pit 3-12 small to moderate tubercles.
Acrostichals 5-12, 8; dorsocentrals 14-20, 18; scutellars 8-10, 9 (6); prealars 9-12, 10.
WING. Length 1.55-1.68, 1.63 mm (6); width 0.46-0.50, 0.48 (6) mm. FCu distal to RM.
VR 0.85-0.87, 0.88 (5). Setae: brachiolum 2-3, 2; squama 2-3, 2; R 13-18, 15; R, 5-10, 8;
Ry4,5 6-13, 10.
LEGS. Foretarsal beard absent. Palmate sensilla chaetica: 10-13, 11 on middle metatarsus,
7-11, 9 on hind metatarsus. Lengths and proportions of legs:
P, P, P3
fe 880-960, 760-790, 855-890,
907 78 874
ti 580-625, 670-695, 870-910,
601 685 900
ta; 1070-1100 380-410, 580-610,
(2) 401 589
tay 510-520 190-210, 295-310,
(2) 203 304
ta; 440-450 135-155, 234-245,
(2) 145 240
tag 390-395 55-60, 120-130,
(2) 59 124
tas 180 60-70, 90-100,
(2) 65 96
LR 1.75-1.82 0.57-0.60, 0.64-0.68,
(2) 0.59 0.66
BV 1.69-1.70 3.78-4.10, 3.01-3.18,
2) 3.94 3.10
SV 1.37-1.42 3.52-3.79, 2.92-3.10,
(2) 3.63 3.01
ABDOMEN. 5 ventral accessory setae on S VI.
HYPOPYGIUM (Figs. 19A, B) with 4-8, 6 medial setae. Gonostylus apically expanded,
curved medially, with 5-6 preapical setae. Superior volsella (Fig. 19C) length 78-95, 85 (5);
width 13-18, 16 (5); LWR 4.9-6.2, 5.5; elongate-cylindrical with pointed apex and a dorsal
basal lobe from which volsella is suspended; dorsal portion with 2-3 sensilla chaetica, elongate
suspended portion with 1 sensilla chaetica. Inferior volsella length 140-163, 153 (6); deeply
bifid, with 2-4 sensilla chaetica in single row on proximal lobe, 2-5 sensilla chaetica in single
row on distal lobe; distal lobe with 1 well developed ventral preapical seta. Anal point bare
dorsally; with 3-6, 4 dorsal basal setae and 4-8, 6 lateral basal setae.
FEMALE IMAGO (n =3)
COLOR (slide mounted specimens). Similar to male.
LENGTH. Total 3.28-3.84, 3.58mm. Thorax 1.08-2.63, 1.18 mm. Abdomen 2.20-2.58, 2.40
mm.
MEM. AMER. ENT. SOC., 36
70 GENUS DICROTENDIPES
HEAD. Setae: temporal 31-41, 36; clypeal 27-33, 30; cibarial 5-10, 8. Palpomere lengths:
40-48, 44; 48-55, 52; 85-108, 97; 100-118, 111; 180-198, 189 (6). Frontal tubercles 12-15, 13
long, 11-17, 14 wide. AR 0.46-0.56 (2).
THORAX. Scutal tubercle well developed; humeral pit with 1 small tubercle. Acrostichals
11-14, 12; dorsocentrals 27-38, 31; scutellars 12-14, 13; prealars 10-12, 11.
WING. Length 1.76-1.94; 1.88 mm; width 0.60-0.68, 0.64 mm. FCu distal to RM. VR 0.83-
0.85, 0.84. Setae: brachiolum 2-3, 2; squama 6-9, 7; R 24-31, 26; R, 15-18, 16; R4+5 23-28,
26.
LEGS. Palmate sensilla chaetica: 18-21, 20 on middle metatarsus, 16-18, 17 on hind me-
tatarsus. Lengths and proportions of legs:
P, P, P;
fe 910-1015, 780-960, 910-1000,
975 867 970
ti 630-705, 715-885, 940-1035,
677 787 1002
tay = 430-465, 580-660,
448 627
tay = 205-215, 295-320,
208 308
tay a 140-150, 240-260,
147 250
tay = 70 120-140,
132
tas = 80 95-105,
102
LR = 0.53-0.60, 0.62-0.64,
0.57 0.63
BV = 3.89-4.49, 3.24-3.31,
4.16 3.28
SV ae 3.48-3.97, 3.08-3.19,
3.68 3.15
ABDOMEN. 2-8 ventral accessory setae on S VI. Notum 190-193, 191; cerci 110-130, 119.
S VIII with 25-28, 27 setae/side; X with 8-9, 8 setae; Gc IX with 2-3, 3 setae/side. DmL,
VIL and ApL as in Fig. 19D.
Dicrotendipes fittkaui sp. nov. (Figs. 20, 21)
TYPE LOCALITY: Brazil, Reserva Duke nr Manaus.
TYPE MATERIAL: Holotype: male/Pex, BRAZIL, Amazonas: Reserva Duke, nr Manaus,
reared from a ground water puddle in the forest, 9-10-V-1961, leg. E.J. Fittkau (ZS). Paratypes
(9): Brazil, Amazonas, Lago do Calado, from vegetation in central part of floating meadow,
10-VIII-1968, leg. W. Junk, 1 pharate male pupa (ZS); Lago do Calado, lower Rio Solimoes,
I a I I LS
JOHN H. EPLER 71
nr village Manacapuru, experimental box filled with wood wool as artificial substrate, exposed
free floating at border of marginal floating meadows, Autumn 1968, leg. W. Junk, 1 pharate
male pupa (ZS); Lago Cabaliana, lower Rio SolimG6es, drift, leg. F. Reiss, 1 male (ZS). Para:
Alemquer, no porto, as 20 horas, na luz, 15-VII-1946, leg. Sioli, 4 males (ZS); Rio Cururt,
Missao Cururu, right tributary of Rio Tapajos, light trap, 16-I-1961, leg. E.J. Fittkau, 2 males
(ZS); same locality & collector, 19-I-1961, 1 male (ZS). Holotype to be deposited in IN; par-
atypes in ZS, JE.
DIAGNOSIS: The immaculate wings, deeply bifid inferior volsella, dis-
tinctive superior volsella, clavate gonostylus and bulbous swelling beneath
the anal point will distinguish this species. The female and larva are
unknown.
ETYMOLOGY. I take great pleasure in naming this species in honor of
Prof. Dr. E.J. Fittkau.
MALE IMAGO (n=6)
COLOR (slide mounted specimens). Head, body and legs light brown. Wing immaculate,
slightly dusky brown, with yellow-brown veins.
LENGTH. Total 3.13-3.45, 3.32 (4) mm. Thorax 0.78-0.96, 0.85 (4) mm. Abdomen 2.35-
2.65, 2.46 (5) mm.
P, P, P;
fe 820-850, 700-745, 770-810,
835 (4) 721 (5) 796 (4)
ti 520-560, 630-665, 810-830,
532 (5) 646 (5) 820 (2)
ta 1090-1140 350-360, 530-540,
(2) 354 (5) 533 (3)
ta) 480-550 170-190, 250-270,
(2) 179 (5) 260 (3)
ta; 400-485 115-130, 190-210,
(2) 122 (5) 202 (3)
ta, 355-430 55-60, 100-110,
(2) 57 (5) 105 (3)
tas 160-175 55-60 80-85,
(2) 58 (5) 82 (3)
LR 2.06-2.19 0.54-0.57, 0.64-0.66,
(2) 0.55 (5) 0.65 (3)
BV 1.53-1.76 4.02-4.22, 3.22-3.39,
(2) 4.14 (5) 3.31 (3)
SV 1.20-1.25 3.71-3.92 3.00-3.08,
(2) 3.86 (5) 3.03 (3)
HEAD. Setae: temporal 29-35, 32 (4); clypeal 8-13, 12; cibarial 2-11, 8 (4). Palpomere
lengths: 30-50, 37 (4); 38-45, 42 (4); 70-95, 81 (4); 105-115, 108 (3); 175-178, 177 (3). Frontal
tubercles 8-20, 13 long, 5-8, 7 wide. AR 1.84-2.08, 1.94 (5).
MEM. AMER. ENT. SOC., 36
72 GENUS DICROTENDIPES
THORAX. Scutal tubercle well developed; humeral pit 1-3 moderate tubercles. Acrostichals
6-11, 8 (5); dorsocentrals 15-20, 17 (4); scutellars 8-11, 9 (4); prealars 7-10, 9 (4).
WING. Length 1.30-1.55, 1.47 mm (5); width 0.37-0.43, 0.41 mm (5). FCu distal to RM.
VR 0.86-0.89, 0.88 (5). Setae: brachiolum 2-3, 2; squama 1-3, 2 (5); R 14-18, 16 (5); R, 7-
9, 8 (5); R445 7-15, 12 (5).
LEGS. Foretarsal beard absent. Palmate sensilla chaetica: 7-13, 10 (5) on middle metatarsus,
3-7, 5 (3) on hind metatarsus. Lengths and proportions of legs on p. 71.
ABDOMEN. Ventral accessory setae on S VI not apparent.
HYPOPYGIUM (Figs. 20A, B) with 3-7, 5 medial setae. Gonostylus apically expanded,
curved medially, with 6-10 preapical setae. Superior volsella (Fig. 20C) length 45-58, 53 (3);
width 33-43, 37 (3); LWR 1.3-1.7, 1.4 (3); semi-pediform, with ventral subapical lobe; with
dorsal and ventral fine setae; with 1-4 small sensilla chaetica. Inferior volsella length 125-154,
142 (5); deeply bifid, with 3-5 sensilla chaetica in single row on proximal lobe, 5-6 sensilla
chaetica in single row on distal lobe; distal lobe with 1 well developed ventral preapical seta.
Anal point bare dorsally, with basal peduncle and bulbous ventral extension, deflexed; with
0-2 dorsal basal setae and 8-14, 10 lateral basal setae.
PUPA (n= 3)
COLOR. Clear with light brown margins.
LENGTH. Total 3.40 mm (1). Cephalothorax 0.42 mm (1). Abdomen 2.98-3.40, 3.20 mm.
CEPHALOTHORAX. Cephalic tubercles (Fig. 21A) 40-50 (2) high, 68-83 (2) wide. Dorsum
moderately smoothly pebbled. Dc, closer to Dc3. Thoracic horn base (Fig. 21B) with tracheal
bundles separate.
ABDOMEN (Fig. 21C). Sternites I-IV with fine lateral shagreen bands; S V-VI with posterior
areas of fine shagreen. Tergite I sometimes with small scattered posterolateral spinules; T II
with T-shaped shagreen area, spinules largest posteriorly; T III-V with median quadrilateral
shagreen area; T VI with triangular shagreen area, spinules largest mesally; T VII with an
anterior pair of small ovoid shagreen areas; T VIII with a pair of longitudinal fine shagreen
bands. Tergites IV and V with a posterior band of fine spinules; T V-VI with posterolateral
group of fine spines. Posterior margin of T II with transverse row of 65-72, 68 hooklets. T
VIII with 4 lateral lamellar setae. Caudolateral spurs on T VIII (Figs. 21D, E) single or double,
moderately large. Anal lobes with 39-52, 46 setae. DR 1.69-2.09, 1.93.
Dicrotendipes palearivillosus sp. nov. (Fig. 22)
TYPE LOCALITY: Costa Rica, San Jose Queb. Muerte.
TYPE MATERIAL: Holotype: male, COSTA RICA: San Jose Queb. Muerte, route 2, 3.5
km (air) W Villa Mills, 9.652 N, 83.743 W, 12.vi.1986, el. 3120 m, Holzenthal, Heyn, Armitage
(CU). Paratypes (2): same data as holotype, 1 male (CU); Heredia, Est. Biol. La Selva, Rio
Puerto Viejo, 10.440 N, 84.012 W, 19-VI-1986, el. 30 m, Holzenthal, Heyn, Armitage, 1 male
(CU). The holotype will be deposited in the US, one paratype in CU and the other in JE.
DIAGNOSIS: The immaculate wings, distinctive digitiform superior vol-
sella with small membranous ventral basal lobe and simple inferior volsella
will distinguish this species. The female and immature stages are unknown.
ETYMOLOGY. From the Latin palear, a flap or wattle, and villosus,
hairy; refers to the basal lobe of the superior volsella.
JOHN H. EPLER 73
MALE IMAGO (n =2)
COLOR (slide mounted specimens). Head and body light yellow-brown (probably green in
life); fore femora greenish-stramineous, apex of femur and remainder of leg light brown, mid
and hind legs greenish-stramineous, apical tarsomeres darker. Wing immaculate, slightly dusky
brown, with light yellow-brown veins.
LENGTH. Total 3.38-3.73 mm. Thorax 0.88-0.90 mm. Abdomen 2.50-2.83 mm.
HEAD. Setae: temporal 33-35; clypeal 18-20; cibarial 5-6. Palpomere lengths (1): 43; 43;
113; 143; 210. Frontal tubercles 2-4 long, 2-3 wide. AR 2.06-2.07.
THORAX. Scutal tubercle well developed; humeral pit indiscernible. Acrostichals 5-6; dor-
socentrals, 15-17; scutellars 7-10; prealars 8-9.
WING. Length 1.70-1.73 mm; width 0.48-0.50 mm. FCu distal to RM. VR 0.87-0.89. Setae:
brachiolum 2-3; squama 4-5; R 19; R,; 8-11; R4+5 15-22.
LEGS. Foretarsal beard absent. Palmate sensilla chaetica: 4-8 on middle metatarsus, 0 on
hind metatarsus. Lengths and proportions of legs:
P, P, P;
fe 880-930 760-790 850-885
ti 760-680 700-710 920-940
tay 1190-1290 380-390 630-645
ta 530-540 200-210 295-310
ta; 455-460 140 230
tay 400-410 70 120-130
tas 175-185 55-60 75-90
LR 1.78-1.90 0.54-0.55 0.68-0.69
BV 1.76-1.82 3.94-3.96 Be 25-3555
SV 1.25-1.30 3.84-3.85 2.81-2.83
ABDOMEN. 3-5 ventral accessory setae on S VI; one specimen with 1 ventral accessory
seta on S VII.
HYPOPYGIUM (Fig. 22A) Gonostylus moderate, curved medially, with 4-5 preapical setae.
Superior volsella (Fig. 22B) length 67-68, width 16-18; digitiform with bare, apparently slightly
membranous apex and with basal membranous ventral lobe or flap covered with long, fine
setae; with 4-6 ventromedial sensilla chaetica. Inferior volsella length 110-113; simply clubbed;
with 3-4 dorsal sensilla chaetica in 2 rows, with 1 well developed ventral preapical seta. Anal
point bare dorsally, on small peduncle, slightly deflexed; with 3-4 dorsal basal setae and 8
lateral basal setae.
Dicrotendipes paradasylabidus sp. nov. (Fig. 23)
TYPE LOCALITY: Brazil, Amazonas, upper Rio SolimGes at mouth of Rio Takana.
TYPE MATERIAL: Holotype: male, BRAZIL: Amazonas: upper Rio Solimoes at the mouth
of Rio Takana (west of Rio Ica), light trap, 15-VIJII-1961, leg. E.J. Fittkau (ZS). Holotype to
be deposited in IN.
MEM. AMER. ENT. SOC., 36
74 GENUS DICROTENDIPES
DIAGNOSIS: The immaculate wings, stout, deeply bifid and densely
setose inferior volsella, distinctive bifid superior volsella and cordiform-
emarginate base of the anal point will distinguish this species. Its small size
and lack of sensilla chaetica on the ventral portion of the bifid superior
volsella will distinguish this species from D. dasylabidus.
ETYMOLOGY. From the Greek para, near, dasys, hairy and /abidos,
forceps; refers to the inferior volsella and the apparent close relationship
between this species and D. dasylabidus.
MALE IMAGO (n=1)
COLOR (slide mounted specimen). Head, body and legs light brown. Wing immaculate,
slightly dusky brown, with light brown veins.
LENGTH. Total 2.95 mm. Thorax 0.75 mm. Abdomen 2.20 mm.
HEAD. Setae: temporal about 30; clypeal 11; cibarial indiscernible. Palpomere lengths: 35;
35; 75; 102; 177. Frontal tubercles 5 long, 6 wide. AR 1.79.
THORAX. Scutal tubercle well developed; humeral pit indiscernible. Acrostichals 8;
dorsocentrals 18; scutellars 8; prealars 10.
WING. Length 1.35 mm; width 0.39 mm. FCu distal to RM. VR 0.86. Setae: brachiolum
2; squama 1; R 14; R, 5; R4+; 7.
LEGS. Foretarsi missing. Palmate sensilla chaetica: 7 on middle metatarsus, 2 on hind
metatarsus. Lengths and proportions of legs:
P, P, P;
fe 740 620 700
ti 470 545 720
ta, ae 340 465
ta) = 160 230
ta; = 110 180
ta, 2 45 90
tas = 50 70
LR = 0.62 0.65
BV = 4.12 33
SV = 3.43 3.05
ABDOMEN. 2 ventral accessory setae on S VI.
HYPOPYGIUM (Fig. 23A) with 17 dorsomedial setae. Gonostylus thin, slightly curved
medially, with 3 preapical setae. Superior volsella (Fig. 23B) bifid; length of dorsal portion
42, ventral portion 49; width of dorsal portion 11, ventral portion 25; dorsal portion small,
digitiform, sclerotized, with 2 sensilla chaetica; ventral portion thin, lamellar, densely setose,
without sensilla chaetica. Inferior volsella length 133; deeply bifid, stout, with 4 sensilla chaetica
each on proximal and distal lobes, densely setose on inner margin, with 2 well developed ventral
preapical setae. Anal point cordiform-emarginate basally, bare dorsally; with 6-8 lateral basal
setae.
Dicrotendipes paterjohni sp. nov. (Fig. 24)
TYPE LOCALITY: Brazil, Amazonas, lower Rio Preto da Eva.
JOHN H. EPLER 75
TYPE MATERIAL: Holotype: male, BRAZIL: Amazonas: lower Rio Preto da Eva, left
tributary of upper Rio Amazonas, village Tiririca, light trap, {no date}, leg. E.J. Fittkau (ZS).
Holotype to be deposited in IN.
DIAGNOSIS: The immaculate wings, deeply bifid inferior volsella,
distinctive pediform superior volsella and bulbous swelling beneath the anal
point will distinguish this species.
ETYMOLOGY. From the Latin father, pater. I take pleasure in naming
this species for ‘‘Father John’’ Kramer, whose assistance, both direct and
indirect, has helped make this study possible.
MALE IMAGO (n= 1)
COLOR (slide mounted specimen). Head, body and legs light brown. Wing immaculate,
slightly dusky brown, with light brown veins.
LENGTH. Total 3.28 mm. Thorax 0.80 mm. Abdomen 2.73 mm.
HEAD. Setae: temporal indiscernible; clypeal 14; cibarial indiscernible. Palpomere lengths:
35; 40; 90; 117; 178. Frontal tubercles 10 long, 7 wide. AR 2.14.
THORAX. Scutal tubercle well developed; humeral pit indiscernible. Acrostichals 8;
dorsocentrals 15; scutellars 8; prealars 9.
WING. Length 1.47 mm; width 0.40 mm. FCu distal to RM. VR 0.88. Setae: brachiolum
2; squama 2; R 16; Ry; 6; R4+; 9.
LEGS. Foretarsal beard absent. Palmate sensilla chaetica: 8 on middle metatarsus, 3 on hind
metatarsus. Lengths and proportions of legs:
P, P, P;
fe 865 710 780
ti 510 625 800
ta, 1180 360 510
ta 550 165 265
ta; 475 110 205
tay 415 50 105
tas 170 50 75
LR 2.31 0.58 0.64
BV 1.59 4.52 3.22
SV 1.17 3.71 3.10
ABDOMEN. Ventral accessory setae on S VI not apparent.
HYPOPYGIUM (Fig. 24A) with 2 medial setae. Gonostylus apically expanded, slightly curved
medially, with 6 preapical setae. Superior volsella (Fig. 24B) length 62; width 40; LWR 1.6;
pediform, with 5 sensilla chaetica. Inferior volsella length 130; deeply bifid, with 4 sensilla
chaetica each on proximal and distal lobes, with 1-2 well developed ventral preapical setae.
Anal point bare dorsally; with basal peduncle and bulbous ventral extension; with 2 dorsal
basal setae and 6-9 lateral basal setae.
MEM. AMER. ENT. SOC., 36
76 GENUS DICROTENDIPES
Dicrotendipes radinovskyi sp. nov. (Fig. 25)
TYPE LOCALITY: Brazil, Amazonas, lower Rio Preto da Eva.
TYPE MATERIAL: Holotype: male, BRAZIL: Amazonas: lower Rio Preto da Eva, left
tributary of upper Rio Amazonas, village Tiririca, light trap, tno date], leg. E.J. Fittkau (ZS).
Paratypes (2): same data as holotype, 2 males (ZS). Holotype to be deposited in IN, paratypes
in ZS.
DIAGNOSIS: The immaculate wings, deeply bifid inferior volsella,
distinctive bifid superior volsella and bulbous swelling beneath the anal point
will distinguish this species.
ETYMOLOGY. I take great pleasure in naming this species for my
undergraduate mentor, Dr. Syd Radinovsky.
MALE IMAGO (n= 3)
COLOR (slide mounted specimens). Head, body and legs light brown. Wing immaculate,
dusky brown, with brown veins.
LENGTH. Total 2.91-3.28, 3.07 mm. Thorax 0.73-0.80, 0.76 mm. Abdomen 2.18-2.48,
2.31 mm. :
HEAD. Setae: temporal 29-34, 31; clypeal 12-14, 13; cibarial 6 (1). Palpomere lengths: 32-
35, 33; 38-47, 43; 74-98, 89; 98-123, 114; 155-183, 168. Frontal tubercles 8-13, 11 long, 7-
8, 8 wide. AR 1.85-1.92, 1.88.
THORAX. Scutal tubercle well developed; humeral pit with 6 medium tubercles. Acrosti-
chals 6-7 (2); dorsocentrals 12-18, 15; scutellars 7-9, 8; prealars 8.
WING. Length 1.48-1.58, 1.52 mm; width 0.41-0.44, 0.43 mm. FCu distal to RM. VR 0.86-
0.89, 0.88. Setae: brachiolum 2; squama 0(?)-4, 2; R 13-18, 15; R, 4-8, 6; R4+5 10-13, 12.
LEGS. Foretarsal beard absent. Palmate sensilla chaetica: 7-13, 9 on middle metatarsus, 3-
4, 4 on hind metatarsus. Lengths and proportions of legs, p. 77.
ABDOMEN. 4-6 ventral accessory setae on S VI.
HYPOPYGIUM (Fig. 25A) with 2-3 medial setae. Gonostylus apically expanded, slightly
curved medially, with 6 preapical setae. Superior volsella (Figs. 25B, C, D) bifid; length of
dorsal portion 43-55, 51, ventral portion 53-80, 66; width of dorsal portion 15-21, 18, ventral
portion 29-43, 36; dorsal portion digitiform with 2 sensilla chaetica, ventral portion expanded,
with 3 sensilla chaetica. Inferior volsella length 138-151, 143; deeply bifid, with 4-5 sensilla
chaetica on proximal lobe, 5-6 sensilla chaetica on distal lobe, with 2 well developed ventral
preapical setae. Anal point bare dorsally; with triangular basal peduncle and bulbous ventral
extension; deflexed; with 1-2 dorsal basal setae and 8-11 lateral basal setae.
Dicrotendipes reissi sp. nov. (Fig. 26)
TYPE LOCALITY: Paranda Madeirinha, lower Rio Madeira, Amazonas, Brazil.
TYPE MATERIAL: Holotype: male, BRAZIL: Amazonas, lower Rio Madeira, Parana
Madeirinha, light trap, 12-[X-1960, leg. E.J. Fittkau (ZS). Paratypes (212): same data as hol-
otype, 202 males (ZS); Amazonas: Ilha do Careiro, upper Rio Amazonas, nr Manaus, Parana
JOHN H. EPLER TT
P, P, P;
fe 810-900, 690-750, 710-865,
847 717 808
ti 480-530, 600-675, 780-880,
500 637 822
ta, 1110 (1) 340-390, 490-560,
360 522
ta 510 (1) 160-185, 245-280,
168 258
ta; 440 (1) 105-110, 185-215,
108 197
ta, 385 (1) 40-50, 47 85-105, 93
tas 185 (1) 50-55, 53 70-85, 78
LR 2.31 (1) 0.55-0.58, 0.61-0.66,
0.57 0.64
BV 1.58 (1) 4.52-4.59, 3.36-3.50,
4.55 3.44
SV 1.16 (1) 3.65-3.84, 3.01-3.26,
3.76 3.13
da Terra Nova at Careiro, ca. 2 km from mouth, light trap, 15-III-1961, leg. E.J. Fittkau, 1
male (ZS); lower Rio Preta da Eva, left tributary of upper Rio Amazonas, village Tiririca,
light trap, tno date}, leg. E.J. Fittkau, 7 males (ZS). Pard: Rio Cururti, Missao Cururu, right
tributary of Rio Tapajés, 12-I-1961, leg. E.J. Fittkau, 1 male (ZS); same locality & collector,
(no date], 1 male (ZS). Holotype to be deposited in IN; paratypes in ZS, JE.
DIAGNOSIS: The immaculate wings, deeply bifid inferior volsella, dis-
tinctive superior volsella, clavate gonostylus and weakly cordiform-emar-
ginate base of the anal point will distinguish this species. The female and
immature stages are unknown.
ETYMOLOGY. I take great pleasure in naming this species in honor of
Dr. F. Reiss, who has been so kind and helpful to me throughout my studies
of the Chironomidae.
MALE IMAGO (n=7)
COLOR (slide mounted specimens). Head, body and legs light brown. Wing immaculate,
slightly dusky brown, with yellow-brown veins.
LENGTH. Total 2.93-3.28, 3.07 mm. Thorax 0.71-0.84, 0.77 mm. Abdomen 2.15-2.55,
2.39 mm.
HEAD. Setae: temporal 32-39, 36 (3); clypeal 11-14, 12; cibarial 7-10, 8 (5). Palpomere
lengths: 27-38, 32 (6); 35-45, 38 (6); 80-101, 88 (6); 89-123, 107 (6); 150-203, 180 (5). Frontal
tubercles 8-13, 11 long, 5-8, 6 wide. AR 1.72-2.00, 1.85.
THORAX. Scutal tubercle well developed; humeral pit with 3 moderate tubercles. Acros-
tichals 7-8, 8 (6); dorsocentrals 14-20, 17 (6); scutellars 7-9, 8 (6); prealars 8-10, 9 (6).
WING. Length 1.31-1.50, 1.41 mm; width 0.38-0.46, 0.41 mm. FCu distal to RM. VR 0.84-
0.88, 0.86. Setae: brachiolum 2; squama 1-3, 2 (5); R 13-16, 14; R; 4-7, 6; R4+s 8-12, 10.
MEM. AMER. ENT. SOC., 595
78 GENUS DICROTENDIPES
LEGS. Foretarsal beard absent. Palmate sensilla chaetica: 6-11, 9 on middle metatarsus, 3-
6, 5 on hind metatarsus. Lengths and proportions of legs:
P, P, PB,
fe 670-840, 580-690, 645-785,
716 649 729
ti 430-520, 495-615, 635-810,
489 564 741
ta, 885-1105, 300-370, 440-520,
1009 (5) 344 483
ta, 420-520, 145-180, 210-265,
473 (5) 344 239
ta; 365-460, 100-135, 170-205,
412 (5) 114 189
ta, 300-395, 40-60, 75-110,
350 (5) 49 (6) 89
tas 145-180, 45-55, 60-75,
159 (5) 53 (6) 67
LR 1.98-2.13, 0.59-0.63, 0.64-0.69,
2.06 (5) 0.61 0.65
BV 1.57-1.70, 3.90-4.32, JAIESSI,
1.63 (5) 4.11 (6) 3.35
SV 1.20-1.30, 3.44-3.68, MBI,
1.25 (6) 3.53 3.04
ABDOMEN. 1-7 ventral accessory setae on S VI.
HYPOPYGIUM (Figs. 26A, B) with 14-30, 21 dorsomedial setae. Gonostylus apically ex-
panded, curved medially, with 3-4, 3 preapical setae. Superior volsella (Figs. 26C, D) length
45-49, 47 (5); width 40-43, 41 (5); LWR 1.1-1.2, 1.1 (5); semi-pediform-triangular viewed
dorsally, with ventral subapical extension; with dorsal and ventral fine setae; with 1-2 small
sensilla chaetica. Inferior volsella length 118-135, 127 (5); deeply bifid, with 4—5 sensilla chae-
tica in single row on proximal lobe, 5-7 sensilla chaetica in single row on distal lobe; distal
lobe with 1 well developed ventral preapical seta. Anal point weakly cordiform-emarginate
basally, bare dorsally, with basal peduncle which slopes down from T IX; with 6-10, 8 lateral
basal setae.
Dicrotendipes soccus sp. nov. (Fig. 27)
TYPE LOCALITY: Igarapé Paracaixi, left tributary of lower Rio Negro, some hours down-
stream from mouth of Rio Branco, Amazonas, Brazil.
TYPE MATERIAL: Holotype: male/Pex, BRAZIL: Amazonas: Igarapé Paracaixi, left trib-
utary of lower Rio Negro, some hours downstream from mouth of Rio Branco, light trap, 14-
II-1962, leg. E.J. Fittkau. Paratypes (42): Amazonas: lower Rio Madeira, Parana Madeirinha,
light trap, 11-IX-1960, leg. E.J. Fittkau, 31 males (ZS); same locality & collector, 12-IX-1960,
2 males (ZS); mouth of Rio Negro at Manaus, light trap, 17-III-1961, leg. E.J. Fittkau, 4 males
(ZS); Upper Rio Solim6es, Igarapé Amataura, ca. 50 km W mouth of Rio Ica, ca. 15 km from
JOHN H. EPLER 719
mouth of Igarapé Amataura, light trap, 27-VIII-1961, leg. E.J. Fittkau, 1 male (ZS); Ilha do
Careiro, upper Rio Amazonas, nr Manaus, Parana da Terra Nova at Careiro, ca. 2 km from
mouth, light trap, 15-III-1961, leg. E.J. Fittkau, 1 male (ZS); lower Rio Negro, Ponta Negra
nr Manaus, light trap, 6-VI-1962, leg. E.J. Fittkau, 1 male (ZS); Rio Negro at Moura, light
trap, 5-II-1962, leg. E.J. Fittkau, 1 male (ZS); Lago Cabaliana, lower Rio SolimGes, drift, 16-
VI-1971, leg. F. Reiss, 1 male (ZS). Holotype to be deposited in IN; paratypes in ZS, JE.
DIAGNOSIS: The immaculate wings, deeply bifid inferior volsella with
greatly enlarged ventral subapical sensilla chaetica, distinctive slipper shaped
superior volsella, clavate gonostylus and strongly cordiform-emarginate base
of the anal point will distinguish this species. The female and larva are
unknown.
ETYMOLOGY. From the Latin soccus, slipper; refers to the slipper
shaped superior volsella.
MALE IMAGO (n=7)
COLOR (slide mounted specimens). Head, body and legs light brown. Wing immaculate,
light dusky brown, with yellow-brown veins.
LENGTH. Total 2.91-3.71, 3.28 (6) mm. Thorax 0.79-0.93, 0.84 (6) mm. Abdomen 2.11-
2.80, 2.45 (6) mm.
HEAD. Setae: temporal 28-40, 35 (4); clypeal 9-14, 12 (6); cibarial 5-8, 7 (4). Palpomere
lengths: 30-40, 34; 35-45, 41; 83-100, 90; 104-123, 113; 153-190, 179. Frontal tubercles 10
long, 5-7, 6 wide. AR 1.88-2.08, 1.99 (6).
THORAX. Scutal tubercle well developed; humeral pit with 3-5 weak to moderate tubercles.
Acrostichals 7-10, 8; dorsocentrals 14-17, 16; scutellars 8-9, 9 (6); prealars 8-10, 8.
WING. Length 1.34-1.63, 1.46 mm; width 0.39-0.44, 0.42 mm. FCu distal to RM. VR 0.84-
0.89, 0.86. Setae: brachiolum 2; squama 1-3, 2 (6); R 11-16, 14; R, 2-7, 5; R4+; 3-15, 10.
LEGS. Foretarsal beard absent. Palmate sensilla chaetica: 6-10, 8 on middle metatarsus, 3-
5, 4 on hind metatarsus. Lengths and proportions of legs, p. 80.
ABDOMEN. 2-7, 5 ventral accessory setae on S VI.
HYPOPYGIUM (Figs. 27A, B) with 22-41, 30 dorsomedial setae. Gonostylus thin, slightly
curved medially, with 2-3, 3 preapical setae. Superior volsella (Fig. 27C) length 50-65, 60;
width 20-25, 22; LWR 2.3-3.1, 2.7; slipper shaped, with 2 preapical sensilla chaetica. Inferior
volsella length 153-165, 155 (6); deeply bifid, with 3-6 sensilla chaetica in single row on prox-
imal lobe, 4-6 sensilla chaetica in single row on distal lobe; distal lobe with 3-8 greatly de-
veloped ventral preapical setae; at least 3 of these setae usually larger than dorsal sensilla
chaetica of same lobe. Anal point strongly cordiform-emarginate basally, bare dorsally, with
basal peduncle which slopes down steeply from a squat T IX; with 4-8, 6 lateral basal setae.
PUPA (n=1)
COLOR. Clear with pale yellow-brown margins.
LENGTH. Total 4.00 mm. Cephalothorax 0.95 mm. Abdomen 3.05 mm.
CEPHALOTHORAX. Cephalic tubercles similar to D. fittkaui (Fig. 20A), moderately de-
veloped. Dorsum moderately smoothly pebbled. Dc, closer to Dc3. Thoracic horn base with
tracheal bundles separate.
ABDOMEN. Similar to D. fittkaui (Fig. 21C). Sternites I-II with fine lateral shagreen bands,
S I also with fine posterior shagreen; S III with scattered fine spinules; S VI with anterior and
posterior areas of fine shagreen. Tergite I bare; T II with T-shaped shagreen area, spinules
largest posteriorly; T III-V with median quadrilateral shagreen area; T VI with broadly tri-
MEM. AMER. ENT. SOC., 36
80 GENUS DICROTENDIPES
P P, P;
fe 725-895, 620-740, 715-840,
799 671 761 (6)
ti 470-585, 540-665, 740-870,
516 599 791 (6)
tay 960-1140, 320-390, 440-540,
1053 (6) 352 493 (6)
tay 450-530, 150-190, 220-270,
494 (6) 170 249 (6)
ta; 385-470, 110-135, 180-210,
428 (6) 121 193 (6)
tay 340-410, 40-55, 80-100,
373 (6) 49 91 (6)
tas 150-180, 45-60, 70-80,
166 (6) 53 73 (6)
LR 2.04-2.18, 0.56-0.63, 0.59-0.67,
2.09 (6) 0.59 0.62 (6)
BV 1.53-1.63, 4.03-4.32, 3.32-3.43,
1.60 (6) 4.15 3.37 (6)
SV 1.18-1.26, 3.41-3.77, 2.93-3.31,
1.22 (6) 3.61 3.16 (6)
angular shagreen area, spinules largest mesally; T VII with an anterior pair of small ovoid
shagreen areas; T VIII with a pair of longitudinal fine shagreen bands. Tergites IV and V with
a posterior band of fine spinules; T V-VII with posterolateral group of fine spines. Posterior
margin of T II with transverse row of 65 hooklets. T VIII with 4 lateral lamellar setae. Cau-
dolateral spurs on T VIII similar to D. fittkaui (Figs. 21D, E) single or double, moderately
large. Anal lobes with 38-39 setae. DR 2.08.
Fic. 13. D. aethiops, adult male. A) Hypopygium, dorsal/ventral. B-D)
Variations of superior volsella, ventral.
JOHN H. EPLER 81
vn
!
MEM. AMER. ENT. SOC., 36
82 GENUS DICROTENDIPES
Fic. 14. D. californicus, adult male. A) Hypopygium, dorsal/ventral.
B) Variation of anal point. C-D) Variations of superior volsella, ventral.
E) Deformed superior volsella, ventral (Utah, U.S.A.).
JOHN H. EPLER 83
MEM. AMER. ENT. SOC., 36
84 GENUS DICROTENDIPES
Fic. 15. D. obrienorum, adult male. A) Hypopygium, dorsal/ventral.
B-C) Variations of superior volsella, ventral.
MEM. AMER. ENT. SOC., 36
JOHN H. EPLER
85
86 GENUS DICROTENDIPES
Fic. 16. D. sinoposus, adult male. A) Hypopygium, dorsal/ventral. B)
Superior volsella, ventral.
MEM. AMER. ENT. SOC., 36
JOHN H. EPLER
87
88 GENUS DICROTENDIPES
Fic. 17. D. amazonicus, adult male and female. A) Hypopygium, dor-
sal/ventral. B) Hypopygium, lateral. C) Superior volsella, lateral. D) Su-
perior volsella, dorsal. E) Female DmL, ApL, VIL.
MEM. AMER. ENT. SOC., 36
EPLER
89
90 GENUS DICROTENDIPES
Fic. 18. D. dasylabidus, adult male. A) Hypopygium, dorsal/ventral.
B) Superior volsella, dorsal.
91
H. EPLER
JOHN
MEM. AMER. ENT. SOC., 36
92 GENUS DICROTENDIPES
Fic. 19. D. demissus, adult male and female. A) Hypopygium, dorsal/
ventral. B) Hypopygium, lateral. C) Superior volsella, dorsal. D) Female
DmL, ApL, VIL.
93
JOHN H. EPLER
MEM. AMER. ENT. SOC., 36
94 GENUS DICROTENDIPES
Fic. 20. D. fittkaui, adult male. A) Hypopygium, dorsal/ventral. B)
Hypopygium, lateral. C) Superior volsella, ventral.
MEM. AMER. ENT. SOC., 36
JOHN H. EPLER
95
GENUS DICROTENDIPES
Fic. 21. D.fittkaui, pupa. A) Cephalic tubercles. B) Thoracic horn base.
C) Abdomen, dorsal. D, E) Caudolateral spurs on T VIII.
oF
JOHN H. EPLER
MEM. AMER. ENT. SOC., 36
98 GENUS DICROTENDIPES
Fic. 22. D. palearivillosus, adult male. A) Hypopygium, dorsal/ven-
tral. B) Superior volsella, ventral.
MEM. AMER. ENT. SOC., 36
JOHN H. EPLER
99
100 GENUS DICROTENDIPES
Fic. 23, D. paradasylabidus, adult male. A) Hypopygium, dorsal. B)
Superior volsella, dorsal.
JOHN H. EPLER 101
MEM. AMER. ENT. SOC., 36
102 GENUS DICROTENDIPES
Fic. 24. D. paterjohni, adult male. A) Hypopygium, dorsal/ventral. B)
Superior volsella, ventral.
103
EPLER
JOHN H.
ee
x
yy
A)
&
~
yy
MEM. AMER. ENT. SOC., 36
104 GENUS DICROTENDIPES
Fic. 25. D. radinovskyi, adult male. A) Hypopygium, dorsal. B, C)
Superior volsella, dorsal. D) Superior volsella, lateral.
JOHN H. EPLER 105
MEM. AMER. ENT. SOC., 36
106 GENUS DICROTENDIPES
Fic. 26. D. reissi, adult male..A) Hypopygium, dorsal/ventral. B) Hy-
popygium, lateral. C) Superior volsella, dorsal. D) Superior volsella, lateral.
JOHN H. EPLER 107
MEM. AMER. ENT. SOC., 36
108 GENUS DICROTENDIPES
Fic. 27. D. soccus, adult male. A) Hypopygium, dorsal/ventral. B) Hy-
popygium, lateral. C) Superior volsella, dorsal.
JOHN H. EPLER 109
MEM. AMER. ENT. SOC., 36
110 GENUS DICROTENDIPES
CHAPTER IV
THE DICROTENDIPES OF THE ORIENTAL-AUSTRALASIAN REGION
The first species of Dicrotendipes described from the regions was named
Chironomus conjunctus Walker (Walker 1856); Skuse (1889) merely re-
peated Walker’s description. Kieffer also described several species from the
regions in several papers (Kieffer 1910; 1911la; 1913a; 1916; 1917; 1921b).
Edwards (1924; 1928) described 2 new species from Fiji and Samoa. Free-
man (1961a) provided the first revision for the genus (as a subgenus of
Chironomus) from the regions, describing 6 species from Australia, 4 of
them new.
Chironomus (Dicrotendipes) canterburyensis Freeman was described from
New Zealand from female specimens (Freeman 1959:425). Forsyth &
McCallum (1978) reared this species (as an inquiline commensal) from a
lammelibranch mussel, Hydridella menziesi (Gray), and on the basis of the
immature stages, placed the species in Xenochironomus Kieffer. The genus
Dicrotendipes remains unknown to me from New Zealand.
Sasa and Hasegawa (1983) provisionally described a species, Dicroten-
dipes sp. ‘‘Yaeyama,”’ from Ishigaki Island in the Ryukyu Islands. The spe-
cies appears to be related to the Holarctic D. nervosus group, not to D.
lobiger as the authors stated. Specimens were not made available to me and
it is not included in this study.
Two additional species of Dicrotendipes have been described from the
region, D. arcistylus and D. canitibialis (Guha et al. 1985). These species
are not included because they are insufficiently described and specimens
were not made available. Dicrotendipes arcistylus may be a synonym of D.
flexus or D. tamaviridis Sasa. Dicrotendipes canitibialis is variously spelled
as canitiibalis and canitibiatis in Figure 5 and the text, respectively. Mention
is made of 3 other species whose names are misspelled and/or misinter-
preted: D. conjunctus (Walker) is misspelled as conjuncts and authorship
of the species name is incorrectly attributed to Freeman; D. incurvus (Sub-
lette), a Nearctic species, is apparently referred to as “‘D. carllus’’ (Guha
et al. 1985:29); and Chironomus (Prochironomus) punctatipennis (a prob-
able synonym of D. septemmaculatus) is misspelled as D. punctipennis.
In this study, I include 17 species from the combined Oriental-Oceanic-
Australian regions. Three species previously considered to be Dicrotendipes
are removed: Ch. blandellus Kieffer, 1906, (new name for blandus Skuse,
1889) is probably a Chironomus (see discussion under D. conjunctus below);
D. paxillus Guha, Chaudhuri et Nandi, 1982, is a junior synonym of Chi-
ronomus glauciventris (Kieffer, 1912); and D. socionotus Guha, Chaudhuri
JOHN H. EPLER 111
et Nandi, 1982, is probably a junior synonym of Chironomus tainanus (Kief-
fer, 1912). It should be noted that Ch. tainanus is also probably a senior
synonym of Nilodorum biroi (Kieffer, 1918).
10.
11.
KEY TO ADULT MALES OF ORIENTAL-AUSTRALASIAN DICROTENDIPES
[D. arcistylus, D. canitibialis and D. semiviridis not included]
Median volsella present (Figs. 28, 35) ..............0002 ee ee eeeee 2
Mediankyolsellayabsentip. 2 awensts b) aee shh coee ae ie itm» Se eeeeee Cle 3
Median volsella short, squat (Fig. 28); wings immaculate . . . . D. balciunasi sp. nov.
Median volsella long, thin (Fig. 35); wings marked with dark clouds and band... . .
a SUA OLE AONE Gl See ICs CAA ARG Pou en eee aeee Me cect eer D. lindae sp. nov.
Inferior volsella with membranous dorsal extension (Fig. 34).............. 4
Inferior volsella without membranous dorsal extension ................. 5
Superior volsella pediform (Figs. 34A, B)............ D. jonmartini sp. nov.
Superior volsella digitiform (Figs. 34C, D) ............. D. sarinae sp. nov.
Anal point sharply deflexed, not visible in dorsal view (Figs. 29, 36) ......... 6
Anal point at most moderately deflexed, visible in dorsal view............. 7
Anal point long and narrow; inferior volsella with broad apical club; gonostylus of
moderate width and dark brown in color (Fig. 29); wings immaculate .......
RY Ra REE hee, oie WT ERAS Ede Saves a Sens D. bilobatus (Kieffer)
Anal point short and broad; inferior volsella with simple apex, not broadly clubbed;
gonostylus broad and usually whitish to light brown (Fig. 36); wings sometimes with
smoky highlights along major veins ............. D. pelochloris (Kieffer)
Wangstwith bandsiorspots <5 2 2 2 2 je ek eee he is eee re ee eh costed a ts 8
Wes TNT WIENSS oc coc oma gm os D oOo oO PUD OOo oe oo D DOD OO OGaD 11
Wing with 6-7 spots; inferior volsella deeply bifid (Freeman 196la:Fig. 20d)......
b GLa OO pee a G EO gh ce OES SEM APE th ache omar D. septemmaculatus (Becker)
Wing with band of dark color; inferior volsella not deeply bifid (although dorsal sensilla
chaetica may be widely separated in D. lJeei [Freeman 1961la:Fig. 20a])...... 9
Superior volsella cylindrical-digitiform with weak membranous apex; Fiji and Samoa
(Rigs 30) eee ee ce as va Sandee 8 te eid Srl ie D. candidibasis (Edwards)
Superior volsella pediform or semi- pediform.................+...-..- 10
Anal cell with spot; apex of superior volsella directed laterad; inferior volsella with
dorsal sensilla chaetica only at or near apex (Freeman 196la:Fig. 20b).......
SaaS loots eae RESTO ORL COR Cuan EaBwarcotet b. oD. bto1 D. taylori (Freeman)
Anal cell without spot; apex of superior volsella directed mesad; inferior volsella with
widely separated rows of dorsal sensilla chaetica (Freeman 196la:Fig. 20a)... . .
58S SLRS Ee aD RES LEORC eon Rec eakdee to tote aroha ama D. leei (Freeman)
Superior volsella slender, usually arched mesad; inferior volsella broad to extremely
broad; medial setae originate from dorsal ovoid area on hypopygium..... . 12
Superior volsella semi-pediform, weakly deltoid or long and slender, arched laterad;
medial setae if present do not originate from dorsal ovoid area of hypopygium . .
SPE are SPP PRN ne ce G7 ok eae enter, alee Basse OS 7 14
MEM. AMER. ENT. SOC., 36
112
12.
13.
14.
15.
GENUS DICROTENDIPES
Inferior volsella extremely broad (Fig. 32); acrostichal setae not present (setae-less pits
PRESENE)), sieicy syrptae poceaseReinn cee ke Manse SeeereaS D. cumberlandensis sp. nov.
Inferior volsella not extremely broad (Fig. 31); acrostichal setae present ...... 13
Superior volsella strongly arched mesad (Figs. 31D, E); foretarsal beard present. . . .
re ee ree ara Sar ie br oar kcuc) cuca coho kone C D. pseudoconjunctus sp. nov.
Superior volsella at most weakly arched mesad (Figs. 31A-C); foretarsal beard absent
RR RS ILI PS oe OME ORO BAT Ge rb pol ch cede 6. Oo D. conjunctus (Walker)
Superior volsella weakly deltoid, densely setose (Fig. 33) ..... D. jobetus sp. nov.
Superior volsella pediform-clubbed or long, slender and arched laterad, not densely
SCUOS Gey so ateienss alee ate stg ete capes eawte cg aa aps WAR State el a 7 15
Superior volsella pediform-clubbed; distal portion not membranous, apex truncate (Fig.
SI) icy nye spice ete wk SIA cp fo. Cte ce RE BRE Oaycc toms eae ae D. tenuiforceps (Kieffer)
Superior volsella thin, slender and arched laterad; distal portion membranous, apex not
truncate (Johannsen 1932:Fig. 18; Hashimoto et al. 1981:Fig. 5D) (similar to
Holarctic! D> nenvosus)in 24) 2 ce eee eee D. flexus (Johannsen)
KEY TO KNOWN PUPAE OF ORIENTAL-AUSTRALASIAN DICROTENDIPES
T VII, VIII and anal lobe with extensive shagreen (Fig. 41B); S VIII with well-developed
posterior shagreen band; cephalic tubercles minute. ..... D. flexus (Johannsen)
T VII with at most an anterolateral pair of ovoid shagreen areas, T VIII with at most
an anterior and posterior pair of ovoid shagreen areas or 2 longitudinal shagreen
bands; anal lobe with or without shagreen; S VIII with faint shagreen; cephalic
tubercles moderately to well developed .......................-. 2
Posterior margin of T V with a row or groups of hooklets............... 3
Posterior margin of T V without hooklets.........................- 4
T V hooklets in 2 groups (Fig. 39K) .................-.-.-- D. conjunctus
(Walker); D. pseudoconjunctus sp. nov. (see text)
T V hooklets in continuous row (Fig. 42B) ........-......-+-+-+-+.+-++-+-:-.
aloe Ses a aE ee D. jonmartini sp. nov.; D. sarinae sp. nov. (see text)
Anal lobe with dorsal shagreen.............. D. septemmaculatus (Becker)
Anal lobe without dorsal shagreen ............-....---2222-2+-0--- 5
Posterior portion of shagreen area on T IV & V with distinctive adjoining area of fine
Spinel bands) (higs159G)\ieneaeae nena ene nenencne D. cumberlandensis sp. nov.
Shagreen areas on T IV & V without such areas ..................... 6
T VIII with 5 lateral lamellar setae; shagreen area on T III with anterior rows of larger
spinules: (Figs 410)iec 2s 5-4 Seeeones eek caer tone D. pelochloris (Kieffer)
T VIII with 4 lateral lamellar setae; shagreen area on T III with shagreen spinules larger
in middle portion of area (Fig. 38C)........... D. candidibasis (Edwards)
—
JOHN H. EPLER 113
KEY TO KNOWN LARVAE OF ORIENTAL-AUSTRALASIAN DICROTENDIPES
1. Frontal apotome with a large anteromesal ovoid or subquadrate area (Figs. 40E, I; 41Q);
frontalgpnrocessiabsent: Ses ico Re Aiey ees, Mindi edly eal CEM ee eS 2
Frontal apotome with small weak to strong frontal pit or process, or without markings
(igs DA Ss 4G s42G) os sislkccws) ee eae GS, 5
2. 6th lateral tooth of mentum rounded and fused/appressed to Sth lateral tooth (Fig.
NUN) G6 se at oO Se CRG) UES et Olle a Nee cre Ae SE AG. Sere D. pelochloris (Kieffer)
6th lateral tooth not rounded or fused/appressed to 5th lateral tooth......... 3
3. Median tooth of mentum broad (Fig. 40F); 29-37, 33 ventromental strial ridges. . . .
Sad setae sara antelope ak ele atch ae ae setae ake RP ey D. cumberlandensis sp. nov.
Median tooth of mentum not broad (Figs. 40B, K); if relatively broad, then ventromental
plates withiaboutr40)straléridges)ss a oe ere eee eee ee eee 4
4. Ventromental plate with 39-45, 42 strial ridges......... D. conjunctus (Walker)
Ventromental plate with about 28 strial ridges ..... D. pseudoconjunctus sp. nov.
5. Frontal apotome with long ventral frontal process (Fig. 7D) ...............
Rn es anc ee Se ee ees ee eee ee D. septemmaculatus (Becker)
Frontal apotome without frontal process; a weakly defined frontal pit may be present
RR Aen Mat ee eee es See Teh GATSee te es ene ee eae SRS) ure, Ke 6
6. Head capsule integument appears grainy at 400X; Ist and 2nd lateral teeth of mentum
S@Panateiectees teeticers deste nccwies ve levess enieuctas Ost ep ee ype a, WS, ec ate teak Goeel et te) Seep sese a eee 7
Head capsule integument not grainy at 400X; 2nd lateral teeth of mentum fused/
appresseditonlistalateralsys = Ai5. = ene; Prema eee gate a eret dep eer wey) Gu sitsmaees is 8
7. Ventromental plate with 35-40 strial ridges ........... D. jonmartini sp. nov.
Ventromental plate with 28-30 strial ridges ............. D. sarinae sp. nov.
8. Median tooth of mentum sunken well below level of Ist lateral teeth (Fig. 38G); 38-40
ventromental strial ridges; frontal pit absent ...... D. candidibasis (Edwards)
Median tooth at same level as Ist lateral teeth; 21-23 ventromental strial ridges; frontal
apotome with weak frontal pit................. D. flexus (Johannsen)
Dicrotendipes balciunasi sp. nov. (Fig. 28)
TYPE LOCALITY: Fogg Dam, NE of Humpty Doo, Northern Territory, Australia.
TYPE MATERIAL: Holotype: male, AUSTRALIA: Northern Territory: Fogg Dam, 15 km
NE Humpty Doo, at UV light, 5-X-1982, leg. J.K. Balciunas & J. Gillett (JB). Paratypes (22):
same data as holotype, 20 males (JB). Western Australia: De Grey River, 80 km NE Port
Hedland, 27-28-XI-1984, leg. B. & M. Baehr, 1 male (ZS); Mary River, 115 km WSW Hall’s
Creek, sandy river bed with some restpools, 17-18-XI-1984, leg. B. & M. Baehr, 1 male (ZS).
Holotype to be deposited in AN; paratypes in AN, BM, JE, and ZS.
DIAGNOSIS: The immaculate wings, somewhat pediform superior vol-
sella and short, squat median volsella will distinguish this species. The fe-
male and immature stages are unknown.
ETYMOLOGY: I am happy to name this species for Dr. Joe Balciunas,
who collected the majority of the type material.
MEM. AMER. ENT. SOC., 36
114 GENUS DICROTENDIPES
MALE IMAGO (n=5S)
COLOR (slide mounted specimens). Head and body brown, with posterior portions of ab-
dominal tergites V-VIII lighter; legs light brown, apical 4 of femora and fore tibiae dark
brown; fore metatarsus light brown at base, gradually darkening distally, tarsomeres dark
brown; mid and hind tibiae light brown with dark brown apices, metatarsi light brown, tar-
somere 2 light brown proximally, darker brown distally, remaining tarsomeres brown. Wing
immaculate, clear; with light yellow-brown veins.
LENGTH (4). Total 3.08-3.98, 3.67 mm. Thorax 0.90-1.05, 0.97 mm. Abdomen 2.10-2.98,
2.70 mm.
HEAD. Setae: temporal 29-40, 33; clypeal 9-17, 13; cibarial 5-16, 11. Palpomere lengths:
40-57, 46; 43-54, 50; 103-135, 112; 150-183, 163; 215-273, 243. Frontal tubercles 5-14, 8 long,
5 wide (3). AR 1.87-2.19, 2.10.
THORAX. Scutal tubercle well developed; humeral pit with 2-5 small to medium tubercles
or a bare spot on cuticle. Acrostichals 12-13 (2); dorsocentrals 17-27, 20; scutellars 6-12, 8;
prealars 6-10, 8.
WING. Length 1.38-1.60, 1.47 mm; width 0.44-0.51, 0.47 mm. FCu slightly distal to or
below RM. VR 0.83-0.95, 0.91. Setae: brachiolum 2; squama 7-11, 8; R 6-14, 10; R,; 0;
Rais 2.
LEGS. Foretarsal beard absent. Palmate sensilla chaetica: 8-16, 11 on middle metatarsus,
0 on hind metatarsus. Lengths and proportions of legs:
Py P, P;
fe 670-810, 610-750, 660-840,
721 654 717
ti 495-620, 565-700, 720-920,
535 605 717
ta, 970-770 260-330, 470-620,
(2) 289 539 (4)
ta. 355-450 150-190, 250-320,
(2) 167 275 (4)
tas 305-360 105-140, 220-275,
(2) 120 240 (4)
ta. 220-280 60-90, 120-165,
(2) 74 136 (4)
tas 130-150 60-80, 80-100,
(2) 68 91 (4)
LR 1.56 0.45-0.50, 0.65-0.76,
(2) 0.48 0.70 (4)
BV 1.92-1.94 3.45-3.80, 2.61-2.85,
(2) 3.62 2.74 (4)
SV 1.47-1.51 4.17-4.63, 2.57-3.01,
(2) 4.36 Al
ABDOMEN. Ventral accessory setae on S VI not apparent.
HYPOPYGIUM (Figs. 28A, B) with 0-3 medial setae. Gonostylus normal, slightly curved
medially, with 4-6, 5 preapical setae. Superior volsella (Fig. 28C) length 43-65, 53; width 34-
JOHN H. EPLER 115
48, 39; LWR 1.1-1.9, 1.4; pediform (somewhat triangular in deformed individuals) with 4-5
sensilla chaetica. A short, squat, membranous median volsella present dorsal to base of inferior
volsella. Inferior volsella length 78-108, 89; simply clubbed, with 8-10 sensilla chaetica scat-
tered on apex; with one well developed ventral preapical seta. Anal point bare dorsally, not
deflexed; with 3-6, 4 dorsal basal setae and 6-8, 7 lateral basal setae.
REMARKS. A most unusual species for the genus; along with D. lindae
sp. nov., D. balciunasi possesses a median volsella. Unfortunately, the im-
mature stages of both species are unknown. I see no reason for establishing
a new genus for this species and /indae; however, this may be necessary once
the immature stages are found.
Dicrotendipes bilobatus Kieffer (Fig. 29)
Dicrotendipes bilobatus Kieffer, 1917:222.
nec Chironomus (Dicrotendipes) conjunctus Walker 1856: Freeman 1961a:695.
DIAGNOSIS: The immaculate wings, digitiform superior volsella and
sharply deflexed anal point, which is not visible from a dorsal view, will
distinguish this species. The female and immature stages are unknown.
MALE IMAGO (n= 3)
COLOR (pinned specimens). Head light brown, antennae darker; thorax light yellow-brown
with red-brown vittae, scuteilum light yellow-brown, postnotum dark red-brown; abdomen
dark red-brown, distally lighter on T III-IV; fore femur stramineous, apex brown, remainder
of leg brown, mid and hind legs stramineous, apex of tibiae brown, metatarsi stramineous
with apices brown, remainder of legs brown. Wings immaculate, light dusky brown; veins light
yellow-brown.
LENGTH. Total 4.52-4.76, 4.63 mm. Thorax 1.14-1.23, 1.19 mm. Abdomen 3.38-3.56,
3.45 mm.
HEAD. Setae: temporal 30-41, 34; clypeal 14-17, 16; cibarial 14-15, 14. Palpomere lengths
(2): 52-62; 65-70; 175-182; 205; 290-295. Frontal tubercles 23-30, 26 long, 10-12, 11 wide.
AR 2.38-2.51, 2.46.
THORAX. Scutal tubercle well developed; humeral pit with 3 large tubercles. Acrostichals
10-14, 12; dorsocentrals 15-19, 17; scutellars 9-13, 11; prealars 8-10, 9.
WING. Length 2.58-2.69, 2.62 mm; width 0.66-0.67, 0.67 mm. FCu slightly distal to or
below RM. VR 0.90-0.97, 0.92. Setae: brachiolum 2; squama 6-11, 9; R 19-25, 22; R, 12-19,
16; R4+,; 24-33, 29.
LEGS. Foretarsal beard apparently absent. Palmate sensilla chaetica: 9-16, 12 on middle
metatarsus, 0 on hind metatarsus. Lengths and proportions of legs:
MEM. AMER. ENT. SOC., 36
116 GENUS DICROTENDIPES
P, P, P;
fe 1230-1240, 1075-1110, 1155-1190,
1237 1090 1172
ti 780-800, 990-1000, 1310-1320,
793 993 1313
ta, 1560-1660, 560-590, 855-905,
1617 573 873
ta, 765-820 295-300, 430-465,
800 298 452
ta, 610-660, 210-215, 350-355,
640 212 352
ta, 475-510, 110-120, 180-190,
498 117 183
tas 200 90 110-115,
112
LR 2.00-2.08, 0.56-0.60, 0.65-0.69,
2.02 0.58 0.67
BV 1.68-1.75, 3.66-3.73, 3.01-3.15,
1.71 3.71 3.06
SV 1.23-1.29, 3.50-3.77, 2.74-2.92,
1.26 3.64 2.85
ABDOMEN. Ventral accessory setae on S VI not apparent.
HYPOPYGIUM (Figs. 29A, B) with 7-9, 8 medial setae. Gonostylus moderately wide, curved
medially, with 5-6, 5 preapical setae. Superior volsella (Fig. 29C) length 80-95 (2); width 30-
33 (2); LWR 2.7-2.9; digitiform with slightly expanded apex, with 6-8, 7 sensilla chaetica.
Inferior volsella length 168-183 (2); with shallow apical notch, with 8-11 sensilla chaetica in
2-3 rows of 2-5 each; one well developed ventral preapical seta. Anal point narrow, bare
dorsally, completely deflexed beneath T IX; with 0 dorsal basal setae and 9-10, 10 lateral basal
setae.
REMARKS. Freeman (1961a) synonymized this species with conjunctus
Walker on the basis of hypopygial similarities. Freeman apparently relied
on Kieffer’s figure (Kieffer 1917:Fig. 15) to do so. The hypopygium of bi-
lobatus is unusual in that the anal point is strongly bent beneath T IX (Fig.
29); Kieffer’s hypopygium figure does not show an anal point. I have seen
pinned and fluid preserved specimens in which the anal point is strongly
deflexed, and I assume that Kieffer had similar specimens before him. The
superior volsellae of the 2 species are basically identical; the only character
separating them is the anal point. Whether the strongly deflexed anal point
is an artifact of preservation, a true species difference or perhaps an indi-
cation of a post-copulating condition may be resolved by examination of
reared specimens. I am returning bilobatus to species status because of this
uncertainty, with the hopes that it might interest workers to rear this unusual
Dicrotendipes.
JOHN H. EPLER 117
The type of bilobatus was apparently in the Hungarian National Museum
and was probably lost in the fire of 1956 (Freeman 1961a).
MATERIAL EXAMINED: AUSTRALIA: Australian Capital Territory: Canberra, pond
margin, 21 Nov. 1956, leg. W.W. Wirth, 1 male (US). New South Wales: Deewhy, South Creek,
27 Sept. 1956, leg. W.W. Wirth, 6 males (US).
Dicrotendipes candidibasis (Edwards) (Figs. 30, 38)
Chironomus (Xenochironomus?) candidibasis Edwards, 1924:573.
Chironomus melanocnemis Edwards, 1928:65. NEW SYNONYMY.
DIAGNOSIS: The adult is recognized by the distinctive wing and leg
markings, the low number of acrostichal setae and the weakly sclerotized
digitiform superior volsella of the male. The pupa can be separated by the
4 lateral lamellar setae on T VIII and the distinctive shagreen pattern. The
unusual larva is recognized by its distinctive mentum with the sunken me-
dian tooth and the almost completely fused first and second lateral teeth.
MALE IMAGO (n=5)
COLOR (pinned specimens). Head and thorax fuscous-light- brown, with scutellum lighter;
abdominal T I-V with greenish base color, T II-IV with median brown saddle; T V with prox-
imal % light green-brown, distal portion brown; T VI-VIII dark brown, hypopygium brown
with gonocoxites, styli and volsellae white to very light brown; legs with femora greenish stra-
mineous, fore and hind femora with brown apices; fore and hind tibiae with proximal 1/5
white, remainder brown or tibiae completely brown, mid tibia greenish-stramineous with light
brown apex; fore metatarsus with extreme proximal light brown band followed by extensive
white area and brown apical band, mid tibia with basal half white with postmedian brown
band and white apex, hind tibia white with brown apex; all remaining tarsomeres brown. Wing
with broad median band beginning at RM and extending distally through % to % of cells r4, 5,
m,,2 and m3,4; band stops at Cu, or continues to lower wing margin and extends forward
or slightly proximal to FCu; also with dark areas over vannal fold and An.
LENGTH. Total 3.75-3.78 (2) mm. Thorax 1.05-1.08 (2) mm. Abdomen 2.65-2.85, 2.77
mm.
HEAD. Setae: temporal 16-32, 22; clypeal 20-39, 26; cibarial 8-10, 9. Palpomere lengths:
42-52, 47; 44-52, 50; 142-183, 166; 195-228, 209; 295-360, 318. Frontal tubercles 2 long, 5
wide (2). AR 1.43-1.73, 1.55.
THORAX. Scutal tubercle moderately developed; humeral pit well developed with many
small to medium tubercles or 2 large tubercles or a pit. Acrostichals 0-2, 1; dorsocentrals 10-
15, 12; scutellars 4-12, 7; prealars 6.
WING. Length 1.75-1.90, 1.84 mm; width 0.49-0.53, 0.51 mm. FCu below RM. VR 0.95-
0.98, 0.96. Setae: brachiolum 1-2, 2; squama 2-6, 4; R 22-33, 27; R, 10-17, 13; R4,5 3-15,
2,
LEGS. Foretarsal beard absent. Palmate sensilla chaetica: 8-16, 11 on middle metatarsus,
0 on hind metatarsus. Lengths and proportions of legs:
MEM. AMER. ENT. SOC., 36
118 GENUS DICROTENDIPES
P, P, P;
fe 1005-1170, 790-950, 980-1180,
1057 850 1051
ti 720-900, 660-800, 950-1120,
774 715 1013
ta, 1130-1220, 410-455, 630-690,
1178 (4) 439 664
ta 555-610, 210-230, 330-410,
580 (4) 218 355
ta; 460-500, 140 275-310,
476 (4) 290
tay 415-450, 55-60, 145-165,
430 (4) 59 151
tas 185-190, 50-60, 75-90,
188 (4) 52 83
LR 1.55-1.64, 0.56-0.64, 0.62-0.68,
1.59 (4) 0.62 0.66
BV 1.73-1.80, 4.00-4.63, 3.08-3.14,
1.76 4.27 3.10
SV 1.48-1.54, 3.44-3.89, 3.00-3.33,
1.50 (4) 3.56 2.51
ABDOMEN. 0-3 ventral accessory setae on S VI.
HYPOPYGIUM (Figs. 30A, B). Gonostylus normal, slightly curved medially, with 5-6
preapical setae. Superior volsella (Figs. 30C, D) length 20-43, 28; width 19-25, 24; LWR
0.8-2.3, 1.3; digitiform with membranous apex; with 4-6, 5 sensilla chaetica. Inferior volsella
length 85-130, 104; notched slightly apically, with 1-5 sensilla chaetica in 1-3 rows, with 1-3
well developed ventral preapical setae. Anal point bare dorsally, not deflexed; with 1-6, 3
dorsal basal setae and 5-8, 7 lateral basal setae.
FEMALE IMAGO (n=2)
COLOR. Similar to male. The band across the wing extends further distad in cell mj, 2.
LENGTH. Total 3.11 (1) mm. Thorax 1.16 (1) mm. Abdomen 1.95-2.03 mm.
HEAD. Setae: temporal 21-22; clypeal 34-36; cibarial 10-12. Palpomere lengths: 48-55, 43-
50; 178-185; 210-245; 330-372. Frontal tubercles minute, not measurable. AR 0.46.
THORAX. Scutal tubercle moderately developed; humeral pit a well developed pit. Acros-
tichals 0-2 (3); dorsocentrals 16; scutellars 8-9; prealars 6.
WING. Length 2.08 mm; width 0.63-0.64 mm. FCu proximal to RM. VR 0.95-0.98. Setae:
brachiolum 2; squama 7; R 30-36; R; 15-16; R4,-5 31-33.
LEGS. Foretarsal beard absent. Palmate sensilla chaetica: 36-42 on middle metatarsus; 0
on hind metatarsus. Lengths and proportions of legs:
JOHN H. EPLER 119
P, P, P;
fe 1100-1105 850-870 1030-1095
ti 760-790 710-750 980-1070
ta, 1270-1310 455-470 660-680
tay 590-625 200-220 310-360
ta3 510-520 135-140 275-295
ta, 460-490 55 130-150
tas 200-205 55-60 80
LR 1.66-1.67 0.63-0.64 0.64-0.67
BV 1.74-1.78 4.40-4.53 3.21-3.36
SV 1.45-1.46 3.43-3.45 3.05-3.18
ABDOMEN. Ventral accessory setae not apparent on S VI. Notum 145-168; cerci 100-120.
S VIII with 25-28 setae/side; X with 4-5 setae; Gc IX with 1-2 setae/side. ApL as in Fig.
30E.
PUPA: (n=9)
COLOR. Clear with pale yellow-brown borders.
LENGTH. Total 4.13-4.79 (2) mm. Cephalothorax 1.00-1.13, 1.06 (3) mm. Abdomen 2.83-
3.68, 3.23 (7) mm.
CEPHALOTHORAX. Cephalic tubercles well developed (Fig. 38A), 25-50 (2) high, 63-108
(2) wide. Dorsum moderately to well pebbled. Dc2 closer to Dc3. Thoracic horn base (Fig.
38B) with tracheal bundles fused.
ABDOMEN (Fig. 38C). Sternite I with very fine shagreen areas, S II-IV with fine lateral
shagreen bands; S VI-VII with anterior pair of oval fine shagreen areas; T II with median
broadly T-shaped shagreen area; T III-IV with median quadrilateral shagreen areas, areas
narrower posteriorly; T VI with roughly V-shaped shagreen area; T VII with an anterior pair
of suboval shagreen areas; T VIII with a pair of longitudinal bands of fine shagreen; shagreen
areas on T II-VI with spines larger in middle portion of area. Tergites IV-V with posterior
band of fine spinules. Posterior margin of T II with transverse row of 64-104, 89 hooklets.
T VIII with 4 lateral setae. Caudolateral spurs on T VIII (Figs. 38D, E) single or with basal
spurs. Anal lobes with 29-44, 37 setae. DR 1.84-2.23, 1.99.
FOURTH INSTAR LARVA: (n=5)
COLOR. Head capsule light brown, postmentum darker.
HEAD. Postmentum length 238-250 (2). Mandible (Fig. 38F) length 188-210, 192, with 3
triangular lateral teeth; two well developed dorsal teeth present. Pecten mandibularis composed
of 7-9, 8 setae. Mentum (Fig. 38G) with 13 teeth, median tooth sunk well below level of 1st
laterals; 1st and 2nd lateral teeth fused, giving appearance of notched Ist lateral tooth; width
95-109, 103 (4); MR 2.24-2.48, 2.34 (4). Ventromental plate with smooth anterior margin;
width 85-92, 88; length 47-54, 49. VPR 1.70-1.90, 1.78; IPD 49-56, 52 (4); PSR 1.64-1.78,
1.72 (4); 38-40, 39 strial ridges. Length of antennal segments (3): 38-40, 39; 51-60, 55; 17-
19, 18; 8-10, 9; 7-8, 7. AR 1.02-1.15, 1.09 (3) (Fig. 38H). Inner blade of premandible (Fig.
381) subequal to outer blade. Pecten epipharyngis (Fig. 38J) with 3 lobes. Anterior margin of
frontal apotome (Fig. 38K) without discernible features, labral sclerite 1 smooth. S I with 6-
9 fringes (Fig. 38L).
BODY. Ventral tubuli absent.
MEM. AMER. ENT. SOC., 36
120 GENUS DICROTENDIPES
REMARKS. A large amount of reared material from Fiji differs from
the holotype specimen of D. candidibasis only by the completely brown fore
femora; in the holotype and one other specimen from Fiji the proximal
portion of the fore femur is white. The superior volsellae of the specimens
are similar; in the specimens with proximally white femora, the superior
volsellae are slightly longer (Figs. 30C, D). Unless future reared material
of specimens similar to the holotype is examined and shown to be radically
different, I am considering all of these specimens to represent D.
candidibasis.
I am here considering Ch. melanocnemis Edwards to be a junior synonym
of D. candidibasis. | have seen a male from Samoa with genitalia similar
to candidibasis but with leg color patterns as in melanocnemis (known only
from the female). The foretibiae are unfortunately missing from this Sa-
moan specimen. Leg color patterns (and other body coloration patterns)
seem to be subject to considerable variation in other species of Dicroten-
dipes, and I do not consider them to be good specific characters.
It has been difficult to find morphological characters which will serve to
separate females to species. The apodome lobe in the female genitalia may
be of use in some instances. Apodome lobes of candidibasis and melan-
ocnemis ‘‘types’’ are quite similar (Fig. 30E). In addition, in both males
and females of both ‘‘types,’’ the number of acrostichal setae is very low,
or they are absent. I believe that melanocnemis is nothing more than a color
variant of candidibasis.
Edwards (1924) described the hypopygium of candidibasis as lacking an
anal point. I have mounted the holotype on a microscope slide (in balsam)
and have observed that the anal point is contracted beneath T IX, somewhat
similar to the anal point of D. bilobatus. I have also mounted the female
holotype of melanocnemis in balsam on a microscope slide. Both types are
in the BM.
MATERIAL EXAMINED: FIJI: Lautoka, 11-V-1921, W. Greenwood, 1 male (holotype
Ch. candidibasis) (BM); Lautoka, 12-VI-1922, R. Veitch, 1 male (BM). Naduruloulou, at light,
8-X-1949, B.A. O’Conner, 5 females (det. melanocnemis) (BM); same locality & collector, 10-
XI-1949, 1 female (det. melanocnemis) (BM). Viti Levu, Laucala Bay (Suva), 13-VI-1985, leg.
J. Martin & C.J. Webb, 4 females (JM); Laucala Bay (Suva), egg mass #2, leg. J. Martin, 1
male/Pex/Lex, 2 males, 4 females, 1 pharate female pupa, 4 Pex (JM); Laucala Bay (Suva),
egg mass #3, coll. from vegetation at edge of a stream (reared at 20° C), 13-VI-1985, leg. J.
Martin, 1 male/Pex, 1 male, 2 pharate male pupae, 1 female/Pex, 2 females, 4 Pex, 2 Lex
(JM); Laucala Bay (Suva), grounds of University of the South Pacific, egg mass #4, 13-VI-
1985, leg. J. Martin, 7 males, 2 pharate male pupae/Lex, 10 females, 13 Pex, 2 larvae, 4 Lex
(JM). SAMOA: Tutuila, Naval Station, at light, 29-VIII-1940, leg. Swezey & Zimmerman, 1
male (det. melanocnemis) (US). Upolu, Apia, II-1924, P.A. Buxton & G.H. Hopkins, | female
(holotype Ch. melanocnemis) (BM); Upolu, Tapatapao, 1000 ft., at light, 22-VII-1940, leg.
Swezey & Zimmerman, | female (US).
2 ————
JOHN H. EPLER 121
Dicrotendipes conjunctus (Walker) (Figs. 31, 39, 40)
nec Chironomus conjunctus Loew 1850, in Keilbach 1982:351 (nomen nudum); Spahr 1985:22.
Chironomus conjunctus Walker, 1856:425; Skuse 1889:253.
nec Chironomus blandus Skuse, 1889:238.
nec Chironomus blandellus Kieffer, 1906:16.
nec Dicrotendipes bilobatus Kieffer, 1917:222.
Orthocladius conjunctus (Walker): Kieffer 1917:228.
Chironomus (Dicrotendipes) conjunctus Walker: Freeman 1961a:695.
DIAGNOSIS: The adult male is recognized by the lack of a foretarsal
beard, the relatively straight digitiform superior volsella, moderately broad
inferior volsella and well developed acrostichal setae. The pupae of con-
junctus and pseudoconjunctus may be inseparable, but in the limited ma-
terial available to me, conjunctus pupae have fewer spines (14-20) on the
posterior margin of T V than psewdoconjunctus (26-31); both species are
distinguished from cumberlandensis by the presence of these spines (lacking
in cumberlandensis). The larva is distinguished by the moderately broad
median tooth of the mentum (broader in cumberlandensis, narrower in pseu-
doconjunctus) and high strial ridge count (39-45, 42 in conjunctus, 29-37,
33 in cumberlandensis, 28 in pseudoconjunctus).
MALE IMAGO (n=5)
COLOR (pinned specimens). Head greenish, pedicels light yellow-red-brown; thorax green
with light red-brown vittae, scutellum green, postscutellum dark red-brown; abdomen brown
with greenish tinge, T VIII and hypopygium brown or abdomen dark green with T VI-IX
brown; fore femora green to greenish-stramineous, apices brown; tibiae and tarsi brown; mid
and hind femora and tibiae greenish stramineous, metatarsi light brown, darkening apically,
remaining tarsomeres brown. Wings immaculate, clear; with light brown veins.
LENGTH (4). Total 4.28-5.73, 4.78 mm. Thorax 1.18-1.58, 1.33 mm. Abdomen 3.10-
4.15, 3.45 mm.
HEAD. Setae: temporal 37-53, 43; clypeal 18-25, 21; cibarial 13-16, 14. Palpomere lengths
(4): 50-73, 56; 55-68, 61; 143-168, 154; 195-230, 206; 277-315, 293. Frontal tubercles 15-45,
32 long, 8-15, 10 wide. AR 2.28-2.74, 2.52 (4).
THORAX. Scutal tubercle moderately to well developed; humeral pit weak, scarlike or with
2-3 small tubercles. Acrostichals 7-17, 11; dorsocentrals 16-34, 21; scutellars 10-19, 13; prea-
lars 10-11, 10.
WING. Length 2.18-2.68, 2.39 mm; width 0.60-0.75, 0.67 mm. FCu below RM. VR 0.94—
0.96, 0.98. Setae: brachiolum 2-4, 3; squama 7-21, 12; R 22-28, 24; R, 15-21, 18; R4,5 18-
30, 24.
LEGS. Foretarsal beard absent. Palmate sensilla chaetica: 9-13, 11 on middle metatarsus,
0 on hind metatarsus. Lengths and proportions of legs:
MEM. AMER. ENT. SOC., 36
122 GENUS DICROTENDIPES
P, P, P;
fe 1005-1150, 950-1040, 1030-1170,
1085 1003 1106
ti 669-910, 850-1030, 1100-1370,
788 932 1224
tay 1305-1500, 480-540, 755-900,
1418 (4) 509 817
tay 590-720, 250-285, 380-480,
670 (4) 264 427
tas 490-580, 170-220, 300-405,
550 (4) 190 343
ta, 410-490, 90-140, 155-230,
449 (4) 109 188
tas 180-220, 70-115, 95-140,
195 (4) 91 116
LR 1.65-2.08, 0.52-0.58, 0.65-0.69,
1.86 (4) 0.55 0.67
BV 1.67-1.85, 3.45-4.11, 2.79-3.09,
1.73 (4) 3.76 2.94
SV 1.24-1.38, 3.71-3.87, 2.79-2.95,
1.31 (4) 3.80 2.85
ABDOMEN. 1-3 ventral accessory setae on S VI.
HYPOPYGIUM (Fig. 31A) with 5-14, 8 medial setae, set within a weakly defined ovoid
area. Gonostylus broad, curved medially, with 5-6 preapical setae. Superior volsella (Figs.
31B, C) length 68-100, 84; width 28-37, 30; LWR 2.5-3.2, 2.8; digitiform with short ventral
extension (similar to that of D. cumberlandensis, cf. Fig. 32B); with 6-8, 7 sensilla chaetica.
Inferior volsella length 145-175, 159; broad and notched slightly apically, with 1-6 sensilla
chaetica in 1-4 rows, with 1-2 well developed ventral preapical setae. Anal point bare dorsally,
slightly deflexed; with 0 dorsal basal setae and 7-12, 11 lateral basal setae.
FEMALE IMAGO (n=2)
COLOR (pinned specimens). Head and thorax yellow-brown, with greenish tinge, scutellum
green, postscutellum red-brown, abdominal T I-VI green, remainder brown; legs with femora
greenish stramineous, tibiae light yellow-brown, metatarsi light yellow-brown proximally, be-
coming brown apically, remaining tarsomeres brown. Wing immaculate, clear; veins light brown.
LENGTH. Total 4.36 (1) mm. Thorax 1.43 (1) mm. Abdomen 2.93 (1) mm.
HEAD. Setae: temporal 33-38; clypeal 34-35; cibarial 15-17. Palpomere lengths: 50-58; 45-
58; 148-172; 198-205; 320-338. Frontal tubercles 10-13 long, 8-9 wide. AR 0.43-0.44.
THORAX. Scutal tubercle well developed; humeral pit a scar with 2-3 tubercles. Acrosti-
chals 14 (1); dorsocentrals 31-32; scutellars 17-20; prealars 8-12.
WING. Length 2.31-3.00 mm; width 0.79 (1) mm. FCu slightly distal to below RM. VR
0.90-0.93. Setae: brachiolum 2-3; squama 13-14; R 23-30; R; 29-35; R4,5 42-59.
LEGS. Foretarsal beard absent. Palmate sensilla chaetica: 33-43 on middle metatarsus; 0-
13 on hind metatarsus. Lengths and proportions of legs:
JOHN H. EPLER 123
P, P, P3
fe 1140 1000-1110 1170 (1)
ti 830-840 940-1050 1200-1440
tay 1390-1490 475-540 790-880
taz 610-680 240-270 370-450
ta3 520 (1) 190 340-360
tag 450 (1) 110-120 170-190
tas 210 (1) 100-110 120-125
LR 1.67-1.77 0.51 0.61-0.66
BV 1.88 (1) 3.72-3.97 3.12 (1)
SV 1.33-1.42 4.00-4.08 2.97 (1)
ABDOMEN. 0-7 ventral accessory setae on S VI. Notum 205 (1); cerci 120-153. S VIII with
28-35 setae/side; X with 3-8 setae; Gc IX with 2-7 setae/side.
PUPA: (n=3)
COLOR. Clear with pale yellow-brown borders.
LENGTH. Total 6.26 (1) mm. Cephalothorax 1.63 (1) mm. Abdomen 3.88-4.63 (2) mm.
CEPHALOTHORAX. Cephalic tubercles well developed (Fig. 39A), 55-70 (2) high, 33-45
(2) wide. Dorsum weakly to well pebbled. Dc, closer to Dc3. Thoracic horn base (similar to
D. pseudoconjunctus, Fig. 39J) with tracheal bundles narrowly joined to barely separated.
ABDOMEN (similar to D. pseudoconjunctus, Fig. 39K). Sternite I with very fine shagreen
areas, S II-IV with scattered fine shagreen; S VI with anterior band of fine shagreen; T I with
very sparse posterolateral shagreen areas; T II-V with median quadrilateral shagreen areas,
with corners somewhat extended laterally; T VI with broadly V-shaped to hourglass-shaped
shagreen area; T VII with an anterior pair of suboval shagreen areas; T VIII with an anterior
and posterior pair of ovoid fine shagreen areas; shagreen areas on T II-V with spines larger
in posterior portion of area, shagreen area on T VI with larger spines anteriorly and posteriorly;
T II-IV with posterior area of fine spines. Posterior margin of T II with transverse row of 62-
88, 73 hooklets; posterior margin of T V with 2 groups of 5-11 spines (14-20 total). A weak
to strong reticulate cuticular pattern present on T IV (VI)-VIII. T VIII with 5 lateral setae.
Caudolateral spurs on T VIII (Fig. 39B) 2-5, well developed, sinuate. Anal lobes with 45-73,
56 setae. DR 2.82-3.20, 3.13.
FOURTH INSTAR LARVA: (n=4)
COLOR. Head capsule light brown to reddish yellow-brown, postmentum darker, postoc-
ciput dark-red to black.
HEAD. Postmentum length 247-378, 306. Mandible (Fig. 40A) length 208-298, 239 (3),
with 3 triangular lateral teeth; two well developed dorsal teeth present. Pecten mandibularis
composed of 14-16, 15 setae. Mentum (Fig. 40B) with 13 teeth, median tooth largest and higher
than Ist lateral teeth; width 140-177, 164 (3); MR 2.33-2.45 (2). Ventromental plate with
smooth anterior margin; width 125-150, 137; length 53-65, 59. VPR 2.31-2.36, 2.34; IPD 56-
64 (2); PSR 2.34-2.64 (2); 39-45, 42 strial ridges. Length of antennal segments: 39-45, 42;
73-107, 91; 21-33, 27; 16-24, 21 (3); 7-8, 7 (3). AR 1.21-1.27, 1.24 (3) (Fig. 40H). Inner blade
of premandible subequal to outer blade. Pecten epipharyngis (Fig. 40D) with 11-13, 12 lobes.
Frontal apotome (Fig. 40E) with large anteromesal ovoid pit, labral sclerite 1 smooth. S I with
about 16 fringes.
BODY. Ventral tubuli absent?
MEM. AMER. ENT. SOC., 36
124 GENUS DICROTENDIPES
REMARKS. The first species belonging to Dicrotendipes described from
Australia, D. conjunctus has been confused with several other species.
Kielbach (1982:351), in a list of amber insect fossils, records a ‘‘Chiron-
omus conjunctus Loew, 1850.’’ The name is a nomen nudum, for it is re-
corded only on the paper in which the amber specimen is wrapped; there
is no publication with such a name (pers. comm., Odwin Hoffrichter to Jon
Martin, 7 May 1984). Spahr (1985) also noted this situation, and listed ‘‘Chi-
ronomus conjunctus’’ as a ‘‘nomenklatorisch unverbindlicher Etiketten-
name’’ (nomenclaturally non-binding label-name).
Freeman (1961la) considered Ch. blandellus Kieffer (nomen novum for
Ch. blandus Skuse, which was a junior objective primary homonym of Ch.
blandus van der Wulp, 1858) a synonym of conjunctus. The only extant
specimen is the female holotype. I have examined this specimen (housed in
the AN) and conclude that it is not a Dicrotendipes. | have mounted the
pinned specimen in balsam on a microscope slide. The specimen’s ventro-
lateral lobe is very reduced, the apodome lobe is well developed with nu-
merous microtrichia and the labia bear microtrichia; the species probably
is a member of Chironomus, Einfeldia or another closely related genus.
Dicrotendipes bilobatus was also considered a synonym of conjunctus by
Freeman (1961la); I am treating bilobatus here as a separate entity. See re-
marks under D. bilobatus.
Freeman (1961la) noted that more than 1 species may have been present
in his conjunctus material. He found differences in the apical width of the
inferior volsella and the extent of the foretarsal beard. He correctly stated
that all existing names applied to specimens with broader appendages and
no beard, i.e., D. conjunctus. Specimens with narrower inferior volsellae
and a tarsal beard are D. pseudoconjunctus. At least a third similar species
occurs with no tarsal beard and much broader (than conjunctus) inferior
volsellae, D. cumberlandensis (q.v.).
In an unpublished Master’s thesis, Martin (1961) briefly described the
larva of D. conjunctus [as Chironomus (Dicrotendipes) conjunctus form C]
with a ‘‘pair of quite long ventral tubuli on the penultimate segment.’’ I
have not examined any material of this species with such tubuli. Ventral
tubuli are often present or absent on several other species of Dicrotendipes
(Epler 1987a). Edward (1964) also described the larva and the pupa of D.
conjunctus. He found the larvae inhabiting ‘‘algal mats and slime in per-
manent and temporary waters.”’
I have examined the type series of conjunctus, housed in the BM. It con-
sists of a male and female, both pinned. The male bears the following labels:
conjunctus n.s./VDL/Type; the female: VDL/68.4/Type. Because both
specimens bear a type label, they must be considered syntypes. I hereby
JOHN H. EPLER 125
designate the male as the lectotype and the female as a paralectotype for
Chironomus conjunctus Walker. | have remounted both specimens in bal-
sam on microscope slides.
MATERIAL EXAMINED: AUSTRALIA: Australian Capital Territory: Canberra, pond
margin, 21 Nov 1956, leg. W.W. Wirth, 1 male (US). New South Wales: The Beardy Waters,
nr Glenn Innes, 9-I-1968, leg. J. Martin, 1 male (JM); Boggy Swamp Creek on the Putty Road,
28-VIII-1981, leg. J. Martin, 2 males/Pex/Lex (JM); Deewhy, South Creek, 27 Sept 1956, leg.
W.W. Wirth, 9 males (US, BM), same locality & collector, 23 Oct 1956, 3 males (US); Hornsby,
at light, 6-8-I-1959, 1 male, 2 females [?) (BM). [Northern Territory]: Van) Diieman’s} Ljand],
1 male, 1 female (lectotype, paralectotype Ch. conjunctus) (BM). Tasmania: Advent Bay, 1
Jan 1922, leg. A. Tonnoir, 1 male (BM). Victoria: Botanic Gardens, Melbourne, 28-VIII-1963,
leg. J. Martin, 1 male (JM); Chiltern, leg. J. Martin, 1 male (JM); Echuca, leg. J. Martin, 1
male (JM); Lorne, 5-III-1958, N. Dobrotworsky, 1 male (BM); Narbethong, 18-III-1958, N.
Dobrotworsky, 3 males (BM); You Yangs, approx. 20 mls. SW Melbourne, 6-VII-1971, leg.
J. Martin, 1 female/Pex/Lex (JM). No data, J. Martin, 1 Pex/Lex (JM).
Dicrotendipes cumberlandensis sp. nov. (Figs. 32, 39, 40)
TYPE LOCALITY: Cumberland River, 8 km S of Lorne, Victoria, Australia.
TYPE MATERIAL: Holotype: male/Pex/Lex, AUSTRALIA: Victoria: Cumberland River
8 km S of Lorne, 4-X-1967, leg. Jon Martin (JM). Paratypes (18): Queensland: Barron River,
100 m below Tinnaroo Dam Spillway, on Hydrilla, 18-X-1982, leg. J.K. Balciunas, 1 female/
Pex (JB). Victoria: same data as holotype, 1 male/Pex/Lex, 1 female/Pex, 1 pharate female
pupa/Lex, 2 larvae (JM); same data as holotype except 16-X-1967, 1 male, 1 female/Pex/Lex,
2 females/Pex, 1 female/Lex, 1 pharate female pupa/Lex, 6 larvae (JM). The holotype will
be deposited in the AN; paratypes will be placed in BM, FS and JE.
ETYMOLOGY. This species is named for the Cumberland River, its type
locality.
DIAGNOSIS: The adult male can be distinguished from the similar D.
conjunctus and D. pseudoconjunctus by the massive inferior volsellae and
by the absence of acrostichal setae. The female can be distinguished by the
absence of acrostichal setae, which are replaced by setae-less pits. See di-
agnosis under conjunctus for pupae and larvae.
MALE IMAGO (n= 3)
COLOR (slide mounted specimens). Head, thorax and abdomen brown; fore femora light
brown with darker distal apex, fore tibiae light brown with darker proximal and distal apices,
mid and hind femora and tibiae light brown, all metatarsi light brown with darker apices,
other tarsomeres brown. Wings immaculate with light brown veins.
LENGTH. Total 5.53-5.98 (2) mm. Thorax 1.48-1.50 (2) mm. Abdomen 3.18-4.48, 3.90
mm.
HEAD. Setae: temporal 35-43, 39; clypeal 14 (2); cibarial 17-19 (2). Palpomere lengths (2):
50-60; 73-95; 177-190; 225; 320-350. Frontal tubercles 18-22, 20 long, 6-8, 7 wide. AR 2.18-
2.43, 2.30.
MEM. AMER. ENT. SOC., 36
126 GENUS DICROTENDIPES
THORAX. Scutal tubercle moderately to well developed; humeral pit weak, scarlike or with
2-3 small tubercles. Acrostichals absent, 8-10 setae-less pits present; dorsocentrals 10-21, 16;
scutellars 9-12, 11; prealars 9-10, 9.
WING. Length 2.38-2.93, 2.73 mm; width 0.67-0.79, 0.72 mm. FCu distal or below RM.
VR 0.92-0.95, 0.94. Setae: brachiolum 2; squama 6-9, 7; R 27-28, 27; R, 20-23, 21; R4,5 27-
38, 34.
LEGS. Foretarsal beard absent. Palmate sensilla chaetica: 14-19, 16 on middle metatarsus,
0 on hind metatarsus. Lengths and proportions of legs:
P, P, P;
fe 1115-1390, 1040-1260, 1110-1360,
1278 VNC 1223
ti 780-1020, 915-1165, 1150-1520,
907 1050 1353
tay 1350-1870, 540-680, 790-1030,
1590 600 907
tay 570-790, 275-340, 390-500,
670 298 447
ta; 475-620, 180-230, 275-380,
547 202 332
tay 380-480, 110-130, 140-190,
303 118 170
tas 180-195, 90-105, 115-130,
190 100 125
LR 1.68-1.83, 0.54-0.59, 0.65-0.69,
1.75 0.57 0.67
BV 2.02-2.07, 3.81-4.14, SESS2,
2.05 3.94 3.25
SV 1.29-1.45, 3.57-3.97, 2.80-2.88,
1.38 3.72 2.84
ABDOMEN. 0-1 ventral accessory setae on S VI.
HYPOPYGIUM (Figs. 32A, B) with 5-12, 8 medial setae, set within a weakly defined ovoid
area. Gonostylus broad, curved medially, with 6-7 preapical setae. Superior volsella (Fig. 32C)
length 93-95 (2); width 28-30 (2); LWR 3.2-3.3 (2); digitiform with short ventral extension,
slightly curved mediad; with 9-10, 9 sensilla chaetica. Inferior volsella length 180-195 (2); apex
extremely broad and notched slightly, with 1-7 sensilla chaetica in 2-3 rows, with 2-3 well
developed ventral preapical setae. Anal point bare dorsally, slightly deflexed; with 0 dorsal
basal setae and 13-20, 16 lateral basal setae.
FEMALE IMAGO (n = 3)
COLOR. Similar to male.
LENGTH. Total 3.80-5.53, 4.95 mm. Thorax 1.05-1.65, 1.44 mm. Abdomen 2.75-3.90,
3.51 mm.
HEAD. Setae: temporal 33-38, 36; clypeal 16-20, 18; cibarial 11-16, 14. Palpomere lengths:
58-75, 68; 55-80, 70; 138-190, 168; 167-247, 218, 173-385, 314. Frontal tubercles 3-15, 10
long, 7-8, 7 wide. AR 0.32-0.39, 0.36.
JOHN H. EPLER 127
THORAX. Scutal tubercle well to moderately developed; humeral pit indiscernible. Acros-
tichals absent, 7-8 setae-less pits present; dorsocentrals 9-17, 14; scutellars 9-12, 11; prealars
8-10, 9.
WING. Length 2.70-3.53, 3.21 mm; width 0.84-1.09 (2) mm. FCu distal to RM. VR 0.86-
0.89 (2). Setae: brachiolum 2-3, 2; squama 8-14, 11; R 21-31, 27; R, 21-35, 29; Ry,5 48-52
(2).
LEGS. Foretarsal beard absent. Palmate sensilla chaetica: 25-27, 26 on middle metatarsus;
2-15, 6 on hind metatarsus. Lengths and proportions of legs:
P, P, P;
fe 1115-1510, 1080-1390, 1160-1470,
1375 1283 1357
ti 790-1090, 990-1240, 1250-1630,
987 1152 1490
ta, 1550-2055, 530-720, 810-1120,
1878 647 993
tay 660-850, 260-350, 420-510,
763 313 474
ta; 530-695, 180-245, 295-380,
623 218 348
tag 410-555, 110-140, 160-190,
495 128 180
tas 170-215, 100-120, 110-140,
193 113 127
LR 1.86-1.96, 0.54-0.58, 0.65-0.69,
1.91 0.56 0.67
BV 1.23-1.26, 3.91-4.06, 3.27-3.49,
1.24 3.99 3.40
SV 1.23-1.28, 3.64-3.91, 2.77-2.98,
1.26 3.78 2.88
ABDOMEN. 0-4 ventral accessory setae on S VI. Notum 170-245, 206 (4); cerci 110-180,
152. S VIII with 24-47, 36 setae/side; X with 1-10, 6 (4) setae; Gc IX with 0-1, 1 (4) setae/
side.
PUPA: (n=8)
COLOR. Light yellow-brown with darker borders.
LENGTH (5) Total 5.13-6.98, 6.23 mm. Cephalothorax 1.43-1.68, 1.57 mm. Abdomen
3.70-5.34, 4.66 mm.
CEPHALOTHORAX. Cephalic tubercles well developed (Fig. 39C), 53-58 (2) high, 100-
133 (2) wide. Dorsum moderately to well pebbled. Dc, closer to Dc; (Fig. 39D). Thoracic horn
base (Figs. 39E, F) with tracheal bundles narrowly joined to barely separated.
ABDOMEN (Fig. 39G). Sternites II-V with very fine shagreen areas; T II with broadly V-
shaped median shagreen area with shagreen larger in posterior portion; T III-IV with median
quadrilateral shagreen areas; T V-VI with somewhat lyrate shagreen areas; T VII with an
anterior pair of suboval shagreen areas; T VIII with an anterior and posterior pair of ovoid
fine shagreen areas; shagreen smallest in anterior portion of areas on T III-VI, subequal over
rest of area; shagreen areas on T IV and V with distinctive adjoining area of fine spine bands
MEM. AMER. ENT. SOC., 36
128 GENUS DICROTENDIPES
in posterior portion of area. Posterior margin of T II with transverse row of 63-92, 74 hooklets.
A weak reticulate cuticular pattern present on posterior portion of T VII. T VIII with 5 lateral
setae. Caudolateral spurs on T VIII (Figs. 39H, I) 1-3, well developed, sinuate, often with
smaller basal spurs. Anal lobes with 39-82, 56 (7) setae. DR 2.48-3.38, 2.88 (7).
FOURTH INSTAR LARVA: (n=5)
COLOR. Head capsule yellow-brown or reddish yellow-brown to brown, postmentum and
genae darker, frontal apotome with darker borders on posterior 4, labral sclerite 1 darker
brown. Head capsule integument with grainy appearance.
HEAD. Postmentum length 240-320, 282 (3). Mandible length 253-265 (2), with 3 triangular
lateral teeth; two dorsal teeth present. Pecten mandibularis composed of 10-15, 12 (3) setae.
Mentum (Fig. 40F) with 13 teeth, median tooth broadly rounded; width 145-200, 173 (4); MR
2.67 (1). Ventromental plate with mostly smooth anterior margin; width 107-124, 115; length
53-70, 60. VPR 1.77-2.09, 1.94; IPD 65-89, 74 (4); PSR 1.39-1.69, 1.55 (4); 29-37, 33 strial
ridges. Length of antennal segments (4): 62-88, 74; 18-20, 19; 12-13, 12; 15-19, 16; 7-8, 7.
AR 1.13-1.60, 1.35 (Fig. 40G). Inner blade of premandible greater than outer blade. Pecten
epipharyngis (Fig. 40H) with 3-6 (2) lobes. Frontal apotome (Fig. 401) with large ventral an-
teromesal ovoid pit, labral sclerite 1 smooth, S I with 6-11, 9 (4) fringes.
BODY. Ventral tubuli absent.
REMARKS. A large species similar to D. conjunctus, D. cumberlandensis
has distinctive large inferior volsellae and lacks acrostichal setae. In their
place is a series of indentations; there is no trace of setae or their sockets.
The pupa lacks the hooklets usually found at the posterior margin of T
V in pupae of this group.
Dicrotendipes flexus (Johannsen) (Fig. 41)
Chironomus (Limnochironomus) flexus Johannsen, 1932:530.
Limnotendipes flexus (Johannsen): Lenz 1937:6.
Dicrotendipes flexus (Johannsen): Sublette & Sublette 1973:403; Hashimoto et al. 1981:14;
Sasa 1985:33.
DIAGNOSIS. The adult male strongly resembles D. nervosus (Staeger),
but can be separated by the apparently disjunct distributions and fewer setae
on R & R,, (21-26 in flexus, over 35 in nervosus). The bizarre pupa is dis-
tinguished by the distinctive shagreen. The larva is separated by its distinc-
tive mentum with the Ist and 2nd laterals almost completely fused.
See adult description in Johannsen (1932:530) and Hashimoto et al.
(1981:14). The female is undescribed.
PUPA: (n=2)
COLOR. Light yellow-brown to light brown.
LENGTH. Total about 4.20 (1) mm and larger. Cephalothorax about 1.20 (1) mm. Abdomen
3.00-4.20 mm.
JOHN H. EPLER 129
CEPHALOTHORAX. Cephalic tubercles very small to essentially absent. Dorsum mod-
erately to roughly pebbled. Dc, closer to Dc;. Thoracic horn base (Fig. 41A) with tracheal
bundles separated.
ABDOMEN. (Fig. 41B). Sternite I with well developed large shagreen spinules; S II-III with
well developed lateral shagreen bands and median shagreen areas; S IV with median fine sha-
green area and posterior area of well developed shagreen spinules; S V-VII with anterior sha-
green areas, S VIII with anterior shagreen area and posterior area of well developed spinules;
T II-VII with median quadrilateral shagreen areas, posterior shagreen spinules larger except
on T V-VI where median group of large spines present immediately posterior to small median
bare area; T VIII with broadly T-shaped shagreen area with posterior lateral extensions from
arms of ‘‘T’’; anal disc with well developed shagreen pattern on anterior 2; T II-VI with
lateral longitudinal shagreen bands; T III (IV)-V with posterior band of very fine spinules.
Posterior margin of T II with transverse row of 70-95 hooklets. T VIII with 4 lateral setae.
Caudolateral spurs on T VIII (Fig. 41C) single, thorn-like. Anal lobes with 97-156 setae. DR
1.80 (1).
FOURTH INSTAR LARVA: (n=3)
COLOR. Head capsule light yellow-brown; postmentum darker.
HEAD. Postmentum length 218-273, 243. Mandible length 168-185 (2), with 3 triangular
lateral teeth; one dorsal tooth present. Pecten mandibularis composed of 8-10, 9 setae. Mentum
(Fig. 41D) with 13 teeth, 2nd lateral teeth reduced and fused to Ist laterals, giving appearance
of notched Ist lateral teeth; width 120-130, 126; MR 2.42-2.73 (2). Ventromental plate with
shallow crenulations; width 82-105, 92; length 44-52, 48; VPR 1.74-2.02, 1.92; IPD 59-61
(2); PSR 0.75-0.85 (2); 21-23, 22 strial ridges. Length of antennal segments: 51-60, 55; 18-
22, 20; 10-11, 11; 11-14, 12; 5-6, 5. AR 1.06-1.20, 1.14 (Fig. 41E). Inner blade of premandible
greater than outer blade. Pecten epipharyngis (Fig. 41F) with 3 lobes. Frontal apotome (Fig.
41G) with small, poorly developed (or barely visible) anteromedian frontal pit; labral sclerite
1 smooth. S I with about 8-13 fringes.
BODY. Ventral tubuli absent.
REMARKS. The pupa of flexus is most unusual for a Dicrotendipes. No
other species in the genus displays such heavy ventral shagreen and spina-
tion, and such extensive shagreen areas on T VII-VIII.
A holotype and paratype are present in the BM. Both specimens are in
alcohol in microvials; their hypopygia are mounted on microscope slides.
There are few data in the microvials or on the slides, but code numbers are
with the specimens. These numbers correspond to the numbers given in Lenz
(1937) in his material examined (Vorkommen) listed at the end of each spe-
cies account. Johannsen (1932) also lists locality data, but does not give the
year in which the specimens were collected. Johannsen’s figure of flexus
(Johannsen 1932:Fig. 18) is of the distorted, upside down paratype’s hy-
popygium; the holotype mount is much better, and one wonders why the
poorer specimen was illustrated.
MATERIAL EXAMINED: AUSTRALIA: Northern Territory: Berry Springs, 57 km S. of
Darwin, on Hydrilla, 4-X-1982, leg. J.K. Balciunas, 1 pharate male pupa/Lex (JB); same
collection data except at UV light, 1 male (JB). INDONESIA): [JAVA): Ostjava, R. Bedali,
Ufer, 15-X-1928, (L4), 1 male (holotype Ch. flexus) (BM); Ostjava, R. Bedali; Lyngbya-Zone
MEM. AMER. ENT. SOC., 36
130 GENUS DICROTENDIPES
(2-6 m Tiefe) 22-XI-1928, leg. Thienemann, (L26), (ZS). Sumatra: N. Sumatra, Lake Toba,
along NE shore Samosir Is., 4 km N of Simanindo, on Potamogeton, 12-IX-1981, leg. J.K.
Balciunas, 1 larva (JB); Lake Toba at Tuk-Tuk, 8 km WSW of Prapat, at UV light, 6-VIII-
1982, leg. J.K. Balciunas, 1 male (JB). Sudsumatra: Ranau-See, Oberflache, 21-I-1929, 1 male,
(R4a), (paratype Ch. flexus) (BM); Heisse Quellen am Ranau-See, 40°, Lyngbya-Zone, 5-II-
1929, leg. A. Thienemann, (R38), 1 pharate male pupa (ZS).
Dicrotendipes jobetus sp. nov. (Fig. 32)
TYPE LOCALITY: Katherine Gorge, Northern Territory, Australia.
TYPE MATERIAL: Holotype: male, AUSTRALIA: Northern Territory: Katherine Gorge,
15 km NE Katherine, open Spinifex-Eucalyptus woodland, 6-8-XI-1984, leg. B. & M. Baehr
(ZS). Paratype (1): AUSTRALIA: W. Australia: Young River Station, outpost camp lake, 11-
12-1959, leg. D.H. Edward, 1 male (BM). Holotype to be deposited in ZS; paratype in BM.
DIAGNOSIS: The adult male is distinguished by its distinctive superior
volsella. The female and immature stages are unknown.
ETYMOLOGY. The name is an anagram constructed from the names of
my parents, John and Betty Epler, for whom I am quite pleased to name
this species.
MALE IMAGO (n= 2)
COLOR (pinned specimen). Head, thorax and abdomen brown. Forelegs light brown, distal
Y% of femur darker, base of metatarsus lighter brown; mid and hind legs light brown, distal
Y% of metatarsi and other tarsomeres darker brown. Wings immaculate, clear; with brown
veins.
LENGTH. Total 3.23-4.75 mm. Thorax 0.88-1.35 mm. Abdomen 3.23-4.75 m.
HEAD. Setae: temporal 30-42; clypeal 15-21; cibarial 11-18. Palpomere lengths: 40-57; 50-
75; 107-140; 145-183; 213-310. Frontal tubercles (1) 20 long, 7 wide. AR 2.11-2.53.
THORAX. Scutal tubercle well developed; humeral pit with 1-5 large tubercles. Acrostichals
7-13; dorsocentrals 13-24; scutellars 11-16; prealars 8-11.
WING. Length 1.45-2.25 mm; width 0.44-0.65 mm. FCu below to slightly distal to RM.
VR 0.91-0.95. Setae: brachiolum 2; squama 6-9; R 8-17; R; 0-1; R45 2.
LEGS. Foretarsal beard absent. Palmate sensilla chaetica: 5-10 on middle metatarsus, 0 on
hind metatarsus. Lengths and proportions of legs, p. 131.
ABDOMEN. Flattened setae not apparent on S VI.
HYPOPYGIUM (Fig. 33A) with 4-5 medial setae. Gonostylus normal, slightly curved me-
dially, with 4-6 preapical setae. Superior volsella (Fig. 33B) length 60-83; width 30-53; LWR
1.6-2.0; obovate, widest apically, with 12 sensilla chaetica. Inferior volsella length 108-175;
apex slightly notched, with 4-5 sensilla chaetica in 2 rows, with 1 well developed ventral preap-
ical setae. Anal point bare dorsally, slightly deflexed; with 6-8 lateral basal setae.
JOHN H. EPLER 131
P, P, P;
fe 720-1010 620-910 670-965
ti 490-730 505-860 730-1100
tay 880 (1) 300-465 470-710
tay 430 (1) 156--240 250-375
ta; 345 (1) 110-185 205-300
ta, 270 (1) 70-120 110-175
tas 135 (1) 65-100 85-120
LR 1.80 (1) 0.54-0.59 0.64-0.65
BV 1.77 (1) 3.47-3.61 2.86-2.88
SV 1.38 (1) 3.75-3.81 2.91-2.98
Dicrotendipes jonmartini sp. nov. (Figs. 34, 42)
TYPE LOCALITY: Cook, South Australia.
TYPE MATERIAL: Holotype: male, (AUSTRALIA): Cook, South Australia, from egg
mass #1, [no date}, leg. Jon Martin & B.T.O. Lee (JM). Paratypes (5): W. Australia: Shark
Lake, Nth Esperance, 10-XII-1959, leg. D.H. Edward, 1 male (BM). Same data as holotype,
2 Pex, 2 larvae (JM). Holotype to be deposited in AN; paratypes in BM, AN, JE.
DIAGNOSIS: The adult male is distinguished by the pediform superior
volsella and by its inferior volsella with its dorsal membranous extension.
The female is unknown. The pupa is very similar to D. sarinae and may be
inseparable; in the material available to me it is distinguished by the lower
number of T II hooklets and higher number of spines on the posterior mar-
gin of T V. The larva possesses a granular head capsule integument and is
distinguished from D. sarinae by the higher ventromental strial count and
higher VPR and PSR.
ETYMOLOGY. I take great pleasure in naming this species for Dr. Jon
Martin, who has been of immeasurable assistance during this study.
MALE IMAGO (n= 2)
COLOR (pinned specimen). Head, thorax, abdomen and legs brown. Wings immaculate;
clear with light yellow-brown veins.
LENGTH. Total 4.03-4.65 mm. Thorax 1.15-1.25 mm. Abdomen 4.03-4.65 mm.
HEAD. Setae: temporal 31-43; clypeal 16; cibarial 15-19. Palpomere lengths: 50-53; 53-
60; 128-135; 155-170; 277-293. Frontal tubercles 10-18 long, 9 wide. AR 2.55-2.66.
THORAX. Scutal tubercle well developed; humeral pit with 5-12 small to large tubercles.
Acrostichals 11-13; dorsocentrals 20-27; scutellars 10-11; prealars 9-11.
WING. Length 1.70-2.00 mm; width 0.52-0.61 mm. FCu below to slightly proximal to RM.
VR 0.98-1.00. Setae: brachiolum 2; squama 12 (1); R 11-12; R; 0-2; Rg, 2.
LEGS. Foretarsal beard absent. Palmate sensilla chaetica: 7-14 on middle metatarsus, 0 on
hind metatarsus. Lengths and proportions of legs:
MEM. AMER. ENT. SOC., 36
132 GENUS DICROTENDIPES
P, P, P;
fe 790-845 760-780 840-890
ti 600-640 720-760 945-990
tay 940-980 330-360 640-650
tap 440-445 190-205 340
ta3 395-400 155-170 290-300
ta, 310 95-100 160
tas 160-165 80-85 110
LR 1.53-1.57 0.46-0.47 0.66-0.68
BV 1.78-1.87 3.39-3.48 2.69-2.78
SV 1.48-1.52 4.28-4.48 2.79-2.89
ABDOMEN. Flattened setae not apparent on S VI.
HYPOPYGIUM (Fig. 34A) with 7-15 dorsomedial setae. Gonostylus broad, almost straight
medially, with 6 preapical setae. Superior volsella (Fig. 34B) length 58-63; width 43-47; LWR
1.2-1.5; pediform, with 5-6 sensilla chaetica. Inferior volsella with membranous dorsal ex-
tension, length 107-110; apex slightly notched, with 2-4 sensilla chaetica in 2-3 rows, with 4
well developed ventral preapical setae. Anal point bare dorsally, slightly deflexed; with 5-8
lateral basal setae.
PUPA: (n=2)
COLOR. Mostly clear, cephalothorax light yellow-brown.
LENGTH. Total 4.68-4.83 mm. Cephalothorax 1.18-1.28 mm. Abdomen 3.50-3.55 mm.
CEPHALOTHORAX. Cephalic tubercles small. Dorsum moderately pebbled. Dc, closer to
Dc,. Thoracic horn base (Fig. 42A) with tracheal bundles narrowly joined.
ABDOMEN (Fig. 42B). Sternite I with weak anterolateral shagreen areas; S II-III with
anterior and lateral fine shagreen areas; S VI-VIII with paired oval areas of fine shagreen; T
I with weak anterolateral shagreen areas, T II with broadly T-shaped median shagreen area,
T II-VI with median quadrilateral shagreen areas; T VII with anterior pair of ovoid shagreen
areas; T VIII with anterior and posterior pairs of ovoid shagreen areas; shagreen on T III-VI
larger medially. Posterior margin of T II with transverse row of 76-78 hooklets; T V with
posterior continuous band of 42-44 spines. A weak reticulate cuticular pattern present on
posterolateral portion of T VII. T VIII with 5 lateral setae. Caudolateral spurs on T VIII (Figs.
42C, D) 1-3, well developed, sinuate. Anal lobes with 31-33 setae. DR 2.11-4.50.
FOURTH INSTAR LARVA: (n=2)
COLOR. Head capsule light brown with no postmental darkening. Head capsule integument
with grainy appearance.
HEAD. Postmentum length 215 (1). Mandible length 188-190, with 3 triangular lateral teeth;
2 dorsal teeth present. Pecten mandibularis composed of 14-15 setae. Mentum (Fig. 42E) with
13 teeth, width 123-128; MR 2.86-3.00. Ventromental plate with smooth anterior margin;
width 119-121; length 52-53; VPR 2.28-2.29; IPD 34-37; PSR 3.27-3.50; 35-40 strial ridges.
Length of antennal segments (1): 69; 18; 13; 14; 7. AR 1.33 (1). (Fig. 42F). Inner blade of
premandible greater than outer blade. Pecten epipharyngis with 3 lobes. Frontal apotome (Fig.
42G) with elongate-oval frontal pit, labral sclerite 1 smooth. S I with 8-9 fringes.
BODY. Ventral tubuli absent.
REMARKS. This species is very similar to D. sarinae in all life stages. It
is possible that the sarinae material (all reared from 1 egg mass) may be
aberrant jonmartini.
JOHN H. EPLER 133
Dicrotendipes leei (Freeman)
Chironomus (Dicrotendipes) leei Freeman, 1961a:691.
See adult description in Freeman (1961a:691). The immature stages are
unknown.
MATERIAL EXAMINED: AUSTRALIA: New South Wales: Hornsby, 6-I-1958, leg. D.J.
Lee, 1 male (holotype) (SP); same data except 9-XII-1958, 1 male (paratype) (BM); same data
except light trap, 7-II-1957, 1 female (without abdomen) (paratype) (SP); same data except 6-
8-I-1959, (no collector data), 1 female (BM).
Dicrotendipes lindae sp. nov. (Fig. 35)
TYPE LOCALITY: Paradise River near Marion, Queensland, Australia.
TYPE MATERIAL: Holotype: male, AUSTRALIA: Queensland: Paradise River bei Mar-
ion, 25 km W Mackay, 22-12-1981, leg. M. Baehr (ZS). Paratype (1): AUSTRALIA: Northern
Territory: 30 km N Adelaide River, tropical savannah woodland, 5-6-11-1984, leg. B. and M.
Baehr, 1 male (ZS). Holotype and paratype in ZS.
DIAGNOSIS: The striking coloration of the body and wings, pediform
superior volsella and long median volsella will distinguish this species. The
female and immature stages are unknown.
ENTYMOLOGY. I take great pleasure in naming this beautiful species
for my beautiful wife, Linda.
MALE IMAGO (n= 2)
COLOR (slide mounted specimens). Head, thorax and abdomen brown, abdominal T II-V
with darker, narrow longitudinal band, wider on T VI and VII; posterior ‘4 of T VII and VIII
lighter brown. Hypopygium brown, gonostyli and inferior volsellae white. Legs with all coxae
dark brown; femora dark brown with light brown extreme distal and proximal apices and
broad light brown median band; tibiae dark brown with light brown extreme proximal apices
and median band; metatarsi light brown proximally, remainder of segment and following tar-
someres dark brown. The proximal dark brown band on the mid and hind tibiae is smaller
than that on the fore tibia. Wings light dusky brown with darker clouds running from RM
along R4,; for approximately *%4 of its length, along Cu and in a band from the vannal fold
to M3,4, along and distal to AN, and along the alula; veins dark brown. The microtrichia
within the dark clouds display a polygonal pattern at 100X.
LENGTH. Total 4.13-4.88 mm. Thorax 1.03-1.25 mm. Abdomen 3.10-3.63 mm.
HEAD. Setae: temporal 30-35; clypeal 18-19; cibarial 10 (1). Palpomere lengths: 48-55; 43-
53; 137-180; 175-213; 243-297. Frontal tubercles 13 long, 10 wide. AR 2.13-2.38.
THORAX. Scutal tubercle well developed; humeral pit with about 8 small to large tubercles.
Acrostichals 10-12; dorsocentrals 23-39; scutellars 11-16; prealars 9.
WING. Length 1.60-2.06 mm; width 0.45-0.62 mm. FCu slightly distal to or below RM.
VR 0.92-0.96. Setae: brachiolum 2; squama 14; R 12-14; R, 11-13; R4,5 5-8.
LEGS. Foretarsal beard absent. Palmate sensilla chaetica: 3-4 on middle metatarsus, 0 on
hind metatarsus. Lengths and proportion of legs:
MEM. AMER. ENT. SOC., 36
134 GENUS DICROTENDIPES
P, P, P3
fe 810-1050 760-915 840-1040
ti 610-775 665-870 895-1100
tay 1040-1350 350-460 610-720
tay 530-625 190-240 340-390
ta; 410-510 115-170 270-305
tag 320-415 55-70 95-115
tas 160-190 60-70 90-95
LR 1.70-1.74 0.53 0.65-0.68
BV 1.73-1.82 4.08-4.23 2.95-3.16
SV 1.35-1.37 3.88-4.07 2.84-2.97
ABDOMEN. Flattened setae on S VI not apparent.
HYPOPYGIUM (Fig. 35A) with 16-18 dorsomedial setae. Gonostylus normal, slightly curved
medially, with 4-7 preapical setae. Superior volsella (Fig. 35B) length 90-98; width 38-53;
LWR 1.8-2.4; pediform, with 4 sensilla chaetica. A long thin membranous median volsella
present laterad to base of superior volsella. Inferior volsella length 148-170; simply clubbed,
with 4 sensilla chaetica in 2 rows; with 1 well developed ventral preapical seta. Anal point bare
dorsally, deflexed; with 7-9 lateral basal setae.
Dicrotendipes pelochloris (Kieffer) (Figs. 36, 41, 50)
Tendipes pelochloris Kieffer, 1912:39; Kieffer 1916:113; Sublette & Sublette 1973:413 (listed as
unplaced species of Chironomini).
Limnochironomus niveicauda Kieffer, 1921b:585. NEW SYNONYMY.
Chironomus (Limnochironomus) niveicauda (Kieffer): Johannsen 1932:528.
Chironomus inferior Johannsen, 1932:534. NEW SYNONYMY.
Cladotendipes inferior (Johannsen): Lenz 1937:7.
Chironomus (Dicrotendipes) wirthi Freeman, 1961a:692. NEW SYNONYMY.
Dicrotendipes inferior (Johannsen): Sublette & Sublette 1973:403.
Dicrotendipes niveicauda (Kieffer): Sublette & Sublette 1973:404; Hashimoto et al. 1981:13.
Kimius hoonsooi Ree, 1981:218; Sasa & Hasegawa 1983:321. NEW SYNONYMY.
Dicrotendipes niveicaudus (Kieffer): Sasa & Hasegawa 1983:321.
Xenochironomus loripes Guha et Chaudhuri, 1981:163. NEW SYNONYMY.
Einfeldia loripes (Guha et Chaudhuri): Chaudhuri & Guha 1987:27. NEW SYNONYMY.
DIAGNOSIS: The adult male is distinguished by the distinctive hypo-
pygium with its sharply deflexed anal point and distinctive superior volsella.
The female can usually be identified by its color pattern, and by the dis-
tinctive apodome lobe with well developed microtrichia. The pupa is sep-
arated by the shagreen pattern and 5 lateral lamellar setae on T VIII (see
key); the larva can be separated by its distinctive mentum.
JOHN H. EPLER 135
MALE IMAGO (n=7)
COLOR (slide and alcohol specimens). Head and thorax brown to dark brown, abdominal
T I-V yellow-green to green to dark brown, T VI-IX dark brown, gonostyli white to light
brown. Legs with coxae yellow-white, femora with proximal 1/8-1/2 yellowish-white to white,
distal portion dark brown; tibiae dark brown; fore metatarsus with proximal 2-% yellowish-
white to white with distal portion dark brown or complete metatarsus dark brown, remaining
fore tarsomeres light brown to brown; ta; and ta of mid and hind legs yellowish-white to
white with distal apices brown, remaining tarsomeres brown, occasionally lighter proximally.
Wings hyaline to dusky brown, some specimens with diffuse brown cloud along Ry, R4,5, M,
Cu and An; veins brown.
LENGTH. Total 3.74-4.40, 4.01 (4) mm. Thorax 1.06-1.25, 1.14 (4) mm. Abdomen 2.65-
3.48, 2.99 (6) mm.
HEAD. Setae: temporal 43-53, 49; clypeal 11-23, 19; cibarial 8-12, 10. Palpomere lengths:
43-50, 47 (6); 47-63, 54 (6); 117-174, 140 (6); 158-195, 176 (5); 220-300, 265 (6). Frontal
tubercles 16-26, 20 long, 7-10, 8 wide. AR 1.95-2.27, 2.09.
THORAX. Scutal tubercle moderately to well developed; humeral pit with 3-5 moderate
tubercles. Acrostichals 9-14, 12; dorsocentrals 15-17, 16; scutellars 6-10, 8; prealars 7-10, 9.
WING. Length 1.73-2.28, 1.96 mm; width 0.51-0.67, 0.57 mm. FCu distal to RM. VR 0.81-
0.92, 0.85. Setae: brachiolum 2-3, 2; squama 3-13, 7; R 14-25, 19; R, 13-18, 15; R4,5 18-24,
20.
LEGS. Foretarsal beard absent or very slight. Palmate sensilla chaetica: 11-18, 16 on middle
metatarsus, 0-9, 4 on hind metatarsus. Lengths and proportions of legs (6):
P; P, P;
fe 885-1080, 850-1030, 970-1120,
962 913 1021
ti 610-770, 790-890, 940-1150,
670 795 1013
ta, 1140-1280, 370-475, 590-770,
1247 (5) All 654
ta) 510-600, 180-260, 290-405,
556 (5) 209 339
ta; 420-520, 130-170, 250-340,
462 (5) 147 286
ta, 320-410, 55-80, 110-160,
357 (5) 74 143
tas 150-180, 55-80, 85-110,
167 (5) 74 102
LR 1.66-2.07, 0.49-0.54, 0.62-0.68,
1.86 (5) 0.52 0.64
BV 1.78-1.98, 4.06-4.74, 2.99-3.44,
1.89 (5) 4.22 3.10
SV 1.20-1.45, 3.98-4.38, 2.93-3.24,
1.33 (5) 4.17 3.12
MEM. AMER. ENT. SOC., 36
136 GENUS DICROTENDIPES
ABDOMEN. 1-3 ventral accessory setae on S VI.
HYPOPYGIUM (Figs. 36A, B, C) with 10-16, 13 medial setae, apparently set within a
weakly defined ovoid area. Gonostylus broad, curved slightly medially, with 6-9, 8 preapical
setae. Superior volsella (Figs. 36D, E, F) length 65-77, 70 (6); width 31-40, 35 (6); LWR 1.8-
2.2, 2.0 (6); somewhat digitiform, often with expanded apex directed mediad; with 2-4, 3
sensilla chaetica. Inferior volsella length 100-125, 111; apex simple, with 2-4 sensilla chaetica
in 3-4 rows, with 2-4 well developed ventral preapical setae. Anal point bare, strongly deflexed
so that it often is not visible dorsally; with 0 dorsal basal setae and 11-16, 13 lateral basal
setae.
FEMALE IMAGO (n = 3)
COLOR. Generally similar to male, abdomen completely dark brown. Wings duskier than
in male and in some specimens with a more distinct, darker cloud over Ry, R4,5, M, Cu and
An.
LENGTH. Total 3.16-4.11, 3.65 mm. Thorax 1.16-1.48, 1.32 mm. Abdomen 2.00-2.63,
2.32 mm.
HEAD. Setae: temporal 38-49, 43; clypeal 16-44, 29; cibarial 9-11, 10. Palpomere lengths:
47-55, 51; 48-60, 55; 118-180, 144; 162-193, 173; 223-300, 255. Frontal tubercles 5-13, 8 long,
5-8, 7 wide. AR 0.46-0.50, 0.48.
THORAX. Scutal tubercle well to moderately developed; humeral pit a low bare area to 5
small scattered tubercles. Acrostichals 10-12, 11; dorsocentrals 16-20, 18; scutellars 10-12, 11;
prealars 8-10, 8.
WING. Length 1.97-2.75, 2.41 mm; width 0.67-0.88, 0.79 mm. FCu distal to RM. VR 0.83-
0.91, 0.86. Setae: brachiolum 2-3, 2; squama 11-19, 16; R 19-23, 22; R, 16-29, 23; R4,5 28-
41, 36.
LEGS. Foretarsal beard absent. Palmate sensilla chaetica: 32-34, 33 on middle metatarsus;
12-23, 18 on hind metatarsus. Lengths and proportions of legs:
P, P, P;
fe 910-1130, 860-1100, 970-1200,
1043 995 1080
ti 600-850, 780-990, 950-1270,
747 897 1128
ta 1400-1585, 375-510, 570-765,
1465 453 688
tay 460-590, 180-240, 290-385,
528 210 343
ta; 375-510, 130-165, 250-320,
445 142 287
ta, 320-390, 75-80, 135-150,
357 78 142
tas 170-185, 75-80, 100-110,
177 WA 105
LR 1.65-2.64, 0.48-0.52, 0.60-0.63,
2.02 0.50 0.61
BV DADAM, 4.33-4.94, 3.19-3.35,
DN) 4.62 3.30
SV 0.95-1.41, 4.09-4.37, 3.06-3.37,
1.23 4.19 3.22
JOHN H. EPLER 137
ABDOMEN. 3-9 ventral accessory setae on S VI. Notum 182-215, 198; cerci 100-148, 132.
S VIII with 21-26, 23 setae/side; X with 6-12, 9 setae; Gc IX with 2-4, 3 setae/side. DmL,
VIL and ApL as in Fig. 25G; ApL with well developed microtrichia.
PUPA: (n=7)
COLOR. Clear with yellow-brown margins; cephalothorax light brown.
LENGTH. Total 4.26-5.28, 4.67 (3) mm. Cephalothorax 1.10-1.25, 1.17 (3) mm. Abdomen
3.10-4.03, 3.54 mm.
CEPHALOTHORAX. Cephalic tubercles well developed (Fig. 41H), 63 (2) high, 75-88 (2)
wide. Dorsum moderately to well pebbled. Dc» closer to Dc; or Dc3. Thoracic horn base (Fig.
411) with tracheal bundles joined or narrowly separated.
ABDOMEN (Fig. 41J). Sternite I with weak posterolateral shagreen, S II-IV with very fine
scattered shagreen; S V-VII with weak anterior band of shagreen; T I sometimes with very
weak anterolateral shagreen areas; T II with broadly V-shaped median shagreen area with
shagreen slightly larger in anterior portion; T III-V with median quadrilateral shagreen areas,
T III-IV shagreen areas with anterior band of well developed spines, T V shagreen with median
area of stronger spines; T VII with an anterior pair of suboval shagreen areas; T VIII with
an anterior and posterior pair of ovoid or elongate fine shagreen areas. Posterior margin of
T II with transverse row of 64-90, 75 hooklets. A weak reticulate cuticular pattern present on
posterior portions of T VII-VIII. T VIII with 5 lateral setae. Caudolateral spurs on T VIII
(Figs. 41K, L, M) 1-4, small to well developed, often with smaller basal spurs. Anal lobes
with 42-69, 59 (6) setae. DR 1.83-3.05, 2.46 (6).
FOURTH INSTAR LARVA: (n=6)
COLOR. Head capsule light brown, postmentum slightly darker.
HEAD. Postmentum length 225-260, 243 (5). Mandible length 195-200, 197 (4), with 3
triangular lateral teeth, a depression proximal to 3rd tooth; two well developed dorsal teeth
present. Pecten mandibularis composed of 8-9, 9 setae. Mentum (Fig. 41N) with 13 teeth, 6th
lateral tooth broadly rounded, Sth reduced; width 123-153, 139 (5); MR 2.14-2.53, 2.39 (4).
Ventromental plate with smooth anterior margin; width 85-106, 96; length 47-58, 53. VPR
1.73-1.88, 1.81; IPD 47-55, 51 (4); PSR 1.96-2.04, 2.01 (4); 30-34, 32 strial ridges. Length
of antennal segments: 55-74, 67 (5); 20-33, 24 (5); 7-10, 9 (4); 11-13, 12 (4); 5-6, 6 (4). AR
0.90-1.51, 1.32 (4) (Fig. 41 O). Inner blade of premandible subequal to outer blade. Pecten
epipharyngis (Fig. 41P) with 4-5, 5 lobes. Frontal apotome (Fig. 41Q) with large dorsal an-
teromesal ovoid depression, labral sclerite 1 smooth. S I with 6-8, 7 fringes.
BODY. Ventral tubuli absent.
REMARKS. A widespread species throughout the Oriental-Australasian
areas. The distinctive hypopygium is very similar to that of the Afrotropical
D. kribiicola (Kieffer); the immature stages of the 2 species differ suffi-
ciently to maintain the 2 as separate species.
I have examined the type series of Tendipes pelochloris, housed in the IP.
It consisted of 3 pinned specimens, apparently the 2 males and 1 female
listed at the end of Kieffer’s (1912) description. Because no holotype was
designated, the 3 specimens must be considered syntypes. The specimens
bear ‘‘type’’ labels that were obviously added at a later date (a viewpoint
agreed to by Dr. H.J. Miiller, director of the IP, in a personal communi-
cation, 23-III-1987). Two specimens bear ‘‘cotypus’’ and “‘paratypus’’ la-
bels, the other specimen bears a ‘‘typus’’ and ‘‘holotypus’’ label. None of
MEM. AMER. ENT. SOC., 36
138 GENUS DICROTENDIPES
these designations are valid, because holotypes and paratypes can only be
fixed by the author in the original publication (International Code of Zo-
ological Nomenclature, Chapter XVI (1985)).
I am designating as lectotype a male specimen, remounted on a micro-
scope slide in Canada balsam, which bears the following labels: ‘‘LEC-
TOTYPE, Tendipes pelochloris Kieffer, 1912/ J.H. Epler 1987/ Paratypus/
COTYPUS (with a line drawn through the word)/ Tainan, Formosa, H.
Sauter, X, 08/ Coll. DEI, Eberswalde/ 87-1’’ (my personal identification
number used in slide preparation) and my determination label. I have des-
ignated the other 2 specimens, a male and female, both still pinned, as par-
alectotypes. The male paralectotype consists only of a head (without an-
tennae), thorax and a wing; it bears the following labels: ‘“Tainan, Formosa,
H. Sauter, X, 08/ Tendipes pelochloris (in Kieffer’s handwriting)/ Holo-
typus/ TYPUS (with a line drawn through the word)/ Coll. DEI, Eber-
swalde/ PARALECTOTYPE” and my determination label. The female par-
alectotype bears similar labels with the exceptions of paratypus and
COTYPUS (with a line drawn through the word) labels, and does not bear
a Kieffer determination label. A non-type female specimen is also present
in the IP collection. It bears a handwritten (in Kieffer’s hand) label “‘Cryp-
tochironomus pelochloris K.’’ The specimen is a female D. pelochloris and
has 3 ventral accessory setae on S VI. I have remounted this specimen in
balsam on a microscope slide.
There is some variation in coloration. Specimens from Japan are strik-
ingly marked, with females possessing wings with extensive smoky-brown
“*clouded’’ areas much darker than those of the male. Specimens from Thai-
land (not examined for this study) were apparently green (Hashimoto, et
al. 1981). Many specimens examined from Australia, such as the type series
of wirthi, are quite dark in general body coloration, and resemble the type
series of pelochloris. I consider wirthi a junior synonym of D. pelochloris;
the immature stages are morphologically inseparable.
The holotype of Chironomus inferior is a female specimen in alcohol with
a wing mounted on a microscope slide. The specimen is bleached and lacks
foretarsi. According to Johannsen (1932:535), this specimen was reared.
Lenz (1937:7) described the larva and pupa of this species, but did not state
that the specimens he described were the exuviae of the holotype, or how
they were associated. I have examined a single larva and a female pupal
exuviae determined by Lenz as inferior. All the specimens were apparently
collected together (code #R38, see also Remarks under D. flexus). I am
assuming that the immature stages are those of inferior; these immature
specimens are inseparable from D. pelochloris and inferior becomes a junior
synonym of D. pelochloris.
JOHN H. EPLER 139
I’ve examined a larva, pupal exuviae and an adult male of Xenochiron-
omus loripes Guha et Chaudhuri (determined as ‘‘Finfeldia loripes’’ by S.K.
Das). Chaudhuri & Guha (1987) later placed this species in Einfeldia. I con-
sider this species to be a junior synonym of D. pelochloris.
I was not able to locate a type specimen of Limnochironomus niveicau-
dus. It is not present in the collection at the University of the Philippines
at Los Banos (V.J. Calilung, pers. comm.), the Hungarian National Mu-
seum (L. Papp, pers. comm.), the IP (H.J. Miiller, pers. comm.) or the US
(R.V. Peterson, pers. comm.).
This is 1 of 2 species known to me outside of the Neotropical region in
which the male bears palmate sensilla chaetica on the metatarsus of the hind
leg (the other species is D. pseudoconjunctus). These sensilla chaetica were
present on all specimens examined except the males from Japan. They were
present on all females examined, including those from Japan.
MATERIAL EXAMINED: AUSTRALIA: New South Wales, Mosman, light trap, 9 March
1957, leg. W.W. Wirth, 2 males (1 male holotype wirthi (US); 1 male paratype wirthi [BM));
same data except 12-I-1957, 2 females (paratypes wirthi [US}). Northern Territory: Goanna
Lagoon, Gulungil Creek, Alligator Rivers region, 3-XI-1979, leg. J. Martin, 1 male/Pex, 2
males (JM). Queensland: Sarina, egg mass Y, laid about 21-I-1969, leg. J. Martin & D.L.
Porter, 4 males/Pex/Lex, 2 females/Pex/Lex, 1 pharate female pupa/Lex, 1 female (laid egg
mass Y), 3 larvae (JM); Somerset Dam, ca. 80 ml. N. Brisbane, 25-V-1969, leg. J. Martin, 7
males, 1 female (JM). INDIA: Tamil Nadu: before Madurai, 25-IX-1985, leg. C.W. & L.B.
O’Brien, 1 male (JE). West Bengal: Kalna, Sept. 1983, leg. S.K. Das, 1 male, 4 Pex, 1 larva
(UB). (INDONESIA): [Sumatra]: Sudsumatra, heisse Quellen am Ranau-See, 40°, Lyngbya-
Zone, 5-II-1929, leg. A. Thienemann, 1 female (holotype Ch. inferior) (BM), 1 female Pex, 1
larva (ZS). JAPAN: Hamamatsu and Shizuoka, Aug-Nov 1984, H. Hashimoto, 6 males, 5
females (HH). PAKISTAN: Sind: Haleji Lake (Indus Delta), 23-VIII-1985, leg. C.W. & L.B.
O’Brien, 3 males (JE). PHILIPPINES: Laguna Prov., Luzon, Laguna de Bay, Los Banos, 31-
V-1983, leg. J.K. Balciunas, 2 males (JB). SOUTH KOREA: leg. H.I. Ree, 2 males (det. K.
hoonsooi) (ZS). [TAIWAN] FORMOSA: Tainan, X-(19}08, H. Sauter, 2 males, 1 female (type
series pelochloris); Taihoku, 7-IX-1912, H. Sauter, 1 female (det. ‘““Cryptochironomus pel-
ochloris K’’).
Dicrotendipes pseudoconjunctus sp. nov. (Figs. 31, 39, 40)
Chironomus (Dicrotendipes) conjunctus Walker. Freeman 1961a:695 (in part).
TYPE LOCALITY: Scotts Lagoon near Lady Barron, Flinders Island, Tasmania, Australia.
TYPE MATERIAL: Holotype: male/Pex/Lex. (AUSTRALIA: Tasmania]: Scotts Lagoon,
near Lady Barron, Flinders Island, 11-II-1976, leg. Jon Martin & B.T.O. Lee (JM). Paratypes
(13); AUSTRALIA: South Australia: Lake Leake, via Kalangadoo, from egg mass #4, [no
date], leg. J. Martin, 1 male, 1 Pex (not associated with male) (JM). Tasmania: Arthurs Lakes,
28-I-1966, leg. G.F. Edmunds, | male (ZS); Interlaken, Lake Sorell, 14-X-1972, leg. J. Martin,
1 male (JM); Lake Leake, 21-X-1972, leg. J. Martin, 1 male (JM); Oatlands, Lake Dulverton,
12-II-1965, 2 males (JM). Victoria: Ocean Grove, 19-VIII-1960, leg. J. Martin, 1 male (BM);
MEM. AMER. ENT. SOC., 36
140 GENUS DICROTENDIPES
same locality & collector, 16-XII-1960, 2 males (BM); South Melbourne, vicinity of Albert
Park Lake, from egg mass A, laid-XII-1967, 2 males (JM). Western Australia: Lake Monger,
18-X-1956, D.H. Edward, 1 male (BM). The holotype will be deposited in the AN, paratypes
in BM, JE, JM, ZS.
DIAGNOSIS: The adult male can be distinguished from the similar D.
conjunctus and D. cumberlandensis by the strongly arched, medially di-
rected superior volsella, by its smaller (than cumberlandensis) inferior vol-
sella and by the presence of acrostichal setae. The female is unknown. See
diagnosis for pupa and larva under D. conjunctus.
ETYMOLOGY: From the Greek pseudos, fallacy. Refers to the confusion
of this species with the similar D. conjunctus.
MALE IMAGO (n=5S)
COLOR (pinned specimens). Head and thorax yellow-brown, scutellum pale green to green;
abdomen green, T VI-VII darker, T VIII-IX brown. Legs light brown to brown, femora lighter
on approximate proximal 2, tibiae sometimes lighter medially. Wings immaculate, very light
dusky brown; veins brown. :
LENGTH. Total 5.46-6.85 (2) mm. Thorax 1.37-1.75, 1.52 (3) mm. Abdomen 4.03-5.10,
4.45 (4) mm.
HEAD. Setae: temporal 45-49, 47; clypeal 17-22, 20; cibarial 8-18, 12. Palpomere lengths:
50-55, 53 (3); 55-75, 63 (4); 140-180, 163 (4); 188-215, 203 (4); 273-315, 285 (4). Frontal
tubercles 23-33, 27 long, 8-13, 10 wide. AR 2.50-2.56, 2.52 (3).
THORAX. Scutal tubercle moderately to well developed; humeral pit weak, scarlike or with
about 5 small tubercles. Acrostichals 8-18, 12 (4); dorsocentrals 18-36, 25; scutellars 11-19,
15; prealars 9-12, 10.
WING. Length 2.38-3.40, 2.75 mm; width 0.66-0.94, 0.78 mm. FCu slightly distal to or
below RM. VR 0.91-0.96, 0.93. Setae: brachiolum 3-5, 4; squama 16-25, 21; R 16-24, 22; R,
2-18, 11; R4,5 11-24, 17.
LEGS. Foretarsal beard well developed. Palmate sensilla chaetica: 9-18, 13 on middle me-
tatarsus, 0-5, 2 (4) on hind metatarsus. Lengths and proportions of legs (4), p. 141.
ABDOMEN. 0-1 ventral accessory setae on S VI.
HYPOPYGIUM (Fig. 31D) with 9-15, 11 medial setae, set within a weakly defined ovoid
area. Gonostylus broad, curved medially, with 4-5, 5 preapical setae. Superior volsella (Fig.
31E) length 80-115, 96; width 33-43, 39; LWR 1.9-2.7, 2.5; somewhat digitiform with ventral
extension, strongly curved mediad; with 7-13, 10 sensilla chaetica. Inferior volsella length 150-
193, 167; apex broad and notched slightly, with 1-8 sensilla chaetica in 2-4 rows, with 2-5
well developed ventral preapical setae. Anal point bare dorsally, slightly deflexed; with 0 dorsal
basal setae and 16-19, 18 lateral basal setae.
PUPA: (n=2)
COLOR. Light yellow-brown with darker borders.
LENGTH. Total 5.60-6.08 mm. Cephalothorax 1.35-1.50 mm. Abdomen 4.25-4.58 mm.
CEPHALOTHORAX. Cephalic tubercles well developed, not measurable in specimens be-
fore me. Dorsum moderately pebbled. Dc, closer to Dc; or De3. Thoracic horn base (Fig. 39J)
with tracheal bundles narrowly joined.
ABDOMEN (Fig. 39K). Sternites I-IV with fine medio-lateral shagreen areas; S V-VIII with
anterior pair of fine shagreen areas; T I with well developed reticulate cuticular pattern; T II
with median quadrilateral shagreen area, shagreen larger in posteromedial portion, and weak
a
JOHN H. EPLER
141
P, P, P3
fe 950-1200, 950-1180, 1015-1320,
1035 1019 1111
ti 770-1010, 900-1170, 1155-1500,
840 980 1259
tay 1190-1450, 470-609, 750-950,
1287 (3) 515 830
taz 535-650, 240-320, 410-510,
588 (3) 271 453
ta; 470-570, 190-260, 320-410,
513 (3) 215 353
ta, 390-470, 120-170, 175-230,
427 (3) 136 195
tas 170-225, 100-140, 110-155,
198 (3) 113 129
LR 1.44-1.58, 0.51-0.54, 0.63-0.69,
1.52 (3) 0.53 0.66
BV 1.75-1.91, 3.31-3.56, 2.73-2.89,
1.84 (3) 3.43 2.83
SV 1.44-1.52, 3.80-3.97, 213-297),
1.47 (3) 3.88 2.85
pair of anterolateral shagreen areas; T III with median quadrilateral shagreen area with an-
terolateral extensions, shagreen larger posteromedially; T II-III also with lateral longitudinal
shagreen bands; T IV-VI with median quadrilateral shagreen areas, shagreen largest poster-
olaterally; T VII with anterior pair of suboval shagreen areas; T VIII with an anterior and
posterior pair of ovoid fine shagreen areas; shagreen areas on T IV-V with distinctive adjoining
area of fine spine bands in posterior portion of area. Posterior margin of T II with transverse
row of 70-80 hooklets; posterior margin of T V with 2 groups of 12-17 spines (26-31 total).
A well developed reticulate cuticular pattern present on T VII-VIII. T VIII with 5 lateral setae.
Caudolateral spurs on T VIII (Fig. 39L) 1-3, well developed, sinuate, often with smaller basal
spurs. Anal lobes with 46-66 setae. DR 3.18-3.76.
FOURTH INSTAR LARVA: (n=1)
COLOR. Head capsule light brown, postmentum darker.
HEAD. Postmentum length 290. Mandible length 238, with 3 triangular lateral teeth; two
well developed dorsal teeth present. Pecten mandibularis composed of 15 setae. Mentum (Fig.
40K) with 13 teeth; not measurable. Ventromental plate with smooth anterior margin; width
119; length 60; VPR 1.98; 28 strial ridges. Length of antennal segments: 90; 26; 14; 18; 6. AR
1.41. Inner blade of premandible greater than outer blade. Pecten epipharyngis with 3 lobes.
Frontal apotome with large anteromesal ovoid pit, labral sclerite 1 smooth. S I with 13 fringes.
BODY. Ventral tubuli absent.
REMARKS. Freeman (1961a:695) first referred to this species as a pos-
sible new species in his discussion of D. conjunctus. He noted that some
specimens had foretarsi with beards, and some variation in the inferior vol-
sellae. Those specimens with bearded foretarsi were D. pseudoconjunctus.
MEM. AMER. ENT. SOC., 36
142 GENUS DICROTENDIPES
The superior volsellae are markedly different (much more so than any dif-
ferences in the breadth of the inferior volsellae noted by Freeman, although
the inferior volsella of pseudoconjunctus is usually narrower apically than
that of conjunctus) in the 2 species. The superior volsella in D. pseudo-
conjunctus is usually strongly arched mediad and has a more pronounced
ventral apical extension (Fig. 31). The superior volsella of D. cumberlan-
densis is intermediate between that of D. conjunctus and D. pseudocon-
junctus; however, D. cumberlandensis is easily distinguished by the huge
apex to the inferior volsella and its lack of a foretarsal beard and acrostichal
setae.
I can not reliably distinguish the pupa of D. pseudoconjunctus from that
of D. conjunctus. The only difference noted between the 2 species was the
number of spines at the posterior margin of T V; 14-20 spines in D. con-
junctus and 26-31 in D. pseudoconjunctus. I have not found the number
of these spines to be a reliable character in the other species of Dicrotendipes
which possess them, but this does not preclude the possibility that these
spines are a useful character in delimiting D. conjunctus and pseudocon-
junctus. This might be tested by examining a larger series of specimens, for
I was limited by an extremely small sample.
The same may be true for the larvae of these 2 species which I examined.
I was able to examine only 1 larva of D. pseudoconjunctus, but it was mark-
edly different from the larva of D. conjunctus. The D. pseudoconjunctus
larva had a low ventromental plate strial ridge count (28) compared to D.
conjunctus (39-45, mean 42).
I have also examined a male/Pex specimen from a pool near Lake Dove,
Cradle Mt. Natl. Park, Tasmania, leg. J. Martin, 18-X-1972, which may
belong here. The superior volsella resembles that of D. pseudoconjunctus,
but the pupa would fit D. conjunctus utilizing the intersegmental hooklet
counts discussed above. Most of the setae on the foretarsi have been lost
and it is not possible to discern if the specimen possessed a foretarsal beard.
The specimen may be an intermediate between D. conjunctus and D. pseu-
doconjunctus, and may indicate that D. pseudoconjunctus may be a higher
latitude variant of D. conjunctus.
Dicrotendipes sarinae sp. nov. (Figs. 34, 42)
TYPE LOCALITY: Sarina, Queensland, Australia.
TYPE MATERIAL: Holotype: Male, AUSTRALIA: Queensland: Sarina, [ex] egg mass Z,
laid about 21-I-1969, Jon Martin & D.L. Porter (JM). Paratypes (6): same locality & collectors,
19-I-1969, 1 female (laid egg mass Z) (JM); same data as holotype, 1 pharate male/Pex, 4
larvae (JM). Holotype to be deposited in AN; paratypes in AN, JE.
SS
JOHN H. EPLER 143
ETYMOLOGY. Named for the type locality.
DIAGNOSIS: The adult male can be distinguished from the similar D.
jonmartini by the smaller, digitiform superior volsella. The pupa has more
T II hooklets and fewer spines on the posterior margin of T V than jon-
martini. The larva has fewer ventromental plate striae, higher VPR and
lower PSR than jonmartini.
MALE IMAGO (n=2)
COLOR (slide mounted specimens). Head, thorax, abdomen and legs brown. Wings im-
maculate, very light dusky brown; veins yellow-brown.
LENGTH (1). Total 4.23 mm. Thorax 1.08 mm. Abdomen 3.15 mm.
HEAD. Setae: temporal 35; clypeal 12-18; cibarial 7-10. Palpomere lengths (1): 52; 63; 108;
172; 243. Frontal tubercles 5-9 long, 8 wide. AR 2.18-2.37.
THORAX. Scutal tubercle moderately developed; humeral pit well developed with 11-12
well developed, large tubercles. Acrostichals 9-11; dorsocentrals 16-18; scutellars 9-12; prea-
lars 5-7.
WING (1). Length 1.70 mm; width 0.57 mm. FCu below RM. VR 0.95. Setae: brachiolum
2; squama 7; R 6; R; 0; R4+; 1.
LEGS. Foretarsal beard very sparse. Palmate sensilla chaetica: 9-10 on middle metatarsus,
0 on hind metatarsus. Lengths and proportions of legs (1):
P, P, P;
fe 770 720 790
ti 570 685 850
ta, 920 310 590
ta) 460 180 290
ta; 375 140 275
ta, 280 90 150
tas 145 70 100
LR 1.61 0.45 0.69
BV 1.79 3.57 2.74
SV 1.46 4.53 2.78
ABDOMEN. Ventral accessory setae not apparent on S VI.
HYPOPYGIUM (Fig. 34C) with 6-8 dorsomedial setae. Gonostylus broad, almost straight
medially, with 4 preapical setae. Superior volsella (Fig. 34D) length 40-43; width 15-18; LWR
2.4-2.7; digitiform, slightly curved mediad; with 4-5 sensilla chaetica. Inferior volsella with
membranous dorsal extension, with 1-3 sensilla chaetica on or near extension; length 75-100;
apex clubbed, with 1-3 sensilla chaetica in 2 rows, with 2 well developed ventral preapical
setae. Anal point bare dorsally, slightly deflexed; with 8-12 lateral basal setae.
FEMALE IMAGO (n= 1)
COLOR. Similar to male.
LENGTH. Total about 3.78 mm. Thorax 1.05 mm. Abdomen about 2.13 mm.
HEAD. Setae: temporal 34; clypeal 23; cibarial 13. Palpomere lengths: 55, 70; 135; 175;
260. Frontal tubercles 5 long, 8 wide. AR 0.46.
MEM. AMER. ENT. SOC., 36
144 GENUS DICROTENDIPES
THORAX. Scutal tubercle well developed; humeral pit well developed with about 12 well
developed tubercles. Acrostichals 7; dorsocentrals 25; scutellars 15; prealars 9.
WING. Length 1.56 mm; width 0.59 mm. FCu below RM. VR 0.87. Setae: brachiolum 2;
squama 9; R 19; R,; 13; R4,5 18.
LEGS. Foretarsal beard absent. Palmate sensilla chaetica: 52 on middle metatarsus; 0 on
hind metatarsus. Lengths and proportions of legs:
P, P, P;
fe 779 710 770
ti 545 650 805
ta 870 300 530
tay 360 150 530
ta; 305 100 =
ta, 245 75 a
tas 135 70 80
LR 1.60 0.46 0.66
BV 2.09 4.20 ee
SV 1.51 4.53 2.97
ABDOMEN. Ventral accessory setae not apparent on S VI. Notum 172; cerci 93. X with
10 setae; Ge IX with 1 seta/side.
PUPA: (n=1)
COLOR. Light yellow-brown.
LENGTH. Not measurable.
CEPHALOTHORAX. Cephalic tubercles well developed, about 43 high, 75 wide. Dorsum
moderately pebbled. Dc, closer to Dc3. Thoracic horn base with tracheal bundles separated.
ABDOMEN. Damaged, but shagreen appears to be similar to D. jonmartini (Fig. 42B).
Posterior margin of T II with transverse row of 95 hooklets. Posterior margin of T V with
about 20 spines, apparently in single row. T VIII with 5 lateral setae. Caudolateral spurs on
T VIII similar to jonmartini (Figs. 42C, D). Anal lobes with 32 setae. DR not measurable.
FOURTH INSTAR LARVA: (n=3)
COLOR. Head capsule light brown, integument with grainy appearance.
HEAD. Postmentum length 193-205, 198. Mandible length 158-175, 165, with 3 triangular
lateral teeth; two well developed dorsal teeth present. Pecten mandibularis composed of 9-10,
10 setae. Mentum (Fig. 42H) with 13 teeth; width 115-125; MR 2.88-2.98 (2). Ventromental
plate with smooth anterior margin; width 85-92, 88 (4); length 47-50, 49; VPR 1.76-1.84,
1.80; IPD 35-41, 37; PSR 2.24-2.46, 2.38; 28-30, 29 (4) strial ridges. Length of antennal
segments (4): 28-30, 29; 52-57, 55; 15-17, 16; 10-11, 10; 12-14, 13. AR 1.15-1.33 (Fig. 42]).
Inner blade of premandible subequal to outer blade. Pecten epipharyngis with 3-4, 3 (4) lobes
(Fig. 42J). Frontal apotome with weak elongate-oval frontal pit; labral sclerite 1 smooth. S I
with 6-11, 8 fringes (Fig. 42K).
BODY. Ventral tubuli absent.
JOHN H. EPLER 145
Dicrotendipes semiviridis (Kieffer)
Chironomus semiviridis Kieffer, 1911a:166; Johannsen 1932:532.
Dicrotendipes semiviridis (Kieffer): Sublette & Sublette 1973:404; Chaudhuri & Guha 1987:27.
I have seen only 1 specimen of this species, a female syntype from the
Brunetti collection in the BM. The female (a pinned specimen) is a washed
out yellow-green with no distinguishing marks; it appears to be a Dicro-
tendipes. There are no ventral accessory setae apparent on S VI.
Kieffer’s figure (191la:Fig. 24) and description contribute little to the
identification of this species. According to Kieffer (1911a:111), the speci-
mens he described in his 1911 paper were kept at the Indian Museum of
Calcutta. These specimens are now apparently with the Zoological Survey
of India, Calcutta. According to M. Datta, Zoological Survey of India (pers.
comm., 12-XI-1986) specimens of semiviridis are in their collection, but ‘‘are
in extremely miserable condition and are suggestive of not being mailed to
anybody so as to save from further deterioration.’’ Until the male of this
species is redescribed (and because the female offers no specific characters),
I consider D. semiviridis a species inquirenda.
MATERIAL EXAMINED: BURMA: Mandalay, 11-IIJ-1908, N. Annandale, ex Brunetti
coll., 1 female (syntype semiviridis) (BM).
Dicrotendipes septemmaculatus (Becker)
See description and remarks in Chapter II.
Dicrotendipes taylori (Freeman)
Chironomus (Dicrotendipes) taylori Freeman, 1961a:692.
See adult description in Freeman (1961a:692); the immature stages are
unknown.
MATERIAL EXAMINED: AUSTRALIA: Queensland: Innisfail, [no date], F.H. Taylor, 2
males (holotype [SP]; paratype [BM]).
Dicrotendipes tenuiforceps (Kieffer) (Figs. 37, 51)
Tendipes tenuiforceps Kieffer, 1913a:136.
Chironomus (Dicrotendipes) innisfailensis Freeman, 1961a:694. NEW SYNONYMY.
Dicrotendipes tenuiforceps (Kieffer): Sublette & Sublette 1973:404; Chaudhuri & Guha 1987:27.
MEM. AMER. ENT. SOC., 36
146 GENUS DICROTENDIPES
DIAGNOSIS: The adult male can be distinguished by its distinctive su-
perior volsella. The immature stages are unknown.
MALE IMAGO (n= 2)
COLOR (pinned specimens). Head light brown, thorax golden yellow to orangish-yellow,
abdomen yellow-orange to light green; legs greenish-stramineous, fore tibia light brown. Wings
immaculate, clear; veins yellow-brown.
LENGTH. Total about 3.80 (1) mm. Thorax 0.78-1.10 mm. Abdomen about 2.70 mm.
HEAD. Setae: temporal 22-46; clypeal 10-16; cibarial 7. Palpomeres not measurable. Fron-
tal tubercles 5-12 long, 5-7 wide. AR 2.03-2.33.
THORAX. Scutal tubercle well developed; humeral pit a scar or with 3-5 small tubercles.
Acrostichals 6-12; dorsocentrals 11-12; scutellars 6-9; prealars 6-10.
WING. Length 1.48-1.63 mm; width 0.42-0.48 mm. FCu distal to RM. VR 0.85-0.86. Setae:
brachiolum 2; squama 1-4; R 9-16; R, 0-3; R4,;5 1.
LEGS. Foretarsal beard absent. Palmate sensilla chaetica: 5 (1) on middle metatarsus, 0 (1)
on hind metatarsus. Lengths and proportions of legs:
P, P, P;
fe 730-920 590-750 700-870
ti 430-610 510-625 740-920
ta, 910 (1) 285 (1) 470 (1)
ta) 430 (1) 145 (1) 230 (1)
ta; 395 (1) 90 (1) 200 (1)
ta, 310 (1) 49 (1) 110 (1)
tas 145 (1) 45 (1) 60 (1)
LR 212 (ab) 0.56 (1) 0.64 (1)
BV 1.62 (1) 4.33 (1) 3.18 (1)
SV 1.27 (1) 3.86 (1) 3.06 (1)
ABDOMEN. Flattened setae not apparent on S VI.
HYPOPYGIUM (Fig. 37A). Gonostylus thin, curved medially, with 5-7 preapical setae.
Superior volsella (Fig. 37B) length 63-75; width 35-50; LWR 1.5-1.8; pediform-clubbed, apex
directed laterad, with 5-8 sensilla chaetica. Inferior volsella length 112-115; apex simply clubbed;
with 3-4 sensilla chaetica in 2 rows, with 1 well developed ventral preapical seta. Anal point
bare dorsally, slightly deflexed; with 7 dorsal basal setae and 7-8 lateral basal setae.
REMARKS. Although not labeled as such, the single male specimen
from the Brunetti collection in the BM may have type status. Kieffer’s
material for his 1913 paper was apparently kept at the Indian Museum of
Calcutta and may now be with the Zoological Survey of India, Calcutta.
I interpret a recent letter from M. Datta (with the Zoological Survey of
India) as stating that no material of tenuiforceps is present in their collec-
tion. The Brunetti specimen was collected at Calcutta in September, which
is in the range of dates Kieffer lists at the end of his description, and may
be one of the original specimens described. Before any type status can be
———
JOHN H. EPLER 147
inferred to this specimen, the collection at the Zoological Survey of India
must be inspected in situ by a competent chironomid worker.
Freeman’s innisfailensis (Freeman 1961a) is clearly a junior synonym of
tenuiforceps, as evidenced by the distinctive hypopygium, and my exam-
ination of the holotype of Ch. innisfailensis.
MATERIAL EXAMINED: [AUSTRALIA]: Queensland: Innisfail, [no date}, FH. Taylor,
1 male (holotype Ch. innisfailensis) (SP). INDIA: Calcutta, 12-IX-1907, ex Brunetti coll., 1
male (BM).
MEM. AMER. ENT. SOC., 36
148 GENUS DICROTENDIPES
Fic. 28. D. balciunasi, adult male. A) Hypopygium, dorsal/ven-
tral. B) Hypopygium, lateral. C) Median and superior volsella, dorsal.
JOHN H. EPLER 149
SEI
"i i
Un
MEM. AMER. ENT. SOC., 36
150 GENUS DICROTENDIPES
Fic. 29. D. bilobatus, adult male. A) Hypopygium, dorsal/ven-
tral. B) Hypopygium, lateral. C) Superior volsella, ventral.
JOHN H. EPLER 151
MEM. AMER. ENT. SOC., 36
152 GENUS DICROTENDIPES
Fic. 30. D. candidibasis, adult male and female. A) Hypopygium,
dorsal/ventral. B) Hypopygium, lateral. C) Superior volsella, ven-
tral. D) Superior volsella, ventral, holotype. E) Female ApL.
153
JOHN H. EPLER
MEM. AMER. ENT. SOC., 36
154 GENUS DICROTENDIPES
Fic. 31. D. conjunctus, adult male (A-C). A) Hypopygium dorsal/
ventral, lectotype. B) Superior volsella, ventral. C) Superior volsella, ven-
tral, lectotype. D. pseudoconjunctus, adult male (D, E). D) Hypopygium,
dorsal/ventral. EE) Superior volsella, ventral.
155
JOHN H. EPLER
MEM. AMER. ENT. SOC., 36
156 GENUS DICROTENDIPES
Fic. 32. D. cumberlandensis, adult male. A) Hypopygium, dorsal/
ventral. B) Hypopygium, lateral. C) Superior volsella, ventral.
Sy)
JOHN H. EPLER
MEM. AMER. ENT. SOC., 36
158 GENUS DICROTENDIPES
Fic. 33. D. jobetus, adult male. A) Hypopygium, dorsal/ventral. B)
Superior volsella, ventral.
159
JOHN H. EPLER
36
MEM. AMER. ENT. SOC.
160 GENUS DICROTENDIPES
Fic. 34. D. jonmartini, adult male (A, B). A) Hypopygium, dorsal/
ventral. B) Superior volsella, ventral. D. sarinae, adult male (C, D). C)
Hypopygium, dorsal/ventral. D) Superior volsella, ventral.
161
EPLER
JOHN H.
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MEM. AMER. ENT. SOC., 36
162 GENUS DICROTENDIPES
Fic. 35. D. lindae, adult male. A) Hypopygium, dorsal/ventral. B)
Median and superior volsella, ventral.
MEM. AMER. ENT. SOC., 36
JOHN H. EPLER
163
164 GENUS DICROTENDIPES
Fic. 36. D. pelochloris, adult male and female. A) Hypopygium, dor-
sal/ventral. B) Anal point variation. C) Anal point, lateral. D) Superior
volsella, ventral, Australia. E) Superior volsella, ventral, Pakistan. F) Su-
perior volsella, ventral, Japan. G) Female DmL, ApL, VIL.
JOHN H. EPLER 165
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MEM. AMER. ENT. SOC., 36
166 GENUS DICROTENDIPES
Fic. 37. D. tenuiforceps, adult male. A) Hypopygium, dorsal/ven-
tral. B) Superior volsella, ventral.
167
JOHN H. EPLER
168 GENUS DICROTENDIPES
Fic. 38. D. candidibasis, pupa (A-E) and larva (F-L). A) Cephalic tu-
bercle. B) Thoracic horn base. C) Abdomen, dorsal. D, E) Caudola-
teral spurs on T VIII. F) Mandible, ventral. G) Mentum and ventro-
mental plate. H)Antenna. I)Premandible. J) Pectenepipharyngis. K)
Anterior portion of frontal apotome and labral sclerites. L) SI.
169
H. EPLER
JOHN
ee
—
Aw
O
MEM. AMER. ENT. SOC., 36
170 GENUS DICROTENDIPES
Fic. 39. D. conjunctus, pupa (A, B). A) Cephalic tubercle. B) Cau-
dolateral spurs on T VIII. D. cumberlandensis, pupa (C-1). C) Cephalic
tubercle. D) Cephalothorax, lateral. EE, F) Thoracic horn bases, showing
variation. G) Abdominal tergites II-VIII, dorsal. H, I) Caudolateral
spurs on T VIII. D. pseudoconjunctus, pupa (J-L). J) Thoracic horn
base. K) Abdomen, dorsal. L) Caudolateral spurs on T VIII.
UA
JOHN H. EPLER
MEM. AMER. ENT. SOC., 36
172 GENUS DICROTENDIPES
Fic. 40. D. conjunctus, larva (A-E). A) Mandible, ventral. B) Men-
tum and ventromental plate. C) Antenna. D) Pecten epipharyngis. E)
Anterior portion of frontal apotome and labral sclerites. D. cumberlan-
densis, larva (F-J). F) Mentum and ventromental plate. G) An-
tenna. HH) Pecten epipharyngis. 1) Anterior margin of frontal apotome
and labral sclerites. J) SI. D. pseudoconjunctus larva. K) Mentum and
ventromental plate.
JOHN H. EPLER 173
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xh yet
sar
MEM. AMER. ENT. SOC., 36
174 GENUS DICROTENDIPES
Fic. 41. D. flexus, pupa (A-C) and larva (D-G). A) Thoracic horn
base. B) Abdomen, dorsal. C) Caudolateral spur on T VIII. D) Men-
tum and ventromental plate. E) Antenna. F) Pecten epipharyngis. G)
Anterior portion of frontal apotome and labral sclerites. D. pelochloris,
pupa (H-M) and larva (N-Q). H) Cephalic tubercle. I) Thoracic horn
base. J) Abdomen, dorsal. K-M) Caudolateral spurs on T VIII. N)
Mentum and ventromental plate. O) Antenna. P) Pecten epipharyn-
gis. Q) Anterior portion of frontal apotome and labral sclerites.
MEM. AMER. ENT. SOC., 36
JOHN H. EPLER
175
176 GENUS DICROTENDIPES
Fic. 42. D. jonmartini, pupa (A-D) and larva (E-G). A) Thoracic horn
base. B) Abdomen, dorsal. C, D) Caudolateral spurs on T VIII. E)
Mentum and ventromental plate. F) Antenna. G) Anterior margin of fron-
tal apotome and labral sclerites. D. sarinae, larva (H-K). HH) Mentum and
ventromental plate. I) Antenna. J) Pecten epipharyngis. K) SI.
7
JOHN H. EPLER
MEM. AMER. ENT. SOC., 36
178 GENUS DICROTENDIPES
CHAPTER V. ZOOGEOGRAPHY
Chironomidae are well known for their usefulness in biogeographic stud-
ies. Brundin’s (1966) classic work demonstrated the hypothesized break up
of Gondwanaland with the present day distribution of podonomine chi-
ronomids. However, many other groups of chironomids, including Dicro-
tendipes, apparently do not lend themselves well to historical biogeographic
work (see Ashe et al. 1987). Brundin (1966) worked mainly with chironomids
in the plesiomorphic subfamily Podonominae (the bulk of his 1966 mon-
ograph is a taxonomic study on the subfamily); most of the species studied
are essentially cold stenotherms restricted to swiftly flowing mountain
streams. In contrast, Dicrotendipes is cosmopolitan, vagile and inhabits
many types of aquatic environments including swiftly flowing streams,
ponds, and lakes, and at least 3 species inhabit brackish water salt marshes
(Epler 1987a, Hashimoto 1984, Prat 1981). The present day distribution of
Dicrotendipes shows no correlation with any tectonic movements.
Fossils may sometimes provide clues to a taxon’s biogeographic history
(Brown & Gibson 1983:249). Although the fossil history of the Chiron-
omidae extends at least to the middle Jurassic to late Cretaceous (sum-
marized by Oliver 1981; see also Ashe et al. 1987), the earliest records of
Dicrotendipes date only from the late Pleistocene (approximately 10-12,000
years b.p.) (Hofmann 1971a, 1971b, 1978), long after any hypothesized ma-
jor plate movements. A larval mentum with ventromental plates illustrated
by Hofmann (1971b:fig. 19) appears to be a D. modestus or D. tritomus.
Thus the fossil record to date provides no indication of the historical bio-
geography of the genus.
DISTRIBUTION PATTERNS
There are some interesting distribution patterns in the genus, which are
summarized and illustrated in the remainder of this chapter. Several papers
have been published which deal with distribution patterns in the Chiron-
omidae (Ashe et al. 1987; Fittkau 1980; Fittkau & Reiss 1978, 1979; Reiss
1977b, 1978; Reiss and Sublette 1985). Distribution records recorded in this
chapter are taken from these papers, specimens examined for this study,
papers with new distribution records (Hashimoto et al. 1981; Reiss 1986;
Sasa & Hasegawa 1983) and previous revisions/descriptions (Contreras-
Lichtenberg 1986; Epler 1987a, 1987b; Freeman 1957, 1961a).
I am utilizing the faunal regions as delimited in Pielou (1979:8), except
that I use Afrotropical instead of Ethiopian (following Freeman & Cranston
1980 and Ashe et al. 1987).
JOHN H. EPLER 179
Holarctic. Four species display a Holarctic distribution: D. lobiger (Fig.
43), D. modestus (Fig. 44), D. nervosus (Fig. 45) and D. tritomus (Fig. 46).
Pan-American. The D. californicus complex (D. californicus, D. cryp-
ticus, D. embalsensis, D. obrienorum & D. pellegriniensis) shows a western
pan-American distribution pattern (Fig. 47). Some other chironomid species
have a distinct pan-American distribution, i.e., Goeldichironomus holo-
prasinus (Goeldi), G. amazonicus (Fittkau), G. carus (Townes) and Cala-
domyia spp. (Reiss & Sublette 1985). If I am correct in my determination
of a solitary larva from Paraguay as D. crypticus (see Chapter III, the spec-
imen may be a D. embalsensis), this species displays an interesting disjunct
distribution (Fig. 47). This distribution, as well as the apparent restriction
of D. californicus to the west of the Andes (D. pellegriniensis may be a form
of D. californicus; see Chapter III), may merely represent the relatively
meager amount of chironomid collecting done in the Neotropics outside of
the Amazon region. See also remarks under each species of the D. califor-
nicus complex in Chapter III.
Dicrotendipes aethiops has a southwestern U.S.-Mexican distribution, but
apparently does not occur in the Neotropical region; D. sinoposus occurs
in Mexico, northern South America and the Caribbean (Dominica), but does
not occur in the Nearctic region (Fig. 48).
Palaearctic-Afrotropical. Several species are found in both of these bio-
geographic regions: D. septemmaculatus (Fig. 49), D. fusconotatus (Reiss
1977b; Contreras-Lichtenberg 1986) and D. peringueyanus (Prat 1981; Con-
treras-Lichtenberg 1986).
Pan-Palaeotropical. Dicrotendipes septemmaculatus (Fig. 49) is found
throughout the tropics of the Old World, extends to southern Europe and
Japan to the north, and is also found in north and south Australia. Along
with D. modestus and D. nervosus, it is among the most widely distributed
species of the genus in the world.
Species with deeply bifid inferior volsellae are distributed pantropically.
However, I believe the Neotropical species with bifid inferior volsellae rep-
resent a different lineage, probably apomorphic, from the Afrotropical spe-
cies with similar inferior volsellae. Until associated larvae of the Neotropical
species are examined, such a hypothesis must remain a conjecture. Neo-
tropical adults (with bifid inferior volsellae) all possess palmate sensilla
chaetica on the hind metatarsus (absent on Afrotropical species). The pupae
of the only 2 Neotropical species possessing bifid inferior volsellae with
known pupae have 4 lateral lamellar setae on T VIII (5 in Afrotropical
species) and lack anal lobe shagreen (present on Afrotropical forms). At
least 2 hypotheses may explain the occurrence in the Neotropics of Dicro-
tendipes with deeply bifid inferior volsellae: 1) a vicariance hypothesis in
MEM. AMER. ENT. SOC., 36
180 GENUS DICROTENDIPES
which the Neotropical lineage is directly descended from the Afrotropical
linege; after the separation of South America and Africa (there are no data
which would indicate that the genus Dicrotendipes had yet evolved at this
period of time) the lineage evolved to the apparent apomorphic state of 4
lateral setae on pupal T VIII and a reversal occurred with the palmate sen-
silla chaetica on the hind metatarsus; 2) a dispersal hypothesis in which the
Neotropical lineage is descended from a more apomorphic Nearctic lineage
which dispersed south and then developed the palmate sensilla chaetica on
the hind metatarsus. Perhaps some environmental factor in the tropics in-
fluences the genome to produce a phenotype with deeply bifid inferior vol-
sellae. More larvae and pupae, currently unknown, of other species from
the neotropics must be examined to test these hypotheses. The V-shaped
median shagreen area on pupal T VI of D. fittkaui and D. soccus may ally
these 2 Neotropical species with deeply bifid inferior volsellae with the Ho-
larctic D. nervosus group. Such an alliance would provide support for the
dispersal hypothesis concerning the distribution of species with deeply bifid
inferior volsellae.
Oriental-Australasian. In addition to the afore-mentioned D. septem-
maculatus, 3 other species range across these 2 faunal regions: D. pelochloris
(Fig. 50), D. flexus (Fig. 51) and D. tenuiforceps (Fig. 51).
ENDEMICS
Many species are apparently endemic to their regions. These are [(?) in-
dicates a questionable species name]:
Nearctic: D. adnilus, D. aethiops, D. botaurus, D. fumidus, D. leucos-
celis, D. lobus, D. lucifer, D. neomodestus, D. simpsoni, D. thanatogratus
(the last species apparently endemic to Florida).
Neotropical: D. alsinensis, D. amazonicus, D. dasylabidus, D. demissus,
D. embalsensis, D. fittkaui, D. nestori, D. palearivillosus, D. paradasyla-
bidus, D. paterjohni, D. pellegriniensis, D. radinovskyi, D. reissi, D. sin-
oposus, D. soccus.
Palaearctic: D. fusciforceps (?), D. inouei, D. notatus, D. pallidicornis,
D. tamaviridis, D. truncatus (?), D. venetus (?).
Afrotropical: D. bredoi, D. chambiensis, D. collarti, D. cordatus, D.
ealae, D. freemani, D. kribiicola, D. leucolabis, D. schoutedeni, D.
sudanicus.
Oriental: D. arcistylus, D. canitibialis, D. semiviridis (?).
Oceanian: D. candidibasis.
Australasian: D. balciunasi, D. bilobatus, D. conjunctus, D. cumberlan-
densis, D. jobetus, D. jonmartini, D. leei, D. lindae, D. pseudoconjunctus,
D. sarinae, D. taylori.
MEM. AMER. ENT. SOC., 36
182 GENUS DICROTENDIPES
Fig. 43: Distribution map for D. lobiger.
Fig. 44: Distribution map for D. modestus.
JOHN H. EPLER 183
MEM. AMER. ENT. SOC., 36
184 GENUS DICROTENDIPES
Fig. 45: Distribution map for D. nervosus.
Fig. 46: Distribution map for D. tritomus.
185
JOHN H. EPLER
MEM. AMER. ENT. SOC., 36
186 GENUS DICROTENDIPES
Fig. 47: Distribution map for D. californicus (@), D. crypticus (A), D.
embalsensis (*), D. obrienorum (@) and D. pellegriniensis. (O).
JOHN H. EPLER 187
MEM. AMER. ENT. SOC., 36
188 GENUS DICROTENDIPES
Fig. 48: Distribution map for D. aethiops (@@) and D. sinoposus (@).
JOHN H. EPLER 189
MEM. AMER. ENT. SOC., 36
190 GENUS DICROTENDIPES
Fig. 49: Distribution map for D. septemmaculatus.
JOHN
H. EPLER
191
MEM. AMER. ENT. SOC., 36
192 GENUS DICROTENDIPES
Fig. 50: Distribution map for D. pelochloris.
Fig. 51: Distribution map for D. flexus (@) and D. tenuiforceps (A).
JOHN H. EPLER 193
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MEM. AMER. ENT. SOC., 36
194 GENUS DICROTENDIPES
CHAPTER VI. PHYLOGENY
A phylogenetic analysis of the species in the genus Dicrotendipes was
performed utilizing a standard Hennigian cladistic analysis as described by
Hennig (1966), Ross (1974) and Wiley (1981).
Saether (1977) provided a cladistic analysis of the genera of the tribe
Chironomini, which includes Dicrotendipes. He proposed that one synapo-
morphy, the presence of microtrichia on the well-developed female genitalic
apodeme lobe, delimited a group of genera (Dicrotendipes, Chironomus,
Glyptotendipes, Kiefferulus and Einfeldia), with the genera Nilodorum and
Goeldichironomus as their sister group. However, Saether used the species
Nilodorum devineyae Beck as the representative of Nilodorum in his analy-
sis; this species is actually a Goeldichironomus (Pinder & Reiss 1983). Saeth-
er’s ‘‘Goeldichironomus-Nilodorum’’ sister-group is actually only
Goeldichironomus. Similarities in the larvae of Kiefferulus and Nilodorum
led me to believe that Nilodorum should actually be included with the same
group as Dicrotendipes, Chironomus, Einfeldia, Kiefferulus, and Glypto-
tendipes. Through the kindness of Dr. L. Hare, females of “‘true’’ Nilodo-
rum sensu Pinder & Reiss (1983) (see Ashe et al. 1987:54 for a discussion
on the taxonomic status of Nilodorum) were made available. Nilodorum
does possess a well-developed apodeme lobe with numerous well-developed
microtrichia, and thus joins the group of genera including Dicrotendipes,
Chironomus, Einfeldia, Glyptotendipes and Kiefferulus (hereafter called the
““Chironomus group’’).
METHODS
Only 27 species known in all 3 life stages were included in the analysis.
This was necessary because in many cases the apomorphy or apomorphies
used to delimit lineages was/were present only in one life stage. It is often
impossible to identify Dicrotendipes to species using only one life stage.
This often necessitates rearing larvae through the pupal stage to the adult
stage to correctly identify some species. Species descriptions from the liter-
ature were not used, because descriptions were incomplete. Many species
are not known in all 3 life stages. The D. californicus species complex and
the D. lucifer complex were treated as single taxa, leaving a total of 25 taxa
to be analyzed. The 25 species used in the cladistic analysis were (abbre-
viations used for the species in Fig. 52 are given in brackets): D. californicus
[CALF], D. candidibasis [CAND], D. conjunctus [CONJ], D. cumberlan-
densis [CUMB], D. flexus [FLEX], D. fumidus [FUMI], D. fusconotatus
[FUSC], D. jonmartini [JONM], D. kribiicola [KRIB], D. leucoscelis
_——————_— sc tt Stt—t—‘CS;C;~™;~™
JOHN H. EPLER 195
[LEUC], D. lobiger [LOBR], D. lobus [LOBS], D. lucifer [LUCI], D.
modestus [MODS], D. neomodestus [NEOM], D. nervosus [NERV], D.
notatus [NOTA], D. pallidicornis [PALL], D. pelochloris [PELO], D.
pseudoconjunctus [PSEU], D. sarinae [SARI], D. septemmaculatus [SEPT],
D. sudanicus [SUDA], D. thanatogratus [THAN] and D. tritomus [TRIT].
Taxa were grouped together on the basis of shared derived characters,
termed synapomorphies. The polarity of character states (derived or
apomorphic and ancestral or plesiomorphic) was determined by outgroup
analysis. If a character existed in a similar state in both the ingroup (the
group being analyzed, in this case Dicrotendipes) and the outgroup (the
Chironomus group), that character state was termed plesiomorphic and was
not used in the analysis. Symplesiomorphies (shared ancestral character
states) are not used because they do not offer phylogenetic information as
it applies to the ingroup. Only synapomorphies indicate that taxa may share
a direct common ancestor. Symplesiomorphies can be synapomorphies at a
different level of universality (Wiley 1981). The character state ‘‘female
apodeme lobe well developed with numerous microtrichia’’ is symplesio-
morphic for Dicrotendipes in relation to the Chironomus group. However,
it is a Synapomorphy for the Chironomus group in relation to the tribe
Chironomini.
Saether (1977, 1979, 1983, 1986) has postulated that underlying synapo-
morphies (homoiologous characters of Hennig (1966:17); ‘‘the inherited
capacity to develop parallel similarities’? (Saether 1986:5)) are useful in
cladistic analyses, a viewpoint condemned by Farris (1985). Saether (1986)
discusses underlying synapomorphies again. I have not used underlying
synapomorphies as characters in my cladistic analysis. A problem inherent
in using underlying synapomorphies as characters in a cladistic analysis is
that one could justify any grouping one desires by declaring any sporadically
occurring character state as an underlying synapomorphy. Underlying
synapomorphies may provide additional evidence to support hypotheses
based on other synapomorphies, but cannot be used as the sole evidence to
support an hypothesis.
The outgroup used in the analysis was the Chironomus group. Because
of taxonomic uncertainty and probable polyphyletic lineages, it was not
possible to select a single genus as an outgroup. All of these genera are in
great need of revision.
RESULTS AND DISCUSSION
The characters and character states used to construct a hypothetical
phylogeny of Dicrotendipes species known in the adult, pupal and larval
MEM. AMER. ENT. SOC., 36
196 GENUS DICROTENDIPES
stages are listed below. Furcations in the cladogram (Fig. 52) are indicated
by upper case letters; numbers refer to apomorphies (those marked with an
* have apparently arisen more than once in different lineages); lower case
letters on the tree refer to losses, reversals, possible underlying synapo-
morphies, or symplesiomorphies and are placed on the tree only for refer-
ence purposes and were not used as characters in the cladistic analysis. In
the list, lower case letters in parentheses indicate (a) = apomorphic, derived,
and (p) = plesiomorphic, ancestral; upper case letters in brackets: [A] =
adult character, [P] = pupal character, [IL] = larval character.
The first 4 apomorphies listed are synapomorphous for the genus.
1. Pecten epipharyngis with less than 15 lobes (a); more than 15 lobes
(p) [L].
2. Ventromental plate ratio less than 2.5 (a); more than 2.5 (p) [L].
3. Thoracic horn base with 2 tracheal bundles (a); 1 tracheal bundle (p)
[P].
4. Sternite VI with ventral accessory setae (a); without ventral accessory
setae (p) [A].
Saether (1977: Fig. 62) utilized the short, squat larval ventromental plates
as his Trend 26 to separate Dicrotendipes from the genera Chironomus,
Kiefferulus, Glyptotendipes and Einfeldia. This trend corresponds to
apomorphy 2 in this cladistic analysis. Apomorphies 3 and 4 are apparently
secondarily lost (reversals have taken place) in several species. The 2 tracheal
bundles of the thoracic horn are partially joined by a narrow ‘‘bridge’’ of
tracheoles in several species, and in at least 2 species only 1 tracheal bundle
is present. Both of these conditions can vary from one side to the other in
the same specimen. The ventral accessory setae on S VI have apparently
been lost in the californicus complex and the septemmaculatus group. This
apparent lack of setae may be a result of the small sample examined in the
septemmaculatus group; it is also possible that some populations of a single
species lack these setae. This last situation is apparent in several other species
(D. candidibasis, D. lobiger, D. modestus). However, I have examined
hundreds of specimens of the californicus complex and have never observed
ventral accessory setae on them.
A 5. Acrostichal setal number reduced (a); normal (p) [A].
6. Frontal apotome with large ventral pit present (a); frontoclypeal
apotome with large ventral pit present (p) [L].
7. Superior volsella without heavily sclerotized apical projection (a); with
heavily sclerotized apical projection (p) [A].
a. S III with needlelike setae [P].
This furcation separates D. lobiger from the remaining species in the
genus. The presence of a frontoclypeal apotome is plesiomorphous through-
JOHN H. EPLER 197
out the Chironominae and apparently in the sister-group, the Orthocladiinae
(Cranston, et al. 1983). The presence of a large ventral pit may indicate that
Einfeldia species group A of Pinder & Reiss (1983) may be the ‘‘closest’’
sister-group to Dicrotendipes. Einfeldia is in drastic need of revision and is
probably polyphyletic.
The needlelike setae on pupal S III are also found in the D. modestus
group (furcation U), and in some species in at least one genus in the
outgroup, Kiefferulus. Kiefferulus is also in need of revision. This character
may be an underlying synapomorphy for the group of genera closely related
to Dicrotendipes.
B 8. Frontal apotome with frontal process (a); frontal apotome with large
ventral pit (p) [L].
9. Posterior margin of T V with 2 groups of hooklets (a); without
hooklets (p) [P].
37. Shagreen on T VI broadly V-shaped (a); shagreen quadrilateral (p)
[P].
Furcations A and B define the 3 main groups (I, IJ and III, Fig. 52) of
species in the genus based on larval and pupal characters. Although other
genera in the outgroup have larvae which also possess frontal or fronto-
clypeal apotomes with variously sized ventral pits, those present in Dicro-
tendipes are shaped differently and are often located in slightly different
areas. The presence of hooklets at the posterior margin of T V is unique
among the group of genera.
C*10. Mentum with 6th tooth fused/appressed to Sth (a); mentum without
fused/appressed teeth (p) [L].
11. Superior volsella digitiform with ventrally directed membranous apex
(a); digitiform without membranous apex (p) [A].
b. T IX with dorsal ovoid area [A].
Apomorphy 10 defines the /eucoscelis-group. A somewhat similar mental
appression/fusion occurs in 2 Nearctic members of the nervosus group.
Apomorphy 11 is distinctive for the conjunctus-group.
The dorsal ovoid area on T IX is a symplesiomorphy shared with many
species of Chironomus, Einfeldia and Kiefferulus. Its presence on the clado-
gram is only for reference purposes. In addition to the 3 species of the
conjunctus group, this character state is also found in D. pelochloris in the
pelochloris group. It occurs sporadically throughout the Chironomus group.
D 12. Acrostichal setae absent (a); present (p) [A].
13. Inferior volsella with extremely wide apex (a); normal apex (p) [A].
c. Loss of hooklets on posterior margin of T V [P].
MEM. AMER. ENT. SOC., 36
198 GENUS DICROTENDIPES
Apomorphies 12 and 13 are autapomorphies for D. cumberlandensis,
which has apparently undergone a reversal with respect to the posterior
hooklets on T V.
E 14. Less than 30 ventromental striae (a); more than 30 striae (p) [L].
15. Superior volsella strongly arched mediad (a); weakly arched or almost
straight (p) [A].
d. Palmate sensilla chaetica present on hind metatarsus [A].
Apomorphies 14 and 15 are autapomorphies for D. pseudoconjunctus.
This species is one of 2 species known outside of the Neotropical region in
which palmate sensilla chaetica are present on the hind metatarsus of the
male. These sensilla are also present in males of Chironomus, Einfeldia,
Kiefferulus and Nilodorum. Their presence may be an underlying synapo-
morphy for the ‘‘Chironomus group.’’ More data are needed.
F*16. Head capsule with grainy integument (a); without grainy integument
(p) [L].
17. Superior volsella stout, with expanded apex (a); superior volsella
digitiform (p) [A].
18. Anal point wide and strongly deflexed (a); anal point normal (p) [A].
Furcation F splits the /ewcoscelis and jonmartini groups from the pelo-
chloris group.
G 19. Mentum with 6th, Sth and 4th lateral teeth fused/appressed (a); with
only 6th and Sth lateral teeth fused/appressed (p) [L].
e. Palmate sensilla chaetica present on hind metatarsus [A].
f. T LX with dorsal ovoid area [A].
g. Loss of spines on posterior margin of T V [P].
Apomorphy 19 is an autapomorphy for D. kribiicola. Apparently 3 rever-
sals have taken place with D. pelochloris. It and D. pseudoconjunctus possess
palmate sensilla chaetica on the male hind metatarsus (see above). The dorsal
ovoid area on T IX is also found on the conjunctus group and in the
outgroup, where it occurs sporadically. D. pelochloris has apparently lost
the posterior hooklets on T V.
H 20. Inferior volsella with membranous dorsal extension (a); without (p)
[A].
I*21. Posterior margin of labral sclerite 1 with low tubercles (a); labral
sclerite 1 smooth (p) [L].
*22. T VIII with 4 lateral lamellar setae (a); with 5 setae (p) [P].
23. T V hooklets in continuous row (a); in 2 groups (p) [P].
24. Frontal apotome with reduced ventral pit (a); with large ventral pit
(p) [L].
h. Mentum normal.
Apomorphy 21 is autapomorphic (within the /eucoscelis group) for D.
notatus. This character state is also found in the modestus-fumidus groups.
JOHN H. EPLER 199
Apomorphies 22-24 define the jonmartini group. Placement of this group
is somewhat arbitrary, and is based mainly on the assumption that the 2
posterior groups of hooklets on T V transformed to the single band found
in this group, and that the large frontal pit has been reduced to the faint,
wide frontal pit found in jonmartini and sarinae. Based on superior volsella
morphology, the 2 unusual species with median volsellae (D. balciunasi and
D. lindae) may belong here, but without the immature stages it is not possi-
ble to accurately place them.
I must also assume here that the apomorphic state of 4 lateral lamellar
setae on T VIII has evolved twice in the genus, because the character state
occurs in 2 different lineages. All other species with 4 setae also possess a
frontal projection, sometimes surrounded by a pit. This pit is not similar
to, and probably not homologous to, the weak pit found in the jonmartini
group. I believe it is more likely that the pupa would lose a seta and add a
few posterior hooklets on T V (as in the jonmartini group), than for the
posterior band of hooklets to have arisen twice within the genus.
The presence of a membranous dorsal extension (apomorphy 20), similar
to that found on the Palaearctic D. notatus, may ally the jonmartini group
with the /eucoscelis group. Both groups also share the larval head capsule
with a grainy integument, a character which has arisen at least 2 other times
in the genus (fumidus and the californicus complex). Placement of the
jonmartini group is to be considered tentative. It is also possible that the
jonmartini group may be more closely allied with the conjunctus group, for
both groups are exclusively Australian. More data are needed, as the
jonmartini-sarinae sample size was small (n = 6 larvae and 3 pupae).
J 25. Superior volsella pediform, apex directed mediad (a); superior volsella
digitiform (p) [A].
26. Less than 30 ventromental striae (a); more than 30 striae (p) [L].
Apomorphies 25 and 26 are autapomorphies for D. jonmartini and D.
Sarinae, respectively.
K*27. T VIII with 4 lateral lamellar setae (a); with 5 setae (p) [P].
28. Anal lobe with shagreen (a); without shagreen (p) [P].
29. Wings with spots (a); wings immaculate (p) [A].
30. Apex of inferior volsella deeply bifid (a); apex clubbed,emarginate-
cordiform (p) [A].
31. Superior volsella mostly bare, long cylindrical, curved mediad, with
moderately sclerotized apex (a); superior volsella digitiform (p) [A].
i. Loss of S VI ventral accessory setae [A].
Apomorphies 28-31 define the septemmaculatus group. I have not
observed S VI ventral accessory setae on any members of this group. The
MEM. AMER. ENT. SOC., 36
200 GENUS DICROTENDIPES
superior volsellar type is unique to this group. Furcations L, M and N split
the group into the 4 species whose immature stages are known.
L*32. Mentum with some lateral teeth fused (a); mentum without fusions
(p) [L].
j. More than 50 ventromental striae [L].
M 33. Hypopygium with accessory lobes (a); without lobes (p) [A].
N*34. Mandible with lateral teeth modifications (a); without modifications
(p) [L].
35. Wing with clouds along veins (a); wing with spots (p) [A].
Apomorphies 32-35 are autapomorphies for species within the septem-
maculatus group.
O*36. Posterior margin of labral sclerite 1 with low tubercles (a); labral
sclerite 1 smooth (p) [L].
38. Superior volsella cylindrical, with membranous apex (a); superior
volsella cylindrical, without membranous apex (p) [A].
39. T V with separate anterolateral shagreen areas (a); without separate
areas (p) [P].
Furcation 0 is the splitting point for the 2 main lineages with 4 lateral
setae on pupal T VIII (with the exception of the jonmartini group, which
is apparently more closely related to the conjunctus-leucoscelis groups). This
refers only to those species whose larvae are known. The Neotropical forms
with bifid inferior volsellae may also belong here, for the pupae of the two
known species have 4 lateral setae on T VIII and the shagreen on T VI is
broadly V-shaped. These Neotropical species probably are a sister group to
the nervosus group.
P*40. Ist and 2nd lateral teeth of mentum fused (a); not fused (p) [L].
41. Proximal inner tooth of mandible modified (a); unmodified (p) [L].
Q 42. Superior volsella deltoid (a); superior volsella cylindrical with
membranous apex (p) [A].
Furcations P and Q are to be considered arbitrary, due to the presence
of D. lobus. This species is difficult to place due to its unusual superior
volsella. The V-shaped shagreen on T VI indicates that this species belongs
with the nervosus group. I have placed D. lobus closer to D. candidibasis
and D. flexus (furcation Q) because these 3 species share an apomorphy,
the fusion of the Ist and 2nd lateral teeth of the mentum. However, this
character is apparently homoplasious, for it appears again in the modestus
group (D. neomodestus) and in the septemmaculatus group. It does appear
that the fusion in /Jobus, candidibasis and flexus is not similar or homologous
to the fusion found in the other groups; the fusion of the teeth is more
complete in /obus, etc. Obviously, more data are needed here.
R 43. Mentum with median tooth sunken well beneath level of Ist lateral
teeth (a); median tooth subequal to Ist lateral teeth (p) [L].
JOHN H. EPLER 201
44. Wing with bands/spots (a); wing immaculate (p) [A].
*45. Apodeme lobe with few, weak microtrichia (a); with numerous, well
developed microtrichia (p) [A].
46. Cephalic tubercles minute (a); well developed (p) [P].
S 47. Superior volsella rotated 90° around longitudinal axis (a); not rotated
(p) [A].
Apomorphies 43-47 are autapomorphies for species within the nervosus
group; 43-45 define candidibasis, 46 flexus, 47 the lucifer complex.
T 48. Superior volsella cylindrical with lightly sclerotized apex (a); with
membranous apex (p) [A].
*49. Head capsule with grainy integument (a); without grainy integument
(p) [L].
*k. S III with needlelike setae [P].
50. Apotome with small frontal pit surrounding frontal projection (a);
with frontal projection only (p) [L].
Apomorphies 48 and 49 are apomorphic for D. fumidus, a species inter-
mediate in characters between the modestus and nervosus groups. Apomor-
phy 50 is a synapomorphy for the modestus group. All species in the
modestus group also have needlelike spines on pupal S III, a character also
found in D. lobiger and in the genus Kiefferulus. Similar groups of spines
are also found on at least one genus outside of the outgroup, Cladopelma.
This character is not used in this analysis, and is placed on the cladogram
for informational purposes only.
U*51. Apodeme lobe with few, weak microtrichia (a); with many, well
developed microtrichia (p) [A].
52. Superior volsella pediform (a); cylindrical (p) [A].
Apomorphy 51 delimits the species D. tritomus in the modestus group.
This species does not have a pediform superior volsella (apomorphy 52),
but possesses a superior volsella somewhat similar to that of D. nervosus.
However, closer inspection reveals that the volsellae of the 2 species are not
similar (Epler 1987a). Larvae and pupae of D. tritomus are almost identical
to D. modestus, and often are inseparable without associated adults.
V 53. Caudolateral spurs on T VIII located anterior to posterior corner (a);
spurs located on posterior corner (p) [P].
*54. Head capsule with grainy integument (a); without grainy integument
(p) [L].
*55. Anal lobe with shagreen (a); without shagreen (p) [P].
56. Fewer than 20 ventromental striae (a); more than 20 ventromental
striae (p) [L].
*57. 1st and 2nd lateral teeth of mentum fused (a); normal (p) [L].
Apomorphies 53 and 54 are apomorphies for the D. californicus complex.
MEM. AMER. ENT. SOC., 36
202 GENUS DICROTENDIPES
Apomorphies 55 and 56 are autapomorphic for D. thanatogratus. Apomor-
phy 57, a homoplasy, is autapomorphic within the modestus group for D.
neomodestus. There is no unambiguous way to delineate the cladogenesis
of these species with current data.
The cladistic analysis indicates that the species fall into the following 9
groups (denoted by numbers above the species names in the cladogram, Fig.
52): 1) lobiger group: lobiger; 2) conjunctus group: conjunctus, cumber-
landensis, pseudoconjunctus; 3) pelochloris group: kribiicola, pelochloris;
4) leucoscelis group: leucoscelis, notatus; 5) jonmartini group: jonmartini,
sarinae; 6) septemmaculatus group: fusconotatus, pallidicornis, septem-
maculatus, sudanicus; 7) nervosus group: candidibasis, flexus, lobus, lucifer,
nervosus; 8) fumidus group: fumidus; 9) modestus group: californicus,
tritomus, modestus, neomodestus, thanatogratus.
Epler (1987c) used the computer program PAUP (Swofford 1985) as a
supplemental tool in phylogenetic analysis. The PAUP analysis basically
agreed with the Hennigian cladistic analysis. PAUP constructs trees utilizing
the Wagner method and a maximum parsimony algorithm, i.e., the tree
with the fewest steps (character state transformations) is the most parsi-
monious. If one assumes that the Hennigian cladogram that invokes the
smallest number of hypotheses to deal with homoplasy (convergence/paral-
lelism: the occurrence of similar character states in species which do not
share an immediate common ancestor possessing that character state) is the
most parsimonious, trees constructed from the same data should yield similar
results (Swofford 1985). Some workers contend that the use of Wagner trees
and computer programs reduces bias and promotes objectivity (Farris 1985;
see also Saether 1986). However, one begins to introduce bias when the
characters for the analysis are chosen (all the potential characters of an
organism are not used), and, as Saether (1986:3) stated: ‘‘there is no biolog-
ical evidence that minimum length trees or equally parsimonious trees are
most in accordance with the ‘‘true’’ tree.’’ To invoke strict parsimony to
explain evolutionary relationships could mean that evolution as a process
“‘knows’’ the shortest path to take to reach a given goal. Hennig (1966) did
not advocate the use of parsimony to resolve conflicts in a cladogram, but
instead recommended that the full set of characters be reexamined to deter-
mine if characters had been misinterpreted. I believe that it is the taxonomic
precision of the worker and his/her knowledge of the group(s) in question
that can decide the usefulness and ‘‘correctness’’ of the hypothesized
cladogram(s).
The 3 major lineages (I, II and III in Fig. 52) could be considered subge-
nera. However, these are based chiefly on characters of the immature stages
(mainly the larval frontal/frontoclypeal apotome). It is impossible in many
——
JOHN H. EPLER 203
cases to place adults in subgenera, and the majority of Dicrotendipes species
are known only as adults. For this reason I am not establishing subgenera
at this time.
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Fig. 52: Cladogram depicting the hypothesized relationships within
Dicrotendipes.
MEM. AMER. ENT. SOC., 36
204 GENUS DICROTENDIPES
It is possible to subjectively place some species in some species groups.
Dicrotendipes adnilus, D. sinoposus and the other 3 members of the califor-
nicus group (D. embalsensis, D. obrienorum, D. pellegriniensis) belong in
the modestus group; D. peringueyanus to the septemmaculatus group; D.
balciunasi and D. lindae to the jonmartini group; D. aethiops, D. freemani,
D. chambiensis, D. inouei and D. tamaviridis in the nervosus group. The
majority of the Amazonian species described in Chapter III probably repre-
sent another lineage near or derived from the nervosus group, based on the
pupal characters of D. fittkaui and D. soccus.
The discovery of the presently unknown immature stages of most Dicro-
tendipes species, especially the Afrotropical and Neotropical species, and
revisions of the genera in the Chironomus group will greatly improve the
taxonomy of the genus.
ACKNOWLEDGEMENTS
I am especially thankful to Drs. W.L. Peters, M.L. Pescador and R.W.
Flowers for providing laboratory space and equipment, and minor funding,
despite the attitude of some administrators towards basic taxonomic re-
search at Florida A&M University. Dr. A.R. Soponis is due special thanks
for her friendship, advice, assistance and critical review of an earlier version
of this manuscript. Drs. W.H. Heard and G.J. Wibmer were extremely help-
ful to me by reviewing portions of this manuscript. I wish to thank Drs.
C.W. and L.B. O’Brien (Florida A&M University) and Dr. P.H. Langton
(March, Cambridgeshire, Great Britain) for the gift and loan of specimens.
Mrs. Evelyn Sellars is also due my thanks for the assistance she has rendered.
I wish to express my gratitude to those individuals listed in the Meth-
odology section who made specimens available to me. Several went above
and beyond the call of duty: Dr. P.S. Cranston made acquisition of type
specimens from the British Museum (Natural History) a painless procedure
and reviewed portions of the manuscript; Drs. E.J. Fittkau and F. Reiss
were extremely helpful and most hospitable during my all too brief visit at
the Zoologische Staatssammlung, Munich; Dr. L. Hare provided extremely
important Afrotropical material and was instrumental in unraveling the mess
concerning Carteronica; and Dr. J. Martin provided the bulk of associated
Australian material, supplied valuable taxonomic information and reviewed
portions of the manuscript. Mr. B.A. Caldwell provided valuable comments
and performed trial runs on my keys. I am grateful to Dr. S.S. Roback for
his assistance in publishing this monograph.
Special thanks are due Dr. H.W. Barry Merrill and his wife Judy, Merrill
Consultants, Dallas, TX, for their assistance in bringing this monograph to
publication.
JOHN H. EPLER 205
Above all, Iam extremely grateful and forever thankful to my wife Linda,
whose secretarial and organizational skills, encouragement, support and love
have made this whole project a reality.
LITERATURE CITED
Ault, A. & M.S. MULLA, 1980. Activity of organophosphate and synthetic pyrethroid insecticides
against pestiferous midges in some southern California flood control channels. Mosq.
News 40:593-597.
ARISTOVSKAYA, G.V. 1935. Materialy k faune khironomid vodoemov Tatarespubliki. Trudy
Tatarsk. Otdel. Vses n. — issl. inst. ozerno-rechn. rybn. Khoz. 2:109-158.
ASHE, P. 1983. A catalogue of chironomid genera and subgenera of the world including syn-
onyms (Diptera:Chironomidae). Ent. scand. Suppl. 17:1-68.
-D.A. Murray & F. REISs. 1987. The zoogeographical distribution of Chiron-
omidae (Insecta:Diptera). Annls. Limnol. 23:27-60.
BEcK, W.M., Jr. 1977. Environmental requirements and pollution tolerance of common fresh-
water Chironomidae. Environmental Monitoring Series, U.S.E.P.A., 261 pp.
BECKER, T. 1908. Dipteren der Kanarischen Inseln. Mitt. zool. Mus. Berl. 4:1-180.
Brown, J.H. & A.C. GIBSON. 1983. Biogeography. C.V. Mosby Co., St. Louis, Mo. 643 pp.
BRUNDIN, L. 1966. Transantarctic relationships and their significance, as evidenced by chiron-
omid midges. With a monograph of the subfamilies Podonominae and Aphroteniinae
and the austral Heptagyiae. K. svenska Vetensk Akad. Handl. 11:1-472.
CHAUDHURI, P.K. & D.K. GUHA. 1987. A conspectus of chironomid midges (Diptera: Chiron-
omidae) of India and Bhutan. Ent. scand. Suppl. 29:23-33.
CHERNOVSKII, A.A. 1949. Opredelitel lichinok komarov semeistva Tendipedidae. Izd. Akad.
Nauk. SSSR 31:1-186.
COFFMAN, W.P. & L.C. FERRINGTON, JR. 1984. Chironomidae, pp. 551-652. In: Merritt, R.W.
& K.W. Cummins (eds.): An Introduction to the Aquatic Insects of North America.
Second Edition. Kendall/Hunt Publishing Co., Dubuque, Iowa. 772 pp.
CONTRERAS-LICHTENBERG, R. 1986. Revision der in der Westpaldarktis verbreiteten Arten des
Genus Dicrotendipes Kieffer, 1913 (Diptera, Nematocera, Chironomidae). Ann. Na-
turhist. Mus. Wien 88/89:663-726.
CRANSTON, P.S. & P.D. ARMITAGE. 1988. The Canary Islands Chironomidae described by T.
Becker and by E. Santos Abreu. Dt. ent. Z. (in press).
.D.R. OLIVER & O.A. SAETHER. 1983. The larvae of Orthocladiinae (Dip-
tera:Chironomidae) of the Holarctic region — Keys and diagnoses. Ent. scand. Suppl.
19:149-291.
.R.D. TEE, P.F. CREDLAND & A.B. Kay. 1983. Chironomid haemoglobins: Their
detection and role in allergy to midges in the Sudan and elsewhere. Mem. Amer. Ent.
Soc. 34:71-87.
Darpy, R.E. 1962. Midges associated with California rice fields, with special reference to their
ecology (Dipt., Chir.). Hilgardia 32:1-206.
DEJoux, C. 1968. Contribution a l’étude des insects aquatiques du Tchad. Cah. O.R.S.T.O.M.,
ser. Hydrobiol., II:51-78.
.1970. Contribution a |’étude des premiers états des Chironomides du Tchad
(Insectes, Diptéres) (3e Note). — Description comparée des nymphes de Chironomus
(Nilodorum) brevibucca, Ch. (N.) brevipalpis et Ch. (N.) fractilobus. Bull. Mus. natn.
Hist. Nat. Paris 42:175-184.
MEM. AMER. ENT. SOC., 36
206 GENUS DICROTENDIPES
.1977. Chironomides du Lac de Bam (Haute-Volta). Cah. O.R.S.T.O.M., ser.
Hydrobiol. X1I:291-295.
.1984. Contribution to the knowledge of West African Chironomidae (Diptera —
Nematocera). Chironomids from the Guinean Republic. Aquatic Insects 6:157-167.
Disney, R.H.L. 1975. Notes of crab-phoretic Diptera (Chironomidae and Simuliidae) and their
hosts in Cameroon. Ent. Monthly Magazine 111:131-136.
EDWARD, D.H.D. 1964. The biology and taxonomy of the chironomids of southwestern Aus-
tralia. Ph.D. Thesis, University of Western Australia.
EDWARDS, F.W. 1924. New species of nematocerous Diptera from Fiji and Trinidad. Ann. Mag.
Nat. Hist. Ser. 9, 14:568-574.
.1928. Insects of Samoa. Part VI. Fasc. 2. Nematocera, pp. 23-102. Jn: Insects
of Samoa and Other Samoan Terrestrial Arthropoda. London:BM(NH). 1-108.
.1929. British non-biting midges (Diptera, Chironomidae). Trans. R. ent. Soc.
Lond. 77:279-430.
.1931. Diptera of Patagonia and South Chile. Part II. Fascicle 5. Chironomidae,
pp. 233-331. Trustees of the British Museum, London.
EPLerR, J.H. 1983. Taxonomic revision of the Nearctic Dicrotendipes Kieffer, 1913 (Dip-
tera:Chironomidae). M.Sc. Thesis, Florida State University, Tallahassee, FL, 283 pp.
.1987a. Revision of the Nearctic Dicrotendipes Kieffer, 1913 (Diptera: Chiron-
omidae). Evol. Monogr. 9: 102 pp + 37 plates.
.1987b. Notes on the Dicrotendipes (Diptera: Chironomidae) of Mexico, with
descriptions of two new species. Ent. scand. Suppl. 29:147-154.
.1987c. Biosystematics of the genus Dicrotendipes Kieffer, 1913 (Diptera: Chi-
ronomidae) of the World. Ph.D. dissertation, Florida State University, Tallahassee, 316
+ Xlv pp.
FARRIS, J.S. 1985. The pattern of cladistics. Cladistics 1:190-201.
FITTKAU, E.J. 1962. Die Tanypodinae (Diptera, Chironomidae). Die Tribus Anatopyniini, Ma-
cropelopiini und Pentaneurini. Abh. Larvalsyst. Insekten 6:1-453.
.1980. Ein Zoogeographischer Vergleich der Chironomiden der Westpalaearktis
und der Aethiopis. Jn: Murray, D.A. (Ed.): Chironomidae pp. 139-143.
.& F. Reiss. 1978. Chironomidae, pp. 404-440. In: Illies, J. (Ed.): Limnofauna
Europaea, 2 Aufl. G. Fischer Verlag, Stuttgart.
.& F. Reiss. 1979. Die zoogeographische Sonderstellung der neotropischen Chi-
ronomiden (Diptera). Spixiana 2:273-280.
ForsyTH, D.J. & I.D. MCCALLUM. 1978. Xenochironomus canterburyensis (Dip-
tera:Chironomidae) an insectan inquiline commensal of Hyridella menziesi (Mollusca:
Lamellibranchia). J. Zool., Lond. 186:331-334.
FREEMAN, P. 1954a. East African Chironomidae and Ceratopogonidae (Dipt.). Arch. Hydro-
biol. 48:441-446.
.1954b. Chironomidae (Diptera) from Western Cape Province-III. Proc. Roy.
Ent. Soc. Lond. (B) 23:17-25.
.1955a. Chironomidae (Diptera Nematocera). Explor. Parc natn. Albert Miss.
G.F. de Witte 83:3-41.
.1955b. Diptera (Nematocera) Chironomidae. South African Animal Life. Br.
Mus. Nat. Hist. 2:361-381.
.1957. A study of the Chironomidae (Diptera) of Africa south of the Sahara.
Part III. Bull. Br. Mus. nat. Hist., Ent. 5:321-426.
.1959. A study of the New Zealand Chironomidae (Diptera, Nematocera). Bull.
Br. Mus. Nat. Hist., Ent. 7:395-437.
.1961a. The Chironomidae (Diptera) of Australia. Aust. J. Zool. 9:611-737.
JOHN H. EPLER 207
.1961b. A collection of Chironomidae and Culicidae subfamily Dixinae (Diptera,
Nematocera) from Madagascar. Mem. Inst. Scient. Madagascar, Ser E 12:237-274.
.& P.S. Cranston. 1980. Family Chironomidae, pp. 175-202. In: Crosskey, R.W.
(Ed.): Catalogue of the Diptera of the Afrotropical Region. British Museum (Nat. Hist.),
London, 1,437 pp.
FROMMER, S.I. & P.A. RAUCH. 1971. Pupal duration, adult emergence and oviposition periods
for the midge Dicrotendipes californicus (Johannsen) (Diptera: Chironomidae). Calif.
Vector Views 18:33-39.
GOETGHEBUER, M. 1928. Diptéres (Nématocéres). Chironomidae. II]. Chironomariae. Faune
Fr. 5:1-174.
.1930. Chironomides Palearctiques (Diptéres) conservés au Musée d’ Histoire Na-
turelle de Vienne. Annls. hist.-nat. Mus. Wien 46:91-115.
.1934. Ceratopogonides et Chironomides du Congo Belge (deuxiéme note). Re-
vue Zool. Bot. afr. 25:191-205.
.1936. Chironomides du Congo Belge. Revue Zool. Bot. afr. 28:453-492.
.1937-1954. Tendipedidae (Chironomidae) b) Subfamilie Tendipedinae (Chiron-
ominae). A. Die Imagines, pp. 1-138. Jn: Lindner, E. (Ed.): Die Fliegen der palaeark-
tischen Region 13c.
Guna, D.K. & P.K. CHAUDHURI. 1981. Record of three genera of Chironominae (Chiron-
omidae:Diptera) from India. Bull. zool. Surv. India 3:159-165.
GuHA, D.K., P.K. CHAUDHURI & S.K. NANDI. 1982. Taxonomic studies of Chironominae (Chi-
ronomidae:Diptera) from West Bengal: Genus Dicrotendipes Kieffer. Proc. zool. Soc.
Calcutta 33:29-38 (1980).
GuHA, D.K., S.K. DAs, P.K. CHAUDHURI & D.K. CHOUDHURI. 1985. Chironomid midges of
the Andaman islands (Diptera:Chironomidae). Proc. Nat. Acad. Sci. India 55(B):22-38.
HAMILTON, A.L., O.A. SAETHER & D.R. OLIVER. 1969. A classification of the nearctic Chi-
ronomidae. Fish. Res. Bd Can. Tech. Rep. 124:42 pp.
Hare, L. & J.C.H. CARTER. 1987. Chironomidae (Diptera, Insecta) from the environs of a
natural West African lake. Ent. scand. Suppl. 29:65-74.
HASHIMOTO, H. 1984. A halophilous chironomid, Dicrotendipes inouei n. sp. (Dip-
tera:Chironomidae). Bull. Fac. Ed., Shizuoka Univ. Nat. Sci. Ser. 35:45-51.
-T. WONGSIRI, N. WONGSIRI, C. TIRAWAT, A. LEWVANICH & K. YASUMATSU. 1981.
Chironominae from rice fields of Thailand with descriptions of 7 new species. Taxonomy
Branch, Entomology & Zoology Division, Dept. Agr., Bangkok, Tech. Bull., 7:1-47.
Hauser, U.A. & T. MorRISSEY. 1945. Limnochironomids in Iowa including their life-histories.
Proc. Iowa Acad. Sci. 52:287-292.
HENNIG, W. 1966. Phylogenetic Systematics. University of Illinois Press, Urbana. 263 pp.
HOFMANN, W. 1971a. Die postglaziale Entwicklung der Chironomiden- und Chaoborus- Fauna
(Dipt.) des Schohsees. Arch. Hydrobiol. Suppl. 40:1-74.
.1971b. Zur Taxonomie und Pal6kologie subfossiler Chironomiden (Dipt.) in
Seesedimenten. Arch. Hydrobiol., Beih. 6:1-50.
.1978. Analysis of animal microfossils from the Grosser Segeberger See (F.R.G.).
Arch Hydrobiol. 82:316-346.
Hupson, P.L. 1987. Unusual larval habitats and life history of chironomid (Diptera) genera.
Ent. scand. Suppl. 29:369-373.
INTERNATIONAL CODE OF ZOOLOGICAL NOMENCLATURE. 3rd Edition adopted by the XX General
Assembly of the International Union of Biological Sciences. 1985. Ride, W.D.L., Sa-
brosky, C.W., Bernardi, G. & Melville, R.V. (eds.). International Trust for Zoological
Nomenclature, London. xx + 338 pp.
MEM. AMER. ENT. SOC., 36
208 GENUS DICROTENDIPES
JOHANNSEN, O.A. 1905. Aquatic nematocerous Diptera, pp. 76-327. In: Needham, J.G., K.I.
Morton & O.A. Johannsen (Eds.): Mayflies and midges of New York. Bull. N.Y. St. Mus.
86:76-327.
. 1932. Chironominae of the Malayan Subregion of the Dutch East Indies. Arch.
Hydrobiol. Suppl. 11:503-552.
KEILBACH, R. 1982. Bibliographie und Liste der Arten tierischer Einschliisse in fossilen Harzen
Sowie ihrer Aufbewahrungsorte. Dt. ent. 2, N.F. 29:301-491.
KIEFFER, J.J. 1906. Diptera Fam. Chironomidae. Jn: Wytsman, P. (Ed.): Genera insectorum
42:78 pp. (reference not seen).
.1910. Etude sur les Chironomides des Indes Orientales, avec description de quel-
ques nouvelles espéces d’Egypte. Mem. Indian Mus. 2:181-242.
.191la. Descriptions de nouveaux Chironomides de |’Indian Museum de Cal-
cutta. Rec. Indian Mus. 6:113-177.
.1911b. Diptera, Chironomidae der Seychellen-Inseln, aus der Sammlung von
Mr H. Scott. Trans. Linn. Soc. Lond., 2nd Ser. Zool. 14:331-366.
.1912. Tendipedidae (Chironomidae) [Dipt.], pp. 27-43. In: Formosa-Ausbeute.
Supplta. ent. 1.
.1913a. Nouvelle étude sur les Chironomides de 1’Indian Museum de Calcutta.
Rec. Indian Mus. 9:119-197.
.1913b. Chironomidae et Cecidomyidae. Résult. scient. Voy. Ch. Alluaud et R.
Jeannel en Afrique orientale (1911-1912) (Dipt.) I:1-43.
.1914. South African Chironomidae (Diptera). Ann. S. Afr. Mus. 10:259-270.
.1916. Tendipedidae de Formose conservés au Museum National Hongrois de
Budapest et déterminés par J.J. Kieffer. Ann. Mus. Nat. Hung. 14:81-121.
.1917. Chironomides d’ Australie conservés au Musée National Hongrois de Bu-
dapest. Annls. hist.-nat. Mus. natn. hung. 15:175-228.
.1918. Chironomides d’Afriques et d’Asie conservés au Museum National Hon-
grois de Budapest. Annls. hist.-nat. Mus. natn. hung. 16:31-139.
.1920. Tableau synoptique suivant des Chironomides paléarctiques appartenant
aux genres Polypedilum et Limnochironomus. Annls. Soc. scient. Brux. 39:159-167.
.1921a. Synopse de la tribu des Chironomariae (Diptéres). Annls. Soc. scient.
Brux. 40:269-276.
.1921b. Chironomides des Philippines et de Formose. Philipp. J. Sci. 18:557-
593.
.1922. Chironomides de l’Afrique Equatoriale (2e partie). Annls Soc. ent. Fr.
91:1-72.
.1923. Chironomides de |’Afrique Equatoriale (3e partie). Annls Soc. ent. Fr.
92:149-204.
.1924. Six nouveaux Chironomides d’Afrique. Annls. Soc. scient. Brux. 43:255-
261.
.1925. Chironomides d’Egypte (Dipt.). Bull. Soc. ent. Egypte 8:244-313.
KRUSEMAN, G. 1949. Note on Tendipedidae of the Suez Canal. Bijdr. Dierkunde 28:249-254.
LANGTON, P.H. 1984. A key to the pupal exuviae of British Chironomidae. P.H. Langton,
March, Cambridgeshire, 324 pp. (Private publication.)
LENZ, F. 1937. Chironomarie aus Niederlandisch-Indien. Larven und Puppen. Arch. Hydro-
biol. Suppl. 15:1-29.
.1954-1962. Tendipedidae (Chironomidae). b) Subfamilie Tendipedinae (Chi-
ronominae). B. Die Metamorphose der Tendipedinae, pp. 139-260. Jn: Lindner, E. (ed.):
Die Fliegen der palaearktischen Region, 13c.
JOHN H. EPLER 209
ManrtTIn, J. 1961. Chromosomal polymorphism in Victorian chironomids. M.Sc. Thesis, Uni-
versity of Melbourne, 224 pp.
OLIVER, D.R. 1981. Chironomidae, pp. 423-458. In: McAlpine, J.F., B.V. Peterson, G.E.
Shewell, H.J. Teskey, J.R. Vockeroth & D.M. Wood (coordinators): Manual of Nearctic
Diptera, Vol. 1. Agric. Canada Monog. 27, 674 pp.
Pacai, A.C. 1975. Formas imaginales y preimaginales de Chironomus (Dicrotendipes) alsi-
nensis sp. nov. Neotropica 21:149-156.
.1978. Formas imaginales y preimaginales de quironomidos (Diptera) IV. Dicro-
tendipes nestori sp. nov. Limnobios 1:235-241.
.1987. Formas imaginales y preimaginales de Quironomidos (Diptera Chiron-
omidae) VI. Dicrotendipes pellegriniensis sp. nov. y D. embalsensis sp. nov. Limnobios
2:695-706.
PIELOU, E.C. 1979. Biogeography. John Wiley & Sons, New York, N.Y. 351 pp.
PINDER, L.C.V. 1978. A key to the adult males of the British Chironomidae (Diptera), the non-
biting midges. Vols. 1, 2. Freshw. Biol. Assoc. Sci. Publ. 37:169 pp, 189 figs.
.& F. REISS. 1983. The larvae of Chironominae (Diptera: Chironomidae) of the
Holarctic region. Keys and diagnoses. Ent. scand. Suppl. 19:293-435.
PRAT, N. 1981. Quironémidos de Catalunya. (2.a nota). Mediterranea 5:43-66.
REE, H.I. 1981. Studies on Korean Chironomidae (Diptera). 2. Description of a new genus
and a new species of Chironomidae. Korean J. Zool. 24:217-220.
REISS, F. 1972. Die Tanytarsini (Chironomidae, Diptera) Stidchiles und Westpatagoniens. Mit
Hinweisen auf die Tanytarsini-Fauna der Neotropis. Stud. Neotrop. Fauna 7:49-94.
.1974. Die in stehenden Gewdassern der Neotropis verbreitete Chironomidengat-
tung Goeldichironomus Fittkau (Diptera, Insecta). Stud. Neotrop. Fauna 9:95-122.
.1977a. Chironomidae. pp. 277-280. In: Hurlbert, S.H. (Ed.) Aquatic biota of
Southern South America, being a compilation of taxonomic bibliographies for the fauna
and flora of inland waters of Southern South America. San Diego, State Univ. California.
.1977b. Verbreitungsmuster bei paldarktischen Chironomidernarten (Diptera,
Chironomidae). Spixiana 1:85-97.
.1978. Sich abzeichnende Verbreitungsmuster in der palaarktischen — palaotro-
pischen Chironomidenfauna (Dipt.). Mitt. dtsch. Ges. allg. angew. Ent. 1:72-76.
.1982. Chironomidae. pp. 433-438. In: Aquatic Biota of Mexico, Central Amer-
ica and the West Indies, 1982, S.H. Hulbert & A. Villalobos — Figueroa (Ed.).
.1986. Ein Beitrag zur Chironomidenfauna Syriens (Diptera, Chironomidae). En-
tomofauna 7:153-168.
.& J.E. Sublette. 1985. Beardius new genus with notes on additional Pan-Amer-
ican taxa (Diptera, Chironomidae). Spixiana 11:179-193.
REMPEL, J.G. 1939. Neue Chironomiden aus Nordostbrasilien. Zool. Anz. 127:209-216.
ROBACK, S.S. 1957. The immature tendipedids of the Philadelphia area (Diptera: Tendipedi-
dae). Monogr. Acad. nat. Sci. Philad. 9:1-152 + 28 plates.
Ross, H.H. 1974. Biological Systematics. Addison-Wesley Publishing Co., Reading, MS. 345
pp.
SAETHER, O.A. 1977. Female genitalia in Chironomidae and other Nematocera:morphology,
phylogenies, keys. Bull. Fish. Res. Bd Can. 197:1-211.
. 1979. Underlying synapomorphies and anagenetic anaylsis. Zool. Scr. 8:305-
312.
.1980. A glossary of chironomid morphology terminology (Diptera:
Chironomidae). Ent. scand. Suppl. 14:1-51.
MEM. AMER. ENT. SOC., 36
210 GENUS DICROTENDIPES
.1983. The canalized evolutionary potential: Inconsistencies in phylogenetic rea-
soning. Syst. Zool. 32:343-359.
.1986. The myth of objectivity—post-Hennigian deviations. Cladistics 2:1-13.
SASA, M. 1981. Studies on the chironomid midges of the Tama River. Part 4. Chironomidae
recorded at a winter survey. Res. Rep. Nat. Inst. Environ. Stud. 29:79-148.
.1985. Studies on the chironomids collected from lakes in southern Kyushu (Dip-
tera: Chironomidae), pp. 25-97. In: Studies on chironomid midges of some lakes in Japan.
Res. Rpt. from Natl. Inst. for Environ. Studies, No. 83.
.& H. Hasegawa. 1983. Chironomid midges of the tribe Chironomini collected
from sewage ditches, eutrophicated ponds, and some clean streams in the Ryukyu Islands,
southern Japan. Jap. J. Sanit. Zool. 34:305-341.
SINGH, S. & A.K. KULSHRESTHA. 1977. Dicrotendipes rajasthani n. sp. from India (Diptera,
Chironomidae). Ent. scand. 8(3):233-235.
SKUSE, F.A.A. 1889. Diptera of Australia. Part VI. The Chironomidae. Proc. Linn. Soc. N.S.W.
(2)4:215-311.
SPAHR, U. 1985. Erganzungen und Berichtigungen zu R. Keilbachs Bibliographie und Liste der
Bernsteinfossilien — Ordnung Diptera. Stuttgarter Beitr. Naturk. 111:1-146.
STAEGER, C. 1839. Systematisk fortegnelser over de i Danmark hidtil fundne Diptera. Krojer;
Naturhist. Tidsskr. 2:549-600.
STRAND, E. 1928. Miscellanea Nomenclatoria zoologica et palaeontogica, I-II. Arch. Natur-
gesch. 92A:30-75.
SUBLETTE, J.E. 1964. Chironomidae (Diptera) of Louisiana I. Systematics and immature stages
of some lentic chironomids of west-central Louisiana. Tulane Stud. Zool. 11:109-150.
.& M.S. SUBLETTE. 1973. Family Chironomidae, pp. 389-422. In: Delfinado,
M.D. & Hardy, D.E. (Eds): Catalogue of the Diptera of the Oriental Region. Vol. I.
Nematocera. The University Press of Hawaii, Honolulu.
SWOFFORD, D.L. 1985. PAUP (Phylogenetic Analysis Using Parsimony) version 2.4.1, user’s
manual. D.L. Swofford, Illinois nat. Hist. Surv., 607 E. Peabody Dr., Champaign, IL
61820, U.S.A.
THIENEMANN, A. & J.J. KIEFFER. 1916. Schwedische Chironomiden. Arch. Hydrobiol. Plank-
tonk. Suppl. 2 (1921):483-554.
Townes, H.K. 1945. The nearctic species of Tendipedini (Diptera: Tendipedidae (= Chiron-
omidae)). Am. Midl. Nat. 34:1-206.
VARGAS, L. 1952. Tendipes (Limnochironomus) californicus y Tendipes (Limnochironomus)
figueroai n. sp. (Diptera, Tendipedidae). Rev. Soc. mex. Hist. nat. 13:47-51.
WALKER, F. 1856. Diptera Pt. V, pp. 415-464. In: Insecta Saundersiana, London.
WILEY, E.O. 1981. Phylogenetics. The Theory and Practice of Phylogenetic Systematics. John
Wiley & Sons, New York. 439 pp.
JOHN H. EPLER 211
APPENDIX 1
List of recognized species’ names in the genus Dicrotendipes, and their
distribution. (?) indicates a questionable species name. Abbreviations for
zoogeographical regions are as follows: NE = Nearctic, NT = Neotropical,
PA = Palaearctic, AF = Afrotropical, OR = Oriental, AU = Australian,
OC = Oceanian.
. adnilus Epler NE
. aethiops (Townes) NE
. amazonicus Epler NT
. alsinensis (Paggi) NT
. arcistylus Guha,
Das, Chaudhuri & Choudhuri OR
balciunasi Epler AU
bilobatus Kieffer AU
botaurus (Townes) NE
bredoi (Goetghebuer) AF
. californicus (Johannsen) NE, NT
candidibasis (Edwards) OC
canitibialis Guha,
Das, Chaudhuri & Choudhuri OR
chambiensis (Goetghebuer) AF
collarti (Goetghebuer) AF
conjunctus (Walker) AU
cordatus Kieffer AF
. crypticus Epler NE, NT?
cumberlandensis Epler AU
dasylabidus Epler NT
demissus Epler NT
ealae (Freeman) AF
embalsensis Paggi NT
. fittkaui Epler NT
flexus (Johannsen) OR, AU
freemani Epler AF
fumidus (Johannsen) NE
fusciforceps (Kieffer) (?) PA
fusconotatus (Kieffer) AF, PA
inouei Hashimoto PA
. jobetus Epler AU
Jonmartini Epler AU
. kribiicola (Kieffer) AF
. leei (Freeman) AU
. leucolabis Kieffer AF
. leucoscelis (Townes) NE
. lindae Epler AU
. lobiger (Kieffer) NE, PA
. lobus (Beck) NE
. lucifer (Johannsen) NE
milleri (Townes) NE
modestus (Say) NE, PA
neomodestus (Malloch) NE
nervosus (Staeger) NE, PA
nestori Paggi NT
nigrolineatus (Freeman) AF
notatus (Meigen) PA
obrienorum Epler NT
. palearivillosus Epler NT
pallidicornis Goetghebuer PA
. paradasylabidus Epler NT
paterjohni Epler NT
pellegriniensis Paggi NT
pelochloris (Kieffer) OR, PA, AU
peringueyanus Kieffer AF, PA
pseudoconjunctus Epler AU
radinovskyi Epler NT
reissi Epler NT
. sarinae Epler AU
schoutedeni (Goetghebuer) AF
semiviridis (Kieffer) (?) OR
septemmaculatus (Becker) PA, AF, OR,
AU
simpsoni Epler NE
. sinoposus Epler NT
soccus Epler NT
. sudanicus (Freeman) AF
tamaviridis Sasa PA
taylori (Freeman) AU
. tenuiforceps (Kieffer) OR, AU
thanatogratus Epler NE
. tritomus (Kieffer) PA, NE
. truncatus (Kieffer) (?) PA
. venetus (Marcuzzi) (?) PA
SSSSSSSSSsSsSSS$h$SsSsssSSS$h$H SSESEESESEH YESS
SSSSSSSSSSS SHSSSSSSSSSSSSSSSSSSSESSESSSE
MEM. AMER. ENT. SOC., 36
212 GENUS DICROTENDIPES
APPENDIX 2
List of recent name changes and current name as recognized in this paper
Previous Name Current Name
binotatus Kieffer, 1911 D. freemani, nom. nov.
(Chironomus)
figueroai Vargas, 1952 D. aethiops (Townes, 1945)
(Tendipes (Limnochironomus))
formosanus Kieffer, 1916 D. septemmaculatus (Becker, 1908)
(Dicrotendipes)
frontalis Kieffer, 1916 D. septemmaculatus (Becker, 1908)
(Dicrotendipes)
hirtitarsis Johannsen, 1932 D. septemmaculatus (Becker, 1908)
(Chironomus)
hoonsooi Ree, 1981 D. pelochloris (Kieffer, 1912)
(Kimius)
incurvus Sublette, 1964 D. tritomus Kieffer, 1916
(Chironomus (Dicrotendipes))
inferior Johannsen, 1932 D. pelochloris (Kieffer, 1912)
(Chironomus)
innisfailensis Freeman, 1961 D. tenuiforceps (Kieffer, 1913)
(Chironomus (Dicrotendipes))
loripes Guha & Chaudhuri, 1981 D. pelochloris (Kieffer, 1912)
(Xenochironomus)
melanocnemis Edwards, 1928 D. candidibasis (Edwards, 1924)
(Chironomus)
niveicauda Kieffer, 1921 D. pelochloris (Kieffer, 1912)
(Limnochironomus)
paxillus Guha, Chaud, & Nandi, 1982 Chironomus glauciventris
(Dicrotendipes) (Kieffer, 1912)
pulsus Walker, 1856 D. modestus (Say, 1823)
(Chironomus)
? punctatipennis Kieffer, 1910 D. septemmaculatus (Becker, 1908)
[Chironomus (Prochironomus)]|
rajasthani Singh & Kulshrestha, 1977 D. septemmaculatus (Becker, 1908)
(Dicrotendipes)
socionotus Guha, Chaud. & Nandi, 1982 Chironomus tainanus
(Dicrotendipes) (Kieffer, 1912)
wirthi Freeman, 1961 D. pelochloris (Kieffer, 1912)
(Chironomus (Dicrotendipes))
The species Dicrotendipes crispi (Freeman, 1957), D. multispinosus |
(Freeman, 1957), D. penicillatus (Freeman, 1957) D. regalis |
(Goetghebuer, 1936) are no longer considered members of Dicrotendipes, |
and must be assigned to a new, as yet undescribed, genus. |
213
Synonyms in italics, new species in boldface
? aequatoris, Poly. — 40
aethiops, Di. — 60
alsinensis, Di. — 60
amazonicus, Di. — 65
balciunasi, Di. — 113
bilobatus, Di. — 115
binotatus, Ch. — 36
bredoi, Di. — 33
californicus, Di. — 61
Calochironomus — 9
candidibasis, Di. — 117
chambiensis, Di. — 34
Cladotendipes — 9
collarti, Di. — 35
conjunctus, Di. — 121
cordatus, Di. — 35
crypticus, Di. — 61
cumberlandensis, Di. — 125
dasylabidus, Di. — 67
demissus, Di. — 68
Dicranotendipes — 9
Dicrotendipes — 9
ealae, Di. — 36
embalsensis, Di. — 62
figueroai, Tend. (Limn.) — 60
fittkaui, Di. — 70
flexus, Di. — 128
forficula, Di. — 37
formosanus, Di. — 42
freemani, Di. — 36
frontalis, Di. — 42
fusconotatum, Calo. — 37
fuscanotatus, Di. — 37
griseonotatus, Calo. — 37
griseoparsus, Calo. — 37
griseovittatum, Poly. — 41
hirtitarsis, Ch. — 42
hoonsooi, Kim. — 134
inferior, Ch. — 134
innisfailensis, Ch. (Di.) — 145
MEM. AMER. ENT. SOC., 36
jobetus, Di. — 130
johnmartini, Di. — 131
Kimius — 9
kribiicola, Di. — 39
leei, Di. — 133
leucolabis, Di. — 40
Limnochironomus — 9
Limnotendipes — 9
lindae, Di. — 133
loripes, Einf. — 134
loripes, Xen. — 134
melanochnemis, Ch. — 117
nervosus, Di. — 63
nestori, Di. — 63
nigrolineatus, Di. — 40
nilicola, Di. — 37
niveicauda, Limn. — 134
obrienorum, Di. — 63
palearivillosus, Di. — 72
paradasylabidus, Di. — 73
paterjohni, Di. — 74
pellegriniensis, Di. — 64
pelochloxis, Di. — 134
peringueyanus, Di. — 41
pictipennis, Di. — 42
pictus, Parat. — 35
pilosimanus, Di. — 42
pseudoconjunctus, Di. — 139
punctatipennis, Ch. (Pro.) — 42
quatuordecimpunctatum, Poly. — 42
quatuordecimpunctatus, Di. — 42
quatuorpunctatum, Poly. — 37
radinovskyi, Di. — 76
rajasthani, Di. — 42
reissi, Di. — 76
sarinae, Di. — 142
schoutedeni, Di. — 41
semiviridis, Di. — 145
septmmaculatus, Di. — 42, 145
214
Sernowia — 9
sexnotatus, Stict. — 42
seychelleanus, Ch. — 36
sinoposus, Di. — 64
soccus, Di. — 78
speciosus, Di. — 42
sudanicus, Di. — 46
tenuiforceps, Di. — 145
?trilabis, Di. — 37
wirthi, Ch. (Di.) — 134
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Memoms oF THE AMERICAN Entomovoaicat Sociery | yee ae ~
; The Cresson Types of Hymenoptera. ‘Ezra T. Cresson. 1916. $12.00, —
A Venational Study of the Suborder Zygoptera (Odonata), with Keys et
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William T. Keeton. 1960. $15.00.
The Genus Bucculatrix in America North of Mexico (Microlepidoptera), Annette Fe
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. The Butterflies of Libeia. Richard M. Fox, Arthur W. Lindsey, Jr., Harry K. Clench —
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. Proceedings of the 8th International Symposium on ‘Ceirooonsidas, Jacksonville,
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. Arreconsideration of the Milliped Genus Sigmoria, with a revision of Deltotaria and an
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Biosystematics of the Genus Dicrote + Kieffer, 1913 (Diptera: Chironomidae:
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AMERICAN ENTOMOLOGICAL SOCIETY
NUMBER 37
REVISION OF THE
MILLIPED FAMILY EURYMERODESMIDAE
(POLYDESMIDA: CHELODESMIDEA)
By
ROWLAND M. SHELLEY
PUBLISHED BY THE AMERICAN ENTOMOLOGICAL SOCIETY
AT THE ACADEMY OF NATURAL SCIENCES
PHILADELPHIA
1989
PauL M. MaArsH
EDITOR
(Issued, January 29, 1990)
PRINTED ON ACID FREE PAPER IN THE UNITED STATES OF AMERICA
MEMOIRS
OF THE
AMERICAN ENTOMOLOGICAL SOCIETY
NUMBER 37
REVISION OF THE MILLIPED FAMILY EURYMERODESMIDAE
(POLYDESMIDA: CHELODESMIDEA)
BY
ROWLAND M. SHELLEY
North Carolina State Museum of Natural Science
P.O. Box 27647, Raleigh, North Carolina 27611
ABSTRACT — Eurymerodesmus Brolemann, the sole genus in
the chelodesmoid milliped family Eurymerodesmidae, is a
speciose mosaic complex that covers a substantial part of the
central, south-central, and southeastern United States. It is the
dominant polydesmoid in prairie ecosystems, and forms occur
in diverse biotopes in both mesic and arid environments.
Autapomorphies include a ventral mandibular ridge and
medial projection in both sexes; elaborate, hirsute gonopodal
apertures; laterally oriented gonopods; simple, unmodified
telopodites; and long, silken prefemoral hairs. Eury-
merodesmus contains 25 species, five of which, e/evatus,
caesariatus, crassatus, dactylocyphus, and clavatus, are newly
described. Kewanius and Paresmus, both authored by
Chamberlin, are placed in the generic synonymy, and the
following species-group names are assigned to specific
synonymies: Paresmus columbus Causey; E. bentonus,
plishneri, spectabilis, and wellesleybentoni, all by Causey;
creolus and schmidti, both by Chamberlin; and hamatilis and
minimus, both by Loomis. Eurymerodesmus booneus
Chamberlin is a synonym of Auturus evides (Bollman).
Eurymerodesmus christianus and louisianus, both by
Chamberlin, are reduced to subspecies of E. varius (McNeill);
E. planus Causey is a race of E. birdi Chamberlin. The genus is
divided into four lineages, three of which, the hispidipes, birdi,
and melacis lines, are composed of one species group each; the
Kewanius lineage is comprised of four species groups. Rela-
tionships among the lineages are melacis + (hispidipes +
(birdi + Kewanius)). Along with Sigmoria, Eurymerodesmus
directs attention to the evolutionary dynamics of mosaic com-
plexes in the Diplopoda and their potential utility as new in-
vestigative subjects that may hold more inherent appeal to
other biologists than traditional milliped taxonomy.
INTRODUCTION
The central, south-central, and southeastern
United States — from northeastern Nebraska, cen-
tral Illinois, and southeastern North Carolina to the
Rio Grande and north Florida (Fig. 1) — is home to
the Eurymerodesmidae (Polydesmida: Chelodes-
midea). Sympatric through much of this area with
eastern Nearctic components of the chelodesmoid
families Xystodesmidae and Platyrhacidae, some of
whose species they superficially resemble in external
body form, eurymerodesmids inhabit a diversity of
biotopes including mesic deciduous forest litter in II-
linois, the Ozark Plateau, and the Ouachita Physio-
graphic Province; under logs in xeric, sandy,
southeastern environments; and under logs and dung
in grasslands of the Central Plains. They are the
dominant polydesmoid diplopods in _ prairie
ecosystems, ranging westward onto the Edwards
Plateau and across the Pecos River in central and
southern Texas.
The Eurymerodesmidae is one of the best defined
chelodesmoid families, but its affinities within the
suborder are obscure. Autapomorphies include a
ventral ridge along the mandibular stipes in both
sexes that extends into a medial projection, usually
larger in males; laterally oriented gonopods with the
cannulas located ventrolaterad on the coxa instead
of mediad; simple, unmodified, and caudally, in-
stead of anteriorly, projecting telopodites; highly
modified and variable gonopodal apertures with
relatively unmodified, simple gonopods, instead of
vice versa; a variably hirsute aperture indented
anteromediad, instead of glabrous and not indented;
and extremely long, silken prefemoral hairs, in some
forms nearly half as long as the telopodite, extending
in variable numbers and arrangements along the
stem of the structure, rather than short hairs chiefly
restricted to the basal region. The most striking
features of eurymerodesmids are the variable aper-
(1)
EURYMERODESMID MILLIPEDS
Fig. 1.
tures, many of which are elaborately complex.
Eurymerodesmus mundus and western populations
of E. birdi birdi, both authored by Chamberlin
(1931), exhibit enormous lobes on the caudal aper-
ture margins that extend well below the levels of the
ambulatory coxae and must therefore affect either
locomotion, posture, or both (Figs. 149-151,
169-171). From spatial considerations, these species
must walk ‘“‘higher’’ for the lobes to clear the
substrate. In some species lacking lobes, for example
E. oliphantus Chamberlin, newtonus Chamberlin,
amplus Causey, varius (McNeill), and angularis
Causey, the sides of the apertures are divided near
midlength thus forming two lateral margins along
the caudal halves. The ‘‘inner lateral margin’’ ex-
tends variably into the opening and the ‘‘outer
lateral margin’ flares variably laterad thereby
enclosing a cavity or ‘‘pouch’’ at the caudolateral
Distribution of Eurymerodesmus and the Eurymerodesmidae.
corners (Figs. 18, 21, 23, 25, 56, 59, 62, 64, 68,
73, 76-78, 84, 91). The pouches vary from open to
closed depending upon the degree that the outer
lateral margin leans mediad over them, and they give
rise to variable tufts of hairs. Another species, E.
dactylocyphus n. sp., lacks both of these features but
possesses an exceptionally broad aperture with
variably elongate flanges at the caudolateral corners;
in some individuals the opening covers nearly the en-
tire ventral breadth of the segment (Fig. 184). The
anatomical wonders are not limited to males;
females of EF. dactylocyphus, angularis, compressus
Causey, impurus (Wood), and crassatus n. sp.
possess dactyliform projections on the cyphopod
valves (Figs. 14, 27, 30, 35, 130, 187) that extend
caudad in dactylocyphus and compressus_ well
beyond the aperture margin, overlapping the suc-
ceeding two or three segments. They are so long in
ROWLAND M. SHELLEY 3
some forms of dactylocyphus as to resemble super-
numerary walking legs. When I first viewed them, I
thought the specimen was a mutant. These extreme
genitalic modifications in both sexes are so bizarre as
to compel speculation on evolution and function,
which I think relate to the recent theory of sexual
selection by females propounded by Eberhardt
(1985) (see sexual selection section, p. 41). Beyond
these striking autapomorphies, I am equally im-
pressed with the general hairiness of eury-
merodesmids, some forms being extremely setose
ventrally. The lobes in mundus and b. birdi are so
densely hirsute as to obliterate the surface, and in
some forms of the latter the hairs become pro-
gressively longer distad with the apical hairs being
fully as long as the lobes themselves, thus effectually
doubling the lengths of the structures (Fig. 155).
Likewise, the tufts from the caudolateral pouches
are extremely dense in some species, and in others
the entire aperture margins are variably hirsute,
sometimes densely so. Thus in some individuals of
oliphantus, there are long hairs arising from the
anterior and lateral rims, the outer surfaces of the
sides extending well onto the metazonum, dense
tufts from within the pouches, additional hairs from
the inner surfaces of the outer lateral and caudal
margins, and still more arising from beneath the
division points on the sides (Fig. 91). In addition
to these aperture hairs, there are variable and often
dense tufts from sternal projections on _pre-
gonopodal and postgonopodal segments, even on
relatively flat caudal segments and on females.
Coupled with those on the gonopods, the overall
effect of these hairs and their variations in lengths
and densities is so impressive as to be a fitting source
for the family-group and type genus names if the
millipeds were newly discovered. Eurymerodesmus
(-idae), literally meaning broad, unit, chain, does not
begin to do justice to the anatomical marvels of these
arthropods.
In addition to their unique anatomical features,
eurymerodesmids are of exceptional ecological in-
terest, because of the aforementioned diversity of
their biotopes and their unusual ability to tolerate ex-
tremely arid conditions. I encountered a sizeable
population of varius varius under a sun-baked log on
dry sandy soil in Gilchrist County, Florida, and have
visited areas in central and southern Texas known to
harbor epigean populations of E. medlacis
Chamberlin and Mulaik that can only be described
as desert. Had I not known of the presence of
melacis, 1 would have sworn that the only possible
epigean diplopods in these sandy, sagebrush en-
MEM. AMER. ENT. SOC., 37
vironments would be desert-adapted spirostreptoids
and spiroboloids. At the opposite moisture extreme,
I have also collected v. varius from wet soil in hard-
wood hammocks in north Florida and oliphantus
from a stump in a mesic deciduous woodland in
southern Illinois. Likewise, I have visited areas in the
Ozark and Ouachita Provinces that would be ex-
pected to contain many eurymerodesmids and
diverse diplopod faunas because of the moist condi-
tions and abundance of deciduous trees. Suffice it to
say that eurymerodesmids have broad ecological
tolerances, and of particular interest to North
America is their abundance in the Central Plains,
where they are the dominant polydesmoids.
Eurymerodesmids are among the earliest de-
scribed American millipeds. The type species of the
type genus, FE. hispidipes (Wood 1864), proposed for
a form from Illinois, was among the first millipeds
named by an American author and one of the first
species described from the New World. With over
2,000 preserved specimens, eurymerodesmids are
abundantly represented in major institutional reposi-
tories in the United States, and many colleges and
universities within the range have samples in insect
and invertebrate collections. Despite this material
and the considerable attention it has received in over
40 publications, the Eurymerodesmidae is also one
of the most poorly understood of the major Nearctic
milliped taxa. Thirty-two species-group names have
been proposed, but only two can be confidently
identified from published diagnoses. The descriptive
accounts are among the worst ever published on the
North American fauna, revealing little effort to iden-
tify taxonomically important structures or assess
their conditions throughout the family. Likewise, the
illustrations do not begin to portray diagnostic
features, particularly of the apertures. Three species
— E. simplex Chamberlin, impurus, and melacis —
have not been figured, and most drawings of other
species are meaningless sketches. A host of
synonyms therefore exists, as few specimens or ac-
counts were consulted before new names were
validated. Finally, the general commentary is so
riddled with contradictions, conflicting opinions,
and outright errors that eurymerodesmid literature is
not only useless but actually hinders study. It would
be far easier to start with nothing than to have to
determine synonymies and untangle such
nomenclatorial confusion.
The taxonomic disarray results from two fac-
tors. First, like Abacion (Shelley 1984a) and
several other Nearctic taxa, only one species was
assumed to exist for many years, and this name
4 EURYMERODESMID MILLIPEDS
was routinely applied to forms throughout the
generic range. Thus, hispidipes has been
erroneously assigned to forms from Arkansas,
Kansas, Louisiana, Texas, and even Illinois as
recently as 1969. Since much of this material is
lost, we can only infer which species actually were
represented. Secondly as stated earlier, the male
gonopods alone are not the principle taxonomic
structures at the specific level, unlike most
polydesmoid families. Specific details obtain
chiefly either in the configuration of the
gonopodal aperture, which extends anteriad and
compresses the prozonum into a narrow band, or
in that plus aspects of the gonopods, which lack a
prefemoral process. Somatically, the collum is
moderately enlarged; the caudolateral corners of
the paranota are broadly rounded; and the
epiproct is pointed and subtriangular as in the
Xystodesmidae. Eurymerodesmids vary in size
from minute specimens of v. varius and melacis to
such large species as FE. goodi Causey and dubius
Chamberlin. They tend to be smaller than most
sympatric xystodesmids, but the general facies are
so similar that mature specimens can easily be
mistaken for juveniles of the latter. Juveniles of
both families are practically indistinguishable.
Despite this resemblance, the families are rarely
mixed in the same sample because of their stag-
gered life histories. Eurymerodesmids are de-
cidedly ‘‘cold-adapted,’’ most collections occurr-
ing between late October and early April (Fig.
210). Conversely, most xystodesmids prefer the
warmer weather of late spring, summer, and early
autumn. Thus, although sympatric through much
of the midwestern and southeastern states, the two
families tend to appear at different times of the
year.
A curious feature of eurymerodesmids is their
stiff, inflexible body; most I have found were
tightly coiled in upper humus layers. They die and
are preserved in this position, and in contrast to
xystodesmids and platyrhacids, are nearly im-
possible to uncurl without breakage at one or
more segmental junctures. Few specimens remain
intact after handling or can be unrolled without
breakage. Indeed, the exoskeleton will often tear
or shatter before adjacent segments can be
separated, and length measurements are difficult
to obtain. Many workers obviously did not try to
uncurl specimens, as evidenced by the unusually
high percentage of sexing errors. Segment 7 is
overlapped by legs from other segments, and since
the gonopodal telopodites are linear and project
almost directly caudad, they are inconspicuous
laterally in a tightly coiled male. Hence, these
specimens are easily mistaken for females, because
every segment appears to possess two pairs of legs
and there are no obvious gonopods or gaps for the
same. Consequently, many ‘‘female allotypes’’
are really male paratypes, and I know of no other
helminthomorph milliped group in which so many
sex notations are wrong. However, with ex-
perience and close observation, one can detect a
slight gap between the legs on segment 7 and the
raised aperture inside. Males also have a decidedly
‘flatter’? appearance than females, as the
paranota are broader and less depressed.
As currently understood (Hoffman 1979), the
Eurymerodesmidae consists of two valid genera,
Eurymerodesmus Brolemann (1900) and Paresmus
Chamberlin (1942a). A third genus-group name,
Kewanius Chamberlin (1938), was correctly
synonymized with Eurymerodesmus by Causey
(1963) and accepted by Hoffman (1979). The two
valid genera are distinguished solely by the stouter
telopodite of Paresmus, and Eurymerodesmus
therefore contains everything that cannot be placed
in the former. This distinction is unsatisfactory,
since eurymerodesmid telopodites vary considerably
in both length and breadth. Thus, major objectives
of this revision are detailed analyses of the tax-
onomically important characters, and decisions on
the number of component genera and the status of
Paresmus. This genus is distinguished more by a
subterminal acropodite than by breadth of the
telopodite, but this condition blends into the
smoothly continuous, broadly terminal acropodite
of Eurymerodesmus through the intermediate condi-
tion represented by the type species of Kewanius, a
narrowly terminal, discontinuous, and _ sharply
demarcated acropodite. Hence, I regard the family
as monotypic and place Paresmus and Kewanius in
synonymy under Eurymerodesmus, which I divide
into four lineages based on gonopodal features and
the configurations of the apertures. These categories
correspond to the subgenera or major components
of Sigmoria (Shelley and Whitehead 1986), but I do
not officially designate them as subgenera because
genus-group names are only available for two. Since
many new forms obviously await detection, it seems
best not to validate names that may be overturned by
future discoveries. Three of the lineages contain a
single species group and are named for the oldest
component species, which is anatomically
characteristic for the assemblage; hispidipes con-
stitutes a separate species group and lineage by itself,
ROWLAND M. SHELLEY 5
since I cannot relate it to another congener. The
fourth lineage, containing 17 species in four species
groups, includes those assigned to both Kewanius
and Paresmus, and hence is named the Kewanius
lineage since there is no characteristic species and this
name has priority by four years. Species groups
within the Kewanius lineage are named for the oldest
component species.
I present (Fig. 215) a diagram that I think reflects
relationships among the lineages, but none of the
sister branches can be unequivocally defined by
autapomorphies, demonstrated by all component
forms. Unique features can only be suggested for the
melacis line. Decisions on polarities are hampered by
insufficient knowledge of potential out-groups, but I
think these traits are plesiomorphies rather than
apomorphies. However with the possible exception
of the hispidipes branch, the lineages are all cohesive
geographic entities, partly sympatric with each other.
The birdi and Kewanius lineages are incompletely
detached and connect through polkensis, cae-
sariatus, pulaski, and serratus, all showing acro-
podal features of the latter and aperture traits of the
former. They demonstrate the phenomenon termed
“incomplete synapomorphy”’ in Sigmoria (Shelley
and Whitehead 1986). The melacis branch, however,
appears completely segregated, and I consider it
sister to the rest of the family.
In conducting this revision, I have attempted to
apply the definitions of taxonomic categories
employed by Shelley and Whitehead (1986) for
Sigmoria. However, I have not personally col-
lected many eurymerodesmids, and despite the
substantial material, significant collecting gaps oc-
cur within the range. Consequently, I cannot
determine the nature of the interfaces between
many species nor the degree of incipient partition-
ing. Thus there are problematical areas that can-
not be fully resolved in this first revision, for ex-
ample between varius and amplus Causey, and
among forms of angularis. However beyond this
purely technical matter, looms the larger question
of the actual status of the mostly parapatric
forms, both here and within Sigmoria. There is no
question that birdi and mundus, broadly sym-
patric through Kansas, Oklahoma, Texas, and
Arkansas, are reproductively isolated and hence
full species, but is this true for the parapatric
forms in the Kewanis and melacis lineages, and
should they be accorded the same status? Resolu-
tion of this question is beyond the scope of this
work and requires the attention of someone more
versed in theoretical concerns or in reproductive
MEM. AMER. ENT. SOC., 37
biology of these forms. However, the ‘‘species’’ of
the melacis and Kewanius lineages may actually be
only semispecies, with the parent lineage becom-
ing a superspecies. These categories cannot be
handled nomenclaturally, so for the present I refer
to all parapatric or narrowly sympatric entities
that are not interconnected as species. This ques-
tion bears on the composition of the genus, since I
recognize 25 species. If the localized entities are
only semispecies, Eurymerodesmus may contain
as few as five species.
Like Sigmoria, Eurymerodesmus is a giant
mosaic complex that blankets much of central and
southeastern North America. Past workers have
extracted pieces of this large puzzle that appear
distinct because they possess different combina-
tions of the features of the apertures and
gonopods. Consequently, they assigned them dif-
ferent names, thereby adding nomenclatural con-
fusion to an already intricate fauna. I have tried to
resolve this difficult problem to the extent possible
and leave a stable base for future students.
However, the overriding significance of Eury-
merodesmus and Sigmoria is their illumination of
mosaics, a widespread phenomenon in the Diplo-
poda (Shelley 1989). The concluding section
therefore attempts to integrate these revisions with
recent treatments of other genera and suggest a
different emphasis in systematic research that may
stimulate interest in diplopodology.
This revision, then, provides complete descrip-
tions or redescriptions of the family Eury-
merodesmidae, the genus Eurymerodesmus, and
all species and subspecies considered valid along
with synonymies and pertinent anatomical illus-
trations. A key has also been formulated to
facilitate determinations. A literature review and
sections on anatomy and taxonomic characters,
and sexual selection precede the systematic ac-
counts, which are followed by sections on ecology,
distribution, relationships, and the aforemen-
tioned conclusion. All pertinent holotypes or syn-
types were examined except those of compressus
and Paresmus columbus and P. pulaski, all
authored by Causey, which are not at the pub-
lished repositories and are lost. Also lost is the
holotype of P. paroicus, whose vial actually con-
tains a male of Pseudopolydesmus pinetorum
(Bollman). These names were interpreted from
near topotypical material, and neotypes are
designated for all except P. columbus, assigned to
synonymy under dubius.
6 EURYMERODESMID MILLIPEDS
In conducting this revision I was fortunate in
receiving material from many large and small collec-
tions, both institutional and private. Acronyms of
sources of preserved study material cited in the text
are as follows:
AMNH — American Museum of Natural History, New
York, NY.
ANSP — Academy of Natural Sciences, Philadelphia, PA.
CAS — California Academy of Sciences, San Francisco,
CA.
EIU — Zoology Department, Eastern Illinois University,
Charleston, IL.
FMNH — Field Museum of Natural History, Chicago, IL.
FSCA — Florida State Collection of Arthropods,
Gainesville, FL.
ILNHS — Illinois Natural History Survey, Urbana, IL.
MCZ — Museum of Comparative Zoology, Harvard
University, Cambridge, MA.
MEM — Mississippi Entomological Museum, Mississippi
State University, Starkville, MS.
MNHP — Museum National d’Histoire Naturelle, Paris,
France.
MWSU — Zoology Department, Midwestern State
University, Wichita Falls, TX.
NCSM — North Carolina State Museum of Natural
Science, Raleigh, NC.
NMNH — National Museum of Natural History, Smith-
sonian Institution, Washington, DC.
OKSU — Biology Department, Oklahoma State University,
Stillwater, OK.
RLH — Private collection of Richard L. Hoffman,
Martinsville, VA.
SFAU — Biology Department, Stephen F. Austin State
University, Nacogdoches, TX.
TAI — Biology Department, Texas A & I University,
Kingsville, TX.
TMM — Texas Memorial Museum, University of Texas,
Austin, TX.
UGA — University of Georgia Museum of Natural
History, Athens, GA.
UMO — Enns Entomological Museum, University of
Missouri, Columbia, MO.
UOK — Stovall Museum of Science and History,
University of Oklahoma, Norman, OK.
WAS — Private collection of William A. Shear,
Hampden-Sydney, VA.
WASU — James Entomological Museum, Washington State
University, Pullman, WA.
LITERATURE REVIEW
Although eurymerodesmid millipeds have been
known since 1864, when Wood described
Polydesmus hispidipes from an unspecified site in
Illinois, the family Eurymerodesmidae was estab-
lished only 38 years ago (Causey 1951), nine years
after Chamberlin (1942a) proposed the last genus-
group name, Paresmus. Eurymerodesmus Brole-
mann, 1900, was designated the type-genus. In the
original account, Polydesmus hispidipes was placed
in the nominate subgenus, but the next year Wood
(1865) transferred it to the subgenus Paradesmus
while repeating the verbal description and publishing
a gonopod drawing showing the long prefemoral
hairs. Wood (1867) described Polydesmus (Poly-
desmus) impurus from material collected at an
unspecified locality in Texas by Dr. Gideon
Lincecum, a pioneer physician and naturalist.
Causey (1952a) reported that Lincecum traveled
widely, that his last home was Bonham, Washington
County, and that the type locality could not be deter-
mined with certainty. However, Bonham is in Fan-
nin County, over 230 miles north of Brenham, the
county seat of Washington County, and her error
could be in either the town or county. By studying
correspondence at the University of Texas library
between Dr. Lincecum and Wood, James Reddell
(pers. comm.) concluded that the type locality was
Long Point, Washington County, near Brenham.
McNeil (1887) described the third eury-
merodesmid, Polydesmus varius, from a female col-
lected at Pensacola, Escambia County, Florida, and
published two cyphopod drawings. Bollman (1888a)
transferred hispidipes to Leptodesmus and noted
that it is common throughout Arkansas, but as
shown by material at the NMNH,, this record refers
to E. pulaski (Causey), serratus n. sp., and perhaps
other species. Bollman (1888b) transferred varius to
Leptodesmus and recorded it from Macon, Bibb
County, Georgia. In 1893 he included ll
three previously described species and several
xystodesmids as components of Leptodesmus and
repeated the type localities of each. Two years later,
Brolemann (1895) included hispidipes in his list of
American myriapods and reported it from Loui-
siana, where it does not occur. As shown by his
material at the MNHP, Brolemann’s record refers to
E. amplus.
In 1900, Brolemann proposed Eurymerodesmus
as a subgenus of Fontaria. He illustrated the
gonopod of amplus, which he misidentified as
hispidipes, and noted that it did not belong in Lep-
todesmus as previously reported. Since the structure
resembled a ‘‘secondary branch’’ of a Fontaria
gonopod, particularly that of E. simoni Brolemann,
a synonym of Harpaphe haydeniana (Wood), he
provisionally included the species in Fontaria under
the new subgenus. Brolemann listed development of
the femur, absence of a ‘‘secondary ramus,” and the
presence of rows of long silken hairs along the
“seminal canal’’ (= prostatic groove) nearly to the
tip as diagnostic characters of the new subgenus.
Thus, hispidipes is the type species of EFury-
ROWLAND M. SHELLEY 7
merodesmus by monotypy, as noted by Jeekel
(1971).
In the ensuing decade, Gunthorp (1913) reported
hispidipes from Cowley, Douglas, and Jefferson
counties, Kansas. Since his specimens are lost, I sur-
mise from known records that the Cowley specimen
was probably mundus and the others were probably
birdi birdi. Brolemann (1915) elevated Eury-
merodesmus to generic status and placed it in the
tribe Rhysodesmini, subfamily Leptodesminae,
family Leptodesmidae. Apparently unaware of
Brolemann’s work, Chamberlin (1918) recorded L.
hispidipes from Creston, Natchitoches Parish, Loui-
siana, a citation based on specimens of amplus and
birdi. Two years later, Chamberlin (1920) ac-
knowledged Eurymerodesmus by publishing a brief
diagnosis and adding the fourth species, simplex,
from New Orleans, a locality that is probably wrong.
The period of intensive descriptive work began in
1931, when Chamberlin described birdi and mundus
from Oklahoma, illustrating the gonopods and aper-
ture lobes of each. Chamberlin (1938) established the
genus Kewanius for simplex because of its ‘‘glabrous
telopodite and larger basal gonopodal division.”’
However, the telopodite, or specifically the
prefemur, is really only less densely hirsute than that
of hispidipes, not glabrous, and the ‘“‘larger basal
gonopodal division’’ probably refers to the broad
clavate condition of the prefemoral stem. Causey
(1963) did not think these features warranted generic
distinction and placed Kewanius in synonymy under
Eurymerodesmus, an action accepted by Hoffman
(1979) and one that I can confirm.
In the decade of the ’40’s, Chamberlin and Mulaik
(1941) described, but did not illustrate, melacis from
Kerr, Boerne, Hidalgo, Kendall, Bandera, and Con-
cho counties, Texas, without clearly specifying the
type locality. Chamberlin (1942a) described and il-
lustrated the gonopods of two new species of
Eurymerodesmus from both Arkansas and Loui-
siana — newtonus and oliphantus, and creolus and
louisianae, respectively — and erected Paresmus for
a third new species from Louisiana, paroicus.
Chamberlin (1942b) proposed EF. booneus for
unillustrated specimens from Boone County, Iowa,
which are actually juvenile platyrhacids. Thus, E.
booneus is a junior subjective synonym of Auturus
evides (Bollman) [syn. nov.!], which is the only
platyrhacid in central Iowa and is common in Boone
County. Chamberlin (1943) described and illustrated
E. schmidti and dubius from Arkansas, and Loomis
(1943) did likewise for E. minimus, from Jackson
County, Florida. In 1946 Chamberlin proposed and
MEM. AMER. ENT. SOC., 37
figured E. christianus, from Harrison County,
Mississippi.
Descriptions of eurymerodesmids increased in the
1950’s and were led by six papers by Causey, two the
first year. She (1950a) described and illustrated five
new Arkansas eurymerodesmids, E. bentonus, spec-
tabilis, and plishneri, and Paresmus pulaski and
columbus, and reported E. hispidipes from Dixon
Springs, Pope County, Illinois, a misidentification
of oliphantus. Later that year Causey (1950b) pro-
posed FE. planus for a form from Rankin County,
Mississippi, and provided a lateral view of segment 7
with the gonopods projecting above the segmental
margin. Causey (1951) established the family
Eurymerodesmidae, listed several autapomorphies,
and described and illustrated angularis from Arkan-
sas. The next year Causey (1952a) published
diagnoses and drawings of FE. compressus and
wellesleybentoni, from Arkansas, and E. sanbernar-
diensis, from Texas, along with a key to the species
of Paresmus and notes on P. impurus. The last
species was transferred from Leptodesmus, where it
had been assigned by Bollman (1893). Causey
(1952b) described and illustrated the fifth and final
species of Paresmus, polkensis, from Polk County,
Arkansas, along with EF. goodi and amplus from
Arkansas and Louisiana, respectively. This work
also included new localities for birdi, schmidti, mun-
dus, spectabilis, and dubius, a sketch of the aperture
of oliphantus, mislabeled as hispidipes, and the
placement of plishneri in synonymy under schmidti.
Chamberlin (1952) reported melacis from Uvalde
County, Texas, and reiterated the type localities for
dubius and schmidti. Causey (1954) transferred
varius into Eurymerodesmus from Leptodesmus,
recorded it from Mobile County, Alabama, and
placed minimus in synonymy, thereby making
Loomis’ gonopod drawing (1943) the first of a male
of varius. Chamberlin and Hoffman (1958) listed all
eurymerodesmid species to that date with statements
of type localities and distributions, and accepted the
synonymies of plishneri with schmidti and minimus
with varius. The last work of the ’50’s recorded new
localities for melacis in south Texas (Loomis 1959).
Interest in eurymerodesmids waned after the
1950’s. Causey (1963) synonymized Kewanius with
Eurymerodesmus and spectabilis with christianus,
recording new localities for the latter from Loui-
siana, Alabama, Mississippi, and Arkansas. Next to
hispidipes, spectabilis was the most widely known
species at that time. Loomis (1969) described and
illustrated E. hamatilis from Angelina County,
Texas, and Stewart (1969) recorded birdi and
8 EURYMERODESMID MILLIPEDS
hispidipes from several northeastern Texas counties,
the latter being misidentifications of amplus. Reddell
(1970) reported Eurymerodesmus from caves in
Bexar, Kerr, and Uvalde counties, Texas, all records
of melacis. Loomis (1976) proposed the most recent
new species, E. digitatus, from Burnet County,
Texas, and appended a key to species in that state.
Hoffman (1978, 1979) considered the Eurymerodes-
midae a derivative of the Xystodesmidae and placed
both in the superfamily Xystodesmoidea along with
the Gomphodesmidae, Campodesmidae, and Oxy-
desmidae. In the latter work he estimated 22 species
for Eurymerodesmus, ranging from Georgia to
Arkansas and Illinois; Paresmus contained five
species distributed from Arkansas to Texas.
Thus at this writing, the Eurymerodesmidae con-
sists of two genera, Eurymerodesmus and Paresmus,
with 23 and 5 nominal species, respectively, listed
below in chronological order with synonymies, type
localities, and other reported counties and states of
occurrence. It is noteworthy that 12 species of Eury-
merodesmus and three of Paresmus supposedly oc-
cur in Arkansas, while six and one, respectively,
have been recorded from Louisiana. Causey (1940)
reported hispidipes from Duke Forest, Durham
County, in central North Carolina, a record deleted
by Shelley (1978). Eurymerodesmus varius does oc-
cur in the southeastern corner of that state but is im-
probable farther inland as is hispidipes anywhere
along the Atlantic Coast.
Genus Eurymerodesmus Brolemann 1900
(= Kewanius Chamberlin 1938)
E. hispidipes (Wood 1864). Illinois without fur-
ther specification. Erroneously reported from
Arkansas and Louisiana in general (Bollman 1888a,
Brolemann 1895), and the following states, counties,
and parishes: Natchitoches Par., LA (Chamberlin
1918); Cowley, Douglas, and Jefferson cos., KS
(Gunthorp 1913); Pope Co., IL (Causey 1950a); and
Angelina, Nacogdoches, Rusk, Sabine, Shelby,
Smith, and Upshur cos., TX (Stewart 1969).
E. varius (McNeill 1887) (=£. minimus Loomis
1943). Pensacola, Escambia Co., FL. Also recorded
from Bibb Co., GA (Bollman 1888b); Jackson Co.,
FL (Loomis 1943); and Mobile Co., AL (Causey
1954).
E. simplex Chamberlin 1920. Reputedly New
Orleans, Orleans Par., LA, but probably some 145
miles to the west northwest around Evangeline and
Rapides parishes.
E. birdi Chamberlin 1931. Unspecified site in
Murray Co., OK. Also reported from Seminole,
Pittsburg, and Hughes cos., OK, and Sebastian,
Logan, and Miller cos., AR (Causey 1952b); and
Red River and Lamar cos., TX (Stewart 1969).
E. mundus Chamberlin 1931. University (of
Oklahoma), Norman, Cleveland Co., OK. Also
recorded from the Wichita National Forest, now
National Wildlife Refuge, Comanche Co., OK
(Chamberlin 1931); and Latimer, Caddo, and Mc-
Clain cos., OK, and Sevier Co., AR (Causey 1952b).
E. melacis Chamberlin and Mulaik 1941. Type
locality not specified in description, and two samples
are labeled ‘‘holotype’’: a female from Raven
Ranch, Kerr Co., TX, the first site listed in
the account and which was recorded as the type
locality by Chamberlin and Hoffman (1958), and a
male from Edinburgh, Hidalgo Co., TX. Also
reported from Kendall, Bandera, and Concho cos.,
TX (Chamberlin and Mulaik 1941); Uvalde Co.
(Chamberlin 1952); Comal and Guadalupe cos.
(Loomis 1959); and caves in Bexar, Kerr, and Uvalde
cos. (Reddell 1970).
E. newtonus Chamberlin 1942a. 1.2 mi. S Jasper,
Newton Co., AR.
E. oliphantus Chamberlin 1942a. 15 mi. SW
Oliphant, Jackson Co., AR.
E. creolus Chamberlin
Shreveport, Caddo Par., LA.
E. louisianae Chamberlin 1942a. 2 mi. S Saline,
Natchitoches Par., LA.
E. schmidti Chamberlin 1943 (=E. plishneri
Causey 1950a). Rich Mtn., Polk Co., AR. Also
recorded from Carroll and Washington cos., AR
(Causey 1950a); and Franklin and Benton cos., AR
(Causey 1952b).
E. dubius Chamberlin 1943. Delight, Pike Co.,
AR. Also reported from Clark, Dallas, Hot Springs,
Saline, and Sevier cos., AR (Causey 1952b).
E. christianus Chamberlin 1946 (=E. spectabilis
Causey 1950a). Pass Christian, Harrison Co., MS.
Also cited from Columbia Co., AR (Causey 1950a);
Union Co., AR, and Claiborne Par., LA (Causey
1952b); and Washington and East Baton Rouge
Pars., LA, Mobile Co., AL, and Jackson Co., MS,
(Causey 1963).
E. bentonus Causey 1950a. Monte Ne, Benton
Co., AR. Also recorded from Carroll Co., AR
(Causey 1950a).
E. planus Causey 1950b. Piney Woods, Rankin
Co., MS.
E. angularis Causey 1951. DeValls Bluff, Prairie
Co., AR.
1942a. 5 mi. NW
ROWLAND M. SHELLEY 9
E. compressus Causey 1952a. Junction City,
Union Co., AR.
E. wellesleybentoni
Phillips Co., AR.
E. sanbernardiensis Causey 1952a. San Bernard
River at US hwy. 59, Fort Bend Co., TX.
E. goodi Causey 1952b. 16 mi. S Mena, Polk Co.,
AR.
E. amplus Causey 1952b. Ruston, Lincoln Par.,
LA.
E. hamatilis Loomis 1969. 8 mi. N Zavalla,
Angelina Co., TX.
E. digitatus Loomis 1976. 5 mi. ESE Marble Falls,
Burnet Co., TX.
Causey 1952a. Helena,
Genus Paresmus Chamberlin 1942a.
P. impurus (Wood 1867). Long Point, near
Brenham, Washington Co., TX.
P. paroicus Chamberlin 1942a. 1.5 mi. N Clay,
Lincoln/ Jackson Pars., LA.
P. pulaski Causey 1950a. 3 mi. S Sweet Home,
Pulaski Co., AR. Also recorded from Sheridan
Co., AR (Causey 1952b), and Grant Co., AR
(Chamberlin and Hoffman 1958).
P. columbus Causey 1950a. 3 mi. E Magnolia,
Columbia Co., AR.
P. polkensis Causey 1952b. 11 mi. N Mena, Polk
Co., AR.
ANATOMY AND TAXONOMIC CHARACTERS
A family level revision is a formidable challenge,
and particularly in the Eurymerodesmidae, where
none of the nominal species has been characterized
in accordance with modern standards and the tax-
onomically important characters are unknown. Nor
has anyone searched for logical patterns in the dif-
ferent aperture types, which promise to be very
useful in making determinations. Species distribu-
tions are also unknown, so geographic data cannot
facilitate identifications. Finally, the higher taxa are
very poorly defined. Distinctions between the genera
are unclear, and although Causey (1951) listed some
autapomorphies of the family, their taxonomic
utilities at either the generic or specific levels have
not been investigated. The more than 40 publications
dealing in part with eurymerodesmids are useless for
revisionary purposes, since they consist chiefly of
either terse and unreliable locality references or mere
validation statements for scientific names. This
situation is much worse than that of the
Xystodesmidae in 1974 when I began studying that
MEM. AMER. ENT. SOC., 37
family; several genera had been revised, and the
literature was reasonably advanced. Nothing
definitive has been published on the Eury-
merodesmidae, so my task as first reviser is to fill as
many voids as possible, realizing that some will re-
main because of the impoverished initial level of
knowledge. Thus, I first examined the available type
specimens to ascetain the forms in question, since
mest illustrations are meaningless, uninterpretable
sketches. Next came the reservoir of unstudied and
unidentified material in institutional collections that,
as expected, contained new forms and the answers to
taxonomic questions, particularly which characters
hold diagnostic utility. Because it was impossible to
thoroughly investigate every potentially useful
character, I review general eurymerodesmid external
anatomy in this section as well as features found to
be taxonomically important. Development of other
potentially important characters is left to future
workers.
At the outset it is informative to digress and
review the historical development of taxonomic
characters per se. Only two of the diagnostic
autapomorphies were known before family status
was proposed in 1951, and only one was recorded
before 1943. Despite the elaborately ornamented
apertures, the hairs on the prefemoral stem were the
only unique feature observed by early authors.
Wood (1864, 1865) noted them in the original
descriptions of hispidipes, stating that the male
genital appendages were ‘“‘beset with very numerous
long hairs,’’ which are shown extending to the tip of
the telopodite in his illustration (1865, fig. 48).
Although the account is internally contradictory,
Wood (1867) also mentioned the hairs in the descrip-
tion of impurus. McNeill (1887) did not allude to
hairs in the description of varius because the types
were females, but Brolemann (1900) used them along
with the absence of a ‘“‘secondary ramus,’’ or
prefemoral process, as justification for proposing
Eurymerodesmus as a new subgenus of Fontaria. He
provided an illustration of amplus, the Louisiana
species that he (Brolemann 1895) recorded as
hispidipes, and noted the presence of a row of long
silken hairs along the length of the seminal branch
almost to its tip. Curiously, in elevating Eurymero-
desmus to generic status fifteen years later,
Brolemann (1915) ignored the hairs, giving as his
criteria a large femur supporting a short, slightly
bent seminal branch without an apical opening, and
adding that the secondary branch was absent. In a
brief generic account and 11 subsequent species
descriptions, Chamberlin (1920, 1931, 1942a, 1943)
10 EURYMERODESMID MILLIPEDS
and Chamberlin and Mulaik (1941) continued to
mention the prefemoral hairs but did not cite any
other autapomorphies, although Chamberlin (1931)
did report and illustrate the lobes of birdi and mun-
dus. Consequently, Loomis (1943) became the first
author to note another feature when he mentioned in
the description of minimus, a synonym of v. varius,
that the mandibular stipes ended simply rather than
in an “‘incurved process’’ as in males of mundus
from Dallas County, Texas. This statement ap-
parently did not register with Chamberlin (1946), as
he said nothing about the mandible in the descrip-
tion of christianus. Thus, the recognition of the
uniqueness of Eurymerodesmus must be attributed
to Causey (1950a), who noted in a brief generic ac-
count that the gonopodal opening was variously
modified posteriorly and laterally with setose lobes,
that the telopodite was a ‘“‘single, unbranched,
slender blade with setae usually in rows,” and that
the leg femora, presumably meaning prefemora,
were unspined. She did not include the mandibular
projection in the generic account but characterized
and/or illustrated it and the second leg coxa in all six
descriptions that year (Causey 1950a, b). The follow-
ing year, Causey (1951) erected the family and listed
most of the autapomorphous features, which she
characterized in the six species proposed the next
year (Causey 1952a, b). Thus, 79 years elapsed from
the date of the first species until a second family
character was recognized, and seven more passed
before other unique features were recorded. Only 87
years later was Eurymerodesmus properly assigned
to a separate family. The prefemoral hairs have
always been associated with Eurymerodesmus, but
the other diagnostic characters have only been
recognized in the past three to four decades.
Developing these as fully as possible is a major ob-
jective of this work.
Body size. — Individual body size is a subjective
criterion at best, and particularly in the Eurymero-
desmidae because of the brittleness and rigidity of
most preserved specimens. Females are generally
larger and more vaulted than males, but both sexes
tend to fragment or even shatter with handling. One
should therefore measure the length while the animal
is alive or freshly killed, a task for future workers
since this study is based primarily on preserved
specimens. Most eurymerodesmids that I en-
countered alive were tightly coiled under leaves or in
upper soil layers. They die and are preserved in this
posture, and hence must be uncoiled for measure-
ment and examination of the genitalia. They usually
fragment during this process beyond the state where
they can be reassembled for reasonably accurate
length measurements, although large species tend to
be more flexible than small ones. However, despite
the difficulty in measuring lengths, some species are
so obviously larger or smaller than others that size
alone is a clue to identity, even in the field, although
it should always be confirmed by examining the
genitalia. For example, because varius is so much
shorter, one could usually distinguish it from such
sympatric species as birdi and angularis in Missis-
sippi and southeastern Louisiana. Likewise in Texas,
the components of the melacis lineage are generally
smaller than parapatric or sympatric forms of birdi
and amplus. Conversely in southern Arkansas and
northern Louisiana, angularis, compressus, goodi,
dubius, and polkensis, the largest eurymerodesmids,
stand out in contrast to such smaller species as birdi,
mundus, and amplus. Averaging all measurements
of each species, I propose the following general size
or length categories, to be modified by measure-
ments of fresh material. They are useful when forms
at opposing extremes, such as small or small to
moderate and large, are found sympatrically.
TABLE 1. Size Categories for Species and Subspecies
of Eurymerodesmus.
Length
Category (mm) Species
Small under 18.0 vy. varius, varius christianus, varius
intergrades, melacis, clavatus.
Small to ca. 19-23 hispidipes, varius louisianae,
Moderate oliphantus, elevatus, dactylocy-
Dhus, sanbernardiensis, digitatus.
Moderate ca. 25-30 impurus, amplus, newtonus,
simplex, caesariatus, birdi,
mundus.
Moderately ca. 30-32 paroicus, crassatus, pulaski, ser-
Large ratus.
Large above 32 angularis, compressus, goodi,
dubius, polkensis.
Head. — The head of eurymerodesmids, which is
smooth and glossy with few facial setae, can easily
be mistaken for that of a xystodesmid. The
epicranial suture is distinct, bifid, and terminates in
the interantennal region. The epicranial and in-
terantennal setal series are usually absent; subanten-
nal setae, one per side, are usually present; and when
present, frontal and genal setae are few in number.
The genae may or may not have faint central impres-
sions, and the ends are rounded and extend beyond
the adjacent cranial margins. As in xystodesmids,
the antennae are long and slender, becoming pro-
gressively more hirsute distally, with antennomeres
ROWLAND M. SHELLEY 11
2-6 being clavate. The four terminal cones are the
only apparent sensory structures.
Projection of the mandibular stipes. — In both
sexes the mandibular stipes possesses a mid-ventral
ridge that becomes broader distad, extending beyond
the corner of the stipes into a medially directed
process. The projection is distinct and extends well
beyond the stipes in most males, whereas it is a
rounded indistinct vestige in nearly all females. The
projection in males may have more taxonomic utility
than currently realized, and future workers may
devise a quantifying index for identifications. Ob-
vious differences in length, breadth, degree of
sclerotization, apical configuration, and whether it is
linear or bowed are potentially useful, but I can only
note them and characterize the structure for descrip-
tions. However, the nubbinlike projection in males
of vy. varius, varius christianus, and varius inter-
grades, which resembles the minute projections of
females, is so much shorter as to be diagnostic for
these forms. Combined with size, this structure
distinguishes these forms of varius without exami-
ning the gonopods or apertures, but unlike other
eurymerodesmids, their sexes cannot be recognized
by the size and shape of the process. The other
subspecies, varius louisianae, cannot be distin-
guished unerringly from sympatric congeners
because its process is longer, broader, and more
typically male-like. Since the projection is short and
subtriangular in clavatus, it may also distinguish this
Texas species from sympatric or parapatric con-
geners. However, more material is needed of
clavatus, known only from the type locality, before
this can be stated with confidence.
Terga. — The dorsum of eurymerodesmids close-
ly resembles those of xystodesmids, and the two
families have been confused on several occasions
probably because of the triangular epiprocts and
generally parallel-sided segments that taper slightly
caudad. The collum is considerably broader than the
succeeding terga but does not always extend below
them (Fig. 89). The anterior surfaces of the paranota
are excavated below the anterior corners, which are
broadly rounded on all segments thus distinguishing
eurymerodesmids from platyrhacids, with which
they are also confused as evidenced by Chamberlin’s
proposal (1942a) of booneus for juveniles of
Auturus evides. American platyrhacids have body
dimensions similar to moderate-size eurymero-
desmids, but they have a broad, truncate epiproct
and small but distinct teeth on the anterior paranotal
corners, which are less rounded than those of eury-
merodesmids. The teeth and truncate epiproct are
MEM. AMER. ENT. SOC., 37
clearly visible on juvenile platyrhacids like the types
of booneus and even early instars, so these two
features invariably distinguish the two families. To
separate eurymerodesmids and xystodesmids, one
must examine the mandibles, the legs, the pre-
gonopodal sternal projections, or the genitalia of
both sexes.
Sterna. — The projections of the sterna in male
eurymerodesmids may also possess taxonomic utili-
ty, a decision left to future workers. In contrast to
xystodesmids, the sterna of male eurymerodesmids
are extremely hairy, particularly those on pre-
gonopodal segments. Eurymerodesmids lack a pro-
jection on segment 4, but segments 5-6 display a
variety of elevations ranging from broad flattened
areas to moderately long knob-like projections, all
shorter than the adjacent coxal widths and apically
hirsute. Likewise between the 9th legs,’ there are a
variety of apically setose elevations and knobs,
which coalesce medially in some forms of dubius
(Figs. 49-51) into a single large projection that can be
longer than the adjacent coxal widths (Fig. 50). The
remaining sterna become progressively flatter
posteriorly with progressively sparser hair patches
subtending the coxae. This is also the condition in
females, which generally lack sternal projections and
have at most only slight elevations on the anterior-
most segments.
Legs. — Eurymerodesmid legs are subsimilar in
size on all segments of both sexes; none are reduced
or crassate. In addition to short, cylindrical, and
truncate gonapophyses, the 2nd coxae of males ex-
hibit variable distal lobes at the anterior and/or
caudal margins, with the former usually being larger
(Fig. 3). Otherwise the legs lack ventral spines and
tubercles, and eurymerodesmids can thus be
distinguished from sympatric xystodesmids by the
absence of the ventrodistal prefemoral spines
characteristic of the latter. However, the prefemora
of males do have large rounded lobes on the dorsal
surfaces (Fig. 4), which differ in size between species
and extend caudad for varying numbers of segments.
In angularis, compressus, goodi, and crassatus the
lobes extend nearly to the caudalmost legs, although
diminishing caudal to midlength, whereas in
clavatus, digitatus, simplex, and varius, they are
restricted to the anterior half of the body, the pre-
gonopodal segments on the last species. In the other
' The gonopods in polydesmoids, located at the anterior posi-
tion of segment 7, represent the highly modified 8th legs. I
therefore refer to the posterior legs on segment 7, the first
postgonopodal pair, as the 9th legs even though they are technical-
ly the 8th ambulatory appendages in adult males.
12 EURYMERODESMID MILLIPEDS
eurymerodesmids, the prefemoral lobes terminate
from segments 10-16, 10-12 on most species. Female
eurymerodesmids and both sexes of xystodesmids
lack these lobes. Eurymerodesmid tarsal claws are
slightly hooked, with a hint of bisinuate curvature in
some species, and arise from a thickened, globose
basal mass.
Male genitalia. — Although the female cypho-
pods distinguish compressus, dactylocyphus, and
some forms of amplus and angularis, most of the
specific characters in Eurymerodesmus involve
aspects of the male genitalia, which in this family
refers collectively to both the aperture and
gonopods.
1. Aperture. — The configuration of the aper-
ture, elaborately ornamented in some species, is the
most obvious characteristic of the Eurymerodes-
midae. Features such as the caudolateral pouches of
oliphantus, paroicus, angularis, and some forms of
amplus, and the lobes of mundus and the western-
most population of b. birdi, are so striking as to
compel attention to their taxonomic utility and func-
tion. Aside from hispidipes, in which the aperture is
uniformly elevated, it is hard to understand how
early workers could have overlooked them. Con-
versely, the gonopods at first glance appear so
uniform as to be virtually devoid of information, but
detailed study reveals that they contain subtle tax-
onomic characters in contrast to the obvious features
of the apertures. Thus, eurymerodesmid determina-
tions rely chiefly on a combination of gonopodal
and aperture characters.
Ironically, the intricacies of the apertures
hampered revision because there are no applicable
precedents. To my knowledge they are unique in the
Chelodesmidea, and perhaps also in the order Poly-
desmida, and therefore require a new format and
terminology. After pondering how to verbally por-
tray the opening, I decided to detail sequentially the
anterior, anterolateral, lateral, caudolateral, and
caudal margins. In addition to structural details,
they also vary in the degree of hirsuteness, which in
some forms is almost as striking as the marginal con-
figurations. Consequently, the aperture descriptions
in the species accounts adhere to the following for-
mat: a) an overview of the aperture describing its
basic type, the general configuration of the opening
itself, width and length measurements, and if dif-
ferent from the opening, the general shape of the en-
tire aperture including marginal adornments; b) the
sequential description of the anterior margin, sides,
caudolateral corner(s), and caudal margin; and c) a
characterization of the aperture hairs in the same
sequence. I recognize two basic types of apertures,
with and without divided lateral margins and
caudolateral pouches. The latter category is further
divided into forms with or without lobes projecting
above the general marginal elevations and located
from the caudolateral corner and the adjoining part
of the sides to near midlength of the caudal margin.
The species displaying these aperture conditions are
listed in table 2. Two general areas are circumscribed
in forms with caudolateral pouches — that of the
opening itself, and that of the opening plus the folds,
flares, and marginal adornments, referred to as the
“aperture apparatus.’’ Both types of apertures
possess indentations at the midline of the anterior
margin, and both extend anteriad in the metazonum,
thereby compressing the prozonum into a narrow
band. This is most apparent by comparing the stric-
ture lines in ambulatory and gonopodal segments, as
shown in figures 5-6 for hispidipes. On midbody
segments, the stricture curves smoothly and
bisinuately, but on the reproductive segment, it
angles anteriad laterally, disappears near the
anterolateral corner, and reappears as a short,
curved, basal line on the anterior indentation. Hoff-
man (1979) stated that the apertures in both
eurymerodesmids and holistophallids protrude into
the prozonum, but my examinations show that this
region is compressed in the Eurymerodesmidae. I
have never been able to trace the stricture com-
pletely, but it is particularly well defined in a few
TaBLE 2. Aperture conditions in Eurymerodesmus.
Sides divided,
with pouches Sides entire, without pouches
With lobes Without lobes
angularis polkensis hispidipes
compressus serratus impurus
varius birdi birdi goodi
amplus mundus dubius
newtonus caesariatus
oliphantus pulaski
elevatus birdi planus
simplex melacis
paroicus dactylocyphus
crassatus sanbernardiensis
digitatus
clavatus
males and can be followed around the antero-
lateral corners to almost join the line on the
anterior indention. The broadest aperture occurs
in dactylocyphus, where it fills the segment width
in ventral view (Fig. 184), and in a few individuals
the aperture extends so far anteriad that the pro-
zonum is reduced to a narrow Sliver, the minimum
necessary to maintain structural integrity of the
ROWLAND M. SHELLEY 13
segment. The stricture is invisible on the anterior
indentation in these males, but its greater angle
laterally further suggests that the prozonum is
compressed rather than penetrated.
a) Anterior margin. — In every species ex-
cept dactylocyphus, where it is elevated into a nar-
row rim that curves around the anterolateral cor-
ners, the anterior margin is flush with the
metazonal surface. All species except caesariatus
possess a midline indentation that varies in length
and breadth, and may be apically rounded to
acute. In goodi, polkensis, paroicus, crassatus,
pulaski, serratus, and sanbernardiensis it is a long,
distinct projection, extending well into the open-
ing proper (Figs. 39, 114, 123, 127, 134, 142, 191);
the opposite extreme occurs in some forms of
dubius, varius, and b. birdi, where it is slight and
barely detectable (Figs. 50-51, 56, 154).
b) Anterolateral corner. — The anterolateral
corner, broadly rounded, poorly defined, and sub-
similar in nearly all species, is more abstract than
real, but is nevertheless a convenient reference
point for the origins of the sides or lateral
margins. Except for dactylocyphus, where it is
elevated, the corner is flush with the metazonal
surface in all species, at least on the anterior part
of the curvature. The lateral elevations begin near
the corner in many species, rising so close in some
that they can truly be said to originate there. Since
this corner is unadorned in eurymerodesmids, it is
omitted from the species descriptions.
c) Lateral margins. — The sides or lateral
margins are the most variable and complex parts
of the aperture. There are two basic types — di-
vided, and undivided or entire — and the eleva-
tions begin at varying distances behind the
anterolateral corner in both. In undivided forms
the elevation usually continues rising to the
caudolateral corner, where it can terminate as in
some forms of dubius (Fig. 46), continue around
the corner onto the caudal margin as in hispidipes
(Fig. 6), or continue caudad, tapering to the
metazonum behind the corner as in impurus (Fig.
11). The rate of elevation varies from uniformly
flattened as in hispidipes (Fig. 6), to gradual as in
digitatus, sanbernardiensis, and clavatus (Figs.
191, 198, 205), to steep as in impurus (Fig. 11).
The elevation may also be upright as in hispidipes
or lean slightly mediad, thus narrowing the open-
ing, as in serratus and some forms of b. birdi
(Figs. 142, 149).
In the type population of birdi planus, the sides
are essentially flush with the metazonum throughout
MEM. AMER. ENT. SOC., 37
their lengths (Fig. 162), and they angle caudomediad
forming a heart-shaped opening in conjunction with
the anterior indentation. Thus in undivided species,
the angle of the sides varies from the condition in b.
planus to nearly linear as in hispidipes (Fig. 6), to
multiangular as in dactylocyphus (Fig. 184). In un-
divided forms the configuration of the apparatus
and the opening are the same, and hence the former
is omitted from species descriptions. Variation ob-
tains in the angle or direction of the sides and the
rate and lean of the elevation. The elevations, par-
ticularly slight ones, are difficult to show from above
in line drawings, some of which are close approxima-
tions of the actual condition.
In species with divided lateral margins, the con-
figuration of the apparatus usually differs from that
of the opening itself; consequently, both are
characterized in descriptions. The divisions occur in
the caudal halves of the sides, the anterior halves
resembling the condition in undivided species, with a
variable elevation, which may be upright or lean
over the opening. At the division point, usually the
highest part of the side, the margin divides into two
lateral margins that continue around the caudo-
lateral corners to form two caudal margins. In most
species the division point is clearly visible in ventral
view, and the ‘‘inner lateral margin’’ angles into the
opening, curves through the ‘‘inner caudolateral cor-
ner,’ and blends into the “‘inner caudal margin.”’
The more variable ‘‘outer lateral margin’’ can con-
tinue the upward slope of the anterior part of the
sides as in elevatus (Fig. 97), descend to a lower level
as in some forms of angularis, varius, and newtonus
(Figs. 21, 56, 84), or flare or angle sublaterad to
varying degrees, in some species extending well
beyond the level of the stigmata, as in oliphantus
and some forms of amplus (Figs. 78, 91). The ‘‘outer
caudolateral corner’’ is more distinct than its inner
counterpart and connects with the ‘‘outer caudal
margin.’’ Up to division point the rim varies from
essentially smooth to slightly irregular. After divi-
sion point, the inner lateral margin is always smooth,
but the outer lateral margin can be as variable as the
undivided part of the sides.
ad) Caudolateral corners. — The caudolateral
corner, in many species imperceptible and poorly de-
fined, is the reference point separating the sides and
caudal margin; there are inner and outer corners in
forms with divided sides. The inner caudolateral cor-
ner is almost always a smoothly rounded blending of
the inner lateral and caudal margins. The outer cor-
ner, however, varies from sharp as in some forms of
amplus and oliphantus (Figs. 77-78, 91), to blunt in
14 EURYMERODESMID MILLIPEDS
some forms of angularis and amplus (Figs. 18, 21,
76), to poorly defined as in newfonus (Fig. 84). It is
usually elevated above the metazonal surface, merg-
ing with the latter a short distance along the outer
caudal margin. The outer and inner lateral margins
are separated to varying degrees, forming a ‘‘pouch’’
or cavity between them at the caudolateral corner.
The cavity likewise varies in breadth, depth, and the
degree to which it is open or closed, or obscured in
ventral view, in turn reflecting the degree of lean in
the outer lateral margin. For example, the pouch in
varius is closed (Figs. 56, 64, 68), whereas that in
olilphantus is open (Fig. 91), and the outer lateral
margin angles strongly sublaterad to form the
broadest pouch in the genus, with the inner margin
visible throughout its length. The pouch in the
variant of amplus named hamatilis by Loomis (1969)
(Fig. 78) is slightly less open than that of oliphantus.
Forms with broadly open caudolateral pouches often
exhibit bizarre configurations of the aperature ap-
paratus, for example trapezoidal in some forms of
varius and oliphantus (Figs. 59, 91), while the actual
opening is broadly ovoid.
Thus, the caudolateral pouch is the cavity between
the outer and inner margins at the caudolateral cor-
ner; it usually gives rise to variable tufts of short
hairs. In most species, the two margins form the in-
ner and outer boundaries of the cavity, and this con-
dition is termed a “‘true pouch.”’ Both e/evatus and
impurus, divided and undivided species, respectively,
display a different pouch (Figs. 11, 97), in which the
lateral elevations continue to rise caudally, and the
inner surfaces are cupped and give rise to hairs.
These are not true pouches because the cavity is
primarily on the inner surface of the outer margin.
The inner lateral margin of e/evatus and the caudal
part of the lateral margin in impurus form the base
of the cavity rather than its inner boundary. Conse-
quently, these pouches are anatomically different
and are termed ‘‘false pouches.”’
The caudolateral corner in undivided species
varies as in divided ones. The only other noteworthy
point is that the lobe is located there in some forms
of b. birdi.
e) Caudal margin. — In divided species, the
caudal margins are essentially parallel and do not
display striking modifications. Occasionally the
outer or both margins may angle slightly caudad in
the midline, even merging with the sternal projec-
tion(s) between the 9th legs as in some forms of
dubius (Figs. 49-50). The outer caudolateral corner
may be slightly elevated, but the outer caudal margin
drops rapidly to the metazonal surface. The caudal
margin is more variable in undivided forms, par-
ticularly b. birdi and mundus, where extremely long
lobes extend above the general marginal elevation,
the locations varying from the caudolateral corner to
near midline of the caudal margin. In mundus and
the westernmost population of b. birdi these lobes
project well below the adjacent coxal margins (Figs.
149-150, 169-170), and similar but smaller lobes oc-
cur in polkensis, pulaski, and serratus. The clavate
lobes of mundus (Figs. 169-171) are always well
removed from the caudolateral corner and located
about 1/3 of the distance along the caudal margin.
The broadly subtriangular lobes of b. birdi can occur
from the caudolateral corner and adjacent part of
the sides to well along the caudal margin. As seen in
caudal profile, they can be in alignment with the suc-
ceeding sternal lobes (Fig. 151) or lateral to these
(Fig. 156). Another undivided species with a striking
feature at the caudolateral corner and caudal margin
is dactylocyphus. Here the sides begin to rise near
midlength, accelerate and extend well ventrad near
the caudolateral corner, and expand onto the caudal
margin. This structure is more of a cupped flange
than a lobe, and hairs arise from the concave inner
surface.
Jf) Aperture hairs. — Aperture hairs are highly
variable throughout Eurymerodesmus. In forms like
b. planus, digitatus, and clavatus (Figs. 162, 164,
168, 198, 205), hairs occur chiefly at the caudolateral
corner and adjacent parts of the sides and caudal
margin. In goodi, hairs arise at the anterolateral cor-
ner and continue onto the caudal margin (Fig. 39),
while in angularis, compressus, oliphantus, simplex,
paroicus, crassatus, b. birdi, mundus, and melacis,
the anterior margin is covered to varying degrees
(Figs. 18, 32, 91, 103, 123, 127, 149, 154, 169, 178).
Hairs tend to be somewhat thicker on the sides of the
anterior indentation, then they taper off and occur
sporadically or regularly along the anterolateral cor-
ners and anterior parts of the sides, arising both
from the rims and from the outer surfaces of the
sides. Densities tend to increase at or near the
caudolateral corners, with tufts in the pouches com-
ing from either the under surfaces of the outer lateral
margins or from the pouches proper. In some forms
of angularis and varius, the division point extends in-
ward into the opening before dividing into inner and
outer margins, and often there will be a hair tuft
under this extended division point (Figs. 23, 59). On
undivided species without lobes, the hair pattern
tapers off on the caudal margins with at most only a
few scattered hairs, usually arranged linearly.
However, the lobes of mundus and b. birdi are so
ROWLAND M. SHELLEY 15
densely pilose, that the surface is obliterated. Hairs
begin basally, become denser and longer distad, and
are directed distad. In some populations of the latter
with moderate-size lobes, the distal hairs are as long
as the lobes themselves (Fig. 155).
2. Gonopods. — Eurymerodesmid gonopods are
undivided, linear to gently curved structures without
a solenomerite or prefemoral process, and feature
long hairs that extend distad along the prefemoral
stem. They are oriented laterally in situ with the can-
nula on the lateral rather than the medial side as in
other chelodesmoids, and the telopodites project
caudad from the aperture instead of anteriad as in
other chelodesmoids. The condition in the Eury-
merodesmidae therefore represents a 180° rotation
in that both the cannula and telopodite have moved
to opposite sides of the aperture; the coxa is also
twisted so that the cannula is exposed in situ instead
of being only slightly visible on the side of the coxa.
In eurymerodesmids, the telopodites overhang and
extend beyond the caudal aperture margin(s) (Figs.
6, 11, 21, 59, etc.), in many overlapping the 9th cox-
ae or the intervening sternum. However, this also
partly reflects compression of the body segments;
occasionally, the anterior segments are so tightly
coiled that the telopodites overhang segment 8. In
dactylocyphus, goodi, and a few other species,
especially those in the melacis lineage, the telopodites
terminate before the caudal margin and are wholly
enclosed by the aperture (Figs. 39, 184, 191, 205), a
condition caused by a very broad aperture and short
telopodites. The only exception to the caudal orien-
tation is caesariatus (Fig. 117), in which the short
telopodites extend directly ventrad, although in a
two dimensional drawing they appear to be directed
anteriad.
In situ configurations vary from parallel and nar-
rowly to widely separated (Figs. 18,32, 91, 123, 127),
to overlapping in the midline (Figs. 21, 59, 73, 76,
149). These arrangements seem to have little tax-
onomic utility, but paroicus and crassatus, with the
broadest, heaviest telopodites, are the most widely
segregated. Telopodite apices are directed variably
laterad to dorsad and are sometimes obscured by
more proximal parts of the structure.
a) Coxae. — The coxae have no taxonomically
significant features as far as I can determine. They
are loosely joined by membrane with no trace of a
sternal remnant and in fact are loosely held in the
aperture on all sides. The membrane holding them in
place tears easily, with that between the coxae being
thicker and tougher. Thus to remove a single
gonopod, I recommend tearing the intercoxal mem-
MEM. AMER. ENT. SOC., 37
brane first while the structures are still attached to
the inside of the aperture.
The coxae are large and globular, lack apophyses,
have a single macroseta on the dorsal surface below
the base of the cannula, hence invisible in situ, and
carry a short, broad sternal apodeme. Their lateral
orientation, representing a 180° rotation from the
condition in the rest of the suborder, is an important
autapomorphy for the family, and the coxa has also
been twisted to bring the cannula to the open or ven-
tral surface. The unpaired macroseta constitutes
another difference from the Xystodesmidae, where
there are two paired marcosetae per coxa. The term,
macroseta, applies to the Xystodesmidae, where it is
longer and stouter than the hairs on the base of the
prefemur, but it is actually a misnomer in the Eury-
merodesmidae because the seta is subsimilar to the
larger ones on the prefemur. However, since it car-
ries no taxonomic value, there is no reason to coin a
new term.
b) Telopodite. — In the ensuing descriptions, I
treat the prefemur and acropodite separately, saying
little about the telopodite, which contains little tax-
onomic information. However, the telopodites do
vary greatly in length, from the short ones of the
melacis lineage to the long ones of the birdi lineage.
The telopodites may be essentially linear, with at
most only a slight curve on the acropodite or the
distal extremity of the prefemur (Figs. 47, 199), or it
may curve or lean to varying degrees over the coxa
(Fig. 185). It may terminate short of (Figs. 47, 86,
93, 111), or at (Figs. 98, 124) the level of the distal
extremities of the hairs. In most species the prefemur
comprises as much as 7/8 of the length of the
telopodite (Figs. 8, 47, 119, 172, 179), but in the im-
purus group, the prefemur is much shorter, con-
stituting from 1/2 to 3/4 of the telopodite length
(Figs. 12, 24, 40). In paroicus, crassatus, pulaski,
and serratus, which have subterminal acropodites,
the prefemur comprises the entire length of the
telopodite.
c) Prefemur. — The prefemur arises centrally
on the ventral surface of the coxa and consists of two
parts — a broad basal mass, closely appressed to,
and partly submerged in, the coxa, and the distal
stem, which is directed caudoventrad to ventrad.
The basal mass extends mediad along the coxa,
bending abruptly ventrad just before the medial
coxal margin, which therefore extends beyond the
bend as seen in lateral profile of some species (Figs.
69, 85, 92, 98). The basal mass is invested with long
matted hairs that are half as long as the prefemoral
stem in many species and are so dense that they
16 EURYMERODESMID MILLIPEDS
obscure the shape of the mass itself. The latter
appears as a silhouette through the hairs, its exact
boundary uncertain. The basal hairs also obscure the
proximal part of the prefemoral stem in lateral view,
so for drawings from this perspective one must ex-
trapolate lines from the visible, distal part of the
prefemur.
The prefemur can be of subequal width
throughout, with sides roughly parallel, or it can
taper or expand distad. The last condition occurs in
simplex and clavatus (Figs. 104-105, 206-207), in
which the prefemur expands distal to midlength into
broad shoulders on the outer and inner margins,
respectively. Eurymerodesmus varius contains in-
termediate forms with strong distal lobes on the
outer surfaces in v. christianus (Fig. 65), intergrade
populations with small lobes at this position (Fig.
63), and forms without a trace of a swelling,
represented by some populations of the nominate
subspecies (Figs. 57-58). No precursor forms are
known for the clavate condition on the inner surface
shown by clavatus.
The most striking characteristic of the prefemoral
stem is the long hairs along the inner surface, a con-
tinuation of those on the basal mass. They are the
only autapomorphy of the family to have been
observed by early authors and have long been used
to distinguish Eurymerodesmus. The hairs are of
several different lengths and breadths, a distinction
that may also possess taxonomic utility. The most
conspicuous hairs on the prefemoral stem are long
and broad, and I like Brolemann’s characterization
(1900) of long, silken hairs. Some species, for exam-
ple serratus (Fig. 145) also display shorter, denser
hairs on the stem. The broadest hairs occur in the
corona of caesariatus (Figs. 118-119), where they are
packed so tightly as to appear sclerotized. The hairs
extend, usually in two rows, along the inner
prefemoral surface for most of its length, or to the
level of the acropodite when this structure is subter-
minal, thereby marking the course of the prostatic
groove. In all species except varius, the hairs are
relatively continuous and regularly arranged, with
only occasional small gaps. In varius, however, they
are sparse and sporadic (Figs. 57-58, 63, 65, 69-71).
All species also have a variable tuft on the outer or
medial margin at the distal extremity of the
prefemur, distal to the inner hairs. The density of
this distomedial tuft, the number of hairs, and the
length it extends proximad vary. Some species also
have two or three scattered hairs proximal to this tuft
on the outer surface, occasionally extending to the
basal prefemoral bend.
d) Acropodite. — In most species of Eury-
merodesmus, the acropodite is very short, only
about 1/4 to 1/8 of the total telopodite length.
However, as the prefemur becomes shorter in the
impurus group, the acropodite becomes corre-
spondingly longer, constituting 1/2 to 3/4 of the
length of the telopodite (Figs. 12-13, 19-20, 22, 24,
33-34, 40-41). The position and basal breadth of the
acropodite are also taxonomically important. It is
broadly terminal in most species, its basal width
being essentially the same as the distal width of the
prefemur, so that the structures blend smoothly
together with an indistinct juncture. Such
acropodites are described as smoothly continuous
with, and poorly demarcated from, the prefemur
(Fig. 47). In varius christianus, the acropodite is
broadly terminal, but is sharply demarcated from
the prefemur because of the latter’s distal lobe (Fig.
65). In simplex, polkensis, and caesariatus, the
acropodite is narrowly terminal, arising solely from
the inner prefemoral margin. The outer margin ter-
minates bluntly in polkensis and caesariatus (Figs.
111-113, 115, 118-119), and expands into a shoulder
in simplex (Figs. 104-105). Eurymerodesmus
clavatus is the obverse in that the narrowly terminal
acropodite arises from the outer prefemoral margin
with the inner surface expanded into a shoulder
(Figs. 206-207). Finally, there are four species in
which the acropodite is subterminal and arises from
the inner margin of the prefemur at varying
distances from the tip — paroicus, crassatus,
pulaski, and serratus. These species demonstrate the
most bizarre, apomorphic gonopods in the family,
because the acropodite is also usually short, and the
prefemur displays modified margins with ir-
regularities, serrations, and/or concavities. In
pulaski and serratus, the acropodite arises in a cavity
on the inner prefemoral surface, which expands into
a hood shielding the structure. In pu/aski the expan-
sion is long and narrow (Figs. 135-136, 138), but in
serratus it is short and broad, with an intricately ser-
rated margin (Figs. 143-145). The acropodite in
pulaski is little more than a thickened boss, but it is
longer in serratus, and the combined effect with the
expanded hood resembles the distal configurations in
the xystodesmids Sigmoria (Cheiropus) planca
(Loomis) and S. (C.) serrata (Shelley) (see Shelley
1984b). Consequently, the acropodite varies from
broadly terminal and poorly demarcated from the
prefemur, to broadly terminal and sharply demar-
cated, to narrowly terminal, to subterminal. These
traits and its length confer more taxonomic utility on
the acropodite than the prefemur, and one’s initial
ROWLAND M. SHELLEY 17
observations of a eurymerodesmid should be of the
acropodite and aperture. Even in broadly con-
tinuous, poorly demarcated forms, the configuration
of the aperture is vitally important in distinguishing
between such species as newfonus and amplus, and
melacis and digitatus. The principle distinction be-
tween the first pair is the uncinate acropodite of
newtonus (Figs. 85-86). Likewise, the acropodite
bends sharply laterad from the prefemur in mel/acis,
whereas it curves broadly dorsad in digitatus (com-
pare Figs. 178-180 and 198-200).
In all species, the prostatic groove runs along the
inner surface of the prefemur onto the acropodite
and opens terminally.
Female genitalia. — In contrast to the Xysto-
desmidae, female cyphopods display several tax-
onomically important features. Some features allow
accurate assignment to species, whereas others allow
placement in a species group. Beyond the cyphopods
themselves, the apertures display elevations, flares,
and other marginal modifications that may also have
taxonomic utility, even though they are not as spec-
tacular as those of males. I did not thoroughly
examine female apertures, and investigations of this
structure are left to future workers.
The cyphopods display the basic chelodesmoid
form, with two variably hirsute valves, open basally
and joined distad, attached to a glabrous receptacle
on the closed, usually lateral, side, with a much
smaller, densely hirsute operculum basally. The in
situ position of the cyphopods varies from transverse
(Fig. 114), to along the body axis (Figs. 42, 201), to
oblique (Fig. 35), and may also have taxonomic im-
portance. Eurymerodesmid cyphopods are com-
paratively large, and in relation to body size are pro-
portionally much larger than those of eastern
xystodesmids. Their various parts are also propor-
tionally large, particularly the receptacle and oper-
culum. The latter lacks taxonomic significance, but
is oblong in profile, wider centrally in head-on view,
and gives rise to long curved hairs from its outer sur-
face. Although the operculum is the smallest compo-
nent of the cyphopods, it is not so minute as to be
practically invisible. It is always evident in front or
head-on view, and except for species in which the
receptacle curves partly around it, the operculum is
also clearly visible in profile.
1. Valves. — The valves are the dominant feature
of the cyphopods, the most visible structures in situ
and larger than the other parts combined. They are
highly variable, equal or unequal in size, and the
open surface is variably hirsute. The joined distal
margin is elevated in many species, diagnostically so
MEM. AMER. ENT. SOC., 37
in a few. In species like varius, the elevation is in-
significant (Fig. 61), but in others it extends into a
distinct rim that protrudes through the aperture.
Such a rim is displayed by amplus, in which it varies
from linear, to angled toward the distal corner, to
angled with a short terminal papilla (Fig. 82). In
some species, the distal margins of the valves are
prolonged into broadly rounded lobes as in dubius,
goodi, and digitatus (Figs. 42-43, 52, 201-202). In
others, the distal corner extends into long dac-
tyliform projections of two main types. In impurus,
angularis, and possibly also crassatus, the projec-
tions are broad, moderately sclerotized, subequal,
and bowed toward each other, the tips touching or
nearly so. The stiff, rigid projections resemble op-
posable digits (Figs. 14, 16, 27-30, 130-132). Valves
exhibiting these projections usually lie transversely in
the aperture, with the projections from each valve
either overlapping or lying next to each other in the
midline (Figs. 14, 27, 130). In compressus, where the
valves lie obliquely in the aperture, the corners ex-
tend into unequal, closely appressed projections.
That of the inner or medial valve is shorter, sub-
triangular, and hidden under the very long, rigid,
dactyliform extension of the outer valve, which pro-
trudes well beyond the aperture and overhangs the
succeeding segment (Figs. 35-37). A similar condi-
tion occurs in some populations of dactylocyphus,
where each valve extends into a long, flexible, dac-
tyliform projection, widely separated from the
other, which projects caudad for two more segments
(Figs. 187-189). Not all populations of dactylo-
cyphus show this extreme development; forms in
Victoria County, Texas, possess variable projections
ranging from short papillae to longer, digitiform
lobes. Regardless of length, the projections of dac-
tylocyphus are flexible and widely separated, as
opposed to the stiff, rigid ones of angularis and
compressus.
2. Receptacle. — Named the ‘‘neckpiece’’ by
Causey (1952a) in descriptions of compressus and
sanbernardiensis, the receptacle is located on the
closed side of the valves, which is usually lateral. It is
also somewhat lower than, or proximal to, the valves
and thus does not extend as far ventrad. The recep-
tacle is glabrous, and in many species the corners of
its sides curve partly around the operculum, thus
partly obscuring it in profile (Figs. 15, 28, 95). In
pulaski the sides are expanded but are not curved
(Fig. 140). Since in other species the sides are neither
expanded nor curved, I judge their condition to have
taxonomic utility and include it in the diagnoses. In
the melacis lineage, the receptacle is typically narrow
18 EURYMERODESMID MILLIPEDS
and compressed into the valves, so as to be barely
visible in profile. In dubius and goodi the receptacle
is absent.
SEXUAL SELECTION
I have not observed any eurymerodesmids copu-
lating and have no direct knowledge of associated
behaviors. Neither are there any published state-
ments, so this section is purely conjectural.
However, several aspects of the male and female
genitalia differ significantly from those in other
chelodesmoids, particularly the Xystodesmidae, and
I think bear on this topic. Genetic variation is ex-
pressed in most male eurymerodesmids through
aspects of the aperture rather than through details of
the gonopods as in xystodesmids. Except for varius,
the gonopods within a species are relatively constant
in the Eurymerodesmidae, and amplus, angularis,
birdi, dactylocyphus, and digitatus exhibit a spec-
trum of aperture variants while maintaining
reasonably uniform copulatory appendages. Like-
wise, females of impurus, angularis, compressus,
crassatus, goodi, dubius, amplus, dactylocyphus,
and digitatus possess cyphopods in which the distal
margins or corners of the valves are markedly ex-
tended, protruding through the cyphopodal aperture
in situ. Even on species like varius, the margins are
elevated into a variable rim. Such conditions do not
occur in the Xystodesmidae, whose valves are blunt
to rounded but never extended. The papillate corner
of the rim in some forms of amplus and the long
dactyliform projections of compressus and dac-
tylocyphus have the appearance of sensory structures
or ‘‘feelers,’’ particularly those of dactylocyphus,
which are flexible. Aside from these structural
features, the entire cyphopods of many preserved
eurymerodesmid females were extruded through the
apertures, with the valves, receptacle, and oper-
culum, lying outside the segment, not just the
valvular projections if present. I have rarely ob-
served this condition in xystodesmids.
With these observations of preserved specimens, I
turn to the concept of sexual selection by females,
advanced by Eberhard (1985) to explain why the
male genitalia have species-specific forms in many
animals and hence are the source of most taxonomic
characters. This hypothesis holds that females
discriminate among males of their own species by the
structure of the latter’s genitalia and that males with
favored morphologies produce more offspring, thus
promoting rapid, divergent evolution of the male
copulatory structures. In some spiders and other
organisms the male organ is first inserted into the
female before being loaded with sperm; it is then
withdrawn, charged with sperm, and reinserted. This
process suggests courtship via the male intromittant
organ and a “‘testing’’ of its configuration by the
female. The above observations of eurymerodesmids
suggest a different procedure, whereby the female
feels or tests the configuration of the male aperture
with the cyphopod valves or the valvular projections
prior to mating, since individual variation in males is
expressed through the aperture. Perhaps the dac-
tyliform appendages actually are ‘‘feelers,’’ to test
the aperture configuration in advance of mating,
and the various distal modifications of the valves
represent the variety of female structures that have
evolved to accomplish this function.
This hypothesis can be tested in two ways. First, it
implies an abundance of sensory cells and neurons in
the valvular projections, which can be searched for
histologically or through transmission electron
microscopy. Secondly, it should be possible to
visually observe some sort of pre-mating testing of
the male aperture by conspecific females, particular-
ly those of dactylocyphus and compressus, which
have the longest valvular projections. Such observa-
tions should be made on captive eurymerodesmids in
the laboratory, where they can be watched con-
tinuously over time. Observations in the field are
more of a chance proposition dependent upon en-
countering a mating pair at this exact moment, and
the mere act of discovery would probably interrupt
mating behavior. To date no one has attempted to
keep eurymerodesmids alive in a laboratory and rear
them, and indeed this has been attempted on very
few larger millipeds. Consequently, maintaining live
colonies of eurymerodesmids and studying repro-
ductive behavior seem fruitful subjects for future
research.
Family EURYMERODESMIDAE Causey
Eurymerodesmidae Causey, 1951:69; 1952b:1; 1954:67. Chamber-
lin, 1952:573. Chamberlin and Hoffman, 1958:78. Loomis,
1959:161. Stewart, 1969:384. Reddell, 1970:399. Hoffman,
1979:159.
Type Genus. — Eurymerodesmus Brolemann,
1900.
Diagnosis. — Relatively stiff, inflexible, small to
moderate-size Chelodesmidea with the following
characteristics: mandibular stipes with basal ridge
extending into variable projection on distal corner,
usually larger in males; sterna unspined, caudal
margins usually straight to slightly curved, with or
without variably hirsute elevated areas, lobes, or
ROWLAND M. SHELLEY 19
knob-like projections adjacent to leg coxae, nar-
rowly or widely segregated in midline, sometimes
coalesced into single large structure, projections
more pronounced in males, becoming progressively
smaller and less hirsute caudally; legs of normal
appearance and proportions for suborder, tarsal
claws uncinate to slightly bisinuate, with globose
basal mass, prefemora without spines and tubercles,
with broadly rounded lobes on outer surfaces of
pregonopodal legs and varying numbers of post-
gonopodal legs in males, 2nd coxae of males with
variable ventrodistal lobes on anterior and/or caudal
margins; gonopodal aperture variably hirsute, con-
figuration highly variable, often elaborately or-
namented, anterior margin usually with variable
midline indentation, sides elevated to varying
degrees, simple or divided to midlength into inner
and outer margins, these continuing into inner and
outer caudal margins and forming setose pouch or
cavity at caudolateral corners, caudal margin with or
without variably subtriangular to clavate, densely
hirsute lobes at varying distances from caudolateral
corners, lobes sometimes enormous and extending
well below levels of adjacent coxae; gonopods struc-
turally simple, oriented laterally in aperture with
cannula located ventrad or ventrolaterad, connected
by membrane only, no sternal remnant; coxae
relatively large, without apophyses, with single
macroseta located dorsad to cannula; telopodite
without prefemoral process, a simple structure
directed caudad to caudoventrad from aperture,
usually curving near midlength to distal extremity of
prefemur; latter varying from 1/3 to 7/8 of
telopodite length, comprising entire structure in a
few species, with long, dense, matted filiform hairs
basally, usually with two to three rows of such hairs
along inner margin, and variable distomedial tuft;
acropodite not divided, usually varying from about
1/8 to 1/2 of total telopodite length, a very short
projection or boss in a few species, usually located
apically and poorly demarcated from prefemur, aris-
ing from inner prefemoral margin in some forms
and subterminally in others; prostatic groove arising
in basal pit in prefemur, opening terminally on
acropodite; cyphopod valves usually relatively large
and moderately hirsute, often with modifications
on distal corners ranging from slight ridge to
variable dactyliform projections protruding through
aperture.
Distribution. — The central, southcentral, and
southeastern United States, ranging from north-
eastern Nebraska, the Missouri River in Missouri,
central Illinois, the Fall Zone region of the southeast,
MEM. AMER. ENT. SOC., 37
and southeastern North Carolina south to northern
Florida and the Rio Grande, and westward to the
central Great Plains of Kansas and Oklahoma and
the Edwards Plateau of Texas (Fig. 1).
Component. — Eurymerodesmus Brolemann.
Genus Eurymerodesmus Brolemann
Eurymerodesmus Brolemann, 1900:101; 1915:554. Chamberlin,
1920:97-98. Attems, 1938:185. Causey, 1950a:267;
1952a:169; 1952b:1; 1954;67; 1963:77. Chamberlin and
Hoffman, 1958:78. Jeekel, 1971:262. Loomis, 1976:287.
Hoffman, 1979:159.
Kewanius Chamberlin, 1938:208. Chamberlin and Hoffman,
1958:82. Jeekel, 1971:268.
Paresmus Chamberlin, 1942b:7. Causey, 1950a:271; 1952a:174;
1952b:4. Chamberlin and Hoffman, 1958:82. Jeekel,
1971:278. Hoffman, 1979:159. NEW SYNONYMY.
Type species. — Of Eurymerodesmus, Poly-
desmus hispidipes Wood, by monotypy; of Ke-
wanius, Eurymerodesmus simplex Chamberlin, by
original designation; of Paresmus, P. paroicus
Chamberlin, by original designation.
Diagnosis. — With the characters of the family.
Color in Life. — Variable; most forms apparently
with pink, orange, or red peritremata, color extend-
ing slightly onto flattened paranotal surfaces;
metaterga varying from speckled black to dark olive-
green with concolorous pink, orange, or red stripes
along caudal margins connecting peritrematal mark-
ings, stripes varying in width, intensity of pigmenta-
tion, and partly or completely interrupted to varying
degrees by base color; collum with concolorous
pink, orange, or red stripes along anterior and
posterior margins.
Description. — A genus of small to large eurymerodesmids with
the following characteristics:
Body composed of head and 20 segments in both sexes; size of
mature adults varying from large, robust forms of about 6.6 mm
wide and 39.5 mm long to minute ones of about 1.6 mm in width
and 14.0 mm in length; W/L ratio similarly varying from about
11.3-19.7%. Body essentially parallel sided in midbody region,
tapering at both ends, more so caudally.
Head of normal appearance, usually smooth and polished, oc-
casionally partly granular. Epicranial suture distinct, terminating
in interantennal region, apically bifid. Interantennal isthmus of
varying width. Genae not margined laterally, with or without
shallow central impressions, ends broadly rounded and projecting
slightly beyond adjacent cranial margins, width variable. Antennae
moderately slender, varying in length, becoming progressively
more hirsute distally, with four short, conical sensory cones on
ultimate article, no other sensory structures apparent. Facial setae
reduced; epicranial absent, clypeal and labral present, with or
without interantennal, subantennal, frontal, and/or genal series.
Mandibular stipes with basal ridge extending into variable projec-
tion ranging from minute, rounded, nubbinlike vestige to long,
heavily sclerotized structure overhanging gnathochilarium, straight
or slightly bowed inward.
20 EURYMERODESMID MILLIPEDS
Terga smooth and polished, occasionally finely granular, with
only trace of wrinkling. Collum considerably longer than remain-
ing tergites, width variable, ends subequal to or extending slightly
below those of following tergite. Paranota distinct but relatively
narrow; varying from flattened, subparallel to substrate, and thus
interrupting slope of dorsum, to variably depressed and relatively
continuous with latter; anterior corners broadly rounded on all
segments, anterior surfaces excavated to varying degrees, more so
in anterior half of body; posterior margins angling anteriad
through about segments 5-6, becoming straight in midbody region
and angling caudad on posteriormost segments; posterior corners
rounded on anteriormost segments, blunt in midbody region,
becoming progressively prolonged and acute caudally. Peritremata
relatively narrow but thick and conspicuous, strongly set off from
paranotal surface, broader caudal to midlength; ozopores located
caudal to midlength, opening laterad. Epiproct moderately long
and subtriangular, apically narrow and blunt.
Sides of metazonites granular or smooth and polished, with oc-
casional shallow, curved impressions and short, acute projections
on first few segments near coxae. Strictures sharp, distinct except
on gonopodal segment. Stigmata small, usually rounded.
Pregonopodal sterna variously modified; those of segments 2-4
normal; 5th and 6th sterna with variably elevated, flattened areas
to narrower paramedian knobs, segregated to varying degrees,
elevations and knobs usually higher and more pronounced on 6th
sternum, all projections variably hirsute. Postgonopodal sterna
with depressions or variably hirsute elevations or projections be-
tween 9th legs, segregated to varying degrees or coalesced in
midline, usually shorter than widths of adjacent coxae, occasional-
ly subequal to or longer than latter, remaining sterna usually flat,
with variably short hirsute projections, or with variable hair
patches subtending coxae, becoming progressively less dense
caudally, occasionally gently rounded, becoming more plate-like
posteriorly, with variable transverse impressions between leg
coxae, caudal margin straight in midbody region. Gonapophyses
short, truncate, cylindrical, located proximad on 2nd coxae. Legs
subequal, of normal length and appearance, 2nd coxae with
variable distal projections on anterior and/or caudal surfaces;
prefemora without ventrodistal spines, pregonopodal prefemora
with variably broad, rounded lobes on outer surfaces, continuing
onto postgonopodal legs in most species and terminating around
midbody region or on caudal legs, 7th and 9th prefemora occa-
sionally with accessory hooks on anterior surfaces; other
podomeres of normal size and appearance on all legs; tarsal claws
hooked or slightly bisinute, with slight to distinct basal globose
enlargement. Hypoproct broadly rounded, extending slightly
mediad; paraprocts with margins narrowly rounded.
Gonopodal aperture extremely variable, often elaborately
ornamented, compressing prozonum into variably narrow, in-
conspicuous band, opening broadly ovoid to elliptical, entire ap-
paratus, including additional space of marginal enlargements,
extending laterad to varying distances over pleural regions, con-
figuration varying from essentially a simple oval or rectangle to
elaborately trapezoidal; anterior margin usually flush with
metazonal surface, almost always with variably broad sub-
triangular indentation in midline; anterolateral corner rounded,
usually flush with metazonum; sides highly variable, usually flush
with metazonal surface at anterolateral corner, occasionally flush
throughout without modifications or elevations, usually with rim
elevated beginning at various distances from anterolateral corner,
upright or leaning mediad to varying degrees and obscuring part of
opening, raised part either of continuous height throughout or
rising slowly or steeply to peak at varying distances caudal to
midlength, thereafter either dropping steeply or slowly to
metazonal surface, lower elevation on sides, or curving around
caudolateral corner for varying distances onto caudal margin, or
with densely hirsute, variably broad, and in some forms enormous
triangular or clavate lobes arising caudally on sides, at caudolateral
corner, or on caudal margin, or divided near midlength with
caudal parts of sides and most of caudal margin consisting of two
margins, inner lateral margin either extending inward into opening
then curving downward, or extending downward and curving and
blending into inner caudal margin, thus essentially continuing cur-
vature of anterolateral margin, outer lateral margin angling or flar-
ing caudolaterad to varying degrees at varying distances from divi-
sion point, either leaning over and obscuring opening or flaring
outward immediately, angle of flare varying from imperceptibly
slight to around 45°, extending laterad, and in some cases caudad,
well beyond levels of stigmata; inner and outer margins forming
variable cavities or pouches at caudolateral corners, either partly or
completely obscured by angle of flare and/or lean of outer lateral
margin or open with strongly flared outer lateral margin;
caudolateral corner variably rounded, blunt or angled, rim flush or
elevated to varying degrees and with or without lobes, usually con-
tinuous with lateral margin, forms with two margins also with in-
ner and outer caudolateral corners, former flush with metazonal
surface and usually smoothly curved, visible or obscured to vary-
ing degrees by outer lateral margin, outer caudolateral corner
rounded, blunt or angled, elevated to varying degrees above
metazonal surface, usually continuous with outer lateral margin;
caudal margin linear or curving smoothly on anterior side of 9th
legs, frequently extending caudad medially and terminating be-
tween 9th legs, occasionally merging with sternal projections
between latter, margin usually elevated laterally, continuous with
sides, elevations tapering or dropping abruptly to metazonal sur-
face at varying distances from caudolateral corner, increasing in
some forms into variably short to enormous triangular or clavate
lobes; forms with pouches and inner and outer lateral margins also
with inner and outer caudal margins, former smoothly continuous
with inner lateral margin, flush with metazonal surface, either
completely inside and essentially parallel to outer caudal margin or
merging with latter near midline, outer caudal margin usually
elevated laterad, continuous with flare or outer lateral margin at
caudolateral corner, tapering to metazonal surface and continuing
along anterior side of 9th legs; aperture margins variably hirsute,
with or without long filiform hairs arising from rim and overhang-
ing opening, shorter and stouter hairs on outer surfaces of lateral
and/or outer lateral margins, and variably dense tufts from
pouches; anterior margin often with 3-6 long hairs along indenta-
tion and scattered hairs elsewhere; lateral margin with sporadic
hairs from rims on anterior halves, hairs on outer surface usually
restricted to caudal halves of sides; marginal lobes densely pilose
with very short hairs arising from base to tip or with hairs becom-
ing progressively denser and longer distad; caudal margin lightly
hirsute or glabrous, usually with at most only a single row of
sporadic hairs. Gonopods in situ with telopodites curving variably
caudoventrad from coxa, touching, overlapping, or narrowly or
widely segregated in midline, extending over caudal margin(s) to
varying degrees with apices directed sublaterad or subdorsad.
Coxae relatively large, usually attaching under lateral margins of
aperture and directed caudomediad, cannulas located ven-
trolaterad to ventrad, curving ventrad and inserting into pit of pro-
static groove in base of prefemur, without apophyses but often ex-
tending slightly ventrad behind base of prefemur, connected by
membrane only, without sternal remnant. Telopodite of variable
length, terminating before, at, or beyond distal extremities of
ROWLAND M. SHELLEY 21
hairs. Prefemur of variable length and width, usually comprising
from 1/3 to 7/8 of telopodite length, in some forms constituting
all of latter, arising ventrally on coxa and extending mediad basal-
ly, closely appressed to and slightly submerged in coxa, curving or
bending ventrad at or just before medial margin of latter and con-
tinuing linearly or in slightly bowed or curved configurations, with
or without slight to moderate distal lobe or shoulder on outer or in-
ner surfaces; usually with dense mass of filamentous hairs, usually
with two to three rows of long, variably spaced filamentous hairs
along inner and lateral surfaces of stem demarcating prostatic
groove, some forms with nearly glabrous stems or with only a few
scattered hairs, usually with distomedial tuft of from 4 to 15-18
hairs. Acropodite usually subtriangular to long, slender, and
subacicular, arising apically or subterminally on prefemur, when
apical smoothly continuous with and poorly demarcated from
prefemur, or discontinuous and sharply demarcated, restricted to
inside or outside of prefemur, length varying from about 1/8 to
2/3 of telopodite length, smoothly curved to sublinear, sides taper-
ing smoothly and continuously to variably acuminate to blunt tip;
acropodite occasionally very short, nubbinlike to a rudimentary
boss, arising subterminally on inner surface of prefemur, shielded
by distal lobe of latter and/or submerged in cavity along inner
margin. Prostatic groove arising in basal pit in prefemur, running
along inner surface of latter to terminal opening on acropodite.
Females usually slightly larger than conspecific males. Agreeing
essentially with latter in somatic features with following excep-
tions: Process of mandibular stipes usually much shorter, a small
nubbinlike vestige to slightly larger rounded lobe. Paranota nar-
rower and more strongly depressed, continuing slope of dorsum,
creating appearance of more highly arched or vaulted body. Sterna
usually flattened and nearly glabrous, transverse grooves less pro-
nounced and becoming even fainter caudally, without lobes, with
sparse hair patches subtending coxae, hairs becoming even sparser
caudally. 2nd coxa without lobes or projections. Prefemora
without lobes on outer surfaces.
Cyphopodal aperture variably rectangular, ovoid, or elliptical,
sides flush with metazonal surface or moderately to strongly
elevated, occasionally flared. Cyphopods in situ either transverse,
oblique, or perpendicular to body axis, valves usually
anterior / posterior to each other with edges or valvular projections
visible in, or protruding through opening, valves occasionally dor-
soventrad to each other in transverse arrangement with edges in-
visible, or arranged along body axis with valves laterad/mediad to
each other and edges visible in opening; corner (tip) of receptacle
often visible in situ. Valves varying in size, subequal to unequal
with either valve larger, distal surfaces variably hirsute, hairs in-
creasing proximad, with or without variable lobes or linear,
angular, papillate, or dactyliform projections usually arising from
distal margins, usually directed ventrad. Receptacle generally sub-
triangular, located laterad, caudad, or dorsad to valves, occa-
sionally submerged in latter to varying degrees, usually smaller
than valves, glabrous, sides either curving distad and partly enclos-
ing or obscuring operculum or not. Operculum varying in size,
small to moderately large, triangular in profile, oblong in frontal
view, with long, curved hairs arising from outer surface.
Synonymy. — Causey (1963) placed Kewanius in
synonymy under Eurymerodesmus based on an ex-
amination of the holotype of E. simplex, the type
species. However, she gave no reasons or illustra-
tions, merely stating that she found no characters
justifying a new taxon. Hoffman (1979) retained this
synonymy. Paresmus was proposed largely because
MEM. AMER. ENT. SOC., 37
of the stouter gonopod of paroicus, the type species
(Chamberlin 1942b, Causey 1950a), and Chamberlin
also indicated that the structure was distally notched
instead of acuminate. However, the most significant
feature of paroicus is the subterminal acropodite
(Figs. 124-125), also displayed by pulaski (Figs. 136,
138), described in Paresmus by Causey (1950a). In
polkensis, assigned to Paresmus in the original
description (Causey 1952b), the acropodite arises
terminally but only from the inner prefemoral
margin. The outer prefemoral surface forms a
shoulder above the acropodite (Figs. 111-113, 115).
The two other species currently in Paresmus, im-
purus and columbus, have broadly terminal acropo-
dites, moderately wide telopodites, and are readily
accommodated by Eurymerodesmus; the latter is
placed in synonymy under dubius, although its type
is lost and this action should be corroborated by
topotypes. Thus paroicus, pulaski, and polkensis
differ primarily in the size of the apical lobe on the
outer surface of the prefemur. In the former two
species it is very large, overhanging and extending
well beyond the level of the acropodite, which
therefore arises subterminally. In the latter species,
the lobe is not expanded, does not overhang the
acropodite, and therefore constitutes a shoulder.
Neither Chamberlin (1920) nor Causey (1963) il-
lustrated simplex, but as shown in figures 104-105, it
displays the configuration of polkensis with a
longer acropodite and a broader, clavate, and more
rounded shoulder. Consequently, the form repre-
sented by paroicus and referrable to Paresmus, with
a subterminal acropodite and an expanded distal
prefemoral lobe, can be derived from that
represented by simplex and referrable to Kewanius
through enlargement of the prefemoral shoulder
such that it extends beyond the acropodite and
causes the latter to arise subterminally. Therefore,
Paresmus should at least be considered a synonym of
Kewanius, which has four years of priority for forms
with distal prefemoral lobes or shoulders. As shown
in figure 65, populations of varius christianus in
Louisiana also display an apical lobe on the outer
surface of the prefemur. This feature is less distinct
in intergrade forms in Mississippi (Fig. 63), and it
disappears entirely in the nominate race on the
Atlantic Coast (Figs. 57-58), which is clearly refer-
rable to Eurymerodesmus. Thus, a transformation
series exists from the subterminal acropodites of
Paroicus and pulaski, to the terminal ones on the in-
ner prefemoral margins of po/kensis and simplex, to
the lobed forms of varius christianus in Louisiana, to
the eastern populations of varius varius with broadly
DP, EURYMERODESMID MILLIPEDS
terminal, poorly demarcated acropodites, subequal
in width to the distal extremity of the prefemur.
Consequently, it follows that Paresmus as well as
Kewanius must fall as synonyms of FEurymero-
desmus.
Distribution. — Same as that of the family.
Species. — 25, arranged into four lineages. Two
species are subdivided into a total of five geographic
races. Future studies based on significantly more
material may show that angularis and amplus should
be divided into additional localized forms,
recognizable at the specific level, and undoubtedly
many more localized forms remain to be discovered
in the Kewanius and melacis lineages. As stated in
the introduction, however, these localized forms
may not possess the full properties of species and
may actually constitute semispecies, in contrast to
birdi and mundus, which are unquestionably
reproductively isolated. Resolution of this theo-
retical matter is beyond the scope of the present revi-
sion, where for consistency with recent studies, par-
ticularly that on Sigmoria (Shelley and Whitehead
1986), I recognize all apparently distinct, localized
forms as well as birdi and mundus at the specific
level. Depending upon how this question is resolved,
Eurymerodesmus will either have many more or less
than 25 species, the former if the localized forms are
evaluated as reproductively isolated, and the latter if
they are found to be less than full species. Con-
ceivably, Eurymerodesmus may have as few as five
species: hispidipes, birdi, and mundus, plus the
melacis and Kewanius lineages collectively.
Remarks. — Although this study is based on ex-
aminations of over 2,000 preserved specimens,
Eurymerodesmus is far from thoroughly sampled.
This is particularly apparent in southern Arkansas
and northern Louisiana, the area with the most
diverse fauna. Fourteen species (mundus, birdi,
angularis, amplus, dubius, paroicus, pulaski,
crassatus, compressus, polkensis, goodi, hispidipes,
simplex, and varius), 56% of the total known fauna,
occur there, with birdi represented by both
subspecies and varius represented by v. christianus
and v. louisianae (Fig. 211). Consequently, this
region should be meticulously sampled to determine
actual distributions. My brief forays into southern
and central Illinois, the New Orleans region of Loui-
siana, and Texas south of Houston and Austin were
not very productive and convinced me that adequate
collecting in the areas of high diversity would take
years and could only be accomplished by someone
based there. The forested part of east Texas, general-
ly north of Houston and east of Dallas, is another
region needing attention.
Without more material and refined knowledge of
distributions, particularly in these states, decisions
on affinities and compositions of lineages and
species groups are tentative. Anatomical patterns
can be detected in both the gonopods and the aper-
ture, but without better knowledge of their distribu-
tions, decisions on relationships are imperfect.
Another difficulty is matching sexes, since some
samples may contain males of one species and
females of another, for example the type series of
angularis and crassatus, and the neotype sample of
pulaski, which also contained specimens of serratus.
Careful field work is needed to check for and record
subtle differences in behavior, color patterns, and
microhabitat, which may indicate different species.
Some questionable samples may contain forms
found close together but in different microhabitats,
which cannot be determined from faded, blanched
specimens and without detailed habitat information
on vial labels; others may contain forms from
relatively close, but nevertheless distinct sites. Thus,
females may not be matched with conspecific males
in some species accounts, and the following descrip-
tions and species groupings are only approximations
based on the incomplete available material. There is
ample room for additional investigation by someone
with the time to conduct meticulous field surveys,
particularly along range peripheries, and record
detailed habitat and color data.
KEY TO SPECIES OF Eurymerodesmus
(based primarily on adult males)
Because of great variation in such species as
angularis, amplus, birdi, and varius, and phenotypic
similarities between variants of others, for example
between oliphantus and forms of amplus, a key to
species of Eurymerodesmus is very difficult to
devise. Few comments can be made about any one
species that do not also apply to another, and the
total range of variation in many species is such that
little can be said that will not exclude an important
variant Added to these standard difficulties with
speciose milliped genera are the complexities of the
aperture and the inherent difficulties in drawing the
structure, verbalizing it, and combining these into
meaningful couplets. Consequently, this key has
limited utility and is best viewed as an introduction,
perhaps more suited to eliminating taxa than to ob-
taining absolute determinations. Use it to reach one
or two final couplets then compare the form in ques-
ROWLAND M. SHELLEY 23
tion with the appropriate illustrations and descriptive
accounts for final determinations. For this purpose,
I have incorporated figure references throughout the
key and attempted to provide more descriptive detail
on variation than usual. The distinctive cyphopods
of dactylocyphus, impurus, angularis, compressus,
and amplus are also incorporated into the key, and
females of these species may be easier to recognize
than males. Eurymerodesmus varius appears twice,
to cover forms with and without prefemoral lobes,
and its three subspecies are keyed in the species ac-
count. Subspecies appear herein for birdi. General-
ized range descriptions are also provided since the
aforementioned difficulties indicate the probability
of more geographically based determinations for
Eurymerodesmus than for most revised milliped
taxa.
1. Acropodite very long, comprising at least half of telopodite length (Figs. 12-13, 19-20, 22, 24, 26, 40-41)...................... 2
Acropodite much shorter, comprising much less than half of telopodite (Figs. 7-8, 33-34, 47-48, 57-58, 63, 65, 69-70, 74-75,
104-105, 111-112, 124-125, 128-129, 135-136, 143-145, 152-153, 206-207)... 1.2... ccc ce ttt eens 4
2. Sides of aperture entire, elevating slowly to peak at or near caudolateral corner (Fig. 39); corners of cyphopod valves rounded but
not extending into segregated projections (Figs. 42-44); Polk and Montgomery cos., AR..............-.005- goodi Causey
Sides of aperture either slightly to moderately elevated and divided caudally into inner and outer margins forming caudo-
lateral pouches (Figs. 18, 21, 23, 25), or undivided but greatly elevated with cupulate flange extending beyond caudolateral
corner and forming an apparent but false pouch between inner surface and metazonum (Fig. 11); distal corners of cyphopod
valves extending into rigid segregated projections, usually bowed towards each other with apices touching or nearly so, occa-
sionally directed oppositely (Figs. 14, 16, 27-30).........
3. Sides of aperture slightly to moderately elevated and divided, with caudolateral pouches (Figs. 18, 21, 23 25); projections of cypho-
pod valves relatively narrow, dactyliform (Figs. 27-30); central MO to northern LA and eastern MS........ angularis Causey
Sides of aperture strongly elevated but not divided, without true caudolateral pouches (Fig. 11); projections of cyphopod valves
relatively broad, not dactyliform (Figs. 14-16); Brazos and Washington cos., TX..............+.e+eee0: impurus (Wood)
4. Sides of aperture divided caudally into inner and outer margins, with variably open to closed caudolateral pouches (Figs. 56, 59,
VTS BA Oils 7/5 TORN rarer a crt ceeto Ree. gare) Sra taE Ioan o Gloria ict boots Glael Mo-cly pet Onna oa onto Male carina Geo oe che tee arnes 5
Sides of aperture entire, without pouches (Figs. 6, 46, 109, 117, 134, 142, 149, 157, 162, 169, 178, 184, 191).................. 18
5. Acropodite essentially smoothly continuous with, and poorly demarcated from prefemur; latter with at most only very slight distal
swelling on outer margin (Figs. 33-34, 47-48, 57-58, 63, 74-75, 85-86) ..... 0... cece cece cece eee eee e eee eens 6
Acropodite not smoothly continuous with prefemur, either subterminal (Figs. 124-125, 128-129, 135-136, 143-145); terminal and
discontinuous, arising from inner prefemoral margin (Figs. 104-105, 111-113, 118-119); or terminal and continuous but sharply
demarcated from prefemur, latter with strong distal lobe on outer margin (Figs. 64, 71)............ cece eee eee e eee 11
6. Prefemur sparsely or lightly hirsute, with light distomedial tuft and at most scattered, sporadic hairs along stem, hairs arranged
irregularly, with variable and sometimes sizeable intervening gaps (Figs. 57-58, 63); southern AR and western LA to FL and
IN Scene gene Pee ope e oy techies TSTOCET Hey sus saat eve Fouls e roel Nen oie
sg RGA ENSIGN AGI dee RSME a SA Sav en adn Dea ae EEN ALS varius (McNeill)
Prefemur moderately to densely hirsute, with variable distomedial tuft and relatively continuous hairs along stem, hairs arranged
regularly or irregularly, but without sizeable gaps (Figs. 33-34, 74-75, 85-86, 92-93, 98-99) .. 00... . ccc cece eee ene iT
7. Acropodite moderately long, longer than 1/4 of telopodite length, broad for most of length, compressed laterally; prefemur without
trace of distal swelling (Figs. 33-34); distal corners of cyphopod valves extended into rigid, unequal, and closely appressed pro-
jections, that of outer (lateral) valve very long and narrow, dactyliform, protruding well beyond aperture and overhanging
succeeding segment (Figs. 35-37); Union Co., AR.......
Cece css Sue NR Ca Oe Re ee eae ee totes eae moskeusie compressus Causey
Acropodite short, no more than 1/4 of telopodite length, relatively narrow with sides usually tapering rapidly, not compressed; pre-
femur usually with slight but detectable distal swelling (Figs. 74-75, 85-86, 92-93, 98-99); corners of cyphopod valves without
dactyliform projections, either not or indistinctly extended into angular ridge, with or without terminal papilla............. 8
8. Acropodite distinctly uncinate, curving downward or dorsolaterad throughout length (Figs. 85-86); caudolateral pouch partly
closed, covered by lean of outer lateral margin (Fig. 84); Benton, Washington, and Newton cos., AR..................-8-
Acropodite either straight or curving distal to midlength (Figs. 74-75, 92-93, 98-99); caudolateral pouch closed to open (Figs. 73,
TSS DSi) doe cage Soe oumac asic ap Han omnis necrne
9. Outer lateral margin rising continuously into broad, elevated lobe at caudolateral corner, not flaring laterad; inner lateral margin
forming floor of pouch, cavity mostly on cupped inner surface of elevation (Fig. 97); Cole and Morgancos.,MO...........
Piao neO aaa ror nere ea hc. crciccrtcacas Ie ea eae elevatus, new species
Outer lateral margin variably rising but not forming elevated lobe, flaring variably laterad; inner lateral margin forming inner edge
of pouch, cavity between outer and inner margins (Figs. 73, 76-78, 91).... 2.2... 6 see cece eee eee eee 10
10. Caudolateral pouch broadly open, outer lateral margin flaring strongly caudolaterad, revealing, entire inner margin in ventral view,
outer surface of outer lateral margin only slightly visible (Fig. 91); corners of cyphopod valves not produced (Figs. 94-95);
southern IL to north-central AR .....................
Bein Rn aoe mics cao OS oliphantus Chamberlin
Caudolateral pouch variable, partly open to completely closed, inner margin usually partly obscured by lean of outer lateral margin,
latter with considerable part of outer surface visible in ventral view (Figs. 73, 76-78); corners of cyphopod valves either im-
perceptibly extended or produced into angular ridge of variable length, with or without terminal papilla (Figs. 80-82); west of
Mississippi River in LA to Wood and Grimes cos., TX, with allopatric population in Mason Co, TX. ........ amplus Causey
MEM. AMER. ENT. SOC., 37
24
11.
12.
13.
14.
15.
16.
17.
18.
19.
20.
21.
2
23.
24.
25.
EURYMERODESMID MILLIPEDS
Acropodite arising subterminally from prefemur; latter variable distad but forming terminal part of telopodite (Figs. 124-125,
1282129: 1352136; 138i VASSV AS): «oct ised, ears Pe Sees cA eae cette ER ee Re ey aoe Cree nee trace ae 12
Acropodite arising terminally, either exclusively from inner prefemoral margin (Figs. 104-105, 111-113, 115, 118-119) or sharply
demarcated by strong distal lobe on outer prefemoral surface (Figs. 65, 71)............ 0.0 cee cece eee ee eee eee 15
Terminal part of prefemur broadly expanded, margin irregularly serrate to jagged (Figs. 143-145); Pulaski Co., AR, and Alachua
0) a) eee ene ere eis sald 5 cia Gu icl & cRNA cick ctid-6, co Uidtenenindet old O's a: AIA AO ovacatiaistp:Saniece Sieve serratus, new species
Terminal part of prefemur variably elongate but not expanded or wide, tapering to blunt or subacuminate tip, margins smooth
(Figs. 124-125,.128-129: 135=136 5 138) socuecousy soecertyoues otto hw apenas meneleren eae le Seen eae ee aes Ieee Ree 13
Acropodite a thickened inconspicuous boss hidden in excavation or concavity on inner surface of prefemur, terminal part of latter
either sublinear or apically uncinate (Figs. 135-136, 138); PulaskiCo., AR........................0.0-- pulaski (Causey)
Acropodite varying from a minute nubbinlike projection to a short spur extending clearly from inner surface of prefemur; latter not
excavated! (Figs: 124-125, 128-129)... sascusonisSeeheveyaecn ree vete ce crests eae ig ave sieyme caine ay nie ereters Danone ee eee 14
Acropodite nubbinlike, indistinct; prefemur with inner margin irregular, terminal part broadly rounded (Figs. 128-129) ..........
Sean tr wn area er eer etme Gre 7 a old cog nar GSO OOo. 0S .00 Sood ood aod crassatus, new species
Acropodite a short, distinct spur; prefemur with inner margin smooth, terminal part tapering to subacuminate tip (Figs. 124-125)
Ae ere ee eet eee nite cea ener cy oan a ROT a oe GES Soe So Uae Uo Be bela aE coe .granos paroicus (Chamberlin)
Acropodite broadly terminal but strongly demarcated from prefemur; latter with outer margin expanding distad into rounded lobe
(Figs. 65, 71), southern AR and western LA to FL and NC............. 0... c ccc e eee e eee es varius (McNeill)
Acropodite narrowly terminal, arising exclusively from inner margin of prefemur (Figs. 104-105, 111-113, 115, 118-119)........ 16
Prefemur distally clavate, sides expanding throughout length to distinct shoulder on outer margin (Figs. 104-105); Evangeline
Pat STA ccc csc ecccaueeactae eke Roe PR Sees RTO ose Sg ee eee simplex Chamberlin
Prefemur with sides relatively parallel throughout (Figs. 111-113, 118-119) ........ 20... ce ee cee eee 17
Terminal margin of prefemur with closely packed, thick hairs forming semicircular corona, lying parallel to and partly obscuring
acropodite, telopodite thus appearing to terminate in calyx; aperture without anterior indentation (Figs. 117-119); Franklin
(rohan ener eetet tetera Ciccone ae ei ea to onlaarS Slanocons RU aoU Oo dho toed eo saaud caesariatus, new species
Terminal margin of prefemur without hairs, acropodite clearly visible; aperture with anterior indentation (Figs. 109-115); Polk
Oo 54 Sere a ee aCe eer eee ey en eee ne en eH PME Cio oOo oomdoS Oto no ooca code dou polkensis (Causey)
Sides of aperture and/or caudolateral corner distinctly elevated above metazonal surface, with or without lobes at corner or on
caudal: margin (Figs. 6, 149) .154, 1517, 169) ieco.<, ue wucueses sree vs aRqe rec cyreucveey cusserm eserere aise lor oaues se eh» eyes) cseeeecue; oLe Ree 19
Sides of aperture and caudolateral corner essentially flush with metazonal surface (Figs. 162, 165, 178, 184, 191).............. 21
Aperture without distinct lobes, marginal elevation of relatively uniform height, extending around caudolateral corner onto caudal
margin (Fig. 6); Coles Co., IL, and Ashley Co., AR ... 2... 0... ccc cece eee neces hispidipes (Wood)
Aperture with densely hirsute, variable lobes on caudal margin or at caudolateral corner (Figs. 149, 154, 157, 169)............. 20
Aperture lobes clavate, very large, clearly located on caudal margin, distinctly removed from caudolateral corner (Figs. 169-171);
northeastern NE to western AR and Johnson CO., TX... 1.2.2.2... 0. c cece ee eee ee eee mundus Chamberlin
Aperture lobes short to long, variably triangular, never clavate, variably higher than marginal elevation, located at caudolateral
corner or on caudal margin at varying distances from latter (Figs. 149-151, 154-158); eastern KS and central MO to San
Patricio Co., TX, Warren Co., MS, and Jefferson Par., LA .............. 0.0 cee eee e eee eee eee birdi birdi Chamberlin
Acropodite narrowly terminal, arising from outer surface of prefemur; latter distally clavate with shoulder on inner margin
Figs..206-207);: Austin @oe.. 1X6. ic siege ccoasaxspeieafeqaueetags. Stasis One Go eo eR eee clavata, new species
Acropodite broadly terminal, smoothly continuous with, and poorly demarcated from prefemur; latter of more or less continuous
width throughout, with or without slight distal swellings but without distinct shoulder, not clavate (Figs. 163-164, 179-180,
185-1865 192-193 ) 199-200) \..)2.cche bo secs See Stee i ces ocr s Sole SD Ohare ae cae ee aoe Pe eee tee eee 22
Telopodites relatively long, extending distinctly beyond caudal margin of aperture and overhanging 9th coxae or sternum between
same (Figs. 162, 165); Rankin Co., MS, to Orleans Par., LA ........... 00.0 cece cece e eee eee eee eee birdi planus Causey
Telopodites relatively short, either wholly enclosed by aperture or barely overhanging caudal margin (Fig. 178, 184,191, 198)... . 23
Aperture very broad, covering essentially entire breadth of segment in ventral view and wholly encompassing telopodites, caudo-
lateral corners elevated into variable flange (Fig. 184); distal corners of cyphopod valves extending into variable papillate to
dactyliform projections, latter flexible, protruding well beyond aperture and overhanging succeeding segments (Fig. 187);
Bosque'to Victoriaicos: TX sin dois races 7 ho eo ron aes Sete Ieee EL ea dactylocyphus, new species
Aperture much smaller, extending to or only slightly beyond levels of stigmata, not nearly covering entire breadth of segment,
telopodites either enclosed within aperture or slightly overlapping caudal margin, caudolateral corners at most only slightly
elevated, not forming flange (Figs. 178, 191, 198); cyphopod valves variable but without dactyliform projections
(Figs... 181-182; W94=195 , 2O2=203) ssc sscosccaeeseoa. a: Foeve- sh ovelsGt Aaron seenece Gs eae OENE IE Le Ck RETEST Oe enc Tara Cee 24
Acropodite bending strongly laterad (Figs. 178-180); Taylor to Terrell and Hidalgo cos., TX...... melacis Chamberlin and Mulaik
Acropodite curving gently or strongly dorsad (Figs. 191-193, 198-200)... 1.0.0... ccc e eee e teen eee eee eens 25
Caudal margin of aperture extending caudad in midline (Fig. 191); acropodite strongly curved (Figs. 192-193); Colorado to
BEAZOPIA COS 5) TX yc openikejes onsets cay eseuendnc ote sd noha GRRE meu REE SOE EEE Le oO ee sanbernadiensis Causey
Caudal margin of aperture linear (Fig. 198); acropodite gently curved, nearly upright (Figs. 199-200); Comanche Co., OK, to
Blanco: COs, TX ed sia. 5cz, 4g eeysceyna ae edee coerce tina: Sat eek OO OE ER Ee digitatus Loomis
ROWLAND M. SHELLEY 25
The Hispidipes Lineage
With raised, vertical, laminate lateral margins that
curve around the caudolateral corner for about 1/3
of the length of the caudal margin, hispidipes differs
from all congeners and merits a separate species
group and lineage, which will become the nominate
subgenus if these categories are shown by future
studies to warrant taxonomic recognition. In other
species lacking caudolateral pouches, the aperture
elevations are thicker and not laminate. The slightly
higher and more densely hirsute terminus of the
elevation on the caudal margin faintly suggests a
lobe, a possible indication of affinity to the birdi
lineage, most forms of which possess densely hirsute
lobes in this position. The hispidipes lineage is com-
posed of one species group and one species.
Eurymerodesmus hispidipes (Wood)
Figs. 2-9, 211
Polydesmus (Polydesmus) hispidipes Wood, 1864:7-8.
Polydesmus (Paradesmus) hispidipes Wood, 1865:220, fig. 48.
Leptodesmus hispidipes: Bollman, 1893:122.
Eurymerodesmus hispidipes: Brolemann, 1915:554. Attems,
1938:185, fig. 203. Chamberlin and Hoffman, 1958:191
(in part).
Type specimens. — Two fragmented male syn-
types and segments 4-7 of one female (NMNH) col-
lected by R. Kennicott on an unknown date from an
unknown site in Illinois. Wood (1864, 1865) did not
mention a holotype nor are any of the fragments so
designated, and it is impossible to match pieces to
form a complete individual. I therefore regard this
material as syntypical. There is a jar labeled
“‘naratype’’ at the ANSP, also from Illinois by R.
Kennicott and therefore probably from the same
series, which consists of shards and debris. The
number of specimens and sexes are unknown, but I
noticed nothing resembling gonopods or an aperture
when sifting through it in 1987. The sample is thus
useless. Wood (1864, 1865) refers to 18 males in the
original descriptions, an unknown number of which
is probably among the debris at the ANSP. For prac-
tical purposes only the two fragmented males at the
NMNH survive.
Diagnosis. — A small to moderate-size species;
males characterized by moderate mandibular projec-
tions; sides of aperture undivided, without caudo-
lateral pouches, with vertical, laminate elevation
curving around caudolateral corner and becoming
progressively higher, terminating abruptly about 1/3
of length of caudal margin; telopodite moderately
long; acropodite short, about 1/8 of telopodite
MEM. AMER. ENT. SOC., 37
length, and broadly terminal, poorly demarcated
from prefemur; latter with many hairs arranged
relatively continuously along inner surface of stem;
females unknown.
Color in Life. — Unknown.
Male syntypes. — Bodies highly fragmented,
lengths unmeasurable, maximum width 3.3 mm.
Head capsule smooth, polished or slightly granular, width
across genal apices 2.8 mm; interantennal isthmus 0.9 mm,
smooth; epicranial suture narrow, linear, terminating in inter-
antennal region, apically bifid. Antennae reaching back to caudal
margin of 2nd tergite, moderately hirsute throughout, first anten-
nomere subglobose, 2-6 clavate, 7 short and truncate; relative
lengths of antennomeres 2=6>3=4=5>15>7, with four apical
sensory cones, no other sensory structures evident. Genae not
margined laterally, with distinct central impressions, ends broadly
rounded and projecting slightly beyond adjacent cranial margins.
Facial setae as follows: epicranial, interantennal, frontal, and genal
absent; clypeal about 4-4; labral about 9-9. Mandibular stipes with
broad projection, in lateral view overhanging lingua lamella of
gnathochilarium, tapering slightly distad to rounded tip on inner
corner (Fig. 2).
Terga smooth, polished or slightly granular; paranota with only
faint wrinkling anteriorly on inner surface. Collum broad, con-
siderably longer than succeeding tergites, ends extending slightly
below those of following tergite, corners narrowly rounded.
Paranota distinct but relatively narrow, flattened and subparallel
to substrate, interrupting slope of dorsum; anterior corners broad-
ly rounded with inner dorsal surface strongly excavated in anterior
half of body, becoming progressively narrower, straighter, and less
excavated caudally; posterior margins angling anteriad through
segment 5, corners blunt on anterior segments, becoming pro-
gressively prolonged and acute posteriorly. Peritremata relatively
narrow but thick and conspicuous, strongly set off from paranotal
surface, broader caudal to midlength. Ozopores located caudal to
midlength, opening laterad. Epiproct moderately long and sub-
triangular, apically narrow and blunt.
Sides of metazonites granular, with occasional shallow, curved
impressions. Strictures narrow but distinct, forming bisinuate
curve on ambulatory segments (Fig. 5), displaced anteriad by aper-
ture on segment 7 (Fig. 6). Sternum of segment 5 slightly elevated
and hirsute adjacent to leg coxae; that of segment 6 with elevations
stronger, broader, and more densely hirsute. Postgonopodal sterna
with transverse impressions between leg coxae varying from strong
and completely crossing 8th sternum to indistinct on 18th sternum;
surface otherwise flat to gently rounded, caudal margin straight in
midbody region, becoming progressively more indented medially
caudad. Legs without spines, moderately and relatively equally hir-
sute throughout, prefemora expanding into broadly rounded lobes
on outer margins, lobes present on pregonopodal legs, becoming
smaller in midbody region and diminishing thereafter; femur
generally longer than other podomeres, tarsal claw varying from
straight to moderately hooked (Fig. 4). Gonapophysis short, cylin-
drical, truncate; 2nd coxa with subconical distal lobes on anterior
and posterior margins, former slightly larger with one seta (Fig. 3).
Hypoproct broadly rounded, extending slightly mediad;
paraprocts with margins slightly thickened.
Gonopodal aperture (Fig. 6) broadly ovoid, without
caudolateral pouch, indented anteriad at midline, 1.9 mm wide
and 1.1 mm long at midpoint, outline of entire apparatus
moderately broadly ovoid; anterior margin flush with metazonal
surface, narrowing mediad and extending into opening as short,
26 EURYMERODESMID MILLIPEDS
broadly triangular projection; sides sublinear, not divided, slightly
indented and flush with metazonal surface at anterolateral corners,
becoming progressively more elevated caudally and continuing
without interruption about 1/3 of distance onto caudal margin,
essentially linear, leaning slightly over opening, rims smooth;
caudolateral corner broadly rounded, nearly a right angle; caudal
margin continuing elevation of sides, elevating slightly and becom-
ing more rounded, then terminating abruptly well short of midline
at about 1/3 length; margins irregularly hirsute with long hairs
arising sporadically from anterior and caudal margins and over-
hanging opening, former with small tufts on sides of indentation
and several other marginal hairs, sides relatively glabrous, caudal
margin with moderately dense tufts on elevation, hairs also arising
from outer surface caudally and caudolaterally, continuing in
linear arrangement along recessed part of margin between lobes.
Gonopods in situ (Fig. 6) with telopodites directed caudoventrad,
curving slightly across midline and crossing opposite member, ex-
tending over caudal margin in midline and overhanging sternum
between 9th legs. Gonopod structure as follows (Figs. 7-8):
Telopodite moderately long, terminating before level of distal ex-
tremities of hairs. Prefemur relatively long, about 7/8 of
telopodite length, linear, tapering slightly near midlength and curv-
ing broadly caudoventrad, with two rows of long, regularly spaced
hairs arising from inner margin and distomedial tuft of ten or so
Figs. 2-8. Eurymerodesmus hispidipes. 2, right mandibular projection of syntype, lateral view. 3, left 2nd coxa of syntype, ventral
view. 4, left 7th leg of male from Coles Co., IL, caudal view. 5, segment 8 of the same, ventral view. 6, aperture and gonopods in situ of
syntype, ventral view. 7, left gonopod of the same, lateral view. 8, the same, medial view. Scale lines for figs. 5-6 = 1.00 mm. Line for
other figs. = 0.60 mm for 2-3, 0.50 mm for 4, and 1.00 mm for 7-8.
ROWLAND
hairs. Acropodite short and broadly terminal, about 1/8 of
telopodite length, arising apically at slight constriction of
prefemur, directed dorsad, slightly hooked or bisinuate, tapering
rapidly to subacuminate tip.
Female Syntype. — Among the fragments in the
NMNH sample are segments 4-7 of a female, which
lack structural peculiarities. The sterna resemble
caudal postogonopodal sterna of males with in-
complete transverse grooves between the leg pairs.
Segment 3 is missing, and Wood’s descriptions of
the cyphopods (1864, 1865) do not reveal significant
features. Thus for practical purposes, females of
hispidipes are unknown.
Variation. — Wood (1864, 1865) noted dif-
ferences in color among individuals in the type
series. He suggested that they might represent a
distinct species but did not propose a name because
of similarities in the genitalia. Since all traces of col-
or and pattern have vanished after 125 years in
preservative, I cannot confirm Wood’s observations.
However, the gonopods of the two surviving males
differ in that the acropodites of one are bisinuate
and more strongly demarcated from the prefemur,
while those of the other are slightly curved. The
aperture configurations are practically identical. The
left lateral margin of the Coles County male flares
outward slightly caudolaterad, whereas the margin
on the right is straight. The acropodites of this
specimen are slightly bisinuate and moderately
demarcated from the prefemur. The degree of hir-
suteness of the aperture is subsimilar to that in the
syntypes, and the only facial setae present are the
clypeal (about 11-11) and labral (about 14-14) series..
The Arkansas male possesses a few scattered hairs
on the lateral margins, but otherwise the aperture
and gonopods closely resemble those of the Coles
County male. Facial setae of the Arkansas male are
subantennal 1-1, frontal 1-1, genal 2-2, clypeal about
10-10, and labral about 18-18. The specimen
measures 22.0 mm in length, 3.9 mm in width, W/L
ratio 17.7%.
Ecology. — The Coles County specimen was
taken from a clod of dirt in a ploughed field.
Nothing is known about the habitat of either the
type series or the Arkansas specimen.
Distribution. — Known definitely only from
Coles County, Illinois, and Ashley County, Arkan-
sas (Fig. 9). Wood (1864, 1865) did not specify a site
or general area in Illinois, and the record from Dix-
on Springs, Pope County (Causey 1950a, Chamber-
lin and Hoffman 1958), incorrectly placed in Lee
County by the latter authors, refers to oliphantus.
MEM. AMER. ENT. SOC., 37
M. SHELLEY 2a
Specimens were examined as follows:
ILLINOIS. Unspecified locality and date, 2M,
F, R. Kennicott (NMNH) SYNTYPES and
unknown number of specimens (ANSP) “‘PARA-
TYPES.”’ Coles Co., Charleston, Normal School
(now Eastern Illinois University), M, 13 May 1918,
T. L. H. (EIU, transferred to NCSM).
ARKANSAS. Ashley Co., 6.1 mi. E. Hamburg,
M, 23 April 1965, collector unknown (FSCA).
Remarks. — The two known localities are
separated by about 470 miles, a great distance cast-
ing doubt on the accuracy of the Arkansas record.
However, other eurymerodesmids cover large areas
and have samples from disparate localities known to
be correct, most notably serratus from Pulaski
County, AR, and Alachua County, FL. I therefore
consider the Arkansas record to be correct and show
it in the distribution map (Fig. 9).
On May 20, 1989, 81 years and one week after col-
lection of the Coles County male, I revisited this area
of eastern Illinois to attempt to secure more
material, particularly of females, whose genitalia are
unknown. I spent two days working in wooded areas
along the Fox River and tributaries, and turning logs
and rocks in clearings, all to no avail. I therefore
believe that this male represents a thoroughly for-
tuitous discovery and that future collections are as
likely to occur through chance accidents as through
planned, systematic field work.
For years hispidipes was considered the only or
most abundant species of the genus and family, and
the name was routinely applied to forms in other
parts of the generic range as recently as 1969, after
many other nominal species had been diagnosed. I
have examined the specimens on which some of
these records are based and review them in Table 3
below. Others are lost but can be assigned
reasonably accurately to one or two species from
knowledge of distributions. All references to
hispidipes are cited in the synonymies of the ap-
propriate species. Since they actually refer to
Wood’s original citations, only Bollman (1893),
Brolemann (1915), Attems (1938), and Chamberlin
and Hoffman (1958) cite hispidipes correctly at least
in part.
The Kewanius Lineage
I combine in one lineage all species in which the
sides of the aperture are divided caudally into inner
and outer margins with pouches at the caudolateral
corners, or which appear related to such species
through shared traits of the gonopods or aperture.
The assemblage is heterogeneous, but all com-
28 EURYMERODESMID MILLIPEDS
Table 3. Previous literature citations of hispidipes.
Author Date Locality Species
Wood 1864, 1865 Original descriptions, unspecified site in IL hispidipes
Bollman 1888a Unspecified sites in AR pulaski, serratus, and per-
haps others
Bollman 1893 Refers to original description hispidipes
Brolemann 1895 Unspecified site in LA amplus
Brolemann 1900 None probably amplus
Gunthorp 1913 Cowley, Douglas, & Jefferson cos., KS Cowley probably mun-
dus, others probably birdi
Brolemann 1915 None hispidipes
Chamberlin 1918 Creston, Natchitoches, Par., LA birdi and amplus
Attems 1938 IL and LA IL = hispidipes
LA = amplus
Chamberlin 1942a Louisiana birdi and amplus
Causey 1950a Dixon Springs, Pope Co., IL oliphantus
Causey 1952b aperture drawing probably of Dixon Springs male oliphantus
Chamberlin & Hoffman 1958 refer to IL in general and Dixon Springs hispidipes and oliphantus
Stewart 1969 Angelina, Nacogdoches, Rusk, Shelby, Smith, amplus
Sabine, and Upshur, cos., TX
ponents are linked either through the aperture,
gonopods, or both, and are thus more closely related
to each other than to ones in other lineages. With 17
species this is the largest branch of Eurymero-
desmus, extending from the Atlantic seaboard to
southern Illinois, central Missouri, western Arkan-
sas, and eastern Texas. For convenience, it is divided
into four species groups based on gonopod struc-
ture. It contains the second oldest species in the
family, impurus, and the type species of the genus-
group names, Kewanius and Paresmus. Since the
former has priority by four years and thus is
available for future subgeneric designation, it is
employed here to designate the lineage.
The Impurus Group
This group contains the four species of Eury-
merodesmus with long acropodites, subequal to or
only slightly shorter than the prefemora, or about
1/3 to 2/3 of the telopodite length. Since the
acropodite is longer than in the other groups, the
telopodite appears longer and typically extends
beyond the level of the distal extremities of the hairs.
The distal corners of the cyphopodal valves are
moderately prolonged in goodi and extended into
curved rigid dactyliform projections in impurus and
angularis. Eurymerodesmus compressus has a
shorter acropodite, around 3/8 of the telopodite
length, but it belongs in this group because the struc-
ture is still longer than those of other groups and
because its cyphopod valves are produced into long,
tapered projections. That of the lateral valve is
Fig. 9. Distributions of the hispidipes lineage, and the impurus
and dubius groups. Stars in dots, hispidipes; stars, impurus; dots,
angularis; squares, compressus; asterisks, goodi; triangles, dubius.
Open squares and triangles denote literature records considered
reliable.
ROWLAND M. SHELLEY 29
longer, and that of the medial valve is closely ap-
pressed to its inner surface.
The impurus group occurs primarily in southern
Arkansas and northern Louisiana, with outlying
localities in central Missouri and north-central
Mississippi. Eurymerodesmus impurus itself is allo-
patric, occurring some 240 miles to the southwest in
eastern Texas adjacent to the melacis lineage. Addi-
tional forms with long acropodites may await
discovery in northeast Texas.
Components. — impurus (Wood), angularis
Causey, compressus Causey, goodi Causey.
Eurymerodesmus impurus (Wood),
new combination
Figs. 9-16, 212
Polydesmus impurus Wood, 1867:43. Attems, 1940;493.
Leptodesmus impurus: Bollman, 1893:122.
Paresmus impurus: Causey, 1952a:174-175, figs. 8-9. Chamberlin
and Hoffman, 1958:82.
Type specimens. — Two male syntypes (ANSP)
collected by Gideon Lincecum on an unknown date
in 1866 at Long Point, Washington Co., TX. The
vial contains parts of two highly fragmented, in-
complete males with two epiprocts, two gonopodal
segments, but only one head. Both gonopods are
missing from one gonopodal segment, and their
location is unknown. The vial contains two labels
with partly conflicting messages. One states
**Polydesmus impurus Wood, type,’’ and the other
reads ‘‘Leptodesmus impurus Wood, probably
type.’’ There is a slide at the AMNH labeled ‘‘Lep-
todesmus impurus Wood, Dr. Lincecum, probably
type, 1506 Phila. Acad.,’’ which is empty but may
have once contained the lost gonopods from the
male at the ANSP. However, the number 1506 does
not match those in the vial. The word ‘‘probably”’
on one label indicates some doubt that this is the
type material of impurus, but it must be considered
as such with no other specimens available.
Diagnosis. — A moderate-size species; males
characterized by long mandibular projection; sides
of aperture undivided, without true caudolateral
pouches, elevating steeply and extending beyond
caudolateral corner, inner surface concave thus
forming false pouch; caudal margin flush with
metazonal surface; telopodite long; acropodite very
long, about 2/3 of telopodite length, broadly termi-
nal, moderately demarcated from prefemur; latter
with moderate number of hairs arranged con-
tinuously along inner margin; cyphopod valves of
females with bowed projections from distal corners,
MEM. AMER. ENT. SOC., 37
apices touching or nearly so; receptacle large, sides
curving partly around operculum.
Color in Life. — Peritremata reddish; metaterga
light with darker mottled areas, with reddish stripes
along caudal margins connecting paranotal mark-
ings; collum with reddish stripes along both anterior
and posterior margins (Wood 1867).
Syntypes. — Bodies highly fragmented, lengths
unmeasurable, maximum width 2.8 mm.
Somatic features similar to hispidipes, with following excep-
tions:
Width across genal apices 1.6 mm, interantennal isthmus 0.4
mm. Antennae reaching back to caudal margin of 2nd tergite,
relative lengths of antennomeres 2>5>6>4>3>1>7. Genae
with faint central impressions. Facial setae as follows: epicranial,
interantennal, subantennal, frontal, and genal absent, clypeal
about 8-8, labral about 14-14. Process of mandibular stipes
relatively long, broadly rounded apically (Fig. 10).
Collum extending slightly below ends of following tergite.
Paranota relatively flattened, interrupting slope of dorsum,
posterior corners rounded to midbody segments, becoming blunt
and progressively more acute thereafter.
5th and 6th sterna with low, rounded knob-like projections
between both pairs of leg coxae, narrowly segregated in midline,
caudal projections slightly larger than anterior ones on each seg-
ment. Postgonopodal sterna with two large, widely separated pro-
jections between 9th legs, lengths subequal to widths of adjacent
coxae, with tuft of long hairs arising apically and from medial sur-
face; remaining sterna without projections but with variable
bicruciform impressions and hair patches adjacent to leg coxae,
becoming flatter and more sparsely hirsute caudally. 2nd coxae
with conical anterodistal projection, without caudal projection.
Prefemoral lobes extending through segment 11.
Gonopodal aperture (Fig. 11) ovoid, without true caudolateral
pouch but with concavity on inner surface of lateral flange, with
moderate anterior indentation, 1.7 mm wide and 0.8 mm long at
maxima, outline of entire apparatus rectangular; anterior indenta-
tion broadly rounded, triangular; sides not divided, elevating
steeply in anterior half, leaning slightly over opening and continu-
ing ventrad as laminate triangular flange, extending caudad
beyond caudal margin, terminating abruptly, and dropping to
metzonal surface beyond caudolateral commer, with shallow con-
cavity on inner surface, rims essentially smooth; caudal margin
flush with metazonal surface, discontinuous with sides, arising
from inner surface of lateral flange, curving slightly caudad in
midline; margins glabrous except for moderate hair tufts arising
from inner surfaces of concavities of lateral flanges. Gonopods in
situ (Fig. 11) with telopodites curving mediad, touching opposite
member in midline, then curving laterad, extending beyond caudal
margin of aperture and terminating over 9th coxae. Gonopod
structure as follows (Figs. 12-13): Telopodite relatively long,
extending just beyond level of distal extremities of hairs. Prefemur
short, about 1/3 of telopodite length, terminating in slight swelling
beyond ventral bend, with two rows of continuous, regularly
spaced hairs along inner margin, distal tufts of 8-12 hairs on
medial and lateral surfaces, and one medial hair at level of bend.
Acropodite broadly terminal and very long, much longer than
prefemur and about 2/3 of telopodite length, slightly narrower
than distal extremity of prefemur, inner margin continuous with
latter, outer margin curving inward slightly basally, leaning slightly
over coxa but extending nearly directly caudad, expanding slightly
30 EURYMERODESMID MILLIPEDS
Figs. 10-16. Eurymerodesmus impurus. 10, right mandibular projection of male from Brazos Co., TX, lateral view. 11, aperture and
gonopods in situ of holotype, ventral view. 12, left gonopod of the same, lateral view. 13, telopodite of the same, medial view.
14, cyphopods in situ, ventral view of female from Brazos Co., TX. 15, left cyphopod of the same, caudal view. 16, the same, medial
view. Scale lines for figs. 11 and 14 = 1.00 mm. Line for other figs. = 0.60 mm for 10 and 15, 0.70 mm for 12 and 16, and 1.00 mm for
13.
ROWLAND M. SHELLEY 31
then narrowing abruptly apically and bending sharply sublaterad,
tip subacuminate.
Female from Brazos County, TX. — Length approximately
18.7 mm, maximum width 3.2 mm, W/L ratio 17.1%. Agreeing
closely with types in somatic features, with following exceptions:
Genal setae 3-3. Corner of mandibular stipes slightly produced,
apically blunt. Sterna flat, unmodified, glabrous.
Cyphopodal aperture elliptical, sides strongly elevated above
metazonal surface. Cyphopods in situ (Fig. 14) with valves ex-
truded through aperture, almost completely exposed, oriented
transversely, open side directed dorsad. Valves (Figs. 15-16) large,
unequal, anterior valve larger, distal corners extending into broad,
curved opposing projections, that of anterior valve broader,
apically blunt, and directed subcaudad, that of caudal valve more
acute, curving anteriad. Receptacle large, glabrous, located lateral
to valves, extending well below latter, sides curving broadly
mediad and partly enveloping operculum. Latter small, tucked
under open (dorsal) side of valves.
Variation. — Measurements of a male from
Brazos County are length 27.0 mm, maximum width
4.0 mm, W/L ratio 14.8%. Facial setae include
frontal and subantennal series, 1-1 each. The aper-
tures in these males are wider than those of the types,
since the lateral flanges project directly ventrad or
angle sublaterad instead of leaning over the opening.
The apices of the flanges are squared rather than
pointed, and a few scattered hairs arise from the
anterior margin, the lateral rims, and the outer sur-
faces of the flanges. The apices of the acropodites
curve more strongly than do those of the types.
Ecology. — According to Wood (1867), the types
were collected under dried cow dung, hence the
specific name. As noted by Causey (1952a), it is not
unusual for millipeds to occur under dung in grass-
land ecosystems, where there is little other cover.
Distribution. — Known only from Washington
and Brazos Counties, in the Coastal Plain of south-
eastern Texas (Figs. 9, 212). Material was examined
as follows:
TEXAS: Washington Co., Long Point, 2M, G.
Lincecum (ANSP) TYPE LOCALITY. Brazos Co.,
College Station, 2M, 4F, 23-24 December 1905, col-
lector unknown (MCZ).
Remarks. — Wood’s original description of im-
purus (1867) is internally contradictory about the
gonopods. He mentions ‘‘two spines,’’ the larger
being regularly curved except apically, where it is
abruptly bent. The smaller is “‘slender, curved, and
acute.’’ Since the telopodite is undivided, there is no
prefemoral process, and the ‘“‘larger’’ projection is
clearly the telopodite, I surmise that Wood mistook
one or more hairs as the second projection. Wood
also contradicts himself regarding the distance that
the hairs extend along the telopodite. He first states
that the “blunt distal end is covered with long
hairs,’ then mentions that neither of the “‘two
MEM. AMER. ENT. SOC., 37
spines’’ is hairy. Only with the present illustrations
can this confusion be clarified. The prefemur of im-
purus is relatively short and the acropodite com-
paratively long; thus the telopodite does not appear
as hairy as those in other species groups.
Causey (1952a) reported that Dr. Lincecum, a
pioneer physician and Texas naturalist, traveled
widely in the state, that it was impossible to
determine the collecting site, and that his last home
was ‘“‘Bonham, Washington County.’’ However,
Bonham is in Fannin County, in north Texas ad-
jacent to Oklahoma. The town in Washington
County is Brenham, and one cannot determine
whether her error is in the town or county. However,
the Lincecum papers at the University of Texas
library reveal that, in 1866, he or a few assistants col-
lected all the Texas myriapods described by Wood
(1867) at his home in Long Point, near Brenham (J.
Reddell, pers. comm.). This site is therefore the type
locality.
I visited Washington County in February 1986 to
try to recollect impurus and verify the type locality.
This effort was unsuccessful, but the samples from
adjacent Brazos County confirm its occurrence in
Washington County. Causey (1952a) reported that
the types lacked gonopods, but they were present on
one specimen when I examined them in 1987, so she
did not thoroughly examine the sample. Her
sketches of the aperture from ventral and lateral
views, some 85 years after its description, were the
first illustrations of impurus, but they do not ade-
quately portray the size of the lateral flange. The
gonopods were omitted since she overlooked them,
so the long acropodites were previously unknown,
although as stated above Wood (1867) alludes con-
fusingly to this condition in the original description.
Without illustration, it is not surprising that Causey
did not deduce the nature of the telopodite. The vir-
tual absence of meaningful illustrations of the four
oldest referrable names — hispidipes, impurus,
varius, and simplex — has greatly hampered identi-
fications in Eurymerodesmus and use of these
animals in other studies.
Eurymerodesmus angularis Causey
Figs. 9, 17-30, 211
Eurymerodesmus angularis Causey, 1951:69-71, figs. 1-3. Cham-
berlin and Hoffman, 1958:78.
Eurymerodesmus wellesleybentoni Causey, 1952a:171-174, fig. 5.
NEW SYNONYMY.
Eurymerodesmus wellesleybentonus: Chamberlin and Hoffman,
1958:81.
32 EURYMERODESMID MILLIPEDS
Figs. 17-30. Eurymerodesmus angularis. 17, right mandibular projection of male from Jefferson Co., AR, lateral view. 18, aperture
and gonopods in situ of the same, ventral view. 19, left gonopod of the same, Jateral view. 20, telopodite of the same, medial view.
21, aperture and gonopods in situ of male from Phillips Co., AR, ventral view. 22, telopodite of left gonopod of the same, medial view.
23, aperture and gonopods in situ of male from Union Co., AR, ventral view. 24, telopodite of left gonopod of the same, medial view.
25, aperture and gonopods in situ of male from Caddo Par., LA, ventral view. 26, telopodite of left gonopod of the same, medial view.
27, cyphopods in situ, ventral view of female from Jefferson Co., AR. 28, left cyphopod of the same, caudal view. 29, the same
dorsomedial view. 30, cyphopods in situ, ventral view of female from Ouachita Par., LA. Scale lines for figs. 18, 21, 23, 25, 27, and 30 =
1.00 mm. Line for other figs. = 0.30 mm for 17 and 28-29, and 0.50 mm for 19-20, 22, 24, and 26.
ROWLAND M. SHELLEY 33
Type specimens. — Male holotype and female
allotype (ANSP) collected by L. Gray, 1 April 1950,
at DeValls Bluff, Prairie Co., AR. The allotype
lacks the valvular projections found on females with
other males of angularis and probably is not con-
specific with the holotype. Consequently, a specimen
from adjacent Jefferson County is used for the
female description.
Diagnosis. — A large species; males characterized
by short mandibular projection; sides of aperture
divided, with variably open to closed caudolateral
pouches; telopodite long; acropodite variably long,
1/3 to 3/4 of telopodite length, broadly terminal,
poorly demarcated from prefemur, curving gently to
strongly dorsolaterad, configuration varying from
obtuse angle to falcate; prefemur with moderate
number of hairs arranged continuously along inner
margin; cyphopod valves of females with dactyli-
form projections from distal corners, varying from
bowed with apices touching or nearly so to linear
and directed oppositely; receptacle large, sides curv-
ing partly around operculum.
Color in Life. — Paranota orange; metaterga
dark mottled olive to black-brown, with or without
orange stripes along caudal margins; collum with
orange stripe along anterior margin (Causey 1951,
1952a). :
Holotype. — Length 36.4 mm, maximum width
4.9mm, W/L ratio 13.5%.
Somatic features similar to hispidipes, with following excep-
tions:
Width across genal apices 3.0 mm, interantennal isthmus 1.0
mm. Antennae reaching back to midlength of 3rd tergite, relative
lengths of antennomeres 2>3>4>5=6>1>7. Genae with faint
impressions. Facial setae as follows: epicranial 2-2, interantennal
and genal absent, subantennal 1-1, frontal 1-1, clypeal about
11-11, labral about 23-23. Process of mandibular stipes short,
broadly rounded (Fig. 17).
Collum extending slightly below ends of following tergite.
Paranota moderately flattened, interrupting slope of dorsum;
posterior corners rounded through segment 6, blunt on 7-11,
becoming progressively more acute caudad.
5th sternum with low, sparsely hirsute, paramedian knobs
between 4th legs and larger, more conical ones between Sth legs;
6th sternum with low, rounded, subequal knobs between 6th and
7th legs, each sparsely hirsute. Postgonopodal sterna with low,
slightly hirsute lobes between 9th legs; segments 8-10 with minute,
sparsely hirsute, paramedian knobs between anterior legs and
larger, conical, sparsely hirsute knobs between caudal legs; remain-
ing sterna with knobs between caudal legs becoming progressively
shorter caudad, disappearing on segment 18. 2nd coxae with
distinct, subconical anterodistal lobes and shorter, acute caudal
ones. Prefemora with broad lobes on outer surfaces of all legs,
becoming considerably smaller on caudalmost segments.
Gonopodal aperture (Fig. 18, not this specimen) broadly sub-
ovoid, with broad, open, caudolateral pouch and strong anterior
indentation, 2.6 mm wide and 1.3 mm long at maxima, outline of
MEM. AMER. ENT. SOC., 37
apparatus subtrapezoidal, extending laterad well beyond stigmata
and cephalad nearly to anterior segmental margin, severely con-
stricting prozonum; anterior indentation broadly triangular; sides
divided, angling slightly caudolaterad and becoming increasingly
elevated to division point, inner lateral margin angling dorsad into
Opening then curving into inner caudal margin, outer lateral
margin tapering gradually dorsad, flaring strongly laterad and
curving broadly into blunt caudolateral corner, forming broad,
deep, open pouch with inner lateral margin, rims smooth; inner
caudolateral corner blunt; outer caudolateral corner blunt; inner
caudal margin sublinear, subparallel to outer caudal margin; latter
sloping to metazonum, curving gently along anterior surfaces to
coxal condyles, extending slightly caudad between legs; margins ir-
regularly hiruste with long hairs arising sporadically from anterior
and lateral rims and outer surfaces and overhanging opening, also
with moderate tufts of hairs in caudolateral pouches and scattered
hairs along outer caudal margin. Gonopods in situ (Fig. 18, not
this specimen) with telopodites extending caudoventrad from
medial margins of coxae and touching in midline, apices curving
dorsolaterad over caudal margins of aperture. Gonopod structure
as follows (Figs. 19-20): Telopodite moderately long, terminating
at level of distal extremities of hairs. Prefemur short, approxi-
mately 1/3 of telopodite length, tapering slightly distad into in-
distinct medial shoulder, with two rows of continuous, regularly
arranged hairs along inner margin and dense distomedial tuft of
around a dozen hairs. Acropodite long and broadly terminal,
about 2/3 of telopodite length, continuous with, and poorly
demarcated from, prefemur, curving dorsolaterad in about a right
angle proximal to midlength, narrowing continuously to
subacuminate tip.
Female from Jefferson Co. — Length 29.9 mm, maximum
width 4.1 mm, W/L ratio 13.7%. Agreeing closely with holotype
in somatic features, with following exceptions: Mandibular stipes
with short, acuminate process. Sterna flat, unmodified, with hair
tufts adjacent to leg coxae, becoming progressively sparser
caudally.
Cyphopodal aperture ovoid, sides strongly elevated above
metazonal surface, thickened. Cyphopods in situ (Fig. 27) with
valves submerged and oriented transversely in aperture, valvular
projections directed mediad and abutting or overlying each other
in midline. Valves (Figs. 28-29) moderately large, subequal, cor-
ners extending into moderately long, dactyliform projections,
curving toward each other and nearly touching apically. Receptacle
large, located dorsolaterad to valves, corners curving partly around
operculum. Latter small, located under free, dorsal, end of valves.
Variation. — I include under angularis all forms in
Arkansas, Louisiana, and Mississippi with long
acropodites, at least 1/3 of the length of the
telopodite, that curve or bend by midlength and
whose apertures display divided lateral margins with
caudolateral pouches that vary from open to closed.
The sides are entire and pouches are absent from
goodi, whose acropodites curve apically. Eury-
merodesmus angularis is highly variable in both
sexes, and the only plausible alternative is several dif-
ferent species. The situation is analogous to that with
the xystodesmid Deltotaria brimleii brimleiti Causey
in the southern Appalachian Mountains, where a
host of dissimilar forms, none demonstrating struc-
tural stability over a broad geographical area, con-
34 EURYMERODESMID MILLIPEDS
nect through intermediate populations that bridge
the anatomical gaps (Shelley and Whitehead 1986).
Intervening forms are unknown in angularis, but
future collections are expected to show that the
localized forms blend together into a single tax-
onomic unit. A few, such as the falcate form in
Union County, Arkansas, may warrant taxonomic
recognition as subspecies, but these decisions are left
to future investigators with access to more material.
The name wellesleybentoni Causey (1952a), a cum-
bersome amalgam of the first and last names of the
collector, was proposed for a form from Phillips
County, Arkansas, with a stronger acropodal bend,
and fortunately can be placed in synonymy.
Variation in males of angularis is analyzed geo-
graphically from the type locality, beginning with
forms to the north and east and proceeding to the
south and west. The males from Missouri are nearly
identical to the holotypes, but that from Izard
County, Arkansas, a partial specimen, is similar to
the one from south of Strong, Union County,
Arkansas in possessing a longer, strongly falcate
acropodite, which is about 3/4 of the telopodite
length. The outer lateral margin in the Izard County
male is lower than the anterior part of the side and
does not extend as far laterad thus forming a
shallower pouch.
East of the type locality there is a trend toward
shorter acropodites beginning with the form named
wellesleybentoni from Phillips County, where it is
subequal to the prefemur and bends more strongly
dorsolaterad near midlength (Fig. 22). The outer
lateral margin is the same elevation and nearly linear
with the anterior part of the sides, and since the inner
lateral margin curves, the pouch is comparatively
narrow, shallow, and closed (Fig. 21). Farther east in
Oktibbeha County, Mississippi, the acropodite is
subequal to the prefemur and curves slightly dorso-
laterad. The outer lateral margin is lower than the
anterior part of the sides and flares moderately
laterad, thus forming a moderate, partly open
pouch.
Forms south of the type locality exhibit greater
variability in both the gonopods and apertures,
although those from adjacent Jefferson County
closely resemble the types. The acropodites and
prefemora are subequal in length in all north-central
Louisiana males except those from Ouachita Parish,
where the former is much shorter, about 1/3 of the
telopodite length and resembles the condition in
compressus. The acropodite curves slightly dor-
solaterad in Morehouse Parish and more directly
laterad in the Franklin and Catahoula males. The
outer lateral margin slopes dorsad and flares farther
laterad in these specimens thus forming a larger
pouch.
Three forms occur in Union County, Arkansas.
The males south of Strong display long falcate
acropodites, around 3/4 of the telopodite length
(Fig. 24), that are convergent with the variant in
Izard County. The bases of the acropodites cross in
the midline, then they curl over the outer caudal
margin on the opposite side of the aperture and cross
the midline and the opposite member again over the
sternum between the 9th legs. The outer lateral
margin rises from the division point, and the inner
lateral margin extends well into the opening, thus
forming a deep, open pouch (Fig. 23). The precise
locality of the other Union County form is
unknown, but the acropodite comprises about half
the telopodite length and curves nearly directly dor-
sad over the sternum between the 9th legs; the
prefemur exhibits a distinct distomedial shoulder.
The outer lateral margin is level, flares moderately
laterad, and forms a moderate pouch with the inner
margin.
Forms in southwestern Arkansas and adjacent
Louisiana display moderately long acropodites.
Those from Columbia County, Arkansas, are sub-
equal to the prefemur, or 1/2 of the telopodite
length, and curve distinctly laterad; there is a slight
distal shoulder on the prefemur. The outer lateral
margin flares farther laterad, and the inner margin
curves more into the opening, thus forming a
broader and deeper pouch. The Caddo Parish, Loui-
siana, male has a very long, bisinuate acropodite,
but it is only about half the telopodite length because
the prefemur is also longer (Figs. 25-26). The
telopodite curves strongly laterad, extending into the
pouch, and the outer lateral margin is much higher
than the anterior parts of the sides. The inner margin
curves smoothly into the opening and the inner
caudal margin (Fig. 25).
Less can be said about variation in females
because most samples contain only males. Females
from Franklin and Catahoula Parishes, Louisiana,
resemble those from Jefferson County, Arkansas, in
having subequal dactyliform projections from the
valves that are bowed towards each other and touch
in some individuals, thus resembling opposable
digits or claspers. The projections are shorter and
diverge, extending directly anteriad and caudad, in
the female from Ouachita Parish, Louisiana (Fig.
30).
Ecology. — No habitat comments have appeared
in the literature, and the only notation on sample
ROWLAND M. SHELLEY 35
labels is ‘‘leaf mold in woodland’”’ for the sample
from Jefferson County, Arkansas.
Distribution. — The Coastal Plain of Arkansas
and northern Louisiana, extending eastward to
eastern Mississippi and northward in Missouri to the
fringe of the Ouachita Physiographic Province (Figs.
9, 211). Specimens were examined as follows:
MISSOURI: Johnson Co., Warrensburg, 2M,
2F, 8 April 1962, K. Oyer (FSCA).
ARKANSAS: JIzard Co., Mt. Pleasant, M, 23
December 1954, N. B. Causey (FSCA). Prairie Co..,
DeValls Bluff, M, F, 1 April 1950, L. Gray (ANSP)
TYPE LOCALITY. Jefferson Co., 0.5 mi. S.
Altheimer, west of AR hwy. 88, 5M, 3F, 14 April
1956, collector unknown (FSCA). Phillips Co.,
Helena, M, October-November 1951, W. Benton
(ANSP), and 20 mi. E Pine Bluff, M, F, October
1954, Kirkwood (FSCA). Ouachita Co., 5 mi. SW
Camden, M, F, 1 January 1956, A. B. Jones
(FSCA). Union Co., locality unknown, 3M, 8
January 1958, L. O. Warren (FSCA); and south of
Figs. 31-37. Eurymerodesmus compressus. 31, left mandibular
projection of neotype, lateral view. 32, aperture and gonopods in
situ of neotype, ventral view. 33, left gonopod of the same, lateral
view. 34, telopodite of the same, medial view. 35, cyphopods in
situ, ventral view of female from Union Co., AR. 36, left
cyphopod of the same, medial view. 37, the same, caudal view.
Scale lines for figs. 32 and 35 = 1.00 mm. Line for other figs. =
0.25 mm for 36-37, 0.30 mm for 31, 0.40 mm for 33, and
0.50 mm for 34.
MEM. AMER. ENT. SOC., 37
Strong, 3M, 19 March 1953, collector unknown
(FSCA). Columbia Co., Magnolia, 4M, 28 January
1955, P. Waler (FSCA), M, 19 April 1958, W. F.
Evans (FSCA), and 2M, 26 November 1961, R.
Rogers (FSCA).
LOUISIANA: Caddo Par., ca. 30 mi. NNW
Shreveport, near Myrtis on McCloud-State Line
Rd., 0.3 mi. W LA hwy. 1, M, 11 March 1966, R. E.
Tandy (FSCA). Morehouse Par., 12 mi. N Bastrop,
along LA hwy. 142, 2.1 mi. S AR line, M, 16 March
1967, J. E. and M. R. Cooper (NCSM); and 5 mi. S
Mer Rouge, M, F, 24 December 1954, N. B. Causey
(FSCA). Ouachita Par., Monroe, 4M, 2 April 1969,
collector unknown (FSCA), and 1410 Forsyth Ave,
M, 26 April 1976, and F, 31 December 1974, M. R.
and J. E. Cooper (NCSM). Caldwell Par., along
Lafourche River, 3M, F, date and collector
unknown (FSCA). Franklin Par., Winnsboro, M, F,
May 1953, M. K. Wiggers (FSCA). Catahoula Par.,
ca. 17 mi. NE Jonesville, along LA hwy. 15 between
Peck and Sicily Island, 3M, 3F, 24 December 1954,
N. B. Causey (FSCA); and 4 mi. W Sicily Island,
along LA hwy, 8, F, date unknown, R. E. Tandy
(FSCA).
MISSISSIPPI: Oktibbeha Co., Craig Springs,
M, 22 April 1980, G. Snodgrass (MEM, NCSM).
Remarks. — With known occurrence in the
western fringe of Caddo Parish, Louisiana,
angularis should be expected in Cass and Marion
counties, Texas.
Eurymerodesmus compressus Causey
Figs. 9, 31-37, 211
Eurymerodesmus compressus Causey, 1952a:169-171, figs. 1-4.
Chamberlin and Hoffman, 1958:79.
Type specimen. — Male neotype and one
associated female (FSCA) collected by W. Nesbit on
an unspecified date in 1957, 6 mi. W El Dorado,
Union Co., AR. This site is about 4 mi. N of the type
locality, Junction City, Union County, where 16
males and 2 females were collected in October 1950.
The types are missing from the ANSP, the published
repository (Causey 1952a, Chamberlin and Hoffman
(1958), nor is there any material from Junction City
in the FSCA. According to Causey (1952a), 1 female
topotype was collected at Junction City on 10 May
1950 and 11 males were taken there on 16 January
1952.
Diagnosis. — A large species; males characterized
by long mandibular projection; sides of aperture
divided, with open caudolateral pouches; telopodite
moderately long; acropodite moderately long, about
36 EURYMERODESMID MILLIPEDS
3/8 of telopodite length, broadly terminal, poorly
demarcated from prefemur, laterally compressed;
prefemur with many hairs arranged continuously
along inner margin; cyphopod valves of females with
unequal, closely appressed, rigid projections from
distal corners, that of outer (lateral) valve very long,
dactyliform, overhanging at least segment 4 in situ,
that of inner (medial) valve much shorter, sub-
triangular; receptacle moderately large, sides curving
partly around operculum.
Color in life. — Peritremata orange: metaterga
uniformly dark mottled olive, without stripes; col-
lum with orange stripe along anterior margin
(Causey 1952a).
Neotype. — Length 37.4 mm, maximum width
5.4mm, W/L ratio 14.4%
Somatic features similar to hispidipes, with following excep-
tions:
Width across genal apices 3.1 mm, interantennal isthmus 1.0
mm. Antennae reaching back to anterior half of 3rd tergite,
relative lengths of antennomeres 2>3>5>4=6>1>7. Genae
with faint impressions. Facial setae as follows: epicranial, inter-
antennal, frontal, and genal absent, subantennal 1-1, clypeal about
14-14, labral about 20-20. Process of mandibular stipes (Fig. 31)
moderately long and broad, apically blunt and rounded.
Collum extending slightly below ends of following tergite.
Paranota moderately flattened, interrupting slope of dorsum;
posterior corners rounded through segment 6, blunt on 7-10,
becoming progressively more acute caudad.
5th sternum with low, sparsely hirsute areas between 4th legs,
narrowly segregated in midline, and strong, sparsely hirsute pro-
jections between Sth legs, narrowly segregated in midline; 6th ster-
num with low, sparsely hirsute elevated areas between both leg
pairs, widely separated in midline, those between anterior (6th) legs
higher. Postgonopodal sterna with deep central impression be-
tween 9th legs; segments 8-9 with low, knob-like projections be-
tween both leg pairs, narrowly segregated in midline, those be-
tween caudal legs much higher with a few more hairs; segments
10-15 flat between anterior legs, with moderate knob-like projec-
tions between caudal legs, becoming progressively smaller caudally
with only light hirsute patches on segments 16-18. 2nd coxae with
sharply conical anterodistal lobes, caudal lobes absent. Prefemoral
lobes present on all legs, becoming smaller on caudalmost
segments.
Gonopodal aperture (Fig. 32) broadly trapezoidal, with narrow,
open caudolateral pouches and broad anterior indentation, 2.9
mm wide and 1.3 mm long at maxima, outline of apparatus
trapezoidal; anterior indentation short, broad, apically rounded;
sides divided, angling slightly caudolaterad, elevating just beyond
anterolateral corner and continuing to division point at midlength,
inner lateral margin plunging deeply into opening then curving
broadly into inner caudal margin, outer lateral margin tapering
rapidly dorsad, flaring slightly laterad and curving into outer
caudolateral corner, forming narrow, open pouch with inner
margin, rims essentially smooth; inner caudolateral corner
smoothly rounded; outer caudolateral corner generally broadly
rounded, somewhat irregular; inner caudal margin curving
anteriad, parallel to outer caudal margin; latter curving broadly
anteriad along anterior surfaces of coxal condyles; margins ir-
regularly hirsute with long hairs arising from rims and outer sur-
faces of sides, slightly more hairs on anterior indentation and
moderately dense tufts from pouches, caudal margins glabrous.
Gonopods in situ (Fig. 32) with telopodites extending caudad in
subparallel arrangement, continuing beyond caudal margins of
aperture and terminating over coxal condyles, apices directed dor-
sad. Gonopod structure as follows (Figs. 33-34): Telopodite
moderately long, terminating well before distal extremities of
hairs. Prefemur moderately long, about 5/8 of telopodite length,
curving slightly near midlength and leaning over coxa, margins
parallel, not tapering, without distal lobe or swelling, with two
rows of long hairs along inner surface angling slightly outward,
and distomedial tuft of 8-10 hairs. Acropodite moderately long
and broadly terminal, about 3/8 of telopodite length, continuous
with, and poorly demarcated from, prefemur, compressed lateral-
ly, curving slightly more dorsad, sides tapering rapidly beyond
midlength to subacuminate tip.
Female — Length 29.6 mm, maximum width 4.5 mm, W/L
ratio 15.2%. Agreeing essentially with neotype in somatic features,
with following exceptions: Mandibular stipes with short,
acuminate, lightly sclerotized projection. Sterna flat, unmodified,
with variably impressed transverse grooves originating between leg
pairs and at most only a few scattered hairs per segment adjacent
to caudal coxae.
Cyphopodal aperture elliptical, sides thickened, strongly
elevated above metazonal surface, tapering rapidly to metazonum
on caudal margin. Cyphopods in situ (Fig. 35) lying obliquely in
aperture, open side anteromediad, valvular projections directed
caudomediad over caudal margin of aperture, those of outer valves
extending over sternum of segment 4, crossing in midline, termi-
nating over prozonum of segment 5. Valves (Figs. 36-37) relatively
large, unequal, lateral (outer) valve much larger, situated obliquely
on receptacle, inner margins moderately hirsute, outer corners ex-
tended into unequal projections, that of inner valve relatively short
and subtriangular, closely appressed to inner margin of outer
valvular projection, latter very long, more than twice as long as in-
ner projection, rigid, and digitiform, apically acuminate. Recep-
tacle moderately large, glabrous, oriented dorsoventrally in aper-
ture, sides curving broadly around sides of operculum. Latter
small, situated directly beneath valves.
Ecology. — Unknown.
Distribution. — Known only from western Union
County, Arkansas (Figs. 9, 211). In addition to
Junction City, Causey (1952a) reported a female
from Three Creeks, which is also lost. Thus, the neo-
type and female were the only specimens examined.
Remarks. — With a moderately long acropodite,
3/8 of the telopodite length, compressus spans the
anatomical gap between the long structures dis-
played by the rest of the impurus group and the short
ones, 1/4 of the telopodite length or less, found in
the varius group, thus blending the dissimilar forms
of these groups into one lineage.
Eurymerodesmus goodi Causey
Figs. 9, 38-44, 211
Eurymerodesmus goodi Causey, 1952b:3-4, fig. 1. Chamberlin
and Hoffman, 1958:79-80.
ROWLAND M. SHELLEY 37
Figs. 38-44. Eurymerodesmus goodi. 38, right mandibular
projection of holotype, lateral view. 39, aperture and gonopods in
situ of male from Montgomery Co., AR, ventral view. 40, left
gonopod of holotype, lateral view. 41, telopodite of the same,
medial view. 42, cyphopods in situ, ventral view of female from
Polk Co., AR. 43, left cyphopod of the same, subcaudal view.
44, the same, anterolateral view. Scale lines for figs. 39 and 42 =
1.00 mm. Line for other figs. = 0.30 mm for 38 and 43-44, and
0.50 mm for 40-41.
Type specimen. — Male holotype (AMNH) col-
lected by N. B. Causey, 7 September 1950, along
Bard Springs Rd., 16 mi. SE Mena, Polk Co., AR.
Diagnosis. — A large species; males characterized
by long mandibular projection; sides of aperture un-
divided, without caudolateral pouches, elevating
slowly throughout, without modifications; telopo-
dite long; acropodite long, about 2/3 of telopodite
length, broadly terminal, moderately demarcated
from prefemur; latter with many hairs arranged con-
tinuously along inner margin; cyphopod valves of
females with distal corners extending into long,
broad projections; receptacle absent.
Color in life. — Pertriemata reddish-orange; dor-
sum olive, darker on prozonites, lighter on meta-
zonites; without metatergal stripes (Causey 1952b).
Holotype. — Body fragmented, length un-
measurable, maximum width 5.8 mm.
Somatic features similar to hispidipes, with following excep-
tions:
Width across genal apices 3.7 mm, interantennal isthmus 1.1
mm. Antennae reaching back to middle of 3rd tergite, relative
MEM. AMER. ENT. SOC., 37
lengths of antennomeres 2>5>4=6>3>1>7. Genae without
impressions. Facial setae as follows: epicranial absent, inter-
antennal 1-1, frontal 1-1, genal absent, clypeal about 11-11, labral
about 20-20. Process of mandibular stipes long, angling caudad,
apically blunt (Fig. 38).
Collum extending slightly below ends of following tergite.
Paranota moderately flattened, interrupting slope of dorsum;
posterior corners rounded on anterior half of body, becoming
blunt and progressively more acute on caudalmost segments.
Sth sternum with short broad knob-like projections between
both leg pairs; 6th sternum with broader, flatter elevated areas be-
tween both pairs of coxae. Postgonopodal sterna with slightly
elevated, flattened areas between 9th legs; remaining sterna with
hirsute, elevated areas on segments 8-10, becoming progressively
flatter and less hirsute caudad, with incomplete transverse impres-
sions originating between leg coxae. 2nd coxae with broad, trun-
cate anterodistal lobe and shorter, acute caudal one. Prefemoral
lobes present on all legs, becoming smaller on caudalmost
segments.
Gonopodal aperture (Fig. 39, not this specimen) very wide,
broadly ovoid, without caudolateral pouches, strongly indented
anteriad at midline, 3.6 mm wide and 1.7 mm long at maxima;
anterior indentation strong, broadly rounded; sides not divided,
curving mediad caudally, elevation beginning behind anterolateral
comer, continuing to peak at caudolateral corner, rims essentially
smooth; caudolateral corner broadly rounded, obtuse; caudal
margin continuous with sides at caudolateral corners, dropping
abruptly to metazonal surface and continuing across midline;
anterior margin glabrous, hairs beginning on sides behind
anterolateral corners, becoming progressively denser caudad, aris-
ing from rim or slight distance down outer surface, densest at
caudolateral corners, continuing along midline of caudal margin in
linear arrangement. Gonopods in situ (Fig. 39, not this specimen)
with telopodite angling caudomediad, apices nearly touching,
located wholly within aperture. Gonopod structure as follows
(Figs. 40-41): Telopodite relatively long, terminating before level of
distal extremities of hairs. Prefemur short, about 1/3 of telopodite
length, expanding distad and terminating in swollen area, with two
rows of hairs along inner margin and distomedial tuft of about a
dozen hairs. Acropodite long and broadly terminal, about 2/3 of
telopodite length, moderately demarcated from prefemur, linear
for most of length, curving broadly distal to midlength, sides
tapering gradually then more rapidly apically, tip subacuminate.
Female from Polk Co. — Length 29.6 mm, maximum width
4.8 mm, W/L ratio 16.2%. Agreeing closely with holotype in
somatic features, with following exceptions: Genal setae 1-1. Pro-
cess of mandibular stipes short, blunt. Sterna relatively flat and
unmodified, with faint, incomplete transverse impressions
Originating between coxae and at most only a few randomly scat-
tered setae, especially along caudal margins.
Cyphopodal aperture ovoid, caudal margin extending caudad in
midline, sides and caudal margin distinctly elevated above
metazonal surface, caudal margin strongly flared, overhanging
segment margin. Cyphopods in situ (Fig. 42) with ventral exten-
sions of valves protruding through aperture, directed caudad,
overhanging caudal margin of aperture and coxae of 3rd legs, open
side visible, facing sublaterad. Valves (Figs. 43-44) large,
moderately hirsute, slightly unequal, dorsal one with short, broad
process curving over opening between valves, distal corners extend-
ing into long broad projections, longer than main part of valves,
leaning dorsad, narrowing then expanding into rounded, knob-like
termini. Receptacle absent. Operculum minute, tucked under
anterior side of valves.
38 EURYMERODESMID MILLIPEDS
Variation. — Measurements of the male from
Montgomery County illustrated in Figs. 39-41 are
32.5 mm in length, 5.3 mm in width, W/L ratio
16.3%. Its gonopods agree closely with those of the
holotype, but the caudal aperture margin extends
caudad between the 8th legs to near the sternal eleva-
tions.
Ecology. — Unknown.
Distribution. — Known only from southwestern
Polk and southwestern Montgomery counties,
Arkansas, in the southern fringe of the Ouachita
Physiographic Province (Figs. 9, 211). Specimens
were examined as follows:
ARKANSAS: Polk Co., along Bard Springs
Rd., 16 mi. SE Mena, M, 7 September 1950, N. B.
Causey (AMNH) TYPE LOCALITY; and Shady
Lake Rec. Area, M, F, 18 June 1953, D. Dowling
(FSCA). Montgomery Co., 10 mi. NW Langley (in
Polk Co.), Albert Pike Cpgd., M, F, 1979, H. W.
Robison (NCSM) and M, 24 December 1982, H. W.
Robinson (NCSM).
Remarks. — Eurymerodesmus goodi is a relative-
ly soft, flexible eurymerodesmid, and its large size is
equivalent to that of moderately-large xystodesmids
such as Boraria profuga (Causey), also occurring in
Arkansas. Because of its size goodi is particularly
likely to be misassigned to the Xystodesmidae, and
close attention should be paid to the structures of the
mandibular stipes and leg prefemora when making
determinations.
The Dubius Group
Eurymerodesmus dubius shares a wide aperture
lacking caudolateral pouches with goodi, which is
parapatric on the north. It differs in having a short
acropodite and short, mostly linear telopodites, the
latter distinguishing it from other members of the
Kewanius lineage. It differs from the species of the
varius group in the linear telopodite and the absence
of caudolateral pouches, and is distinguished from
those in the simplex group by its broadly terminal
acropodite.
Component. — dubius Chamberlin.
Eurymerodesmus dubius Chamberlin
Figs. 9, 45-53, 211
Eurymerodesmus dubius Chamberlin, 1943:38, fig. 8; 1952:573.
Causey, 1952b:2-3. Chamberlin and Hoffman, 1958:79.
Paresmus columbus Causey, 1950a:272, figs. 10-11. Chamberlin
and Hoffman, 1958:832. NEW SYNONYMY.
Type specimens. — Male holotype and female
paratype (FMNH) collected by K. P. Schmidt, 16
April 1941, at Delight, Pike Co., AR.
Diagnosis. — A large species; males characterized
by short mandibular projection; sides of aperture
undivided, without caudolateral pouches, elevating
slowly, without modifications; telopodite short,
nearly upright for most of length; acropodite short,
1/8 to 1/4 of telopodite length, broadly terminal,
poorly demarcated from prefemur, gently curved;
prefemur with many hairs arranged continuously
along inner surface; cyphopod valves of females
with distal corners only slightly extended; receptacle
absent.
Color in life. — Unknown. Chamberlin’s descrip-
tion (1943) indicates a stripe across the caudal
margins of the metaterga, but the living colors can-
not be inferred from those in preservative.
Holotype. — Body highly fragmented, length un-
measurable, maximum width 5.8 mm.
Somatic features similar to hispidies, with following exceptions:
Width across genal apices 3.6 mm, interantennal isthmus 1.0
mm. Antennae reaching back to caudal margin of 3rd tergite,
relative lengths of antennomeres 2>6>3=4=5>1>7. Genae
with distinct, central impressions. Facial setae as follows:
epicranial, interantennal, and frontal absent; genal 1-1, clypeal
about 9-9, labral about 16-16. Process of mandibular stipes short
and stout, apically broadly rounded (Fig. 45).
Collum extending slightly below ends of following tergite.
Paranota moderately flattened, interrupting slope of dorsum;
posterior comers rounded through segment 6, blunt on 7-14,
becoming progressively more acute caudally.
Sth sternum with short projections between anterior legs, closely
appressed in midline, and broader, more widely segregated, knob-
like processes between caudal legs; 6th sternum with short, knob-
like projections between anterior legs and broad, flattened, areas
between caudal legs. Postgonopodal sterna with short, broad lobes
between 9th legs; sternum of segment 8 deeply incised transversely,
with hair patches adjacent to leg coxae; remaining sterna becoming
flatter and more plate-like caudally, with hair patches adjacent to
leg coxae through segment 17, becoming progressively less hirsute
caudad. 2nd coxae with short distal lobes on anterior and caudal
surfaces. Prefemoral lobes extending through segment 12.
Gonopodal aperture (Fig. 46) nearly circular, without caudo-
lateral pouches, with slight anterior indentation, 2.0 mm wide and
1.4 mm long at maxima; anterior indentation short and broad,
apically rounded; sides curving gently caudomediad, undivided,
elevating slightly to peak just before caudolateral corner, then
dropping abruptly to metazonal surface, rims becoming slightly ir-
regular caudad; caudolateral corner moderately blunt, sloping
downward to metazonal surface; caudal margin not continuous
with and much lower than lateral, extending slightly caudad in
midline; anterior and caudal margins glabrous, sides with only a
few scattered hairs on rim and outer surface near peak at caudo-
lateral corners. Gonopods in situ (Fig. 46) angling toward midline
with tips nearly touching, telopodites located wholly over aperture.
Gonopod structure as follows (Figs. 47-48): Telopodite relatively
short and upright, terminating well before level of distal ex-
tremities of hairs. Prefemur relatively long, about 7/8 of
ROWLAND
Figs. 45-53.
holotype, ventral view. 47, left gonopod of the same, lateral view. 48, telopodite of the same, medial view. 49, aperture and gonopods in
situ of male from Bradley Co., AR, ventral view. 50-51, apertures and gonopods in situ, ventral views of two males from Clark Co., AR.
52, left cyphopod of female paratype, medial view. 53, the same, anterior view. Scale lines for figs. 46 and 49-51 = 1.00 mm. Line for
other figs. = 0.35 mm for 45, 0.40 mm for 47, and 0.50 mm for 48 and 52-53.
telopodite length, upright but not linear, margins slightly
bisinuately curved, narrowing slightly distad, basal hairs relatively
sparse, hairs denser on stem, with two rows along inner margin
angling outward distad and terminating in tuft of about 12 hairs,
with distomedial tuft of 8 hairs. Acropodite very short, broadly
terminal, about 1/8 of telopodite length, covered by prefemoral
hairs, essentially just a short spur on distal extremity of prefemur,
continuous with, and poorly demarcated from, latter, curving
gently dorsad, sides tapering to acuminate tip.
Female paratype. — Body fragmented, length unmeasurable,
maximum width 4.8 mm. Agreeing essentially with holotype in
somatic features, with following exceptions: Subantennal and
frontal setae present, 1-1 each; genal setae absent. Process of man-
dibular stipes indistinct, broadly rounded. Sterna relatively flat,
unmodified, with at most only a few randomly scattered hairs on
caudal margins.
Cyphopodal aperture small, circular, sides and caudal margin
nearly flush with metazonal surface. Cyphopods in situ with tips of
valves visible in aperture, not protruding, open side directed
anteriad. Valves (Figs. 52-53) moderate-size, moderately hirsute,
slightly unequal, outer (lateral) valve larger, distal corners very
slightly extended, broadly rounded. Receptacle absent. Operculum
relatively large, moderately hirsute, located at bases of valves on
open (anterior) side.
MEM. AMER. ENT. SOC., 37
M. SHELLEY 39
Eurymerodesmus dubius. 45, right mandibular projection of holotype, lateral view. 46, aperture and gonopods in situ of
Variation. — Eurymerodesmus dubius is one of
the more flexible eurymerodesmids, and it was possi-
ble to uncoil without breakage and measure males
from the counties in Table 4:
Table 4. Size variation of E. dubius males.
County length width W/L ratio
Saline 37.8 6.4 1.69%
Garland 33.6 5.6 16.7%
Garland 33.6 5.6 16.7%
Montgomery 32.5 5.3 16.3%
Clark 25.0 4.7 18.8%
Clark 32.7 4.7 14.4%
Pike 34.3 5.5 16.0%
Pike 30.8 4.7 15.3%
Pike 33.6 5.6 16.7%
Pike 29.3 5.3 18.1%
Pike 31.6 5.5 17.4%
Pike 29.8 4.9 16.4%
Nevada 27.5 4.8 17.5%
Bradley 32.5 6.4 19.7%
40 EURYMERODESMID MILLIPEDS
Aside from one male from Clark County that
lacks the distomedial tuft of prefemoral hairs, the
gonopods of dubius are uniform. However, the in
situ configuration varies depending upon the degree
of coiling of the body, the degree to which the coxae
are submerged under the lateral margins of the aper-
ture, and the caudal configuration of the latter.
Thus, the apices of the telopodites may be very close,
touch, or cross, and they may be wholly within the
aperture or slightly overlap the caudal rim. In one
tightly coiled male, the aperture was nearly com-
pletely covered by segment 6, and the telopodites
were correspondingly reoriented, being directed
mediad with the apices extending across the opposite
side of the aperture and the tips touching the oppos-
ing coxae.
As discussed by Causey (1952b), the aperture con-
figuration and the size and shape of the sternal pro-
cess(es) between the 9th legs vary considerably. The
height and length of the lateral elevations varies, and
they may be nearly linear or broadly curved. In
northern populations, the lateral elevations extend
around the caudolateral corners onto the caudal
margin, where they terminate either abruptly or
smoothly by tapering to the metazonal surface.
However, in the southernmost populations from
Nevada, Ouachita, and Bradley counties, the eleva-
tions are limited to the sides, terminating either at or
just before the caudolateral corners.
The caudal margin displays two configurations. It
is essentially straight as in the holotype in males from
Saline, Garland, and Nevada counties, and in these
individuals, the sternal projections between the 9th
legs are separate, short, and knob-like, terminating
well below the margins of the adjacent coxae. How-
ever, in some males from Montgomery, Clark, Pike,
Ouachita, and Bradley counties, the margin extends
caudad in the midline between the 9th legs termina-
ting at, and tapering into, the sternal projection
(Figs. 49-50). Coupled with the anterior indentation,
the aperture in these individuals appears heart-
shaped. In specimens from Montgomery, Pike, and
Bradley counties, the sternal knobs are shorter than
the adjacent coxal widths and are separated to vary-
ing degrees, either directed ventrad or leaning slight-
ly toward the midline. However, in males from
Clark and Ouachita counties, the knobs are co-
alesced in the midline to form a single ventral projec-
tion of varying length (Figs. 50-51). It is longer than
the adjacent coxal widths in one male from Clark
County (Fig. 50). As a further complication, a short
midcaudal spur extends anteriad inside the aperture
in a few males from Ouachita and Bradley counties
(Fig. 49). None of these variants demonstrates
enough consistency or enough of a geographical
component to warrant taxonomic recognition at the
subspecific level. However, they emphasize the need
to examine every specimen; males within some
samples differ.
The cyphopods generally resemble those of the
female paratype and protrude through the aperture
in a few individuals. In one from Garland County,
the valves are oriented laterad/mediad, with the
open side facing forward.
Ecology. — Labels with specimens from Bradley
County state that they were taken in “‘large traps”’ in
pine-oak woods.
Distribution. — The southern fringe of the
Ouachita and the adjacent Coastal Plain Physio-
graphic Provinces of southern Arkansas, with an
allopatric record from Louisiana (Figs. 9, 211).
Specimens were examined as follows:
ARKANSAS: Montgomery Co., along US hwy.
270, 5.4 mi. E Mt. Ida, M, 6 March 1977, R. E.
Woodruff (FSCA). Garland Co., Hot Springs, 3M,
F, 4 January 1957, collector unknown (FSCA); and
Big Goat Island, Lake Hamilton, M, 28 December
1954, Atkins (FSCA). Hot Springs Co., Magnet, F,
1880’s but exact date unknown, Hutcherson
(NMNH). Saline Co., locality unknown, 2M, 2
juvs., 1950, N. B. Causey (FSCA). Clark Co., 4.5
mi. N Arkadelphia, M, 19 January 1953, and 4M,
February 1953, J. E. Sublette (FSCA); 4 mi. E Arka-
delphia, M, F, date and collector unknown (FSCA);
and 8 mi. W Arkadelphia, 3M, 2F, 10 September
1950, collector unknown (FSCA). Pike Co., Delight,
M, F, 16 April 1941, K. P. Schmidt (FMNH) TYPE
LOCALITY; and Bear Cr., 7M, 2F, 18 April 1954,
N. B. Causey (FSCA). Nevada Co., Jackson Twp.,
5M, 3F, 30 December 1954, R. Delaney (FSCA).
Ouachita Co., 5 mi. SW Camden, M, 27 November
1955, A. B. Jones (FSCA). Bradley Co., Prospect,
3M, 29 November 1967, Whitcomb (FSCA); and
Sumpter, 2F, September and 13 October 1965, and
3M, 21 and 28 December 1964, L. Whitcomb
(FSCA).
LOUISIANA; Catahoula Par., Sicily Island
(not town), 2M, 2F, 2 juvs., 12 October 1974, S.
Ziser (FSCA).
In addition to these specimens, the record from
Dallas County, Arkansas, locality not specified
(Causéy 1952b), is considered valid and is so in-
dicated in figures 9 and 211. There is also an Arkan-
sas sample in the FSCA containing two males, six
females, and six juveniles from an unspecified loca-
tion in Washington County. This site is so distant
ROWLAND M. SHELLEY 41
from the others that I think it may represent a label-
ling error and prefer to omit it. For many years Dr.
Causey’s home was in Washington County, where
she and others collected many eurymerodesmids. If
dubius does occur there, I think it would have been
collected more than once.
Since Causey’s sketch of the gonopod of Pares-
mus columbus (1950a, fig. 11) closely resembles that
of dubius, 1 include P. columbus in the synonymy
and show its type locality, 3 miles east of Magnolia,
Columbia County, Arkansas, as a literature record
(Figs. 9, 211). The type series of P. columbus con-
tained the male holotype and a female allotype, both
of which are missing from the ANSP, the reported
repository (Causey 1950a, Chamberlin and Hoffman
1958). No other localities have been recorded for P.
columbus.
Remarks. — Eurymerodesmus dubius is one of
the more flexible congeners, hence the large number
of measurements since it was possible to uncoil many
males without breakage. Like goodi, dubius may be
confused with xystodesmids because of its large size.
The relatively linear, upright telopodites are superfi-
cially similar to those of digitatus (compare figs.
47-48 and 199-200), a resemblance attributable to
convergence. Likewise, their enclosure within the
aperture is convergent with the condition in the
melacis lineage.
The Louisiana locality is about 100 miles from the
closest known site in Arkansas. Field work is there-
fore needed in northern Louisiana to determine
whether these areas connect.
The Varius Group
The varius group is characterized as follows: aper-
tures with divided lateral margins and caudolateral
pouches; acropodites short, no more than 1/4 of the
telopodite length, broadly terminal, and poorly
demarcated from the prefemur at least in some
populations of each component species; prefemora
long, at least 3/4 of telopodite length, without
distinct terminal lobes on outer surfaces in all
populations. The subspecies of varius in south-
western Louisiana between the Pearl and Mississippi
Rivers, v. christianus, possesses strong terminal
prefemoral lobes that clearly demarcate the
acropodite. However, this trait is not shared by all
races of varius. The nominate subspecies, in the
southeast along the Atlantic Coast, and some forms
of v. /ouisianae in northern Louisiana lack distinct
lobes, and the acropodite is therefore continuous
with the prefemur. Other species have at most a
MEM. AMER. ENT. SOC., 37
slight lobe or swelling, so the key feature of the
varius group is the absence of the lobe from some
populations of each species. Since the simplex group
contains forms with strong prefemoral shoulders
and subterminal or narrowly terminal acropodites,
varius itself bridges the anatomical gap between the
two groups through its lobed forms. The two groups
could therefore be combined, but it is convenient to
segregate them based on the acropodal feature. The
varius group covers nearly all of the range of the
Kewanius lineage, extending from the Atlantic
Ocean to eastern Texas, east-central Missouri, and
southern Illinois, with an apparently allopatric
population in Mason County, Texas (Figs. 213). The
only regions that it does not encompass are the area
occupied by impurus, Brazos and Washington coun-
ties, Texas, and west-central Missouri.
Components. — varius (McNeill) [v. varius, v.
christianus Chamberlin, v. louisianae Chamberlin];
newtonus Chamberlin; oliphantus Chamberlin;
amplus Causey; elevatus, new species.
Eurymerodesmus varius (McNeill)
Diagnosis. — A small to moderate-size species;
males characterized by variable mandibular projec-
tion, ranging from minute, nubbinlike, and subequal
to that of females to moderate-size; sides of aperture
divided, with closed caudolateral pouches; telopodite
long; acropodite short, about 1/4 of telopodite
length, broadly terminal, poorly to sharply demar-
cated from prefemur; latter with or without rounded
distal lobe on outer margin, with relatively few hairs
arranged sporadically and discontinously along inner
surface; cyphopod valves of females with distal cor-
ners extending into angular ridge, angling toward
corner; receptacle moderate-size, sides not curving
around operculum.
Remarks. — The third oldest species in the fami-
ly, varius occupies the largest distribution, covering
the entire area east of the Mississippi River, with
scattered, possibly relictual populations in north-
central Louisiana and southern Arkansas (Fig. 54).
The distal end of the gonopod prefemur undergoes
an east-west clinal change ranging from either no, or
only a slight, distal prefemoral swelling in popula-
tions along the Atlantic and Gulf Coast westward to
western Alabama, to a somewhat larger swelling in
those in Mississippi, to a strong, broadly rounded
lobe in ones in southeastern Louisiana between the
Pearl and Mississippi Rivers, to a variably small to
moderate-size swelling in forms in northern Loui-
siana and Arkansas. As the swelling becomes larger,
42 EURYMERODESMID MILLIPEDS
Fig. 54. Distributions of the varius and simplex groups. Squares, varius varius; X’s, varius intergrades; vertical half-shaded squares,
varius christianus; diagonal half-shaded squares, varius louisianae; dots, amplus; diamonds, newtonus; asterisks, oliphantus; stars,
elevatus; upright triangles, simplex; inverted triangles, po/kensis; star in dot, caesariatus; ovals, paroicus; equals sign, crassatus; 8’s,
pulaski; and rectangles, serratus. Open symbols denote literature records considered valid.
the acropodite becomes correspondingly more
sharply demarcated, and in populations in
southeastern Louisiana the latter is discontinuous
with the prefemur, thereby resembling the condition
in simplex. Most Atlantic populations exhibit a trace
of a swelling, but a few lack this and have parallel-
sided prefemora, such as that at the type locality,
Pensacola, Escambia County, Florida. Accompany-
ing these gonopodal changes is variation in the man-
dibular projection. All males from east of the Mis-
sissippi River display minute, rounded, and weakly
sclerotized processes, at most only slight nubbinlike
vestiges or rounded lobes subsimilar to those in
females. The projection is larger and variable west of
this boundary, being subtriangular and apically sub-
acuminate in some males and still larger, more heavi-
ly sclerotized, and apically blunt or truncate in
others. All forms possess sparsely and sporadically
hirsute prefemora indicative of a single species, but
the clinal changes in the distal prefemoral swelling
and those involving the mandibular process are
significant enough to justify taxonomic recognition
at the subspecific level. Since the type locality is in
the east, in the area occupied by forms with only
slight swellings, they become the nominate sub-
species, and /ouisianae Chamberlin was proposed
for those in northern Louisiana and southern Arkan-
sas with larger mandibular projections. No name is
available for the lobed population in southeastern
Louisiana, but christianus Chamberlin was proposed
for an intergrade form with an intermediate swelling
from Harrison County, Mississippi, only about 35
miles to the east. As first reviser, I conserve this
name and assign it to the lobed population. At the
Pearl River, the increase in size of the lobe is sharp
and dramatic, but the change is more gradual to the
east and appears to begin in Mississippi. Conse-
quently, I consider the forms in this state as in-
ROWLAND M. SHELLEY 43
KEY TO SUBSPECIES, BASED ON ADULT MALES
1. Projection of mandibular stipes weakly sclerotized, a rudimentary nubbin or short, rounded lobe, subsimilar to condition in
females!(Hig@ 55) iac cis cesta t crac aet eet esa aeeeeaonees
Projection larger, heavily sclerotized, subtriangular to blunt and truncate (Figs. 66-67)............. varius louisianae Chamberlin
2. Gonopod prefemur with at most only a low, rounded distal swelling, only slightly elevated above outer surface; acropodite poorly
demancatedi(Rigsss7/-58) sree cence
BOTs che CE Roa Acie ea ene reer poe are varius varius (McNeill)
Prefemur with distinct terminal lobe, strongly elevated above outer margin; acropodite sharply demarcated (Fig. 65).............
tergrades between v. christianus and the nominate
subspecies, occurring from Alabama eastward.
Triangular and blunt mandibular projections are in-
termixed in northern Louisiana and southern Arkan-
sas, the area occupied by v. /ouisianae.
The clinal change in varius from parallel-sided to
lobed individuals is significant because it links the
continuous, broadly terminal, and poorly demar-
cated acropodal forms clearly assignable to
Eurymerodesmus to the narrowly terminal, strongly
demarcated, and discontinuous acropodite of
simplex, type species of Kewanius, which in turn
bridges the anatomical gap to the subterminal
acropodal forms assignable to Paresmus. Conse-
quently, this clinal change provides the necessary
evidence to synonymize the last two genus-group
names with Eurymerodesmus. Eurymerodesmus
varius is thus an intermediate species connecting all
these forms into a single assemblage, but for con-
venience and ease of comprehension, I divide it into
two groups based on poorly or sharply demarcated
acropodites. Since this structure is continuous in
most populations, varius is the oldest available name
for the poorly demarcated form and is therefore
used to designate this species group.
There are several very large samples of the
nominate subspecies, v. christianus, and some size-
able ones of intergrades, particularly those taken in
pitfall traps. I could also have collected scores of in-
dividuals at the North Carolina, and Alachua and
Gilchrist counties, Florida, sites. These numbers sug-
gest that studies on population structure and ecology
of varius would be fruitful.
Eurymerodesmus varius varius (McNeill), new status
Figs. 54-61
Polydesmus varius McNeill, 1887:323-324.
Leptodesmus varius: Bollman, 1888b:344.
Eurymerodesmus minimus Loomis, 1943;320, fig. 2.
Eurymerodesmus varius: Causey, 1954:67. Chamberlin and Hoff-
man, 1958:81.
Type specimen. — Female holotype (NMNH)
collected by C. H. Bollman, March-April 1886, at
Pensacola, Escambia Co., FL. The original descrip-
MEM. AMER. ENT. SOC., 37
5 AER REE eT Re A ane varius christianus Chamberlin
tion (McNeill 1887) indicates that the type series con-
tained three females, but the other two are lost.
Housed with the holotype is a vial labeled ‘‘Lep-
todesmus varius McNeill, type specimen,”’ contain-
ing a male of this form supposedly collected at an
unknown site in Indiana. The source of this
specimen is unknown, but it is not part of the type
series and represents a labeling error since no authen-
tic Indiana localities are known for a eury-
merodesmid. It is ignored in this study.
Diagnosis. — A small subspecies; males with pro-
jection of mandibular stipes minute, nubbinlike, in-
distinct; acropodite essentially smoothly continuous
with, and poorly demarcated from, prefemur; latter
with at most only a slight suggestion of distal swell-
ing, weakly elevated above margin.
Color in life. — Peritremata pink; metaterga
mottled brownish olive with narrow pink stripes
along caudal edges connecting peritrematal mark-
ings; collum with pink stripes along both anterior
and posterior margins. Loomis (1943) described the
living color as ‘‘translucent white with a light pinkish
tinge,’’ which fades rapidly in alcohol.
Male topotype. — Length about 14.2 mm, max-
imum width 1.8 mm, W/L ratio 12.7%.
Somatic features similar to hispidipes, with following excep-
tions:
Width across genal apices 1.3 mm, interantennal isthmus 0.4
mm. Antennae reaching back to caudal margin of 2nd tergite,
relative lengths of antennomeres 2>3>4=5=6>1>7. Genae
without impressions. Facial setae as follows: epicranial and
subantennal absent, interantennal 1-1, frontal 1-1, genal 3-3,
clypeal about 10-10, labral about 14-14. Mandibular stipes with
minute, vestigal projection, corner rounded, of general size and
configuration as that of female eurymerodesmids, subsimilar to
females of varius (Fig. 55).
Collum extending slightly below ends of following tergite.
paranota depressed, continuing slope of dorsum; posterior corners
rounded through segment 7, blunt on 8-13, becoming progressively
more acute caudad.
5th and 6th sterna without elevations, with moderate hair
patches adjacent to leg coxae. Postgonopodal sterna with minute
hirsute lobes between 9th legs, remaining sterna flat and un-
modified, with faint transverse grooves and sparse hair patches dis-
appearing in midbody region, caudal sterna glabrous. 2nd coxae
without projections. Prefemoral lobes relatively small, restricted to
pregonopodal legs.
Gonopodal aperture (Fig. 56) broadly rectangular, with variable
caudolateral pouches, slightly indented anteriad at midline, 1.0
44 EURYMERODESMID MILLIPEDS
Figs. 55-71. Eurymerodesmus varius. 55-61, varius varius. 55, projection of right mandible of topotype, lateral view. 56, aperture
and gonopods in situ of topotype, ventral view. 57, left gonopod of the same, lateral view. 58, telopodite of the same, medial view.
59, aperture and gonopods in situ of male from Robeson Co., NC, ventral view. 60, left cyphopod of holotype, caudal view. 61, the
same, medial view. 62-63, varius intergrades. 62, aperture and gonopods in situ of male from Harrison Co., MS, ventral view. 63, telopo-
dite of left gonopod of the same, medial view. 64-65, varius christianus. 64, aperture and gonopods in situ of male from Washington
Par., LA, ventral view. 65, telopodite of left gonopod of male from Ascension Par., LA, medial view. 66-71, varius louisianae. 66, right
mandibular process of holotype, lateral view. 67, right mandibular process of paratype, lateral view. 68, aperture and gonopods in situ of
holotype, ventral view. 69, left gonopod of the same, lateral view. 70, telopodite of the same, medial view. 71, telopodite of left gonopod
of male from Columbia Co., AR, medial view. Scale lines for figs. 56, 59, 62, 64, and 68 = 1.00 mm. Line for other figs. = 0.40 mm for
55, 66-67, and 69; 0.50 mm for 57-58, 60-61, 63, 65, and 70-71.
ROWLAND M. SHELLEY 45
mm wide and 0.5 mm long at maxima, outline of apparatus slight-
ly trapezoidal; anterior indentation minute, broadly rounded,
barely perceptible; sides angling progressively mediad, leaning over
aperture, and becoming progressively more elevated to division
point located caudad over caudal margins, inner lateral margin
plunging into opening and curving into inner caudal margin,
obscured by lean of flare and outer lateral margin, latter leaning
strongly mediad but angling laterad and dropping to metazonal
surface at caudolateral corner, forming closed pouch with inner
margin, rims essentially smooth; inner caudolateral corner
smoothly curved, obscured by lean of flare; outer caudolateral cor-
ner rounded; inner caudal margin curving gently and bisinuately,
subparallel to outer caudal margin; latter flush with metazonal sur-
face, angling slightly caudad in midline; margins glabrous except
for slight hair tufts in cavity. Gonopods in situ (Fig. 56) with
telopodites curving mediad, touching opposite member in midline,
then curving broadly laterad, extending well beyond aperture and
terminating over coxae or prefemora of 9th legs. Gonopod struc-
ture as follows (Figs. 57-58): Telopodite relatively long, ter-
minating at about level of distal extremities of hairs. Prefemur
long, about 3/4 of telopodite length, sides parallel for most of
length, narrowing slightly and curving distad, with only 3-4 widely
separated hairs on inner margin of proximal part of stem and
distomedial tuft of 9 moderately separated hairs. Acropodite short
and broadly terminal, about 1/4 of telopodite length, smoothly
continuous with, and poorly demarcated from, prefemur, continu-
ing broad dorsolateral curvature of latter, narrowing rapidly to
subacuminate tip.
Female Holotype. — Body fragmented, length unmeasurable
(15 mm according to McNeill (1887) ), maximum width 1.7 mm,
W/L ratio 11.3%. Agreeing closely with male topotype in somatic
features, with following exceptions: Genal setae 1-1. Segments 3-8
with moderate elevations lateral to legs, highest on segments 3-4.
Sterna glabrous, flat and unmodified, with incomplete transverse
impressions originating between leg coxae.
Cyphopodal aperture elliptical, sides strongly elevated above
metazonal surface. Cyphopods in situ with distal edges of valves
visible in aperture, open side facing dorsad. Valves (Figs. 60-61)
relatively large, subequal, completely filling respective sides of
aperture, moderately hirsute, distal corners extending ventrad into
distinct ridge, bluntly angular at corners. Receptacle moderate in
size, glabrous, located laterad to valves, corners not extending
partly around operculum, hidden internally under side of aperture.
Operculum minute, located under open, dorsal side of valves.
Variation. — The Escambia County males agree
closely with each other, the only exception being
slight differences in the slope of the flare, or outer
lateral margin, of the aperture. When the angle of
slope is greater, the aperture extends farther laterad
and the caudal edge is folded, as shown by a North
Carolina male (Fig. 59). The slope of the flare also
affects the size and shape of the pouch, and the
degree to which it is open or closed; the cavity is nar-
rower and more closed in males with more gradual
slopes. The corner of the division point, the highest
point on the sides, extends inward and downward or
dorsad in males throughout the range.
The gonopods vary in the distance they project
caudad, which is a measure of how tightly the body
is coiled. Normally, they overhang the sternum be-
MEM. AMER. ENT. SOC., 37
tween the 9th legs or the adjacent coxae, but occa-
sionally they extend over more of segment 8. In
more tightly coiled specimens, the 6th metazonite
may extend over as much as half of the aperture.
Topotypical males from Escambia County lack any
trace of a distal prefemoral swelling, but most in-
dividuals from Georgia, Florida, North Carolina,
and Alabama display a suggestion of an enlarge-
ment.
Ecology. — Causey (1954) collected this sub-
species under live oaks in Escambia County, Florida.
Habitat notations on vial labels include the follow-
ing: in house on floor (Alachua Co., FL), under logs
(Grady Co., GA), in roadside stream (Okaloosa Co.,
FL), in pine hardwood litter (Monroe Co., GA), and
under horse dung (Gilchrist Co., FL). The samples
from Tall Timbers Research Station, Leon County,
and Big Bend Horticultural Lab, Jefferson County,
Florida, were recovered from pitfall traps, and in at
least one sample from the latter site, the trap was
under a pecan tree.
In North Carolina, v. varius is plentiful at a rural
residence in Robeson County, the only known site of
eurymerodesmids in the state. They occur under pine
and oak logs and in pine litter under azaleas and
sesanqua bushes. I thought that this population
might be introduced, but the family has resided there
for over 20 years, and the shrubs were present when
they came. Consequently, this seems a natural
population, suggesting that v. varius will eventually
be found at other sites in southeastern North
Carolina and also in coastal South Carolina, where it
has not been encountered. This population probably
is not the source of the Eurymerodesmus sp. or
“‘Leptodesmus_ hispidipes’’ records from Duke
Forest, Durham County (Brimley 1938, Causey
1940, Wray 1967), which I (1978) deleted from the
North Carolina fauna, since Causey’s illustration
(1940, fig. 8) is of a species with continuous
prefemoral hairs. Thus, Eurymerodesmus is still
unknown from piedmont North Carolina. I have
searched for it repeatedly in Duke Forest and neigh-
boring areas at appropriate times of the year without
success.
In Florida, v. varius inhabits a variety of biotopes.
In Gilchrist County I found a sizeable population in
and under exposed, rotting oak logs and stumps on
sandy substrate. Large populations were also en-
countered in hardwood litter at Fort Caroline, Duval
County, and Marjorie Kinnan Rawlings State
Historic Site, Alachua County. The latter site is a
mesic hardwood hammock dominated by sweet
gum, laurel oak, and cabbage palm without pine.
46 EURYMERODESMID MILLIPEDS
The Hernando County locality also is a hardwood
forest, but it contains more oak trees and is more
xeric.
Distribution. — The Atlantic and Gulf Coastal
Plains of the southeastern United States, from
southeastern North Carolina to Hernando County,
Florida, and inland to the Fall Zone Region of
Alabama and Georgia as far west as Mississippi (Fig.
54). I investigated many sites in peninsular Florida in
January 1988 to determine the southern range limit
for the species and family along the Atlantic Coast
and did not find it east of the St. John’s River. Con-
sequently, Ft. Caroline, less than 10 miles from the
ocean, is the only known locality in eastern Florida.
On the western side of the peninsula I extended the
range southward about 80 miles, from Gainesville to
northern Hernando County, but did not encounter
the milliped farther south, even in hardwood areas
around Brooksville where Sigmoria (Cheiropus)
planca (Loomis) occurs (Shelley 1984b). These hard-
wood communities end in Hernando County, and
the habitat around Tampa Bay and farther south is
unsuitable. Consequently, northern Hernando
County seems to be near the actual southern range
limit for both v. varius and the Eurymerodesmidae
along the Atlantic Coast. No records are available
from South Carolina, but the milliped should be ex-
pected throughout the southeastern Coastal Plain
judging by its occurrence in North Carolina and the
general uniformity of coastal environments in the
Carolinas and Georgia. I doubt if this is an allopatric
locality, as occur in hispidipes and serratus, and the
absence of South Carolina records probably reflects
the absence of winter collecting. Specimens were ex-
amined as follows:
NORTH CAROLINA: Robeson Co., 1.2 mi.
SW Proctorville, along SR 2265, 0.4 mi. W jct. NC
hwy. 130, M, 17 April 1984, D. Hedgepeth (NCSM);
28M, 20F, 17 April 1985, R. M. Shelley and J.
Alderman (NCSM); and 6M, 9F, 4 November 1986,
R. M. Shelley (NCSM).
GEORGIA: Monroe Co., 5 mi. N Lizella, 2M,
F, 19 February 1983, J. A. Payne (RLH). Bibb Co.,
Macon, juv. M, date unknown, L. M. Underwood
(NMNH). Talbot Co., Geneva, 2F, 24 April 1960.
L. Hubricht (RLH). Baker Co., near Newton,
Pineland Plantation, 5M, 4F, 30 March 1959, D. B.
Jester, H. Wyatt (FSCA). Worth Co., 4 mi. E
Sylvester, 6F, 6 juvs., 11 June 1959, N. B. Causey
(FSCA). Seminole Co., 4 mi. NW Donaldson, 4M,
3F, 18 March 1961, L. Hubricht (RLH). Decatur
Co., near Spring Cr. W of Brinson, 2M, F, 18
March 1961, L. Hubricht (RLH). Grady Co., 3 mi.
W Cairo, 3M, 3F, 26 January 1965, N. B. Causey
(FSCA).
FLORIDA: Duval Co., Ft. Carolina, 6M, 7F, 8
November 1985, R. M. Shelley (NCSM). Alachua
Co., Gainesville, F, 29 October 1959, W. J. Platt
(FSCA); and Marjorie K. Rawlings St. Hist. Site,
31M, 22F, 21 January 1988, R. M. Shelley and G. B.
Edwards (NCSM). Gilchrist Co., S. of Trenton, 6
juvs., 2 April 1959, R. E. Woodruff (FSCA); and
along FL hwy. 340 at Suwannee R., 19M, 14F, 6
November 1985, R. M. Shelley (NCSM). Hernando
Co., 7 mi. NNE Brooksville, along US hwy. 40, 2M,
3F, 22 January 1988, R. M. Shelley (NCSM). Leon
Co., Tall Timbers Res. Sta., 34M, 22F, 23 April
1969-24 January 1972, P. Fall, W. W. Baker, and
W. H. Whitcomb (FSCA); and Tallahassee, F, 10
November 1958, R. McFarland (FSCA). Jefferson
Co., locality and collector unknown, 47M, 31F, 20
November 1968 - 7 February 1969 (FSCA); and Big
Bend Hort. Lab near Monticello, 52M, 32F, 2
February 1969 - 12 December 1969, W. H. Whit-
comb (FSCA). Jackson Co., Marianna, F, 6 June
1970, D. C. Bennett (FSCA), M, date and collector
unknown (MCZ), and juv., 27 October 1941, E. M.
Loomis (NMNH); and Florida Caverns St. Pk., 19
juvs., 27 May 1958 - 2 July 1960, N. B. Causey
(FSCA). Okaloosa Co., locality unknown, 5M, 10F,
14 March 1961, H. A. Denmark (FSCA). Escambia
Co., Pensacola, F, March-April 1986, C. H. Boll-
man (NMNH), and 3M, SF, 15 March 1970, C. Seal
(FSCA) TYPE LOCALITY; Cantonment, 5M, F, 1
May 1954, N. B. Causey (FSCA); and 5 mi. S At-
more, AL, F, 27 November 1964, E. Harvey
(FSCA).
ALABAMA: Perry Co., 6 mi. NW Marion, 3F,
6 January 1954, N. B. Causey (FSCA). Lee Co.,
Auburn, M, 4F, date unknown, B. B. Warwick
(NMNBH), 2M, 2F, 1898, F. Farley (NMNH), and
M, 6F, 3 juvs., 13 June 1959, N. B. Causey (FSCA).
Houston Co., ‘Brannon Stand”’ along US hwy. 84,
2M, 3F, 10 juvs., 26 January 1954, N. B. Causey
(FSCA). Mobile Co., ‘Seven Hills,’’ SM, 2F, 6
April 1958, S. Lazell (FSCA); Bellingrath Gardens,
M, 22 January 1965, N. B. Causey (FSCA); and
roadside park along US 90 E of Pascagoula, MS, 3F,
21 January 1965 (FSCA).
Eurymerodesmus varius christianus Chamberlin,
new status
Figs. 54, 64-65, 211
Eurymerodesmus christianus Chamberlin, 1946:140, fig. 5.
Chamberlin and Hoffman, 1958:79. Causey, 1963:77-78.
ROWLAND
Type specimens. — Male holotype, female allo-
type, and two male paratypes (NMNH) collected by
J. and W. Rapp, 16 February 1946, at Pass Chris-
tian, Harrison Co., MS. As stated in the species ac-
count, these specimens actually belong to intergrade
populations, but since no name is available for the
lobed form in eastern Louisiana and this site is only
about 35 miles to the east, it is appropriate to con-
serve christianus and assign it to this race.
Diagnosis. — A small subspecies; males with pro-
jection of mandibular stipes minute, nubbinlike, in-
distinct; acropodite discontinuous with, and sharply
demarcated from prefemur; latter with high,
rounded distal lobe, strongly elevated above distal
margin (Figs. 64-65).
Variation. — In some males, the corner of the
division point of the aperture extends downward
well into the opening, nearly touching the gonopodal
coxae, and in specimens from East Baton Rouge
Parish, the outer caudal margin is rimmed and ex-
tends into the opening. On the gonopods the
prefemoral lobe in some males leans dorsad rather
than being aligned with the stem.
Ecology. — Specimens from East Baton Rouge
Parish were collected in magnolia woods and under
dead leaves and grass around a house foundation.
Distribution. — Southeastern Louisiana between
the Pearl and Mississippi Rivers (Figs. 54, 211).
Specimens were examined as follows:
LOUISIANA: Washington Par., Angie, 2M,
2F, 2 May 1958, J. L. Crain (FSCA) and 68M, 79F,
12 December 1958, collector unknown (FSCA); 7
mi. S Angie, 10M, 4F, 16 and 24 November 1958, J.
L. Crain (FSCA); and 6 mi. SW Bogalusa, 2F, 21
January 1965, N. B. Causey (FSCA). St. Tammany
Par., 5 mi N Hickory, F, 21 February 1965, W.
Longest (FSCA); and 1 mi. E Covington, 2 juvs., 2
December 1980, M. R. and J. E. Cooper (NCSM).
Tangipahoa Par., 1 mi. N Hammond, F. 23
December 1964, R. E. Randy (FSCA); and 6 mi. E
Ponchatula, 24 March 1962, K. A. Arnold (FSCA).
St. Helena Par., Grangeville, M, 11 December 1963,
S. Geauthreaux (FSCA). Livingston Par., 2.8 mi. W
Killian, F, 2 May 1971, D. A. Rossman (FSCA).
East Feliciana Par., nr. S gate LA state prison, 3M,
3F, 1970, collector unknown (FSCA). Ascension
Par., nr. Prairieville, 6M, 3F, 20 January 1965, W.
Longest (FSCA). East Baton Rouge Par., locality
unknown, 2M, 5F, 4 November 1959, collector
unknown (NMNH) and 3M, 3F, 7 November 1964,
collector unknown (FSCA); Baton Rouge, 2M, 3F,
November 1969, and M, 2F, 25 October 1964, col-
lectors unknown (FSCA), 2M, F, 11 November
MEM. AMER. ENT. SOC., 37
M. SHELLEY 47
1963, W. J. Harman (FSCA), and M, F, 2 juvs., 9
January 1971, D. A. and C. E. Rossman (FSCA);
LSU and environs, 3M, 2 February 1964, and M, F,
9 juvs., 12 December 1964, R. E. Tandy (FSCA);
and LSU farm, M, 20 February 1963, B. Hepburn
(FSCA).
Eurymerodesmus varius louisianae Chamberlin,
new status
Figs. 54, 66-71, 211
Eurymerodesmus louisianae Chamberlin, 1942a:6, pl. 2, fig. 17.
Chamberlin and Hoffman, 1958:80.
Eurymerodesmus_ spectabilis Causey, 1950a:270, figs.
1952b:2. Chamberlin and Hoffman, 1958:81.
SYNONYMY.
Type specimens. — Male holotype and one male
paratype (NMNH) collected by L. Hubricht, 12
April 1936, 2 mi. S Saline, Natchitoches Par., LA.
The original description (Chamberlin 1942a) stated
that the type series contained a male and female,
labeled allotype in the vial, an example of sex mis-
determination.
Diagnosis. — A small to moderate-size sub-
species; males with projection of mandibular stipes
distinct, moderately long, triangular to blunt (Figs.
66-67); acropodite poorly to moderately demarcated
from prefemur; latter with variable distal swelling
weakly to moderately elevated above distal margin
(Figs. 69-71).
Color in Life. — Paranota red; metaterga dark
brown with red stripes along caudal margins, prob-
ably connecting with paranotal markings (Chamber-
lin 1942a). Causey (1950a) confirmed the striped pat-
tern, reporting orange rather than red and stripes
along both margins of the collum. She noted further
that the metatergal stripe is interrupted on some
caudal segments, that the stripe is replaced by four
orange dots in some specimens, and that the orange
pigment was so intense in some individuals that they
appeared entirely orange from a distance.
Variation. — The holotype has an oblong, blunt
mandibular process (Fig. 66), but it is triangular
(Fig. 67) in the paratype, presumably collected at the
same place and time. Both conditions are also dis-
played by males in Avoyelles and Catahoula
parishes. The Arkansas male and those from
Rapides and Avoyelles parishes have moderate distal
swellings on the gonopod prefemora (Fig. 71),
resulting in a slightly discontinuous, moderately
demarcated acropodite. The telopodites of in-
dividuals from Natachitoches and Rapides parishes
curve more dorsad than laterad, best seen in situ
6-7;
NEW
48 EURYMERODESMID MILLIPEDS
(Fig. 68). Thus, they resemble the condition in
amplus, but they lack continuous hairs on the
prefemoral stem. Some males of this race have more
prefemoral hairs than do those of the other
subspecies, but the hairs are not continuous or
regularly spaced. Thus, the main distinctions be-
tween amplus and v. louisianae are the regularity of
hairs on the prefemoral stem combined with the in
situ orientation of the gonopods.
Ecology. — According to Causey (1950a), the
Arkansas specimens were taken in a climax pine-
hardwood area. The label with the type specimens
states that they were encountered under logs; that
with the Rapides Parish specimens indicates that
they were found under cow dung.
Distribution. — Known from six scattered
localities from southern Arkansas to central Loui-
siana (Figs. 54, 211). Specimens were examined as
follows:
ARKANSAS: Columbia Co., Magnolia, M, F,
24 December 1949, N. B. Causey (ANSP).
LOUISIANA: Natchitoches Par., 2 mi. S Saline
(in Bienville Par.), 2M, 12 April 1936, L. Hubricht
(NMNH) TYPE LOCALITY. Vernon Par., 2.7 mi.
NE Caney, M, 8 April 1967, L. D. Wilson (FSCA).
Avoyelles Par., Evergreen, M, 28 October 1965, M.
Kordish (FSCA). Rapides Par., Forest Hill, 2M, F,
11 December 1945, R. Wenzel (FMNH). Catahoula
Par., 1 mi. WNW Harrisonburg, 2M, F, 25 April
1971, D. A. Rossman (FSCA).
Remarks. — Causey (1963) recorded this form
from an unspecified locality in Union County,
Arkansas. The specimens were supposedly in her col-
lection, but they are not among the material now at
the FSCA.
With only six scattered samples from an area that
has been reasonably well collected, v. /ouisianae may
be a relictual form, declining naturally toward ex-
tinction. The most obvious cause would be swamp-
ing by amplus, which is widespread and abundant in
Louisiana west of the Mississippi River. However,
much more field work is necessary in this area to
search for v. /ouisianae as well as components of the
simplex group.
Shelley and Whitehead (1986) defined subspecies
in Sigmoria as reasonably homogeneous taxa that
connect with other such taxa through intergrades.
Populations had to be continuous for subspecies to
be applicable; when there were hiatuses, the species
category was employed even if intermediates were
known. Populations of varius are continuous up to
the Mississippi River, then there is a gap of about 40
miles to the closest known site of v. /ouisianae, in
Avoyelles Parish. I therefore debated for some time
the status of this race before reasoning that the
similarities between the gonopods of v. /ouisianae
and those of the other races is so close that specific
status based solely on the mandibular projection was
unreasonable. This decision should be reviewed by
future workers with access to more material and par-
ticularly ones who can sample between West Feli-
ciana and Avoyelles parishes, to try to connect v.
louisianae and y. christianus.
Aside from fewer prefemoral hairs, the
differences between v. /ouisianae and amplus are
subtle. They differ in size, with v. /ouisianae being
small while amplus, though variable, is larger. The
apertures of v. Jouisianae and the form of amplus in
northern Louisiana are closely similar, but there are
gonopodal distinctions. Best seen in situ, the
acropodites bend more strongly from the prefemur
and are directed more laterad in v. Jouisianae,
whereas in amplus they are curved and extend in a
dorsal direction thus being more continuous with the
axis of the prefemur. Finally, the prefemur of v.
louisianae often exhibits a slight but distinct distal
swelling on the outer surface, whereas there is no
trace of this feature in amplus. The combination of
this swelling and the sparsely hirsute condition
resembles that in v. christianus. Causey (1963) pro-
vides a clue to the status of v. /ouisianae by placing
the synonym spectabilis under christianus, indicating
that she too was influenced by the prefemoral
similarities, particularly the sparse hairs.
The similarities between v. /ouisianae and amplus
pose a difficult taxonomic problem. Since some
males of the former have more prefemoral hairs than
occur on other subspecies, they could be regarded as
intergrades between the sparsely and sporadically
hirsute stems and those with dense, continuous hairs.
Furthermore, the acropodites of some males of v.
louisianae are more aligned with the prefemur and
curve dorsad in situ rather than laterad, a condition
that could also be interpreted as intermediate be-
tween varius and amplus. Thus, the question arises
as to whether amplus and varius are indeed
reproductively isolated, and the argument that they
should be combined carries considerable merit.
Others analyzing the available material might con-
sider some of my samples of v. /ouisianae to be
amplus. For clarity, I separate these forms and
describe fhe situation. The complexity in a single
species combining the multitude of forms of varius
and amplus is more than I can handle on first revi-
sion, and additional material may always clarify the
picture. Final resolution is therefore left to future
ROWLAND M. SHELLEY 49
workers, particularly persons in Louisiana who can
meticulously sample its eurymerodesmid fauna.
Eurymerodesmus varius intergrades
Figs. 54, 62-63
I combine under this heading forms with
moderate distal prefemoral swellings (Figs. 62-63),
intermediate between the strong lobes of v. chris-
tianus and the slight enlargements of the nominate
subspecies. As stated in the species account, the
enlargements become more noticeable in Mississippi,
and the boundary of the strongly lobed race is sharp,
the Pearl River. Thus, I consider all records of varius
from Mississippi as intergrades.
Ecology. — The following habitat notes are on
vial labels: under strawberries in strawberry field,
and under boards and rocks, Jackson Co.; ‘‘waste
ground,’’ Lauderdale Co.; and in pitfall traps in cot-
ton field, Oktibbeha Co.
Distribution. — Mississippi (Fig. 54). Specimens
were examined as follows:
MISSISSIPPI: Panola Co., 13 mi. E Como,
5M, 3F, 25 December 1964, W. Longest (FSCA).
Monroe Co., Aberdeen, 3M, 3F, 6 March 1961, R.
E. Hutchins (FSCA). Oktibbeha Co., Craig Springs,
10 mi. S Starkville, 18M, 23F, jv., 1 October - 31
December 1979, W. H. Cross (MEM, NCSM).
Newton Co., Decatur, many juvs., 5 May 1933, H.
Janes (NMNH). Lauderdale Co., Meridian, 1st St.
and 53rd Ave., 5M, 6F, 10 December 1961, L.
Hubricht (RLH). Stone Co., MS hwy. 15 at Har-
rison Co. line, 2F, date unknown, N. B. Causey
(FSCA). Jackson Co., Pascagoula, 2M, 3F, 12
February 1962, T. Sheldon (FSCA); Spanish Fort,
10M, 4F, 21 January 1965, N. B. Causey (FSCA); 3
mi. N Fontanbleau, 11M, 12F, 1 January 1954, N.
B. Causey (FSCA); and Gulf Coast Res. Lab, Ocean
Springs, 3M, 7F, 27 November 1964, C. Guise
(FSCA). Harrison Co., Biloxi, 20 juvs., 12 July
1927, H.A. Robinson (NMNH). Gulfport, M, 2F,
18 November 1922, E. K. Bynum (NMNH); and
Pass Christian, 2M, 2F, 16 February 1945, J. and
W. Rapp (NMNB).
Remarks. — The type collection of christianus
contains a gonopod of angularis that appears to
have been accidentally placed in the wrong vial,
probably by Causey when she was examining and
comparing eurymerodesmids years ago. A gonopod
of the holotype of wellesleybentoni, a synonym of
angularis, is missing, and the extraneous gonopod in
the christianus vial conforms closely to the remain-
ing one in the wellesleybentoni vial.
MEM. AMER. ENT. SOC., 37
Eurymerodesmus amplus Causey
Figs. 54, 72-82, 211-212
Leptodesmus _hispidipes: 1895:67.
1918:370 (in part).
Fontaria (Eurymerodesmus) hispidipes: Brolemann, 1900:129, fig.
32.
Eurymerodesmus amplus Causey, 1952b:4, fig. 3. Chamberlin and
Hoffman, 1958:78.
Eurymerodesmus hamatilis Loomis, 1969:250-251, figs. 11-13.
NEW SYNONYMY.
Eurymerodesmus hispidipes: Stewart, 1969:384.
Brolemann, Chamberlin,
Type specimens. — Male holotype and female
allotype (AMNH) collected by W. J. Harmon in
November 1951 at Ruston, Lincoln Par., LA.
Diagnosis. — A moderate-size species; males
characterized by long mandibular projections; sides
of aperture divided, with variably closed to partly
open pouch; telopodite long; acropodite short,
about 1/8 of telopodite length, broadly terminal,
poorly demarcated from prefemur; latter with many
hairs arranged continuously along inner margin;
cyphopod valves of females with distal corners ex-
tending into distinct ridge of variable length and con-
figuration, sublinear or angling toward corner, with
or without terminal papilla in more angular forms;
receptacle moderate-size, sides curving partly around
operculum.
Color in Life. — Paranota red; metaterga dark
speckled olive with red stripes along caudal margins
connecting paranotal spots, stripes partly obliterated
by intrusions of base color; collum with red stripe
along anterior margin.
Holotype. — Body highly fragmented, length un-
measurable, maximum width 4.2 mm.
Somatic features similar to hispidipes, with following excep-
tions:
Width across genal apices 3.0 mm, interantennal isthmus 0.7
mm. Antennae reaching back to caudal margin of 3rd tergite,
relative lengths of antennomeres 2>3>4>6>5>1>7. Genae
with faint central impressions. Facial setae as follows: epicranial
and genal absent, subantennal 1-1, frontal 1-1, clypeal about
16-16, labral about 22-22. Process of mandibular stipes moderately
long, broadly rounded apically, curving slightly inward at
midlength (Fig. 72).
Collum extending slightly below ends of following tergite.
Paranota moderately depressed, mostly continuing slope of
dorsum, peritremata flattened; posterior corners rounded through
segment 9, blunt on 10-12, becoming progressively more acute
caudally.
5th and 6th sterna with widely separated knobs between leg
pairs. Postgonopodal sterna with narrowly segregated, elevated
patches between 9th legs; remaining sterna becoming progressively
flatter and less hirsute caudally. 2nd coxa with large, subconical
anterior, and smaller rounded posterior lobes. Prefemora with
broad lobes on outer surfaces of legs through segment 11.
Gonopodal aperture (Fig. 73, not this specimen) broadly ovoid,
with closed, caudolateral pouches and moderate anterior indenta-
50 EURYMERODESMID MILLIPEDS
tion, 1.6 mm wide and 1.0 mm long at maxima, outline of entire
apparatus irregularly hexagonal; anterior indentation short, apical-
ly rounded; sides divided, curving broadly mediad, leaning over
opening, elevating gradually to division point near midlength, in-
ner lateral margin curving dorsolaterad then blending smoothly
into inner caudal margin, outer lateral margin leaning over open-
ing, angling moderately laterad then curling mediad to metazonal
surface at caudolateral corner, forming closed pouch with inner
margin, rims slightly irregular; inner caudolateral corner smoothly
curved but hidden by overhanging outer lateral margin; outer
caudolateral corner moderately sharp, forming approximately a
60° angle; inner caudal margin extending linearly across midline,
not parallel with outer caudal margin; latter curving along anterior
surface of coxal condyles, extending caudad in midline; margins ir-
regularly hirsute, with long hairs arising sporadically from anterior
rim and overhanging opening, particularly along indentation and
anterolateral corners, with additional short hairs arising from
outer surface of outer lateral margin and moderate tufts from
pouches. Gonopods in situ (Fig. 73, not this specimen) with
telopodites lying partly over and under one another in midline of
aperture, diverging at level of caudal margins, curving gently sub-
dorsad and terminating over 9th coxae. Gonopod structure as
follows (Figs. 74-75): Telopodite terminating well before level of
distal extremities of hairs. Prefemur long, about 7/8 of telopodite
DL | \i
ILI IN\, |
\ ET Te
Vane (AY)
Figs. 72-82. Eurymerodesmus amplus. 72, projection of right mandible of holotype, lateral view. 73, aperture and gonopods in situ
of topotype, ventral view. 74, left gonopod of holotype, lateral view. 75, telopodite of the same, medial view. 76-78, aperture and
gonopods in situ of males from Evangeline and Calcasieu pars., LA, and Angelina Co., TX, respectively, ventral views. 79, prefemur of
left 7th leg of male from Angelina Co., anterior view. 80, cyphopods in situ of female from Vernon Par., LA, ventral view. 81, left
cyphopod of allotype, ventral view. 82, the same, medial view. Scale lines for figs. 73, 76-78, and 80 = 1.00 mm. Line for other figs. =
0.30 mm for 72 and 79, 0.40 mm for 74 and 81-82, and 0.50 mm for 75.
ROWLAND M. SHELLEY 51
length, stem leaning over coxa for most of length, more so distad,
sides parallel throughout, with two rows of continuous, regularly
spaced hairs along inner margin, angling slightly outward distad,
and distomedial tuft of about 10 hairs. Acropodite short and
broadly terminal, only about 1/8 of telopodite length, continuing
curvature of prefemur and directed dorsolaterad, smoothly con-
tinuous with, and poorly demarcated from prefemur, sides nar-
rowing continuously to acuminate tip.
Allotype. — Length 30.7 mm, maximum width 4.2 mm, W/L
ratio 13.7%. Agreeing closely with holotype in somatic features,
with following exceptions: Process of mandibular stipes relatively
large, extending distinctly from base, broadly triangular. Sterna
with variably broad impressions in midline, with sparse hair tufts
subtending coxae except on segments 17-18.
Cyphopodal aperture broadly rectangular, sides strongly
elevated above metazonal surface. Cyphopods in situ with corners
of valves protruding through opening, valves oriented transversely.
Valves (Figs. 81-82) large, subequal, ventral margin elevated into
rim, angling slightly distad and slightly longer at corner, medial
margin moderately hirsute. Receptacle moderately large, glabrous,
located laterad at bases of valves, sides curving partly around oper-
culum. Latter small, hidden under free (anterior) side of valves.
Variation. — The colors reported above were of
the specimens I collected in Harris County, Texas.
According to Causey (1952b), the paranota of the
types were reddish orange; she provided no addi-
tional color details. The label accompanying the
specimens from Sulphur, Calcasieu Parish, Loui-
siana, states ‘‘orange,’’ presumably referring to the
paranota, so the color of this species may vary.
Eurymerodesmus amplus is difficult to com-
prehend on its own merits, not to mention the close
similarities with v. louisianae. The gonopods exhibit
little intraspecific variation, but there are three main
aperture variants that tend to occupy different sec-
tors of the range thus suggesting that subspecies may
be warranted. However, there are enough instances
where one variant also occurs in another sector or
there are additional sympatric forms, that this
possibility is obviated. Moreover, the anatomical
diversity is greater in Texas, and one or more
heterogeneous races there would conflict with com-
paratively homogeneous ones in Louisiana. Finally,
female variation is not parallel and would therefore
counter subspecies based on males. By not showing
comparable differences at fairly sharp male boun-
daries, females also contraindicate specific statuses
for the principle male variants. Females show a
general clinal trend toward a longer and more
angular valvular ridge in a southerly direction. In-
dividuals from northern areas like Lincoln and Nat-
chitoches Parishes have distinct, but only slightly
elevated, ridges that are either linear or angle slightly
laterad toward the distal corner. In southern Loui-
siana the rim is distinctly longer at this corner and
therefore obviously slanted in this direction. This
MEM. AMER. ENT. SOC., 37
variant extends into western Louisiana and eastern
Texas, and in many individuals the medial corner is
additionally prolonged into a papilla or short
digitiform projection (Fig. 80). As with males,
females are more variable in Texas than in Loui-
siana, and the overall picture in both states is ex-
tremely complex when one considers both sexes.
Conclusions based on one sex are contradicted by
the other, and in general female variation traverses
the boundaries between male forms. I therefore
regard amplus as a single highly variable species with
three principle male morphs, which are characterized
in the ensuing paragraphs along with lesser forms in
Texas.
The first variant is represented by the holotype
and occurs in northern Louisiana (Lincoln, Natchi-
toches, Desoto, Bienville, Red River, Sabine, and
eastern Vernon parishes), extends across the Sabine
River into Harrison County, Texas, and occurs
sporadically to the west in Texas (Wood, Anderson,
Tyler, and to a lesser extent Nacogdoches counties).
In this form (Fig. 73), the aperture sides are
moderately high and lean well over the opening at
division point; the inner lateral margin extends
downward or dorsad into the opening, and the outer
lateral margin flares moderately laterad, all combin-
ing to form a distinct closed pouch. In addition to
the rim, females in this area display a variable inden-
tation on the medial valvular surface that is greater
in individuals from the northern periphery of the
third male variant, thus constituting another trait
traversing a boundary between male forms.
The second male variant (Fig. 76) occurs to the
south in Evangeline, Jefferson Davis, and Iberia
parishes Louisiana. Here the sides of the aperture are
more strongly elevated and upright from the antero-
lateral corner to division point. The outer lateral
margin does not angle as strongly laterad and in
some specimens is nearly colinear with the anterior
part of the sides. The inner lateral margin is directed
downward, and the net effect is a shallower pouch
that is virtually absent from a few males. A similar
form occurs in Montgomery County, Texas, and one
in Nacogdoches County is intermediate between this
and the first morph.
The third variant occurs in western Louisiana
(western Vernon, Beauregard, Calcasieu, and
Cameron parishes) and eastern Texas (Shelby,
Jasper, Rusk, Smith, Polk, Houston, Grimes,
Chambers, Galveston, and Harris counties) where it
intermixes with the other two forms. Here the sides
rise gradually to division point, which leans strongly
mediad over the opening (Fig. 77). The inner lateral
52 EURYMERODESMID MILLIPEDS
margin extends slightly inward into the opening
before curving broadly dorsad, and the outer lateral
margin angles moderately laterad, forming a deep,
moderately open pouch. This morph intermixes in
eastern Texas with the second variant and with that
named hamatilis by Loomis (1969) (Fig. 78), in
which the pouch is even deeper and more open
because the outer lateral margin flares strongly
laterad and the inner margin extends submediad into
the opening before curving dorsad. Thus, although
the sides do not lean strongly over the opening, the
pouch is still broad and deep because of the initial in-
vagination of the inner lateral margin. As noted by
Loomis (1969), this variant also possesses a sclero-
tized hook on the anterior surfaces of the prefemora
on the 7th and 9th legs (Fig. 79). This type of
aperture is known from three widely separated
localities in Angelina, Panola, and Mason counties,
the latter sites respectively about 66 miles north and
280 miles west of the first. The Mason County
record represents an apparently allopatric popula-
tion in the midst of melacis, some 200 miles west of
the closest known site of either amplus, the varius
group, or the Kewanius lineage. In the area of
eastern Texas occupied by the third variant, the
valvular rim in females slopes strongly mediad,
usually culminating in a variably papillate to
digitiform projection on the distal corners (Fig. 80),
the projection tending to be longer and more
digitiform to the south. However, some females in
this area lack papillae and exhibit slanted rims. The
form named hamatilis is recognizably different, but
the situation with females and sympatry with other
males indicates that it. is merely one variant of a
highly variable species. Consequently, hamatilis is a
junior synonym of amplus.
Ecology. — The male I collected in July 1980 in
Harrison County, Texas, the wrong season for eury-
merodesmids, consisted of loose exoskeletal rings
without appendages and was identified from the
aperture configuration. The segments were found
under leaf litter beside a building at Caddo Lake
State Park. The specimens from Harris County were
discovered under logs beside a trail in a
predominantly pine habitat at Memorial Park,
Houston. Habitat notations on other vial labels are
as follows: under logs and boards in a bottomland
meadow adjacent to a pine-hardwood forest (Red
River Par., LA), in a rotting shed (Sabine Par., LA),
at a plant nursery (Jefferson Davis Par., LA),
under sawdust (Vernon Par., LA), under a log
(Nacogdoches Co., TX), in a greenhouse (Smith
Co., TX), and on a prairie (Chambers Co., TX).
The sample from near Steinhagen Lake, Tyler Coun-
ty, Texas, was discovered by ultraviolet light at
night, apparently flourescing.
Distribution. — Louisiana from the Mississippi
River and the Atchafalaya Bayou to the middle
Trinity and lower Brazos Rivers in eastern Texas,
with an allopatric population 200 miles to the west
on the eastern periphery of the Edwards Plateau in
Mason County (Figs. 54, 211-212). Specimens were
examined as follows:
LOUISIANA: Lincoln Par., Ruston, M, F,
November 1951, W. J. Harmon (AMNH) and 4M,
juv., 2 January 1953, F. L. Afeman (FSCA) TYPE
LOCALITY. Bienville Par., Castor, 2M, 21
November 1954, J. E. Sublette (FSCA). Nat-
chitoches Par., Creston, 14M, 8F, 27 February-5
May 1915, K. P. Schmidt (MCZ); Grand Ecore, M,
26 October 1934, J. E. Sublette (FSCA); Nat-
chitoches, M, 13 November 1953, B. Stanberry
(FSCA) and M, 25 February 1954, J. E. Sublette
(FSCA); and Kisatchie, M, F, 30 October 1954, W.
J. Harmon (FSCA). Avoyelles Par., Evergreen, F,
23 December 1965, M. Kordisch (FSCA). Red River
Par., 3.7 mi. NW Coushatta, 4M, 3F, 10 April 1971,
P. J. Kinnick (FSCA); and Coushatta, M, 28
November 1953, Bamburg (FSCA). Desoto Par.,
along US hwy. 171, 2.5 mi. N Mansfield, 4M, 13
March 1966, R. E. Tandy (FSCA). Sabine Par.,
Hodges Garden, along US hwy. 171 N of Sandel,
2M, 7F, juv., 26 March 1965, R. E. Tandy, G. E.
Gates (FSCA). Vernon Par., unknown locality, F,
17 June 1956, J. E. Sublett (FSCA); LA hwy. 111,
1.9 mi. N. jct. LA hwy. 464, F, 19 February 1966, R.
E. Tandy (FSCA); 2.4 mi. ENE Pitkin, 3M, 3F, 20
February 1966, R. E. Tandy (FSCA); near Sabine R.
on LA hwy. 8, 4M, 2F, 25 March 1965, R. E.
Tandy, G. E. Gates (FSCA); and along LA hwy.
464, 11.5 mi. SSW Caney, M, 19 February 1966, R.
E. Tandy (FSCA). Allen Par., West Bay Game
Area, F, 2 October 1965, L. D. Wilson (FSCA).
Beauregard Par., US hwy. 190 at Sabine R., F, 30
April 1966, R. E. Tandy (FSCA); and DeRidder,
8M, 6F, 20 March, 17 October, and 9 December
1969, H. Rhame (FSCA); and 5 mi. S DeRidder,
6M, 3F, 22 juvs., 10 December 1969, H. Rhame
(FSCA). Evangeline Par., Lake Chicot, F. 20 March
1965, M. Kordisch (FSCA); 2.5 mi. S Chicot St. Pk.,
M, 30 January 1968, E. D. Kaiser (FSCA); and
Boggs Bayou, 2.7 mi. W jct. LA hwys. 106 and 13,
M, F, 9 January 1966, R. E. Tandy (FSCA). Iberia
Par., Avery Island, M, F, juv., 7 April 1974, N. B.
Causey (FSCA). Acadia Par., 1-10 at Mermentan
R., 2F, 26 September 1965, R. E. Tandy (FSCA).
ROWLAND M. SHELLEY 53
Jefferson Davis Par., Elton, 14M, 11F, 29
November 1964 and 1 March 1965, C. Steal (FSCA).
Calcasieu Par., Lake Charles, M, juv., 15 April
1965, and 4M, 4F, 28 April 1967, M. Kordish
(FSCA); Sulphur, 4M, 4F, 22 February 1965, collec-
tor unknown (FSCA); and 10 mi. W Lake Charles,
M, F, 18 November 1964, M. Kordish (FSCA).
Cameron Par., 5 mi. W Holly Beach, M, 29 April
1962, K. Arnold (FSCA).
TEXAS: Harrison Co., nr. Waskom, 2F, 25
March 1962, N. B. Causey (FSCA); and Caddo
Lake St. Pk., M (exoskeletal fragments), 11 July
1980, R. M. Shelley (NCSM). Panola Co., Car-
thage, 3M, 3F, 10 March and 24 November 1962, L.
P. Hull (FSCA). Shelby Co., Paxton, 13M, 8F, date
and collector unknown (FSCA). Wood Co., Alba,
5M, 3F, 2 juvs., 7 February 1932, collector unknown
(NMNBH). Smith Co., Tyler, 2M, 28 December 1964,
T. C. Stewart (FSCA). Rusk Co., along US hwy
259, 5 mi. N Mt. Enterprise, M, 19 November 1967,
T. C. Stewart (SFAU); along FM 836, 3 mi. N New
Salem, 4M, 3F, 17 November 1967, T. C. Stewart
(FSCA); along FM 834, 3 mi. S New Salem, F, 3
juvs., 17 November 1967, T. C. Stewart (SFAU);
and Lufkin, juv. M, 22 May 1940, S. and D. Mulaik
(NMNH). Anderson Co., Palestine, 3M, 3F, 3 juvs.,
26 January 1962, J. O’Keefe (FSCA). Houston Co.,
Crockett, 6M, 3F, 4 December 1961, L. Heard
(FSCA). Nacogdoches Co., Nacogdoches, 10M, 6F,
November 1931, collector unknown (NMNH) and
M, 11 October 1966, T. C. Stewart (SFAU); and
Stephen F. Austin Forest, 2M, 16 March 1968, J.
Reddell (FSCA). Angelina Co., 8 mi. N Zavalla,
near Sam Rayburn Res., 4M, F, 4 juvs., 23 April
1967, J. C. Loomis (NMNH, FSCA); and county
airport, F, 13 October 1966, J. A. Kelgey
(SFAU). Jasper Co., Jasper, 26M, 32F, juv., 15
January-25 March 1962, J. DuRard (FSCA). 7yler
Co., Woodville, 12M, 14F, 8 juvs., 22 February
1962, R. Timbrook (FSCA); and along US hwy. 190
at Steinhagen L., 25M, 4F, 20 October 1985, J. C.
and J. E. Cokendolpher (NCSM). Polk Co.,
Dallardsville, 2M, juv., 24 February 1962, J. P. Ken-
nedy (FSCA). Grimes Co., Navasota, 7, January
1931, collector unknown (NMNH) and M, F, 5
juvs., 2 December 1961, W. Watkins (FSCA).
Montgomery Co., Conroe, 5M, 2 juvs., 10 March
1963, B. Ray (FSCA). Harris Co., Houston, F, 20
December 1930, collector unknown (NMNH) and
Memorial Pk., M, 2F, 18 February 1986, R. M.
Shelley (NCSM). Chambers Co., Smith Point, 4M,
2F, 3 November 1918, Hansen and Barber
(NMNH). Galveston Co., La Marque, 4M, 9F, 7
MEM. AMER. ENT. SOC., 37
juvs., date unknown, W. A. Benton (FSCA); and
Galveston Island, 2M, F, 1870, W. Julich (MCZ).
Mason Co., 1 mi. NE Mason, 3M, 8 November
1964, J. Reddell (FSCA).
Remarks. — The specimens from an unspecified
site in Louisiana and reported by Brolemann (1895)
as Leptodesmus hispidipes are amplus. This sample
consists of 11 males and 2 females and is housed at
the MNHP. Brolemann (1900) did not mention a
locality, but I believe he was referring to this sample
when he proposed Fontaria (Eurymerodesmus)
hispidipes. The specimens from Creston, Nat-
chitoches Parish, Louisiana, which Chamberlin
(1918) also recorded as Leptodesmus hispidipes, are
mostly amplus, though they also contain a male of
birdi. An amplus male from this site measured 29.1
mm long, 4.3 mm wide, W/L ratio 14.8%. Finally,
the records of E. hispidipes by Stewart (1969) from
Angelina, Nacogdoches, Rusk and Smith, counties,
Texas, also refer to amplus, and since his records
from Sabine, Shelby, and Upshur counties lie within
the range, they are shown as literature records of
amplus in figures 54 and 212.
Eurymerodesmus newtonus Chamberlin
Figs. 54, 83-88, 211
Eurymerodesmus newtonus Chamberlin, 1942a:5, pl. 2, fig. 14.
Chamberlin and Hoffman, 1958:80.
Eurymerodesmus bentonus Causey, 1950a:268-270, fig. 5. Cham-
berlin and Hoffman, 1958:78. NEW SYNONYMY.
Type specimens. — Male holotype and two
female paratypes (NMNH) collected by L. Hubricht,
29 April 1936, in the Boston Mountains, 1.2 mi.
S Jasper, Newton Co., AR. The head and pre-
gonopodal segments of the holotype are missing,
and the male description is therefore based on the
specimen from Monte Ne, Benton County, Arkan-
sas, whose gonopods and aperture are virtually
identical.
Diagnosis. — A moderate-size species; males
characterized by moderate-size mandibular projec-
tion; sides of aperture divided, with moderately open
caudolateral pouches; telopodite long; acropodite
short, about 1/4 of telopodite length, broadly term-
inal, poorly demarcated from prefemur, uncinate;
prefemur with many hairs arranged continuously
along inner surface; cyphopod valves of females not
modified, without projections; receptacle moderate-
size, sides curving partly around operculum.
Color in Life. — Paranota orange; metaterga
dark olive with orange stripes along caudal margins
connecting paranotal markings; collum with orange
stripes along both anterior and posterior borders
54 EURYMERODESMID MILLIPEDS
Figs. 83-88. Eurymerodesmus newtonus. 83, projection of
right mandible of male from Benton Co., AR, lateral view.
84, aperture and gonopods in situ of male from Washington Co.,
AR, ventral view. 85, left gonopod of holotype, lateral view.
86, telopodite of left gonopod of male from Benton Co., AR,
medial view. 87, left cyphopod of female paratype, caudal view.
88, the same, medial view. Scale line for fig. 84 = 1.00 mm. Line
for other figs. = 0.35 mm for 85, 0.40 mm for 83 and 87-88, and
0.50 mm for 86.
(Causey 1950a). Chamberlin (1942a)
preserved coloration and noted stripes.
Male from Monte Ne, Benton Co., AR. Body
highly fragmented, length unmeasurable, maximum
width 4.8 mm.
Somatic features similar to hispidipes, with following excep-
tions:
Width across genal apices 2.7 mm, interantennal isthmus 0.9
mm. Antennae reaching back to caudal end of 2nd tergite; anten-
nomeres becoming progressively more hirsute distad, relative
lengths 2>3>5>4>6>1>7. Genae with faint central impres-
sions. Facial setae as follows: epicranial and interantennal absent,
subantennal 1-1, frontal 1-1, genal 2-2, clypeal about 9-9, labral
about 14-14. Projection of mandibular stipes moderately long,
apically narrowly rounded (Fig. 83).
Collum not extending below ends of following tergite. Paranota
moderately depressed, continuing slope of dorsum basally, flat-
tened distad; posterior corners rounded through segment 6, blunt
on 7-12, becoming progressively more acute posteriorly.
Sth sternum with minute lobes between 4th legs and broader
elevated areas between Sth legs; 6th sternum with short, para-
reported
median knobs between both leg pairs. Postgonopodal sterna with
minute, widely segregated lobes between 9th legs; remaining sterna
generally flat and plate-like, with variable transverse impressions
originating between leg pairs and moderate hair patches subtend-
ing coxae. 2nd legs absent, coxal configuration unknown.
Prefemoral lobes broadly rounded, extending through segment 11.
Gonopodal aperture (Fig. 84, not this specimen) ovoid, with
partly open caudolateral pouches, slightly indented anteriad at
midline, 1.3 mm wide and 0.4 mm long at maxima, outline of ap-
paratus irregularly hexagonal; anterior indentation short, broad,
apically rounded; sides curving slightly mediad and leaning slightly
over opening, rising slowly and continuously to division point
distal to midlength, inner lateral margin very short, curving im-
mediately into inner caudal margin, outer lateral margin flaring
slightly sublaterad, not leaning over opening, curving and drop-
ping rapidly to metazonal surface just before caudolateral corner,
forming moderate, open pouch with inner margin, rims slightly
emarginate; inner caudolateral corner indistinct, smoothly curved
continuation of inner lateral margin; outer caudolateral corner
broadly curved; inner caudal margin angling caudad, outer third
slightly elevated; outer caudal margin angling slightly caudad,
more so in midline; anterior margin glabrous, sides with scattered
hairs arising from rims beginning along elevation before division
point, with short marginal hairs at caudolateral corners and slight
tufts from within pouches. Gonopods in situ (Fig. 84, not this
specimen) with telopodites angling across midline and crossing op-
posite member, extending to caudal margin then curving laterad,
lying nearly completely over and under each other, tips directed
dorsolaterad. Gonopod structure as follows (Figs. 85-86):
Telopodite terminating just before level of distal extremities of
hairs. Prefemur long, about 3/4 of telopodite length, stem curving
gently and evenly dorsad for most of length, more so distad, ex-
panding slightly near midlength, narrowing distad, with two rows
of continuous, regularly spaced hairs along inner margin and
distomedial tuft of about 6 hairs. Acropodite short and broadly
terminal, about 1/4 of telopodite length, uncinate, curving strong-
ly dorsolaterated, poorly demarcated from, and smoothly con-
tinuous with, prefemur, narrowing smoothly and continuously to
subacuminate tip.
Female paratype. — Length 27.0 mm, maximum width 4.2
mm, W/L ratio 15.6%. Agreeing essentially with holotype in
somatic features, with following exceptions: Mandibular stipes
with short, rounded projection. Sterna flat, with only a few sparse
hairs on anterior segments, glabrous thereafter.
Cyphopodal aperture elliptical, caudal margin indented slightly
mediad, sides slightly elevated above metazonal surface.
Cyphopods in situ with valves oriented transversely in aperture,
central groove visible in opening. Valves (Figs. 87-88) moderately
large, unequal, anterior one slightly larger, without lobes or pro-
jections, moderately hirsute distad. Receptacle oriented dorsoven-
trad at bases of valves, forming right angle with latter, curving
partly around base of operculum, glabrous. Operculum relatively
large, located under valves, partly shielded on anterior and
posterior sides by receptacle.
Variation. — The holotype gonopods lack the
characteristic distomedial tuft of prefemoral hairs
that is present on other males. The male from Wash-
ington County exhibits a slight distal swelling on the
outer prefemoral margin.
Distribution. — Known definitely from only
three counties in the northwest corner of Arkansas
(Figs. 54, 211). Specimens were examined as follows:
ROWLAND M. SHELLEY 55
ARKANSAS: Newton Co., Boston Mts., 1.2
mi. S Jasper, M, F, 29 April 1936, L. Hubricht
(NMNH) TYPE LOCALITY. Benton Co., Monte
Ecology. — The material from Benton County
reported by Causey (1950a) was collected in
predominantly oak woods at the base of a north-
facing slope.
Ne, M, 2F, 19 November 1949, N. B. Causey
(ANSP, FSCA). Washington Co., Cove Creek
Valley, M, Spring 1956, M. Hite (FSCAQ).
Remarks. — According to Chamberlin (1942a),
the type series of newtonus contained the male
holotype and three females, two of which are lost.
Causey (1950a) reported collecting numerous
specimens of newtonus along with birdi at Blue
Springs, Carroll County, Arkansas. This material
also is lost, but the site is within the area circum-
scribed by the three known counties and thus is
plausible for the species. It is shown as a literature
record in figures 54, 211.
One of the specimens Causey (1950a) reported
from Benton County is an ‘“‘intersex,’’ labeled a
“hermaphrodite’”’ in the vial, with typical
cyphopods, gonopods, and genital apertures at the
normal locations. The specimen is present in the
FSCA and is a functional female with eggs and the
female paranota, sterna, and legs. I interpret the
gonopods as a developmental accident reflecting the
ability of appendages on anterior segments in both
sexes to become either legs or gonopods depending
on genetic signals. I (1977) summarized examples of
this phenomenon in the Xystodesmidae, and it is
also exhibited by an accessory gonopod on a male of
b. birdi. Such accidents are rare in females, and it is
noteworthy that the gonopods are at the normal
location, the anterior position of segment 7.
Eurymerodesmus oliphantus Chamberlin
Figs. 54, 89-95, 211
Eurymerodesmus oliphantus Chamberlin, 1942a:6, pl. 2, fig. 15.
Chamberlin and Hoffman, 1958:80.
Eurymerodesmus hispidipes: Causey, 1950a:267-268, figs. 1-4;
1952b, fig. 2. Chamberlin and Hoffman, 1958:191 (in part).
Type specimens. — Male holotype and female
allotype (NMNH) collected by L. Hubricht, 10 April
1936, 15 mi. SW Oliphant, Jackson Co., AR.
Diagnosis. — A small to moderate-size species;
males characterized by long mandibular projection;
sides of aperture divided, with widely open
caudolateral pouches; telopodite long; acropodite
short, about 1/8 of telopodite length, broadly ter-
minal, poorly demarcated from prefemur; latter
MEM. AMER. ENT. SOC., 37
with many hairs arranged continuously along inner
margin; cyphopod valves of females not modified,
without projections; receptacle moderate-size, sides
curving partly around operculum.
Color in Life. — Paranota red; metaterga black
with broad, red stripes along caudal margins
connecting paranotal markings; collum with red
stripes along both anterior and posterior borders
(Fig. 89). Chamberlin (1942a) and Causey (1950a)
also noted the striped pattern in _ preserved
specimens.
Holotype. — Body highly fragmented, length un-
measurable, maximum width 3.8 mm.
Somatic features similar to hispidipes, with following excep-
tions:
Width across genal apices 2.6 mm, interantennal isthmus 0.9
mm. Antennae reaching back to caudal margin of 2nd tergite,
relative lengths of antennomeres 2>3>4=5=6>1>7. Genae
without impressions. Facial setae with only clypeal about 8-8, and
labral, about 12-12, series. Process of mandibular stipes long,
bowed inward, narrowly rounded apically (Fig. 90).
Collum extending slightly below ends of following tergite. Para-
nota moderately flattened, interrupting slope of dorsum; posterior
corners rounded through segment 6, blunt on 7-10, becoming pro-
gressively more acute caudally.
Sth sternum with dense hirsute patches adjacent to leg coxae,
not elevated; 6th sternum with low, elevated, areas adjacent to leg
coxae. Postgonopodal sterna with deep central impression and
low, elevated, hirsute areas between 9th legs; remaining sterna
generally flat and unmodified, with hair patches adjacent to leg
coxae, becoming progressively sparser caudally. 2nd coxa with
broadly rounded, subequal anterior and posterior lobes.
Prefemoral lobes extending through segment 10.
Gonopodal aperture (Fig. 91, not this specimen) ovoid, with
open, caudolateral pouches and short, broad anterior indentation,
2.3 mm wide and 0.9 mm long at maxima, outline of apparatus
trapezoidal; anterior indentation short, broad; sides curving
mediad to division point, upright, not leaning over opening, rising
progressively to division point at midlength, inner lateral margin
continuing curved contours, extending inward, dropping abruptly,
and curving smoothly into inner caudal margin, outer lateral
margin well separated from inner, flaring strongly and obliquely
caudolaterad, lowering progressively to metazonal surface at
caudolateral corner, forming broad, deep, open pouch with inner
margin, rims of anterolateral margins smooth, those of inner and
outer margins slightly irregular; inner caudolateral corners in-
distinct, broadly rounded; outer caudolateral corners broadly
triangular, about 45°; inner caudal margin curving slightly, drop-
ping rapidly to metazonal surface, and extending sublinearly
across midline; outer caudal margin curving slightly along anterior
surfaces of coxal condyles and continuing as faint line across
midline; margins densely and irregularly hirsute, with long hairs
arising sporadically from anterior and lateral rims, particularly
along anterior indentation and from outer surface of outer lateral
margin, and overhanging opening, also with dense tufts of short
hairs arising from within pouches, caudal margins glabrous.
Gonopods in situ (Fig. 91, not this specimen) with telopodites
abutting or lying parallel to each other in midline, curving laterad
over caudal margins and terminating over 9th coxal condyles, tips
directed dorsolaterad. Gonopod structure as follows (Figs. 92-93):
56 EURYMERODESMID MILLIPEDS
Telopodite terminating well before level of distal extremities of
hairs. Prefemur long, about 7/8 of telopodite length, leaning over
coxa, narrowing slightly to around midlength then expanding,
widest at distal extremity, with numerous long hairs arranged con-
tinuously in two rows along inner margin and distomedial tuft of
about 10 hairs. Acropodite short and broadly terminal, about 1/8
of telopodite length, angling dorsolaterad, poorly demarcated
from, and smoothly continuous with, prefemur, narrowing rapidly
near midlength to acuminate, slightly bisinuate tip.
Allotype. — Length 28.4 mm, maximum width 3.8 mm, W/L
ratio 13.4%. Agreeing closely with holotype in somatic features,
Figs. 89-95.
of holotype, lateral view. 91, aperture and gonopods in situ of male from Jackson Co., IL, ventral view. 92, left gonopod of holotype,
lateral view. 93, telopodite of the same, medial view. 94, left cyphopod of allotype, caudal view. 95, the same, medial view. Scale lines for
figs. 89 and 91 = 1.00 mm. Line for other figs. = 0.60 mm for 90 and 92, 1.00 mm for 93-95.
with following exceptions: Mandibular stipes with short, broadly
rounded projection. Sterna flat, nearly glabrous, with at most only
an occasional scattered hair.
Cyphopodal aperture subrectangular, sides elevated above
metazonal surface. Cyphopods in situ with valves oriented
transversely in aperture, distal surface of anterior valve visible in
Opening. Valves (Figs. 94-95) relatively large, unequal, anterior
valve larger, extending slightly above and overlying caudal valve,
moderately hirsute. Receptacle located laterad at bases of valves,
relatively narrow, sides curving partly around operculum. Latter
large, located under free, dorsomedial side of valves.
Euerymerodesmus oliphantus. 89, segments 1-5 of male from Jackson Co., IL, dorsal view. 90, right mandibular process
ROWLAND M. SHELLEY S7/
Variation. — Measurements of a male from
White County, Arkansas, adjacent to Jackson
County, are length 23.7 mm, maximum width 3.7
mm, W/L ratio 15.6%.
The most notable variation is the marked reduc-
tion in hirsuteness in the southernmost males from
Jackson and White counties, Arkansas. All other
specimens are extremely hairy, particularly inside the
pouches and along the rim and outer surfaces of the
outer lateral margins; hence one such male was used
to depict the aperture configuration and the in situ
gonopod arrangement (Fig. 91) rather than the holo-
type.
The aperture configurations are very similar
throughout the range. In males from White County,
Arkansas, and Phelps, Pulaski, and St. Louis coun-
ties, Missouri, the flare extends more obliquely
laterad and tends to lean over the pouch, thus form-
ing a cavity beneath the outer lateral margin from
which more hairs arise. The angle of slope of the in-
ner caudal margin can vary from gradual as in the
holotype to steep as in males from Pope County, II-
linois. The distance between the inner and outer
lateral margins, or the rate at which they diverge,
also varies, and males in which this distance is
greater possess a cavity with hairs beneath the divi-
sion point.
In situ the telopodites may be parallel (Fig. 91) or
they may cross each other basally in the midline of
the aperture (at the bases of the prefemora) and
again distad at the bases of the acropodites above the
sternum between the 9th legs. In northern males the
acropodite is not apically bisinuate.
The female from St. Louis County, Missouri, the
only adult female collected aside from the allotype,
agrees with the latter in that the anterior valve is
larger than the caudal one.
Ecology. — According to vial labels, the males
from Pope County, Illinois, were encountered under
rocks. I discovered that from Jackson County, II-
linois, deeply buried in a rotting oak stump in the
Shawnee National Forest.
Distribution. — The eastern Ozark Plateau of
Arkansas and Missouri and the western Interior Low
Plateaus of southern Illinois (Figs. 54, 211). The
available samples come from four areas, and field
work is needed in the lacunae to determine if they
connect. Specimens were examined as follows:
ILLINOIS: Jackson Co., 1.5 mi. W Pomona,
along rd. to Pomona Nat. Bridge, M, 15 April 1984,
R. M. Shelley (NCSM). Johnson Co., Ferne Clyffe
St. Pk., 2M, 27 October 1967, J. A. Beatty (WAS).
MEM. AMER. ENT. SOC., 37
Johnson/Pope cos., 3 mi. E Ozark, M, 11 March
1950, P. W. Smith (FSCA). Pope Co., Dixon
Springs, 2M, 3 april 1948, Smith, Burks, and Stan-
nard (FSCA, ILNHS).
MISSOURI: St. Louis Co., Ranken, exact loca-
tion unknown, M, F, 2 April 1933 and 17 March
1935, E. P. Meiners (UMO). Phelps Co., 3 mi. NE
Rolla, M, 17 April 1965, G. L. Rotramel (ILNHS);
and 4 mi. S Jerome, Gable Springs Cv., M, 4 April
1979, J. E. Gardner (NCSM). Pulaski Co., 8 mi.
WSW Edgar Springs, Ryden Cv., M, 3 March 1979,
J. E. Gardner (NCSM).
ARKANSAS: Independence Co., 1.3 mi. W
Cushman, 2M, 2 juv. F, 5 March 1973, R. M.
Blaney (FSCA). Jackson Co., 1.5 mi. SW Oliphant,
M, F, 10 April 1936, L. Hubricht (NMNH) TYPE
LOCALITY. White Co., Russell, M, 25 December
1952, C. Pennington (FSCA).
Remarks. — The record of hispidipes from Dixon
Springs, Pope County, Illinois, by Causey (1950a),
subsequently cited by Chamberlin and Hoffman
(1958), actually refers to oliphantus. Since the types
of hispidipes are from an unknown site in Illinois
(Wood 1864, 1865) and oliphantus is the only species
known from forested areas in the southern part of
the state, it is frequently misidentified as the former.
For example, I originally assumed that my specimen
from Jackson County was hispidipes, but two
species occur in this state. The prior confusion is
understandable since no one has attempted to il-
lustrate the apertures of the hispidipes syntypes,
which have resided at the NMNH for a century and
a quarter. To my knowledge they have not been ex-
amined since 1865, and in particular Causey (1950a)
missed a golden opportunity to establish the identity
of hispidipes by examining and illustrating them. In-
stead, she mistakenly assumed that the male of
oliphantus from Dixon Springs (ILNHS) was
hispidipes, so named it, and illustrated the man-
dibular process, the 2nd coxa, the aperture, a
gonopod from lateral, not ventral, view, and the
sternal projections between the 9th legs, 8th by her
count. The aperture drawing from the lateral per-
spective does not show the pouches and is meaning-
less, so none of these drawings are definitive. Causey
(1952a) provided a ventral view of the aperture
showing the pouches and again, erroneously, labeled
it hispidipes. Subsequent confusion can be traced to
these mislabeled drawings, and when I examined the
types of hispidipes, I was shocked to view a different
form. I surmise that hispidipes is more common in
central Illinois, where there has been little recent col-
lecting, whereas oliphantus is abundant in the
58 EURYMERODESMID MILLIPEDS
Figs. 96-101. Eurymerodesmus elevatus. 96, projection of right mandible of holotype, lateral view. 97, aperture and gonopods in situ
of holotype, ventral view. 98, left gonopod of the same, lateral view. 99, telopodite of the same, medial view. 100, left cyphopod of
female paratype, caudal view. 101, the same, medial view. Scale line for fig. 97 = 1.00 mm. Line for other figs. = 0.30 mm for 96, 0.40
mm for 98 and 100-101, and 0.50 mm for 99.
ROWLAND M. SHELLEY 59
southern part of the state, in the mesic woodlands of
the Shawnee National Forest.
Eurymerodesmus oliphantus is one of three
species taken from caves, the others being melacis
and birdi. As indicated later, melacis may seek
refuge in caves, but o/iphantus is surely an accidental
species, since there is abundant, moist epigean
habitat in the Ozarks.
Eurymerodesmus oliphantus has the largest, most
open caudolateral pouches in the family, so much so
that they alone are diagnostic for the species.
Eurymerodesmus elevatus, new species
Figs. 54, 96-101
Type specimens. — Male holotype and female
paratype (FSCA) collected by W. W. Dowdy, 12
February 1966, on Neilhorn Place, Jefferson City,
Cole Co., MO. Male paratype (NCSM) taken by
same collector, 15 April 1965, on Lepage Place, Cole
Co., presumably in Jefferson City.
Diagnosis. — A small to moderate-size species;
males characterized by relatively short mandibular
projection; sides of aperture divided, outer lateral
margin rising beyond division point into cupped
elevated lobe forming false pouch on inner surface;
telopodite long; acropodite short, about 1/4 of the
telopodite length, broadly terminal, poorly demar-
cated from prefemur; latter with many hairs ar-
ranged continuously along inner margin; cyphopod
valves of female not modified, without projections;
receptacle moderate-size, sides not curving around
operculum.
Color in life. — Unknown.
Holotype. — Fragmented into three pieces, ap-
proximate length 19.2 mm, maximum width 3.2
mm, W/L ratio 16.7%.
Somatic features similar to hispidipes, with following excep-
tions:
Width across genal apices 2.3 mm, interantennal isthmus 0.8
mm. Antennae reaching back to middle of 3rd tergite, relative
lengths of antennomeres 2>3>5>6>4>1>7. Genae with faint
central impressions. Facial setae as follows: epicranial and in-
terantennal absent, subantennal 1-1, frontal 1-1, genal 1-1, clypeal
about 7-7, labral about 16-16. Process of mandibular stipes
relatively short, subtriangular, apically blunt (Fig. 96).
Collum not extending below ends of following tergite. Paranota
moderately depressed, continuing slope of dorsum; posterior cor-
ners rounded through segment 10, blunt on 13-15, becoming pro-
gressively more acute caudally.
5th sternum with elevated flattened areas between 4th legs and
larger paramedian knobs between 5th; 6th sternum with broad,
elevated areas between both leg pairs. Postgonopodal sterna with
slightly elevated, widely segregated hirsute areas between 9th legs;
remaining sterna generally flat and unmodified, with variably im-
pressed transverse grooves originating between leg pairs and hair
MEM. AMER. ENT. SOC., 37
patches adjacent to leg coxae, becoming progressively sparser
caudally. 2nd coxa with short, broadly triangular anterior, and
minute caudal, lobes. Prefemora with broad lobes on outer sur-
faces of legs through segment 10.
Gonopodal aperture (Fig. 97) broadly ovoid, with open
caudolateral pouches and short, broad anterior indentation, 1.6
mm wide and 0.7 mm long at maxima, outline of apparatus
subrectangular; anterior indentation short, broad, subacuminate;
sides divided, essentially upright, not leaning over opening, rising
continuously from anterolateral corner to peak on outer lateral
margin at caudolateral corner beyond division point, latter caudal
to midlength, inner lateral margin continuing curvature of anterior
part of sides, blending smoothly into inner caudal margin, outer
lateral margin rising continuously into broad, elevated lobe at
caudolateral corner, not flaring laterad, cupped with slight cavity
on inner surface in conjunction with outer caudal margin, rims of
anterior lateral margins slightly irregular, becoming more so on
elevation; inner caudolateral corners smoothly curved, indistinct,
continuous with inner lateral margin; outer caudolateral corners
relatively sharp, nearly a right angle; inner caudal margin smoothly
continuing curvature of sides, merging with outer caudal margin
well before midline, at about 1/3 length; latter dropping rapidly
from caudolateral corner to level of metazonum, angling inward
and merging with inner caudal margin, the two then continuing as
single, smoothly curved caudal margin across midline; anterior
margin and anterior halves of sides essentially glabrous, a single
hair arising from rim of latter, more hairs arising from rim of
outer caudal margin on elevation beyond division point, with addi-
tional short hairs on outer surface of elevation and moderate tuft
in pouch, caudal margin glabrous. Gonopods in situ (Fig. 97) with
telopodites approaching each other in midline then curving broadly
laterad across caudal margin, terminating over 9th coxal condyles,
apices directed sublaterad. Gonopod structure as follows (Figs.
98-99): Telopodite extending slightly beyond level of distal ex-
tremities of hairs. Prefemur long, about 3/4 of telopodite length,
bowed for most of length, more so distad, narrowing smoothly
and continuously, with two rows of continuous, evenly spaced
hairs along inner margin, angling outward distad, and distomedial
tuft of four hairs. Acropodite short and broadly terminal, about
1/4 of telopodite length, poorly demarcated from, and smoothly
continuous with, prefemur, smoothly continuing curvature of
distal part of latter basally, flattening at midlength, becoming un-
cinate distally, narrowing continuously to acuminate tip.
Female paratype. — Body fragmented, length approximately
25.3 mm, maximumn width 3.9mm, W/L ratio 15.4‘%. Agreeing
essentially with holotype in somatic features, with following excep-
tions: Mandibular stipes with minute, rounded, process. Stema
flat and nearly glabrous, with only a few scattered hairs.
Cyphopodal aperture subrectangular, sides slightly elevated
above metazonal surface, caudal margin with moderate indenta-
tion at midpoint. Cyphopods in situ with edges of valves and tip of
receptacle visible in aperture. Valves (Figs. 100-101) oriented
transversely in aperture, relatively small, subequal, without pro-
jections or modifications, lightly hirsute on medial side. Receptacle
located laterad to valves, relatively small, glabrous, sides not curv-
ing around operculum. Latter small, located under free, dorsal,
side of valves.
Male paratype. — The male paratype agrees with the holotype
in all particulars.
Variation. — The other Cole County males and
females agree closely with the holotype and
paratype, except that the gonopods in situ may cross
60 EURYMERODESMID MILLIPEDS
in the midline of the aperture and again over the
caudal margin, or lie over and under one another in
the midline. In the Morgan County male, the outer
lateral margin is more strongly elevated and flares
slightly laterad, thus forming a higher and slightly
deeper pouch.
Ecology. — According to the vial label, the
holotype and female paratype were collected ‘‘in
soil.’” The male from the flood plain in Cole County
was taken from the ‘‘first inch’’ presumably of the
soil or leaf litter. There are no indications whether
these sites were in forest, pasture, or other environ-
ments.
Distribution. — Known only from two counties
in central Missouri (Fig. 54), the distance between
known sites being only about 50 miles. Specimens
were examined as follows:
MISSOURI: Cole Co., Jefferson City, Neilhorn
Pl., M, F, 12 February 1966, W. W. Dowdy (FSCA)
TYPE LOCALITY; Lepage Pl., probably near Jef-
ferson City, M, 15 April 1965, W. W. Dowdy
(NCSM); flood plain without further data, M, 19
January 1968, W. W. Dowdy (FSCA); and along US
hwy 54 in southern Cole Co., M, 2F, 25 October
1965, W. W. Dowdy (FSCA). Morgan Co., Ver-
sailles, M, 12 April 1959, D. Zusteg (FSCA).
Remarks. — The specimens of elevatus were
either broken when I obtained them or fragmented
upon handling. All measurements are therefore ap-
proximations.
The cavity formed by the elevated outer lateral
and caudal margins, the cupped inner surface of the
elevation, is convergent with that of impurus in that
the inner lateral margin forms the floor instead of
the inner edge. It differs from the true pouches of
other species in the varius group, but the hair tuft in
the cavity and the uncinate acropodite, similar to
that of newtonus, indicate that elevatus properly
belongs here.
The Simplex Group
The simplex group includes forms with short
acropodites that are discontinuous with, and strong-
ly demarcated from, the prefemur in all populations.
It contains the fourth oldest species, simplex. The
acropodite may be subterminal, with the distal ex-
tremity of the prefemur overhanging and extending
beyond it, or narrowly terminal arising exclusively
from the inner prefemoral margin, the outer surface
of which projects above the junction and may be ex-
panded or clavate but does not extend distad beyond
the origin of the acropodite. Apertures vary, some
having divided lateral margins and pouches, as in
simplex, paroicus, and crassatus, others having un-
divided sides and triangular lobes on the caudal
margin as found in birdi, for example polkensis and
pulaski, and still others exhibiting undivided sides
without lobes, for example caesariatus and serratus.
The simplex group occurs sporadically from Florida
to Louisiana and western Arkansas. Most com-
ponents are widely segregated, and serratus even has
allopatric populations some 700 miles apart. Conse-
quently, a number of undiscovered forms probably
exist, particularly in southern Arkansas, northern
Louisiana, and western Mississippi.
As stated previously, varius, with discontinuous
acropodites in v. christianus, links the simplex and
varius groups into a single natural assemblage. For
convenience I separate them based on the degree of
differentation between the acropodite and prefemur,
and the relative number of populations displaying
this trait.
Components: simplex Chamberlin; paroicus
(Chamberlin); pulaski (Causey); polkensis (Causey);
caesariatus, new species; crassatus, new species; and
serratus, new species.
Eurymerodesmus simplex Chamberlin
Figs. 54, 102-107, 211
Eurymerodesmus simplex Chamberlin, 1920:98-99. Attems,
1938:187.
Kewanius simplex: Chamberlin and Hoffman, 1958:82.
Type specimen. — Male holotype (MCZ) and one
male and one female paratype (NMNH) collected by
H. E. Hubert on an unknown date in 1918 from an
unknown locality in Louisiana. The labels in the
vials and Chamberlin (1920) reported New Orleans
as the type locality, but Chamberlin and Hoffman
(1958) stated that the site was ‘“‘probably near”’ this
city and that no definite localities were known. Two
other eurymerodesmids, birdi planus and varius
christianus are known from New Orleans and en-
virons, but the only definite sites for simplex are in
Evangeline and Rapides Parishes, about 145 miles
north northwest of the city. Thus, the type locality is
unknown.
Diagnosis. — A moderate-size species; males
characterized by short mandibular projection; sides
of aperture divided, forming moderately open
pouches; telopodite long; acropodite short, about
1/4 of telopodite length, narrowly terminal, arising
solely from inner prefemoral margin; prefemur with
moderate number of hairs arranged continuously
along inner surface, expanding distad into broad,
ROWLAND M. SHELLEY 61
ee
in
)
104
Figs. 102-107. Eurymerodesmus simplex. 102, projection of
right mandible of holotype, lateral view. 103, aperture and
gonopods in situ of holotype, ventral view. 104, left gonopod of
holotype, lateral view. 105, the same, medial view. 106, left
cyphopod of female from Evangeline Par., LA, medial view.
107, the same, anterior view. Scale line for fig. 103 = 1.00 mm.
Line for other figs. = 0.30 mm for 102, and 0.50 mm for 104-107.
clavate shoulder on outer margin; cyphopod valves
of females with subterminal marginal spur on medial
valve, distal corners not produced; receptacle small,
sides not curving around operculum.
Color in life. — Unknown.
Holotype. — Body highly fragmented, length
unmeasurable, maximum width 4.2 mm.
Somatic features similar to hispidipes, with following excep-
tions:
Width across genal apices 2.2 mm, interantennal isthmus 0.9
mm. Antennae reaching back to mddle of 3rd tergite, relative
lengths of antennomeres 2>3>4>6>5>1>7. Genae without
impressions. Facial setae as follows: epicranial, interantennal, and
subantennal absent, frontal 1-1, genal 1-1, clypeal about 8-8, labral
about 14-14. Process of mandibular stipes short and broad, apical-
ly indented (Fig. 102).
Collum extending below ends of following tergite. Paranota
moderately flattened, interrupting slope of dorsum; posterior cor-
ners rounded through segment 6, blunt on 7-14, becoming pro-
gressively more acute beginning on segment 15.
5th sternum with low, rounded elevated areas between both leg
pairs; 6th sternum with broad elevated areas between anterior legs
and pair of long, conical, caudally directed projections between
caudal legs, narrowly segregated in midline, longer than widths of
adjacent coxae. Postgonopodal sterna with pair of broad knob-
like processes between 9th legs, narrowly segregated in midline,
apices angling anteriad, lengths subequal to widths of adjacent
coxae; remaining sterna with variably elevated areas between both
leg pairs on segments 8 and 9, caudal ones longer and tapering
mediad, sterna becoming flatter and more plate-like with in-
complete transverse impressions thereafter, with slight hair patches
MEM. AMER. ENT. SOC., 37
adjacent to coxae. 2nd coxa with broad anterior, and sub-
acuminate posterior, lobes. Prefemoral lobes continuing through
segment 9.
Gonopodal aperture (Fig. 103) subrectangular, with slight
caudolateral pouches and broad anterior indentation, 1.9 mm wide
and 0.8 mm long at maxima, outline of apparatus rectangular;
anterior indentation short, broadly rounded; sides divided, slightly
wider anteriad then narrowing, angling slightly mediad, and
elevating beyond division point to caudolateral corner, division
point located caudal to midlength, just before caudolateral corner,
inner lateral margin very short, heading directly into inner
caudolateral corner and then into inner caudal margin; outer
caudal margin rising slightly to outer caudolateral corner, forming
slight, open pouch with inner margin, rims slightly irregular; inner
caudolateral corner indistinct; outer caudolateral corner rounded
but sharp, nearly a right angle; inner caudal margin merging with
outer margin just beyond caudolateral corner; outer caudal margin
very short, extending from corner directly into inner margin, the
two then continuing in slight bisinuate curve across midline, flush
with metazonal surface; margins irregularly hirsute, with long hairs
arising from anterior margin, thickest at anterolateral corners,
with denser tufts of short hairs in pouches and a few short stiff
hairs from caudal margin. Gonopods in situ (Fig. 103) with
telopodites angling mediad and extending directly caudad, over-
lying each other in midline and projecting well beyond caudal
margin, apices diverging over sternum between 9 legs. Gonopod
structure as follows (Figs. 104-105): Telopodite terminating just
before level of distal extremities of hairs. Prefemur long, about 3/4
of telopodite length, leaning over coxa, expanding greatly beyond
midlength into broad, clavate shoulder on outer margin, with two
relatively sparse rows of continuous, but irregularly spaced hairs
on inner margin to level of shoulder, culminating in a slight tuft, a
distomedial tuft of about 12 hairs on shoulder at junction of
acropodite, and one additional proximal hair on medial surface.
Acropodite short and narrowly terminal, about 1/4 of telopodite
length, arising from inner prefemoral margin, discontinuous with,
and strongly demarcated from prefemur, much narrower than lat-
ter, bisinuate, apically blunt.
Female from Evangeline Par. — Approximate length 28.0 mm,
maximum width 3.5 mm, W/L ratio 12.5%. Agreeing closely with
holotype in somatic features, with following exceptions: Subanten-
nal setae 1-1, genal 4-4. Mandibular stipes with short process,
apically squared. Sterna relatively flat and unmodified, with
variable, incomplete transverse impressions on all segments.
Cyphopodal aperture elliptical, sides and caudal margin elevated
above metazonal surface. Cyphopods in situ with corners of valves
protruding through aperture, open side facing anteriad. Valves
(Figs. 106-107) moderately large and moderately hirsute, unequal,
lateral larger than medial, oriented dorsoventrally in aperture,
medial valve with subterminal marginal spur, distal corners not
produced. Receptacle relatively small, glabrous, located dorsad to
valves on caudal side, not curving partly around operculum. Latter
moderate-size, tucked under open (anterior) side of valves.
Variation. — The gonopods of all males are near-
ly identical to those of the holotype. The aperture
varies slightly in that it does not lean as far over the
Opening in some individuals, thus forming a
shallower, more open concavity. In the male from
Rapides Parish the caudal margin is distinctly
elevated across the midline. Hairs arise irregularly all
along the lateral rims and down the outer surfaces on
62 EURYMERODESMID MILLIPEDS
specimens from Evangeline and Rapides parishes.
The lobes between the 9th legs are similar to those on
the holotype in the Rapides Parish male and one
from Evangeline Parish. The other Evangeline male
possesses enormous subconical projections, much
longer than the widths of the adjacent coxae, and
they lean anteriad, thus displacing the apices of the
telopodites. One lies between the lobes, and the other
is pushed laterad over the coxa.
Ecology. — The Evangeline specimens were
found under logs on a mixed pine-hardwood hillside.
Distribution. — Known definitely from the
following two sites in central Louisiana (Figs. 54,
211).
LOUISIANA: Rapides Par., Forest Hill, M, 11
October 1945, R. L. Wenzel (FMNH). Evangeline
Par., O.25 mi. S Rapides Par. line near Cocodrie
Bayou, 2M, 3F, 1 February 1968, E. D. Keiser
(FSCA).
Remarks. — The males from Evangeline and
Rapides Parishes were tightly coiled and broke into
several pieces upon handling, rendering length
measurements impossible. The female measurement
is only an approximation. However, simplex is
generally a moderate-size species, smaller than goodi
and dubius, but considerably larger than varius.
In proposing Kewanius for simplex Chamberlin
(1938) did not officially form the new combination.
This was done 20 years later by Chamberlin and
Hoffman (1958), and remains the only formal publi-
cation of K. simplex.
This species is the fourth oldest in the genus, and it
has never been illustrated. Causey (1963) restudied
the types, claimed to find no differences warranting
a new genus, and synonymized Kewanius with
Eurymerodesmus, all without publishing a word of
description or any illustrations. When I first saw the
holotype I was surprised to see the discontinuous
acropodite and clavate prefemur, since none of the
previous works alluded to them. After analyzing
Eurymerodesmus as a whole, I agree that Kewanius
is a Synonym, but it is amazing that Causey (1963)
was so unimpressed by these characters that she did
not even consider them worth mention. The discon-
tinuity is so striking, that without studying all
eurymerodesmids, I think one would have to give
credence to Chamberlin’s generic name.
Eurymerodesmus polkensis (Causey),
new combination
Figs. 54, 108-115, 211
Paresmus polkensis Causey, 1952b:5, fig. 4. Chamberlin and
Hoffman, 1958:83.
Type specimen. — Male holotype (AMNH) col-
lected by N. B. Causey, 6 September 1950, 11 mi. N
Mena, Polk Co., AR.
Diagnosis. — A large species; males characterized
by large, broad mandibular projection; sides of
aperture not divided, without caudolateral pouches,
elevating continuously, extending around caudolat-
eral corners into moderately large, broadly rounded
lobes on caudal margin; telopodite long; acropodite
short, about 1/8 of telopodite length, narrowly ter-
minal, arising from inner prefemoral margin;
prefemur with many hairs arranged continuously
along inner surface, not expanding distad, without
detectable shoulder; females unknown.
Color in life. — Paranota red-orange; metaterga
dark olive, prozonites darker, with red-orange
“medial triangles’? (Causey 1952b). None of the
available specimens show evidence of color pattern,
but from Causey’s remarks, the species apparently
displays middorsal spots instead of stripes.
Holotype. — Body highly fragmented, length un-
measureable, maximum width 6.6 mm.
Somatic features similar to hispidipes, with following excep-
tions:
Width across genal apices 3.0 mm, interantennal isthmus 1.1
mm. Antennae reaching back to caudal margin of 3rd tergite,
relative lengths of antennomeres 2>5>6>3>4>1>7. Genae
with faint central impressions. Facial setae as follows: epicranial,
interantennal, subantennal, and genal absent, frontal 1-1, clypeal
about 12-12, labral about 18-18. Process of mandibular stipes
broad, heavily sclerotized, apically blunt (Fig. 108).
Collum extending slightly below ends of following tergite.
Paranota moderately depressed, mostly continuing slope of dor-
sum, peritremata flattened; posterior corners rounded through
segment 8, blunt on 9-14, becoming progressively more acute
caudally.
5th sternum with knob-like projections between both leg pairs,
those between caudal (Sth legs) larger and more widely segregated;
6th sternum with larger projections, those between caudal (7th)
legs more widely separated. Postgonopodal sterna with short,
knob-like processes between 9th legs; remaining sterna with lower
elevations on segments 8-9, becoming progressively flatter and
more sparsely hirsute caudally. 2nd coxa with strong subconical
anterodistal, and short rounded posterior, lobes. Prefemoral lobes
extending through segment 16.
Gonopodal aperture (Fig. 109, not this specimen) subrectangu-
lar, without caudolateral pouches, with strong anterior indenta-
tion, 2.7 mm wide and 1.0 mm long at maxima; anterior indenta-
tion long, triangular, acuminate; sides not divided, almost linear,
curving very slightly caudomediad, elevating smoothly and con-
tinuously around caudolateral corner onto caudal margin, leaning
only very slightly over opening, rims smooth; caudolateral corners
rounded, obtuse; caudal margin continuing elevation of sides
laterally, extending rapidly into moderately large, broadly rounded
lobes (Fig. 110), tapering rapidly to metazonal surface and ex-
tending caudad in midline; margins densely hirsute, with tufts
apically and along sides of anterior indentation, sporadic hairs
along anterior margin, becoming progressively denser caudally
along sides on rims and outer surfaces, densest on caudal lobes,
ROWLAND M. SHELLEY 63
narrowing into a row along caudal midline. Gonopods in situ (Fig.
114, not this specimen) with telopodites abutting or overlying each
other in midline of aperture, diverging, extending over caudal mar-
gin, and terminating over sternum between 9th legs. Gonopod
structure as follows (Figs. 111-113): Telopodite terminating before
level of distal extremities of hairs. Prefemur long, about 7/8 of
telopodite length, sublinear and of nearly uniform width for most
of length, narrowing distad and curving dorsad, with slight cavity
on apical margin, with two rows of continuous, regularly spaced
hairs along inner surface and distomedial tuft of about 10 hairs.
Acropodite short and narrowly terminal, about 1/8 of telopodite
length, arising from inner distal margin of prefemur, discon-
tinuous with, and sharply demarcated from latter, curving slightly
ventrad, apically blunt.
Variation. — The male from Scott County
measures 39.4 mm in length, 6.2 mm in width, W/L
12
Figs. 108-115. Eurymerodesmus polkensis.
ratio 15.7%. The lobes on the caudal margin of the
aperture appear slightly smaller than those of the
type, and the caudal margin extends farther caudad
in the midline. The left gonopod was displaced and
directed ventrad rather than caudad (Fig. 109). The
unbroken specimen from Montgomery County
measures 33.7 mm long, 5.4 mm wide, W/L ratio
16.0%. The lateral margins of the aperture are more
strongly elevated, and the lobes are distinctly higher
but markedly less hirsute. On the gonopods the
prefemur is bent more strongly distad, and the
acropodite extends around the distal extremity of the
prefemur and is expanded and cupped to form a
definite terminal calyx (Fig. 115) with irregular
margin.
108, projection of right mandible of holotype, lateral view. 109, aperture and gonopods
in situ of male from Scott Co., AR, ventral view. 110, the same, lateral view. 111, left gonopod of holotype, lateral view. 112, telopodite
of the same, medial view. 113, apex of the same, ventral view. 114, aperture and gonopods in situ of male from Montgomery Co., AR,
ventral view. 115, apex of telopodite of left gonopod of the same, lateral view. Scale lines for figs. 109-110 and 114 = 1.00 mm. Line for
other figs. = 0.30 mm for 108 and 111, 0.40 mm for 112, and 0.50 mm for 113 and 115.
MEM. AMER. ENT. SOC., 37
64 EURYMERODESMID MILLIPEDS
Ecology. — Notations on the vial labels indicate
that the male from Scott County was taken on a
wooded hillside; those from Montgomery County
were discovered in a ‘‘pig dung trap.”’
Distribution. — Known only from three adjacent
counties in western Arkansas (Figs. 54, 211).
Specimens were examined as follows:
ARKANSAS: Scott Co., 4 mi. W Y City, M, 24
October 1963, L. Hubricht (RLH). Polk Co., 11 mi.
N Mena, M, 6 September 1950, N. B. Causey
(AMNH) TYPE LOCALITY. Montgomery Co.,
Muddy Cr. Wildlife Management Area, along US
hwy. 270, 5.2 mi. W Pencil Bluff, 2M, 6-8 March
1977, R. E. Woodruff (FSCA).
Remarks. — Eurymerodesmus polkensis is placed
in the Kewanius lineage, simplex group, because of
the discontinuous prefemora, but it could equally
well go in the birdi lineage because of the undivided
aperture sides, the absence of pouches, and the
distinct lobes on the caudal margin. By com-
bining the aperture of birdi with the simplex/-
varius gonopod, polkensis proves that the
undivided /lobed-divided/pouch aperture extremes
are congeneric.
Eurymerodesmus caesariatus, new species
Figs. 54, 116-121
Type specimens. — Male holotype and one
female and one juvenile paratype (FSCA) collected
by N. B. Causey, 23 December 1953, 7 mi. N Mead-
ville, Franklin Co., MS.
Diagnosis. — A moderate-size species; males
characterized by moderately long mandibular pro-
jection; sides of aperture not divided, without
caudolateral pouches, elevating slightly, continuing
around caudolateral corner, and tapering rapidly to
metazonal surface a short distance onto caudal
margin; telopodite short, extending directly ventrad
not caudad, wholly enclosed within aperture;
acropodite short, about 1/4 of telopodite length,
arising solely from inner prefemoral margin;
prefemur not expanding distad, without detectable
shoulder, with many hairs arranged continuously
along inner surface and dense terminal corona lying
parallel to acropodite; cyphopod valves of females
moderately large, distal corners slightly elongate;
receptacle narrow, sides not curving around opercu-
lum.
Color in life. — Unknown. The blanched types
lack any trace of either color or pattern.
Holotype. — Length 27.0 mm, maximum width
4.2 mm, W/L ratio 15.6%.
Figs. 116-121. Eurymerodesmus caesariatus. 116, projection
of right mandible of holotype, lateral view. 117, aperture and
gonopods in situ of holotype, ventral view. 118, left gonopod of
holotype, lateral view. 119, telopodite of the same, medial view.
120, left cyphopod of female paratype, medial view. 121, the same,
anterior view. Scale line for fig. 117 = 1.00 mm. Line for other
figs. = 0.30 mm for 116, 0.40 mm for 118 and 120-121, and 0.50
mm for 119.
Somatic features similar to hispidipes, with following excep-
tions:
Width across genal apices 2.4 mm, interantennal isthmus 0.6
mm. Antennae reaching back to just beyond caudal margin of 2nd
tergite, relative lengths of antennomeres 2>3>4>5=6>1>7.
Genae without impressions. Facial setae as follows: epicranial, in-
terantennal, frontal, and genal absent, subantennal 1-1, clypeal
about 8-8, labral about 14-14. Process of mandibular stipes
moderately long, apically blunt (Fig. 116).
Collum not extending below ends of following tergite. Paranota
moderately depressed, continuing slope of dorsum, peritremata
flattened; posterior corners rounded through segment 10, blunt on
11-14, becoming progressively more acute caudally.
5th sternum with short, subequal, knob-like projections between
leg pairs, those between caudal (Sth) legs more widely separated;
6th sternum with similar knobs between anterior legs and broad,
flattened, elevated areas between caudal legs. Postgonopodal ster-
na with low, densely hirsute, knob-like processes between 9th legs;
remaining sterna with hirsute elevated areas on segments 8-10,
becoming progressively flatter, more plate-like, and less hirsute
caudally, caudalmost sterna glabrous or nearly so. 2nd coxa with
short, conical anterior, and minute rounded caudal, lobes.
Prefemoral lobes extending through legs on segment 10.
Gonopodal aperture (Fig. 117) broadly trapezoidal, without
caudolateral pouches and anterior indentation, 1.5 mm wide and
0.8 mm long at maxima; anterior margin without trace of indenta-
tion; sides not divided, angling caudolaterad, elevating impercep-
ROWLAND M. SHELLEY 65
tibly to caudolateral corner, not leaning over opening, rims
smooth; caudolateral corner broadly rounded; caudal margin con-
tinuing elevation of sides laterally, tapering rapidly to metazonal
surface, extending slightly caudad in midline; margins almost
glabrous, with only very few hairs on caudolateral corners.
Gonopods in situ (Fig. 117) with telopodites extending nearly
directly ventrad, in parallel arrangement, completely enclosed by
Opening, oriented vertically and perpendicular to body axis, not
extending caudad. Gonopod structure as follows (Figs. 118-119):
Telopodite relatively short, terminating well before level of distal
extremities of hairs. Prefemur long, about 3/4 of telopodite
length, configuration linear, widening beyond midlength then nar-
rowing distad, with two dense rows of continuous, regularly
spaced hairs along inner margin, additional hairs between rows,
and dense terminal marginal tuft of 16 or so very broad hairs lying
parallel to and obscuring acropodite, and forming corona.
Acropodite short and narrowly terminal, about 1/4 of telopodite
length, obscured by terminal prefemoral hairs, arising from inner
margin of distal extremity of prefemur, discontinuous with, and
sharply demarcated from, latter, curving slightly ventrad, apically
rounded.
Female paratype. — Body highly fragmented, length un-
measurable, maximum width 3.9 mm. Somatic features agreeing
essentially with holotype, with following exceptions: Process of
mandibular stipes minute, rounded, barely projecting from stipital
surface. 3rd sternum with two minute, hirsute lobes, remaining
sterna flat and essentially glabrous.
Cyphopodal aperture subrectangular, sides elevated above
metazonal surface. Cyphopods in situ with corners of valves pro-
truding through aperture, valves oriented along body axis. Valves
(Figs. 120-121) relatively large, subequal, distal corners slightly
prolonged and rounded, moderately hirsute. Receptacle narrow,
located on caudal surface of valves, closely adhering to latter, sides
not curving partly around operculum. Latter comparatively large,
clearly visible under free, anterodorsal corner of valves.
Ecology. — Unknown.
Distribution. — Known only from the type locali-
ty (Fig. 54).
Remarks. — The terminal prefemoral hairs are
packed so tightly that they superficially resemble a
sclerotized lamella; at first glance there appeared to
be a terminal clayx. They form a semicircular corona
and I think are homologous to the distomedial tuft
on other species, which is situated more on the
medial surface a short distance from the end of the
prefemur.
In general configuration, the telopodite of
caesariatus is closest to that of po/kensis. However,
caesariatus occurs some 280 miles southeast of
polkensis, so the intervening part of Arkansas and
Louisiana probably contains other forms with nar-
rowly terminal acropodites.
Eurymerodesmus paroicus (Chamberlin),
new combination
Figs. 54, 122-125, 211
Paresmus paroicus Chamberlin, 1942a:7, pl. 2, figs. 18-19. Cham-
berlin and Hoffman, 1958:82.
MEM. AMER. ENT. SOC., 37
Type specimen. — Male neotype and one addi-
tional male (FSCA) collected by W. J. Harmon,
December 1954, 900 S. Farmerville St., Ruston, Lin-
coln Par., LA. This site is around 5-6 miles north of
the type locality, 1.5 mi. N Clay, at the border of
Lincoln and Jackson Parishes. The type vial, con-
taining the original label, is at the NMNH, but it
contains four males of Pseudopolydesmus pine-
torum (Bollman) and not even a loose eury-
merodesmid segment. Thus the type series, con-
taining the holotype and an unknown number of
paratypes, is lost.
Diagnosis. — A moderate to large species; males
characterized by short mandibular projection; sides
of aperture divided, forming shallow open pouches;
telopodite long; acropodite short, about 1/8 as long
as prefemur arising subterminally from inner
prefemoral margin; prefemur very long and very
broad, comprising entire length of telopodite, with
rounded, glabrous, distal lobe overhanging and ex-
tending beyond level of acropodite and with modest
number of hairs arranged continuously along inner
margin to near base of acropodite, inner surface
smooth, not excavated; females unknown.
Color in life. — Unknown.
Neotype. — Body highly fragmented, length un-
measurable, maximum width 4.0 mm.
Somatic features similar to hispidipes, with following excep-
tions:
124
Figs. 122-125. Eurymerodesmus paroicus. 122, projection of
right mandible of neotype, lateral view. 123, aperture and
gonopods in situ of neotype, ventral view. 124, left gonopod of the
same, lateral view. 125, telopodite of the same, medial view. Scale
line for fig. 123 = 1.00 mm. Line for other figs. = 0.13 mm for
122, 0.40 mm for 124, and 0.50 mm for 125.
66 EURYMERODESMID MILLIPEDS
Width across genal apices 2.6 mm, interantennal isthmus 0.9
mm. Antennae reaching back to caudal margin of 3rd tergite,
relative lengths of antennomeres 2>4>3>5>6>1>7. Genae
with faint central impressions. Facial setae as follows: epicranial
1-1, interantennal and frontal absent, subantennal 1-1, genal 4-4,
clypeal about 10-10, labral about 13-13. Mandibular stipes with
short, narrowly rounded projection (Fig. 122).
Collum not extending below ends of following tergite. Paranota
moderately flattened, interrupting slope of dorsum; posterior
corners blunt on anterior segments, becoming progressively pro-
longed and acute posteriorly.
Sternum of segment 5 with broad, flattened, elevated areas
between both pairs of coxae, caudal elevations (between Sth legs)
higher and more segregated in midline; 6th sternum with elevations
stronger, more rounded, and more densely hirsute, caudal ones
(between 7th legs) higher. Postgonopodal sterna with low, widely
segregated lobes between 9th legs; remaining sterna becoming pro-
gressively flatter and more plate-like posteriorly, with slight hirsute
lobes adjacent to coxae on segments 8 and 9, tapering into hirsute
patches on 10-11, becoming less dense caudally, with variably
impressed transverse grooves originating between leg pairs. 2nd
coxa with strong, conical anterior, and shorter rounded posterior,
lobes. Prefemoral lobes extending through anterior legs of segment
12.
Gonopodal aperture (Fig. 123) broadly rectangular, with nar-
row, open caudolateral pouches and strong anterior indentation,
2.8 mm wide and 1.7 mm long at maxima, outline of apparatus
subtrapezoidal; anterior indentation long, triangular apically
acute; sides divided, curving slightly mediad, rising to division
point caudal to midlength and leaning imperceptibly over opening,
inner lateral margin plunging directly dorsad into opening then
curving broadly into inner caudolateral corner, outer lateral
margin upright, well separated from inner, flaring caudolaterad
and dropping slightly to caudolateral corner, forming shallow
open pouch with inner margin, rims slightly irregular; inner
caudolateral corner broad, rounded, visible deep inside opening;
outer caudolateral corner broadly rounded but approximately 90°;
inner caudal margin curving smoothly across midline well inside
opening, generally invisible; outer caudal margin dropping slightly
from caudolateral corner to metazonal surface, running along
anterior surfaces of coxal condyles, extending slightly caudad in
midline; margins moderately hirsute with long hairs arising
sporadically from rims and overhanging opening, slightly denser
on anterior indentation, much denser at division points on sides,
with relatively sparse growths from pouches evenly dispersed;
caudal margin with a few widely separated hairs. Gonopods in situ
(Fig. 123) with telopodites diverging and angling caudolaterad over
inner caudal margin of aperture, acropodites directed dorsad.
Gonopod structure as follows (Figs. 124-125): Telopodite termina-
ting at level of distal extremities of hairs. Prefemur very long and
broad, comprising entire length of telopodite, distal third glabrous,
curving gently and leaning over coxa, curve increasing distad, ter-
minating in broadly rounded lobe overhanging and extending
beyond level of acropodite, of nearly subequal width throughout,
narrowing slightly distad, with ridge along inner surface demar-
cating prostatic groove, with two rows of subcontinuous hairs
along inner margin terminating proximal to origin of acropodite,
distolateral tuft of 3 hairs, dense distomedial tuft of about 9-12
hairs, and 2 long proximal hairs on medial surface. Acropodite
short, about 1/8 as long as prefemur, profile subtriangular, arising
subterminally on inner surface of prefemur, sides narrowing rapid-
ly to subacuminate tip, directed dorsad.
Variation. — All males of paroicus are frag-
mented beyond the point where they can be re-
assembled for measurements. The only noticeable
variation is slight differences in the alignment of the
lateral aperture margins. They are linear in the
neotype, angled slightly caudolaterad in another
male, and bowed slightly in the others. The last two
conditions result in slightly larger pouches.
Ecology. — According to Chamberlin (1942a),
the type series was encountered under logs.
Distribution. — Known only from _ Lincoln
Parish, Louisiana (Figs. 54, 211). Specimens were
examined as follows:
LOUISIANA: Lincoln Par., Ruston, 900 S.
Farmerville St., 2M, December 1954, W. J. Harmon
(FSCA) NEOTYPE; and unknown location in
Ruston, 2M, October and November 1951, collec-
tors unknown (FSCA).
Remarks. — The gonopod of paroicus is one step —
removed from the condition in po/lkensis. If the apex
of the latter’s prefemur were swollen and extended
slightly beyond the acropodite, the two would be
practically interchangeable except for the slightly
greater thickness of paroicus. However, the aper-
tures are entirely different. That of paroicus has
divided lateral margins and caudolateral pouches,
whereas that of polkensis displays undivided sides
and caudal lobes.
Eurymerodesmus crassatus, new species
Figs. 54, 126-132, 211
Type specimens. — Male holotype and 6 male
and 2 female paratypes (FSCA) collected by L. D.
Wilson, 24 March 1966, off old US highway 165, 2.6
mi. S Georgetown, Grant Par., LA. One male and
one female paratype deposited in NCSM.
Diagnosis. — A moderate to large species; males
characterized by short mandibular projection; sides
of aperture divided, with open caudolateral pouches;
telopodite long; acropodite minute, no more than
1/10 as long as prefemur, nubbinlike, arising sub-
terminally from inner prefemoral margin; prefemur
very long and very broad, comprising entire length
of telopodite, with broad distal lobe overhanging
and extending well beyond level of acropodite, with
modest number of hairs arranged continuously
along inner margin to near base of acropodite, inner
surface emarginate; cyphopod valves of females with
bowed projections from distal corners, apices
touching or nearly so; receptacle moderate-size, sides
curved partly around operculum.
ROWLAND M. SHELLEY 67
130
Figs. 126-132. Eurymerodesmus crassatus. 126, projection of
right mandible of holotype, lateral view. 127, aperture and
gonopods in situ of paratype, ventral view. 128, left gonopod of
the same, lateral view. 129, telopodite of the same, medial view.
130, cyphopods in situ, ventral view of female paratype. 131, left
cyphopod of the same, caudal view. 132, the same, medial view.
Scale lines for figs. 127 and 130 = 1.00 mm. Line for other figs. =
0.35 mm for 126 and 131, 0.40 mm for 128, and 0.50 mm for 129
and 132.
Color in life. — Unknown. The specimens are
uniformly tawny after 23 years in alcohol and exhibit
no trace of either color or pattern.
Holotype. — Length 25.0 mm, maximum width
4.1 mm, W/L ratio 16.4%.
Somatic features similar to hispidipes, with following excep-
tions:
Width across genal apices 2.7 mm, interantennal isthmus 0.8
mm. Antennae reaching back to middle of 3rd tergite, relative
lengths of antennomeres 2>3>5>4>6>1>7. Genae with faint
central impressions. Facial setae as follows: epicranial and in-
terantennal absent, subantennal 1-1, frontal 1-1, genal 2-2, clypeal
about 9-9, labral about 16-16. Process of mandibular stipes short,
broadly rounded (Fig. 126).
Collum extending slightly below ends of following tergite.
Paranota moderately depressed, somewhat interrupting slope of
dorsum, peritremata flattened; posterior corners rounded through
segment 3, blunt on 4-10, becoming progressively more acute
caudally.
5th sternum with narrowly rounded, slightly elevated areas be-
tween both coxae; 6th sternum with broader, higher, flattened
areas between coxae. Postgonopodal sterna with low, rounded
elevations between 9th coxae; remaining sterna becoming pro-
gressively flatter and more plate-like caudally, with sparsely hirsute
patches adjacent to coxae. 2nd coxa with moderately large, broad-
MEM. AMER. ENT. SOC., 37
ly rounded anterior lobe and slightly shorter, subacute caudal one.
Prefemoral lobes present on all but ultimate two leg pairs.
Gonopodal aperture (Fig. 127, not this specimen) broadly
ovoid, with open caudolateral pouches and strong anterior inden-
tation, 2.4 mm wide and 1.1 mm long at maxima, outline of ap-
paratus broadly trapezoidal; anterior indentation long, triangular;
sides curving gently mediad and leaning progressively over open-
ing, rising to division point caudal to midlength, inner lateral
margin exposed, extending nearly directly dorsad, curving strongly
into inner caudolateral corner, outer lateral margin curling sharply
dorsolaterad, not obscuring inner margin, dropping to outer
caudolateral corner, forming shallow, open pouch with inner
margin; inner caudolateral corner comparatively sharp, approx-
imately a right angle; outer caudolateral corner broadly rounded;
inner caudal margin flush with metazonal surface, extending
slightly inward into opening; outer caudal margin dropping to
metazonal surface shortly after outer caudolateral corner, extend-
ing caudad in midline; margins moderately hirsute, with tufts of 8
or so hairs on sides of anterior indentation, regularly spaced hairs
along rest of anterior margin and sides to division point, moderate
tufts in pouches, and scattered hairs between caudal margins.
Gonopods in situ (Fig. 127, not this specimen) with telopodites
widely separated and diverging, extending caudolaterad over inner
caudal margin, apices curving sublaterad. Gonopod structure as
follows (Figs. 128-129): Telopodite essentially terminating at level
of distal extremities of hairs. Prefemur very long, comprising en-
tire length of telopodite, distal third glabrous, extending linearly
for most of length, curving broadly distad and overhanging and
extending well beyond level of acropodite in broadly rounded ter-
minal lobe, very broad basally, widening into broad shoulder on
medial surface near midlength, narrowing thereafter and tapering
to broadly rounded tip, inner surface emarginate, with row of sub-
continuous hairs on inner surface extending to just beyond
shoulders and dense distal tufts of 15 or so hairs on lateral and
medial surfaces of shoulders, latter located slightly distal to
former. Acropodite minute and indistinct, a short, nubbinlike ex-
tension of inner surface of prefemur near base of distal curvature,
no more than 1/10 as long as prefemur.
Female paratype. — Length 23.2 mm, maximum width 4.0
mm, W/L ratio 17.2%. Agreeing closely with males in structural
details, with following exceptions: Process of mandibular stipes
short, subacute.
Cyphopodal aperture broadly ovoid, sides strongly elevated above
metazonal surface. Cyphopods in situ with edges of valves and
projections visible in aperture (Fig. 130). Valves (Figs. 131-132)
large, subequal, oriented transversely in aperture, leaning over
receptacle, distal corners extending into broad, digitiform lobes,
curving towards each other with apices touching or nearly so, with
scattered hairs on projections. Receptacle relatively large,
glabrous, located directly beneath valves, sides cupped partly
around operculum. Latter small, hidden beneath, and obscured
by, valves.
Male paratypes. — The male paratypes agree with the holotype
in all particulars.
Ecology. — The types were taken at an old mill
site, but the habitat is not indicated on the vial label.
Distribution. — Known only from the type locali-
ty.
Remarks. — This species is similar to paroicus,
and future collecting in the gap of around 40 miles
may show them to connect and be geographic races.
68 EURYMERODESMID MILLIPEDS
The apertures are very close, and the major dif-
ferences involve the much shorter acropodite and ir-
regular inner margin of crassatus.
The female is suspiciously similar to those of
angularis in the valvular projections and may actual-
ly be referrable to this species, occuring only two
counties to the east. However, the position of the
valves directly above the receptacle (Fig. 131) is
unique, so it may be conspecific with the males.
Eurymerodesmus pulaski (Causey),
new combination
Figs. 54, 133-140, 211
Leptodesmus hispidipes: Bollman, 1888a:2, in part.
Paresmus pulaski Causey, 1950a:271-272, fig. 9; 1952b:4-5.
Chamberlin and Hoffman, 1958:83.
Type specimen. — Male neotype, two additional
males, and one female (NMNH) collected by C. H.
Bollman, on an unknown date in the 1880’s before
1888, at Little Rock, Pulaski Co., AR. The type
locality, 3 mi. S Sweet Home, is on the southern
periphery of Little Rock, and the type series con-
sisted of two males collected in 1949. The holotype,
reportedly at the ANSP (Causey 1950a, Chamberlin
and Hoffman 1958), and the paratype, retained in
Causey’s private collection (Causey 1950a), now at
the FSCA, are lost.
Diagnosis. — A moderate to large species; males
characterized by long mandibular projection; sides
of aperture not divided, without caudolateral
pouches, elevating slightly, curving around caudo-
lateral corners, with suggestion of lobe on caudal
margin; telopodite long, acropodite minute, less
than 1/10 as long as prefemur, a blunt spur arising
subterminally from inner prefemoral margin and
situated in cavity in latter; prefemur very long and
broad, comprising entire length of telopodite, ex-
panding at level of acropodite into hood overhang-
ing and shielding latter, broadly sublinear or distally
uncinate, with cavity on inner margin and many
hairs arranged continuously along inner surface;
cyphopod valves of females not modified, distal cor-
ners not extended; receptacle moderately large, sides
expanding but not curving around operculum.
Color in life. — Paranota orange; metaterga dark
olive, prozonites darker, with orange stripes along
caudal margins connecting paranotal markings, ex-
panding slightly mediad; collum with stripes on
anterior and posterior margins (Causey 1950a).
Neotype. — Body highly fragmented, length un-
measurable, maximum width 5.2 mm.
Figs. 133-140. Eurymerodesmus pulaski. 133, projection of
right mandible of neotype, lateral view. 134, aperture and
gonopods in situ of neotype, ventral view. 135, left gonopod of the
same, lateral view. 136, telopodite of the same, medial view. 137,
aperture and gonopods in situ of male from Pulaski Co., AR, ven-
tral view. 138, apex of telopodite of left gonopod of the same,
medial view. 139, left cyphopod of female from Pulaski Co.,
lateral view. 140, the same, medial view. Scale lines for figs. 134
and 137 = 1.00 mm. Line for other figs. = 0.25 mm for 133, 0.37
mm for 135, 0.40 mm for 139-140, and 0.50 mm for 136 and 138.
Somatic features similar to hispidipes, with following excep-
tions:
Width across genal apices 3.2 mm, interantennal isthmus 0.8
mm. Antennae reaching back to middle of 3rd tergite, relative
length of antennomeres 2>3>4>6>5>1>7. Genae with faint
central impressions. Facial setae as follows: epicranial, inter-
antennal, subantennal, frontal, and genal absent, clypeal about
10-10, labral about 15-15. Process of mandibular stipes long, sub-
conical, tapering to blunt tip (Fig. 133).
Collum extending slightly below ends of following tergite.
Paranota moderately depressed, mostly continuing slope of dor-
sum, peritremata flattened; posterior corners rounded through seg-
ment 5, blunt on 6-11, becoming progressively more acute
posteriorly. 2nd coxa with moderate-size subconical anterior lobe,
caudal lobe only slightly smaller. Prefemoral lobes extending
through segment 10.
Gonopodal aperture (Fig. 134) broadly ovoid, without
caudolateral pouches, with strong anterior indentation, 2.7 mm
wide and 1.0 mm long at maxima; anterior indentation long,
triangular, acute; sides not divided, curving gently mediad, rising
very slightly to caudolateral corner, not leaning over opening, rims
ROWLAND M. SHELLEY 69
smooth; caudolateral corner broadly rounded, slightly elevated,
forming a slightly obtuse angle; caudal margin rising slightly into
suggestion of lobe then tapering to metazonal surface, extending
caudad in midline; anterior margin glabrous, hairs beginning at
midlength of sides and continuing onto caudal margin, regularly
spaced. Gonopods in situ (Fig. 134) with telopodites extending
caudad in parallel arrangement, extending well beyond aperture,
overhanging sternum between 9th legs, apices directed dorsad.
Gonopod structure as follows (Figs. 135-136): Telopodite termi-
nating near level of distal extremities of hairs. Prefemur long, com-
prising entire length of telopodite, configuration slightly bisinuate,
bending strongly dorsad distally, narrow basally, expanding
beyond midlength into concave hood shielding acropodite,
narrowing distad, apically blunt, with cavity on inner surface of
hood, two rows of continuous, subregularly spaced hairs on inner
surface to near distal bend, tuft of around 10 hairs on inner surface
just before distal bend, distomedial tuft of around 10 hairs on
hood, and 4 or so scattered hairs proximally on medial margin.
Acropodite minute, a blunt subtriangular spur located subterm-
inally in cavity on undersurface of hood at level of distal bend,
partly shielded by sides of hood.
Female from Pulaski Co. — Body highly fragmented, length
unmeasurable, maximum width about 4.5 mm. Process of mandi-
bular stipes short, truncate. Other somatic features unknown
because of great fragmentation and mixture of male and female
segments in sample; impossible to reliably segregate segments by
sex.
Cyphopodal aperture ovoid, sides flush with metazonal surface.
Cyphopods in situ with edges of valves visible in aperture. Valves
(Figs. 139-140) moderate-size, subequal, oriented transversely in
aperture, medial (open) side densely hirsute, corners not extended.
Receptacle moderately large, glabrous, located laterad at bases of
valves, sides expanding beyond latter but not cupped around oper-
culum. Latter small, located under free (open) end of valves.
Variation. — The neotype was chosen because it
is the only available male matching Causey’s illustra-
tion (1950a, fig. 9) of the holotype; it may have come
from the same general area in south Little Rock. The
NMNH sample contains fragments of two different
males that may occur sympatrically in Little Rock.
One exhibits sufficient gonopodal similarities to the
neotype to be considered conspecific, but the other is
so different as to require another binomial, and is
therefore described in the ensuing account. The
female in this sample may well be assignable to the
following species, the other samples of which con-
tain only males. There being no way this can be
determined at present, I arbitrarily assign it to
pulaski.
The acropodite of the variant is rudimentary, a
thickened boss barely elevated above the prefemoral
surface (Fig. 138). The cavity on the undersurface of
the prefemur is very deep; the hood curves well over
the acropodite as in the neotype; and the tip curls
mediad to a subacuminate tip. The aperture con-
figuration is the same, but the elevations on the
caudal margin are higher, more distinctly lobe-like,
and more densely hirsute (Fig. 137).
MEM. AMER. ENT. SOC., 37
Ecology. — The type series was taken in an oak-
pine association (Causey 1950a).
Distribution. — Known only from Pulaski Coun-
ty, Arkansas (Fig. 54, 211). The neotype and the
associated males are the only available specimens.
Remarks. — Because of the high fragmentation
of the Bollman and Hutcherson samples, it is im-
possible to determine which segment is which and
thus to report the conditions of the sterna.
The neotype is one of Bollman’s original samples
and the label is in his handwriting. Consequently,
this is probably one of the species he (1888a) iden-
tified as “‘Leptodesmus hispidipes’’ and stated was
very common throughout Arkansas.
Pulaski County, and the area around Little Rock
specifically, need to be meticulously sampled, since
at least three forms occur there. At this writing I can-
not determine if they are sympatric or parapatric;
Bollman could well have combined samples from
distant sites in the county under the general heading
of Little Rock.
Eurymerodesmus serratus, new species
Figs. 54, 141-146, 211
Leptodesmus hispidipes: Bollman, 1888a:2, in part.
Type specimen. — Male holotype (FSCA) col-
lected by ‘‘Coffee,’’ 11 October 1955, at Cato,
Pulaski Co., AR. Male paratype (FSCA) collected
G. B. Edwards, 4 November 1978, ca. 10 mi. SE
Gainesville, along FL hwy. 325, 4 mi. S jct. FL hwy.
20, Alachua Co., FL.
Diagnosis. — A moderate to large species; males
characterized by long mandibular projection; sides
of aperture not divided, without caudolateral
pouches, rising gradually to short lobe at caudo-
lateral corners, caudal margin descending rapidly to
metazonal surface; telopodite long; acropodite
minute, less than 1/10 as long as pefemur, arising
subterminally from inner prefemoral margin at base
of expansion, situated in cavity in latter; prefemur
very long, comprising entire length of telopodite, ex-
panding greatly at level of acropodite into short,
broad hood overhanging and shielding latter,
margins of hood irregularly serrate to jagged, with
cavity on inner margin and many hairs arranged
continuously along inner surface; females unknown.
Color in life. — Unknown. Though faded, the
Florida male shows evidence of stripes along the
caudal metatergal margins connecting with the
paranotal markings and expanding in the midline;
the collum is striped along both margins.
70 EURYMERODESMID MILLIPEDS
Holotype. — Body highly fragmented, length un-
measurable, maximum width 5.1 mm.
Somatic features similar to hispidipes, with following exceptions:
Head partly decomposed, facial characters obtained from
paratype.
Collum not extending below ends of following tergite. Paranota
moderately depressed, partly continuing slope of dorsum;
posterior corners rounded through segment 10, blunt on 11-12,
becoming progressively more acute caudally.
5th sternum with short, narrowly segregated, knob-like pro-
cesses between anterior (4th) legs and broader, higher, widely
separated ones between 6th legs; Sth sternum with subsimilar
knob-like projections between both leg pairs, those between caudal
(7th) legs slightly higher. Postgonopodal sterna with strong, sub-
conical, widely separated lobes subtending coxae of 9th legs, only
slightly shorter than widths of latter; remaining sterna with short,
broad lobes between both legs on segment 8, becoming progres-
sively flatter posteriorly, with modest hair patches beside legs on all
segments. 2nd coxa with strong subconical anterior, and slightly
shorter posterior lobes. Prefemoral lobes extending through seg-
ment 15.
Gonopodal aperture (Fig. 142) ovoid, without caudolateral
pouches, with strong anterior indentation, 2.1 mm wide and 1.1
mm long at maxima; anterior indentation relatively long,
triangular, apically acute; sides not divided, angling slightly
mediad, rising gradually and continuously to short lobe at
caudolateral corner, rims smooth; caudolateral corner broad, in-
distinct; caudal margin continuous with lobe at caudolateral cor-
ner, dropping rapidly a short distance from latter to metazonal
Figs. 141-146. Eurymerodesmus serratus. 141, projection of
right mandible of paratype, lateral view. 142, aperture and
gonopods in situ of holotype, ventral view. 143, left gonopod of
the same, lateral view. 144, telopodite of the same, medial view.
145, the same, dorsal view. 146, aperture and gonopods in situ of
male from Alachua Co., FL, ventral view. Scale lines for figs. 142
and 146 = 1.00 mm. Line for other figs. = 0.25 mm for 141, 0.30
mm for 143, and 0.50 mm for 144-145.
surface, curving across midline and extending slightly caudad;
margins irregularly hirsute with scattered hairs on anterior margin,
none on indentation, relatively dense hairs arising on outer sur-
faces of sides and extending rather continuously to end of elevation
on caudal margin, lobes without denser hairs. Gonopods in situ
(Fig. 142) with bases of telopodites touching or nearly so in
midline, diverging and extending well beyond caudal margin of
aperture, overhanging 9th coxae, terminating over caudal margin
of 7th segment, apices directed dorsad. Gonopod structure as
as follows (Figs. 142-145): Telopodite terminating at level of distal
extremities of hairs. Prefemur very long, comprising entire length
of telopodite, apically glabrous, configuration slightly curved with
lengthy linear section in middle, apex bending more strongly dor-
sad, narrow basally, expanding markedly at midlength, narrowing
slightly at distal bend, then expanding in perpendicular plane into
short, broad, marginally serrated hood, inner surface strongly con-
cave, with two rows of continuous, regularly spaced hairs on inner
and lateral margins extending to just short of distal bend and level
of acropodite, distomedial tuft of 6 or so hairs extending to mid-
length of hood, and one or two more proximal hairs medially.
Acropodite a short, blunt spur on inner surface of prefemur,
shielded ventrally but not surrounded by hood, directed outward
or dorsad.
Male paratype. — Fragmented, unmeasurable. Width across
genal apices 3.9 mm, interantennal isthmus 1.1 mm. Antennae
reaching back beyond caudal margin of 3rd tergite, relative lengths
of antennomeres 2>6>4>5>3>1>7. Genae without impres-
sions. Facial setae as follows: epicranial and interantennal absent,
subantennal 1-1, frontal 1-1, genal 2-2, clypeal about 11-11, labral
about 21-21. Process of mandibular stipes relatively long, angling
slightly caudad, apically rounded (Fig. 141).
The male paratype agrees closely with the holotype, which is
remarkable considering the 700 mile gap between them. The ster-
nal projections on segments 4 and 5 are perhaps slightly longer, but
otherwise there are no differences in somatic features. The aper-
ture is considerably less hirsute, but its configuration is very similar
to that of the holotype (Fig. 146). The elevations begin more sud-
denly a short distance back from the anterolateral corner, plateau,
and continue around the caudolateral corner into a less distinct
lobe, located more on the caudal margin. Aside from a slightly dif-
ferent degree of serration on the hood, the gonopods are virtually
identical.
Variation. — The male from Little Rock agrees
closely with the holotype in all particulars.
Ecology. — There are no indications of habitat
on the vial labels with the Arkansas males. That
from Florida was found on top of litter at the base of
a small water oak in a wooded transition zone be-
tween slash pine and water oak.
Distribution. — Known from two widely sepa-
rated sites, about 700 miles apart, in central Arkan-
sas and northern Florida (Fig. 54). It is unknown
whether these reflect truly allopatric populations or
if the hiatus contains numerous additional popula-
tions, such that the range is reasonably continuous.
Specimens were examined as follows:
ARKANSAS: Pulaski Co., Cato, just S of
Faulkner Co. line, M, 11 October 1955, Coffee
(FSCA) TYPE LOCALITY; and Little Rock, M,
1880’s, Hutcherson (NMNH).
ROWLAND M.
FLORIDA: Alachua Co., 10 mi. SE Gainesville,
along FL hwy. 325, 4 mi. S jct. FL hwy. 20, M, 4
November 1978, G. B. Edwards (FSCA).
Remarks. — The three males of this species are so
highly fragmented that length measurements would
be meaningless. However, the animals can be
characterized as ‘‘moderately-large,’’ i.e. smaller
than such species as goodi and dubius, but larger
than other intermediate-size species like hisipdipes.
The male from Little Rock is only a partial
specimen, consisting of segments 6-9, and was mixed
in the same vial with the neotype and two
fragmented male variants of pulaski. Since this sam-
ple was labeled ‘“‘Leptodesmus hispidipes’’ by
Bollman in his handwriting, serratus probably con-
stitutes part of the basis for his published record
(1888a). Included in this vial is part of a female,
which I characterized in the pulaski account but
which may be referrable to serratus. The bulk of the
NMNH sample consists of mixed segments that can-
not be segregated either by sex or species. The holo-
type and paratype were preserved intact, but they
were so rigid and tightly coiled that they shattered
when I tried to open them for examination, deter-
mination, and measurement.
The configuration of the telopodite of serratus is
remarkably similar to those of the xystodesmids
Sigmoria (Cheiropus) planca (Loomis) and S. (C.)
serrata Shelley (see Shelley 1984b, figs. 3-7, 19-22) in
being expanded distally into a marginally serrated
hood, partly shielding the structure carrying the
prostatic groove, the solenomerite in the species of
Sigmoria. This is a remarkable example of con-
vergence, and as in S. (C.) serrata the eury-
merodesmid hood is actually quite jagged. The
margin is thickened, particularly on the lateral side,
and in places there are up to three rows or layers of
sharp, tiny teeth that line the inner margin over-
hanging the cavity as well as the outer border. Seen
in profile from the proper perspective, the apex of
the telopodite of serratus closely resembles a mitten,
with the acropodite being the thumb and the hood
representing the other fingers combined, which was
the basis for Loomis’ name, planca. The jagged
nature of the distal margin is by far the dominant
feature, and except for the inner prefemoral margin
of crassatus, no other eurymerodesmid is even slight-
ly irregular. Smooth gonopodal margins are the rule
in this family.
The Birdi Lineage
The birdi lineage is characterized by continuous,
broadly terminal, and poorly demarcated acro-
MEM. AMER. ENT. SOC., 37
SHELLEY 71
Fig. 147.
half-shaded squares, birdi planus; dots, mundus. Open symbols
denote literature records considered reliable.
Distribution of the birdi lineage. Squares, birdi birdi;
podites; sides of apertures undivided, without
caudolateral pouches; and caudal margins either
with densely hirsute lobes or comparatively dense
hair patches in this location. Females cannot be
reliably distinguished from those in other groups. In
lobed forms the densest marginal hairs are on the
projections, so when the latter are absent, the
densest patches are usually at this general location,
suggesting that pilosity is the sole remaining rem-
nant. Forms with lobes also display variably elevated
lateral margins, which tend to be lower in ones lack-
ing projections; the lobes arise from the elevations,
and in general as one becomes smaller so does the
other. In populations of birdi planus in southeastern
Louisiana, there is only a low, continuous marginal
elevation extending around the caudolateral corner
without a trace of lobes, and in the northernmost
population, the aperture is strikingly unadorned with
only modest hair patches at the caudolateral corners.
The sides and caudal margin of this form are either
flush with the metazonum or so slightly elevated as
to appear flat in ventral view.
The birdi lineage contains two forms with strong
lobes, mundus and the western populations of b.
birdi. The lobes are rigid, inflexible structures, and
72 EURYMERODESMID MILLIPEDS
here they are very large, extending ventrad well
below the levels of the adjacent coxae, suggesting
impairment of motility and promotion of ecological
and/or behavioral adaptations. I have not en-
countered either species, but for clearance, it seems
that the lobes must alter the animal’s posture. If they
do not carry themselves high off the substrate, the
lobes would seemingly become embedded or broken.
I saw no individuals with broken lobes nor any in
which the apical hairs had been scraped off, in-
dicating that the structures are unaffected by
motion. Meaningful observations and ecological
studies on these eurymerodesmids are desirable
because of these prominent structural features.
The birdi lineage, comprised of a single species
group, the birdi group, is the northernmost in the
family, extending from northeastern Nebraska and
central Mississippi to the Gulf Coast of Louisiana
and south-central Texas (Figs. 147, 213-214). Its area
ranks second in size to that of the varius group,
Kewanius lineage, but it contains only two species in-
stead of five.
Components. — birdi Chamberlin [b. birdi, birdi
planus Causey], mundus Chamberlin.
Eurymerodesmus birdi Chamberlin
Diagnosis. — A moderate-size species; males
characterized by moderate to long mandibular pro-
jection; sides of aperture not divided, without
caudolateral pouches, configuration variable, either
flush with metazonal surface throughout, or
elevating slightly, continuing around caudolateral
corners, and descending a short distance along
caudal margin, or elevating more strongly and con-
tinuing into variably long, broadly rounded, sub-
triangular lobes at caudolateral corners or on caudal
margin; telopodite long; acropodite short, about 1/8
of telopodite length, broadly terminal, poorly
demarcated from prefemur; latter with many hairs
arranged continuously along inner margin; cypho-
pod valves of females with distal corners varying
from slightly produced to elevated into short rim;
receptacle large, sides not curved around operculum.
Remarks. — As represented by the types, the
westernmost population of birdi possesses long aper-
ture lobes that extend below the levels of the adja-
cent coxae. The projections are distinctive, easily
recognized, and were adequately illustrated by
Chamberlin (1931, fig. 8). This population occurs in
Murray, Garvin, Seminole, and Pottawatomie coun-
ties, Oklahoma, and ones with equally long lobes oc-
cur sporadically to the east, for example in Vicks-
burg, Warren County, Mississippi. However, most
forms outside these counties display variably shorter
lobes, and there is a general trend toward smaller
projections and lower marginal elevations as one
proceeds southeastward in Louisiana. The lobes are
also short north and south of these counties, but
show no geographical trends. The projections disap-
pear at New Orleans, leaving a low, uniform
marginal elevation extending from the anterior
halves of the sides around the caudolateral corners.
In central Mississippi, the form named planus by
Causey (1950b) exhibits an unadorned aperture
without elevations but with modest hair patches at
the caudolateral corners. The opening is flush with
the metazonum, and the hair patches occur at the
approximate position of the lobes. Southward
towards New Orleans the sides and caudal margin
begin to elevate, but always uniformly, without a
trace of lobes. The evidence is not as convincing as in
varius, where there is clear intergradation between v.
varius and vy. christianus, but I think that these forms
with and without subtriangular lobes blend smoothly
together, since forms in East Baton Rouge and West
Feliciana parishes have very short lobes, barely
detectable above the marginal elevations. Thus,
planus is a form of birdi, the oldest name for this
ensemble, and as first reviser, I assign planus to the
populations lacking lobes, ranging essentially from
New Orleans to near Jackson, Mississippi. The only
other population showing reasonable anatomical
stability is the one in central Oklahoma with large
lobes represented by the types of birdi. However, if it
is designated the nominate subspecies, the intergrade
area will be vastly larger than that of either race,
since the other three available names — creolus,
schmidti, and plishneri — apply to forms in areas of
high instability. This solution is impractical, and I
therefore combine all forms with detectable lobes
under the nominate subspecies, essentially everything
north and west of New Orleans, including a few
localities east of the Mississippi River. In contrast to
varius, I do not recognize an intergrade region per se
in birdi because this is virtually the entire range
between New Orleans and central Oklahoma. Con-
sequently, birdi is the most intricate eurymerodesmid
next to amplus, since it encompasses the second
greatest array of variants. Causey (1950b) was amply
justified in considering planus a valid species, for it
could hardly be more distinct from the types of
birdi. However, these forms are merely opposite
ends of a complex spectrum of lobed eury-
merodesmids that Chamberlin (1931) was the first to
name.
ROWLAND M. SHELLEY
Figs. 148-167. Eurymerodesmus birdi. 148, projections of right mandible of holotype, lateral view. 149, aperture and gonopods in
situ of holotype, ventral view. 150, the same, lateral view. 151, the same, caudal view. 152, left gonopod of the same, lateral view. 153,
telopodite of the same, medial view. 154, aperture and gonopods in situ of male from Caddo Par., LA, ventral view. 155, the same,
lateral view. 156, the same, caudal view. 157, aperture and gonopods in situ of male from Polk Co., AR, ventral view. 158, the same,
lateral view. 159, left cyphopod of allotype, caudal view. 160, the same, lateral view. 161-167, birdi planus. 161, projection of right man-
dible of holotype, lateral view. 162, aperture and gonopods in situ of holotype, ventral view. 163, left gonopods of the same, lateral view.
164, telopodite of the same, medial view. 165, aperture and gonopods in situ of male from Pearl River Co., MS, ventral view. 166, left
cyphopod of female from Simpson Co., MS, caudal view. 167, the same, dorsal view. Scale lines for figs. 149-151, 154-158, 162, and 165
= 1.00 mm. Line for other figs. = 0.30 mm for 148 and 161; 0.40 mm for 152, 163, and 166-167; and 0.50 mm for 153, 159-160, and
164.
MEM. AMER. ENT. SOC., 37
73
74 EURYMERODESMID MILLIPEDS
Eurymerodesmus birdi birdi Chamberlin, new status
Figs. 147-160, 211-232
Leptodesmus hispidipes: Gunthorp, 1913:162 (in part). Chamber-
lin, 1918:370 (in part).
Eurymerodesmus birdi Chamberlin, 1931:101-102, figs. 6-8.
Attems, 1938:185-186, figs. 204-205. Causey, 1952b:1-2.
Chamberlin and Hoffman, 1958:79. Stewart, 1969:384.
Eurymerodesmus creolus Chamberlin, 1942a:6, pl. 2, fig. 16.
Chamberlin and Hoffman, 1958:79. NEW SYNONYMY.
Eurymerodesmus schmidti Chamberlin, 1943:38, fig. 7; 1952:573.
Causey, 1952b:2. Chamberlin and Hoffman, 1958:81. NEW
SYNONYMY.
Eurymerodesmus plishneri Causey, 1950a:271, fig. 8. NEW
SYNONYMY.
Type specimens. — Male holotype, one male
paratype, and female allotype (NMNH) collected by
R. D. Bird, 17 October 1929, from an unknown site
in Murray Co., OK.
Diagnosis. — Apertures with distinct sub-
triangular lobes of varying lengths at varying posi-
tions along caudal margin to caudolateral corners.
Color in Life. — Peritremata orange; metaterga
gray-brown with orange stripes along caudal
margins connecting paranotal markings; collum with
stripes along both anterior and posterior margins
(Causey 1950a). Causey further states that the
metatergal stripes are usually wider mediad, render-
ing a trimaculate appearance resembling the syntopic
platyrhacid Auturus evides (Bollman). Chamberlin
(1931, 1942a, 1943) also recorded the striped pattern,
but from faded preserved specimens.
Holotype. — Body highly fragmented, length un-
measurable, maximum width 4.2 mm.
Somatic features similar to hispidipes, with following excep-
tions:
Width across genal apices 2.7 mm, interantennal isthmus 0.7
mm. Antennae missing, length and relative lengths of antenno-
meres unknown, recorded from paratype. Genae with faint central
impressions. Facial setae as follows: epicranial, interantennal, and
genal absent, subantennal 1-1, frontal 1-1, clypeal about 8-8, labral
about 13-13. Projection of mandibular stipes long, stout, bluntly
rounded (Fig. 148).
Collum extending slightly below ends of following tergite.
Paranota depressed, continuing slope of dorsum; posterior corners
rounded through segment 8, blunt on 9-13, becoming progressively
more acute posteriorly.
5th sternum with broad, flattened, paramedian processes be-
tween both leg pairs; 6th sternum with subconical processes be-
tween both coxae. Postgonopodal sterna with subconical projec-
tions between 9th legs; remaining sterna with shorter subconical
lobes on segment 8, becoming progressively flatter posteriorly with
variable bicruciform impressions and variable hair tufts subtending
coxae, becoming sparser caudally. 2nd coxa with short, rounded
anterior, and longer, subacuminate caudal lobes. Prefemoral lobes
extending through segment 16.
Gonopodal aperture (Figs. 149-151) subrectangular, without
caudolateral pouches, with moderate anterior indentation, 1.8 mm
wide and 0.7 mm long at maxima; anterior indentation short,
broad, rounded; sides entire, nearly linear, becoming progressively
more elevated caudad, leaning slightly over opening, rims essen-
tially smooth; caudolateral corner broad, rounded; caudal margin
continuous laterally with sides, extending immediately into enor-
mous, broadly subtriangular lobe, leaning caudad, narrowly
segregated in midline from opposite lobe and thus occupying near-
ly half of breadth of aperture, lobe extending well below margins
of adjacent coxae, apex broad, rounded, perhaps 1/3 as wide as
base, dropping to metazonal surface just before midline; margins
irregularly hirsute with long hairs arising from rim or short
distances down outer surfaces and overhanging opening, with
several hairs on anterior indentation, scattered hairs on sides and
caudolateral corner, hairs shorter and very dense on all surfaces of
lobes, becoming slightly denser distad. Gonopods in situ (Fig. 149)
with telopodites overlapping opposite member in midline, extend-
ing caudad between aperture lobes and terminating between 9th
legs below level of apices of lobes. Gonopod structure as follows
(Figs. 152-153): Telopodite terminating before level of distal ex-
tremities of hairs. Prefemur long, about 7/8 of telopodite length,
linear for most of length, curving distad, sides parallel for most of
length, tapering smoothly and continuously distad with two rows
of continuous, regularly spaced hairs along inner margin and
distomedial tuft of about six hairs. Acropodite short and broadly
terminal, about 1/8 of telopodite length, smoothly continuous
with, and poorly demarcated from prefemur, continuing curvature
of latter, sides tapering rapidly to acuminate tip.
Male paratype. — Body fragmented, length unmeasurable.
Antennae reaching back to anterior half of 3rd metatergite,
relative lengths of antennomeres 2>4>3>5=6>1>7. Somatic
and gonopodal features closely similar to those of holotype.
Female allotype. — Length 22.2 mm, maximum width 3.6 mm,
W/L ratio 16.2%. Agreeing essentially with holotype in somatic
features, with following exceptions: Subantennal and frontal setal
series absent. Projection of mandibular stipes short, blunt. Sterna
relatively flat and unmodified, without projections, with only a
few randomly scattered setae, especially along caudal margins.
Cyphopodal aperture subrectangular, caudal margin distinctly
elevated above metazonal surface. Cyphopods in situ with valves
oriented transversely, edges of valves and corner of receptacle
visible in aperture, open side facing dorsad. Valves (Figs. 159-160)
large, moderately hirsute, slightly unequal, anterior one larger,
distal margins elevated into short rim. Receptacle large, glabrous,
located laterad to valves, sides curving partly around operculum.
Latter relatively large, located at bases of valves on dorsal side.
Variation. — Size varies markedly, with the
smaller individuals occurring to the north in
Missouri and Kansas and the largest specimens in
Polk County, Arkansas. Elsewhere body propor-
tions seem random and probably reflect habitat con-
ditions. Piecing together males with no more than
two fragments the following measurements were
made:
Table 5. Size variation of E. b. birdi males.
Length Width W/L
Sine Conte) (mm) (mm) ratio (%)
Kansas Douglas 22.0 3.4 15.5
Oklahoma Payne 24.1 4.0 16.6
Oklahoma Garvin 27.0 4.4 16.3
Arkansas Washington 28.7 4.5 15.7
Arkansas Polk 37.8 5.3 14.0
Louisiana Natchitoches 33.9 5.0 14.7
Texas Bowie 33.2 4.8 14.5
ROWLAND M. SHELLEY 75
The gonopods are relatively constant throughout
the range but the angle of projection from the aper-
ture varies from nearly directly ventrad with the tips
leaning slightly between the aperture lobes, to
caudad, with the apices extending directly between
and beyond the lobes, and overhanging the sternum
between the 9th legs. On the apertures, considerable
variation obtains in the configurations of the lateral
margins, the size and degree of separation of the
caudal lobes, and the hirsuteness of the lateral and
caudal margins. Since the lobes are the most promi-
nent feature, differences in their relative proportions
are particularly noticeable, and I analyzed this geo-
graphically. The longest lobes occur primarily in
forms along the western range periphery, around the
type locality in central Oklahoma, where all males
exhibit them. Elsewhere they are random. These
lobes are moderately hirsute, much longer than the
widths of the 9th coxae, usually extend ventrad
below the telopodites, are relatively broad, and are
narrowly segregated in the midline. Together they
comprise most of the caudal margin of the aperture
and are aligned with the sternal processes between
the 9th legs (Fig. 151). Thus, a subspecies could be
recognized from Seminole, Pottawatomie, Murray,
and Garvin counties, Oklahoma, but elsewhere the
lobes vary randomly and capriciously with too little
stability, even within single populations as reflected
by the larger samples, to justify taxonomic recogni-
tion. In the most common configuration (Figs.
154-156), the lobes are subequal to or only slightly
longer than the 9th coxae, do not extend ventrad
below the telopodites, and are more widely
segregated thus being situated more laterad, at or
partly on the caudolateral corners, and ‘‘outside”’
the succeeding sternal processes as seen in caudal
profile (Fig. 156). However, this form intermingles
sporadically with ones with shorter lobes located
either more laterad or mediad (Figs. 157-158), and
the male from Mississippi has long lobes like the
form in central Oklahoma. In the male from East
Baton Rouge Parish, Louisiana, the lobes are
situated at the caudolateral corners and are so widely
segregated that the caudal margin seems to lack
them. A specimen from Grand Ecore, Natchitoches
Parish, exhibits a uniformly elevated caudal margin
without a trace of lobes, as in b. planus, but the
other male in this sample has prominent lobes. Males
from West Feliciana and Jefferson parishes show
very small lobes, only slightly higher than the
marginal elevations, and those in south Texas are
small to moderate-size, with extremely long hairs in
some individuals. Finally, the lobes are not always
equal in a single individual; in a few males, one is
clearly longer than the other. Thus, aside from cen-
tral Oklahoma, b. birdi does not exhibit a consistent
set of features, thus obviating additional races. If
future studies indicate otherwise, creolus has priority
over both schmidti and plishneri.
Aside from the length and position of the lobes,
they can also extend either directly ventrad or lean
anteriad or caudad, accordingly creating narrower or
broader openings. Where they lean anteriad, the
aperture appears wider along the anterior margin.
The degree of hirsuteness of the lobes and the lateral
margins vary, and some males are exceptionally
hairy with long, matted hairs, nearly as long as the
lobes themselves, extending from their tips and
anterior and caudal margins (Fig. 155). On the sides
of the apertures, hairs arise mostly from the rim, but
on sporadic individuals they also extend varying
distances down the outer surface (Figs. 157-158).
These lateral marginal hairs can also be very long,
overhanging up to perhaps 1/4 of the width of the
aperture.
The lateral elevations also vary, in many in-
dividuals arising smoothly and continuously begin-
ning at the anterolateral corners and curving around
the caudolateral corners into the lobes. In other
males the elevations begin a short distance back of
the anterolateral corners, producing a “‘stepped’’ ef-
fect in conjunction with the lobes. in some such in-
dividuals, the elevated parts of the lateral margins
lean inward over the opening, causing it to appear
wider along the anterior border.
Ecology. — The specimen reported by Cham-
berlin (1942a) was collected under a log in Caddo
Parish, Louisiana; Causey (1950a) mentioned that
oaks were dominant at the six recorded localities of
plishneri in Washington and Carroll counties,
Arkansas. The following habitat data was indicated
on vial labels: on a driveway (Payne Co., OK); ona
patio (Garvin Co., OK); under rocks in woods
(Wagoner Co., OK); in an old stump, in rocks, and
in water wells (Natchitoches Par., LA); under logs
(Caddo Par., LA); in hardwoods at a lumber yard
(West Feliciana Par., LA); and in sandy post oak
soil (Wilson Co., TX). The milliped was collected in
caves in Seminole, Cherokee, and LeFlore counties,
Oklahoma, one of three eurymerodesmids known
from subterranean habitats along with oliphantus
and melacis. These areas are either on the western
periphery of the Ozark and Ouachita Provinces or
the eastern fringe of the Central Plains and may or
may not represent accidental records.
76 EURYMERODESMID MILLIPEDS
Distribution. — From the southern Central
Lowland Physiographic Province south of the
Missouri River in northeastern Kansas and central
Missouri, through the western Ozark Plateau and
Ouachita Provinces to northeast Texas and the
southern Mississippi Valley, with additional,
possibly allopatric, populations in south-central
Texas on the western fringe of the Coastal Plain
(Figs. 147, 211-212). Specimens were examined as
follows:
KANSAS: Douglas Co., Lawrence, 3M, Sep-
tember 1977, R. Graham (RLH); and Univ. of Kan-
sas Nat. Hist. Res., M, 2F, 29 September 1961, R.
C. Funk (EIU) and 3M, 4 October 1966, D. Clark
(FSCA). Johnson Co., 3 mi. E. Eudora, M, 4
October 1966, D. Clark (FSCA).
MISSOURI: Johnson Co., Knob Noster, 2M,
TF, 29 September 1962, and 11M, 25 F, November
1962, E. Peterman (FSCA). Vernon Co., Nevada,
M, | October 1961, D. J. McReynolds (MCZ).
OKLAHOMA: Cherokee Co., 3 mi. SE Ft.
Gibson Dam, Dressler Cave, 5M, F, 21 March 1971,
J. H. Black (FSCA). Wagoner Co., Lake Bixhoma,
M, 1 March 1975, J. M. Nelson (NCSM). Payne
Co., Stillwater, M, 15 March 1984, R. M. Edwards
(OKSU). Pottawatomie Co., Dale, M, 3F,
September 1954, Brown (FSCA). Seminole Co.,
locality unknown, M, May 1931, P. Newport
(SMUO); and Whiskey Cave, M, 6 January 1974, J.
H. Black (FSCA). Garvin Co., locality unknown,
4M, F, 2 November 1984, B. Lucas (OKSU). Mur-
ray Co., locality unknown, 2M, F, 17 October 1929,
R. D. Bird (NMNH) TYPE SPECIMENS, and 2M,
F, 3 juvs., 17-26 October 1928, R. D. Bird (SMUO);
Sulphur, M, 18 April 1976, W. Magdych (SMUO);
and near Turner Falls, Arbuckle Mts., M, 23 April
1936, L. Hubricht (NMNH). LeFlore Co., 10 mi.
NE Talihina, Bear Den Cave, 4M, 17 November
1973, J. H. Black (FSCA).
ARKANSAS: Benton Co., 5 mi. E Siloam,
Chamber Spgs. Ranch, 3M, 2 September 1955, C. E.
Hastings (FSCA); and Bella Vista, 2M, 13 April
1962, H. Hart (FSCA). Carroll Co., Blue Springs,
3M, F, 29 October 1949, N. B. Causey (FSCA).
Washington Co., Springdale, Eden’s Bluff, 3M, 8F,
2 juvs., 1 October 1949, N. B. Causey (FSCA);
Fayetteville, M, 8 April 1959, N. B. Causey (FSCA),
Mt. Kessler, M, F, 11 April 1949, N. B. Causey
(ANSP) and 2M, F, 3 November 1949, collector
unknown (FSCA), and Univ. of Arkansas campus,
2M, 10 February 1953, L. Jackson (FSCA); nr.
Prairie Grove, Cove Cr. Valley, 18M, 23F, 4 juvs., 1
November 1955-25 March 1960, M. Hite (FSCA);
and Devil’s Den St. Pk., M, 3F, 2 juvs., 29
September 1948, N. B. Causey (FSCA) and M,
December 1963, N. B. Causey (MCZ). Pope Co.,
locality unknown, M, 6 February 1955, B. Bowie
(FSCA). Logan Co., along AR hwy. 23 nr. Boone-
ville, M, 19 November 1966, M. Hite (FSCA). Polk
Co., Rich Mtn., Mena, 11M, 2F, 21-22 March 1938,
K. P. Schmidt (FMNH, NMNH), and 5M, 7F,
1950-May 1956, A. McWilliams and N. B. Causey
(FSCA).
MISSISSIPPI; Warren Co., Vicksburg, M, 7
February 1933, O. F. Cook (NMNH).
LOUISIANA; Caddo Par., 5 mi. NW Shreve-
port, M, 13 April 1936, L. Hubricht (NMNH); and
Shreveport, 3M, 9F, 3 juvs., 24 March 1962, N. B.
Causey (FSCA). Natchitoches Par., locality
unknown, 25M, 13F, October 1954-February 1956,
various collectors (FSCA); Natchitoches, North-
western State Univ. campus, 7M, 12F, 3 juvs.,
February 1954-March 1956, J. E. Sublette and C.
Morehead (FSCA); and 16 mi. SSE Natchitoches,
2M, 3F, 16 October 1965, D. E. Hahn (FSCA).
Rapides Par., along LA hwy. 28, 8.8 mi. E Gardner,
8M, 7F, 7 November 1965, R. E. Tandy and L. D.
Wilson (FSCA). Concordia par., Tensas R., 2M,
1954, B. Stanberry (FSCA). Avoyelles Par.,
Evergreen, 2M, 4F, 28 October 1965, M. Kordisch
(FSCA). Evangeline Par., 2.5 mi. S Chicot St. Pk.,
M, 2F, juv., 30 January 1968, E. D. Kaiser (FSCA).
St. Landry Par., along LA hwy. 10, 2 mi. W
Mellville, M, 3F, 24 March 1965, R. Tandy (FSCA).
East Baton Rouge Par., Louisiana St. Univ. Cam-
pus, M, November 1971, collector unknown
(FSCA). West Feliciana Par., locality unknown, M,
F, 18 March 1962, L. Hubricht (RLH). Jefferson
Par., Narahan Jct., Elmwood Plantation along
River Rd. at Research Dr., 0.3 mi W jct. US90, 2M,
2F, juv., 17 February 1986, R. M. Shelley (NCSM).
TEXAS: Bowie Co., along US hwy. 259, 5 mi. S
Red River, M, 9 December 1967, T. C. Stewart
(SFAU). Bell Co., Belton, M, January 1931, collec-
tor unknown (NMNH). Bastrop Co., Bastrop St.
Pk., 2M, 2 April 1960, J.C. Loomis (RLH). Wilson
Co., 8 mi. N Floresville, 2M, 29 April 1961, collector
unknown (FSCA); and 8 mi. N Stockdale, M, 6
April 1961, J. F. Quinlan (FSCA). San Patricio Co.,
Lake Corpus Christie St. Pk., M, 2 July 1969, J. R.
Heitzman (FSCA).
The following literature records of additional
localities are believed to refer to b. birdi and are so
indicated in figures 147 and 211-212.
KANSAS (Gunthorp, 1913, cited as Leptodesmus
hispidipes): Jefferson Co., no further data.
ROWLAND M. SHELLEY 7
OKLAHOMA (Causey 1952b); Hughes Co., no
further data.
ARKANSAS (Causey 1952b): Sebastian Co., no
further data. Miller Co., no further data.
LOUISIANA (Chamberlin 1918, cited as Lep-
todesmus hispidipes): Natchitoches Par., Creston.
TEXAS (Stewart 1969): Lamar Co., 8 mi. SE
Paris.
Additional samples with a male from White Rock,
Arkansas (FSCA), and a male and several females
from Pacheco, Texas (FSCA), cannot be located in
maps or gazetters of these states.
Remarks. — The males from Norman, Cleveland
County, Oklahoma (SMUO), which Chamberlin
designated paratypes of birdi are actually specimens
of mundus.
The label with the sample from 8 mi. N Flores-
ville, Wilson Co., TX, states that the milliped was
‘‘very common, usually paired and grouped,” but
does not indicate whether the pairing was between
opposite sexes.
The male from Payne County, Oklahoma
(OKSU), has an accessory gonopod at the right
anterior position of segment 8, the first example of
heteromorphosis in the Eurymerodesmidae. The
structure is shorter than normal and enclosed within
a small circular aperture that does not extend across
the midline. The lateral margin is normally elevated
and extends into a typical caudal lobe, subequal in
length to those on segment 7. A normal leg occurs
opposite the accessory gonopod at the left anterior
position of segment 8.
Eurymerodesmus birdi planus Causey, new status
Figs. 147, 161-167, 211
Eurymerodesmus planus Causey, 1950b:196-197, fig. 5. Chamber-
lin and Hoffman, 1958:80.
Type specimen. — Male holotype (ANSP) col-
lected by S. Bender, 13 April 1950, from Piney
Woods, Rankin Co., MS. According to Causey
(1950b) the type series also contained three females,
now lost.
Diagnosis. — Apertures without lobes at any
position, usually with denser hair patches at
caudolateral corners, with or without uniform
marginal elevations, arising on anterior halves of
sides, curving around caudolateral corners, termi-
nating before midline on caudal margin (Figs. 162,
165).
Color in life. — Paranota orange; metaterga dark
with orange stripes along caudal margins connecting
paranotal markings; collum with orange stripes
MEM. AMER. ENT. SOC., 37
along both anterior and posterior margins (Causey
1950b). Causey also mentioned the presence of a
roughly triangular median orange spot on some
metatergites, perhaps reflecting expansion of the
stripe in the midline, and several small, confluent
orange dots on others. A paratype female had ‘‘two
to six’’ orange dots on the metatergites.
Variation. — All specimens were tightly coiled
and would not unroll without fragmenting into two
or more pieces. A male from Pearl River County,
Mississippi, which only broke into two pieces,
measured approximately 24.0 mm long, 3.4 mm
wide, W/L ratio 14.2%.
As stated, the holotype exhibits an unadorned
aperture, margins being flush with the metazonum.
Its only attributes are modest hair patches at the
caudolateral corners, which really are nonexistant
since the sides curve strongly mediad. Consequently,
the aperture is proportionately the smallest in the
genus and is heart-shaped in overall appearance (Fig.
162). The gonopods are a little more curved than the
linear ones of the nominate subspecies (Figs.
163-164).
The Simpson County males from Braxton are
near topotypes, occurring just across the county line
from the type locality. Here the sides of the aperture
are slightly but distinctly elevated beginning at the
anterolateral corners. In the males from Jefferson
Davis County, the caudolateral corner is angled and
slightly more elevated; thus the caudal margin slopes
downward from the corner, meeting the metazonal
surface in the midline. Essentially the same condition
obtains in the Pearl River County males (Fig. 165),
except the caudolateral corner is rounded and leans
over the aperture in one male. All Mississippi males
exhibit hairs at the caudolateral corners, their den-
sities and lengths varying, being longest in the last
sample.
The Louisiana males display essentially the same
conditions as those from southern Mississippi. In the
St. Tammany Parish specimens, the moderately
elevated sides continue around a squared caudo-
lateral corner, then slope to the metazonal surface
near the midline. In that from New Orleans, the
sides are essentially flush with the metazonal surface,
with the caudolateral corner being slightly elevated
and rounded. The caudal margin extends slightly in
the midline in all Louisiana males, and the hairs con-
tinue a short distance beyond the caudolateral corner
in the New Orleans male.
The mandibular projection of the holotype is
shorter and broader than in the nominate race (Fig.
78 EURYMERODESMID MILLIPEDS
161). The female genitalia are closely similar to those
of b. birdi (Figs. 166-167).
Ecology. — The samples from Pearl River
County Mississippi, were collected in ‘‘flood plain,
water oaks.’’ Those from 2.5 miles south of Pren-
tiss, Jefferson Davis County, were retrieved from a
pitfall trap on a hillside in pine woods.
Distribution. — The southern Gulf Coastal Plain
of Mississippi and southeastern Louisiana, ranging
from Jackson to New Orleans (Figs. 147, 211).
Specimens were examined as follows:
MISSISSIPPI: Rankin Co., Piney Woods, M,
13 April 1950, S. Bender (ANSP) TYPE LOCALI-
TY. Simpson Co., near Braxton, 2M, 5F, 4 juvs., 1
April 1958, S. Lazell (FSCA); and Mendenhall, M, 2
January 1960, L. Hubricht (RLH). Jefferson Davis
Co., 3 mi. SSW Prentiss, along White Sand Cr., 2M,
25 December 1959, L. Hubricht (RLH); and 2.5 mi.
SW Prentiss, 4M, 27 February 1987, P. K. Lago
(RLH). Pearl River Co., 11 mi. E Bogalousa, LA,
6M, 4F, 21 January 1965, N. B. Causey (FSCA).
LOUISIANA: St. Tammany Par., Talisheek,
along Talisheek Cr., 2M, 12 December 1965, L. D.
Wilson (FSCA); and 1 mi. W Talisheek, 2M, 9
February 1939, collector unknown (NMNH).
Orleans Par., New Orleans, city park, M, 20
February 1950, collector unknown (FSCA).
Eurymerodesmus mundus Chamberlin
Figs. 147, 168-175, 211-212
Leptodesmus floridus: Kenyon, 1893:16.
Leptodesmus hispidipes: Gunthorp, 1913:162 (in part).
Eurymerodesmus mundus Chamberlin, 1931:102-103, figs. 3-5.
Attems, 1938:186-187, fig. 206. Causey, 1952b;2. Chamber-
lin and Hoffman, 1958:80.
Type specimens. — Male holotype and male
paratype (NMNH) collected by D. Zeigler, 26
February 1930, at University (Norman), Cleveland
Co., OK. One male paratype (NMNH) collected by
R. D. Bird, 5 April 1929, at an unknown site in
Cleveland County.
Diagnosis. — A moderate-size species; males
characterized by long mandibular projection; sides
of aperture not divided, without caudolateral
pouches, elevating slowly around caudolateral cor-
ners onto caudal margin, extending into very large
clavate lobes; telopodite long; acropodite short,
about 1/8 of acropodite length, broadly terminal,
poorly demarcated from prefemur; latter with many
hairs arranged continuously along inner surface;
cyphopod valves of females with distal corners
slightly produced; receptacle moderate size, sides
curving around operculum.
Color in Life. — Paranota red; metaterga brown
with concolorous stripes along posterior margins
connecting paranotal markings; collum with brick
red stripes along both anterior and posterior margins
(Chamberlin 1931). All specimens available to me
were preserved with faded colors, but many still
showed clear evidence of this striped pattern.
Holotype. — Body fragmented, length un-
measurable, maximum width 4.1 mm.
Somatic features similar to those of E. hispidipes, with follow-
ing exceptions:
Width across genal apices 3.2 mm, interantennal isthmus 0.8
mm. Relative lengths of antennomeres 2>3>5>4>6>1>7.
Genae with faint central impressions. Facial setae as follows:
epicranial, interantennal, frontal, and genal absent; clypeal about
11-11, labral about 14-14. Projection of mandibular stripes long
and broad, ventral margin indented, tip smoothly irregular (Fig.
168).
Collum extending slightly below ends of following tergite.
paranota moderately flattened, interrupting slope of dorsum;
posterior corners rounded through segment 8, blunt on 9-14,
becoming progressively more acute posteriorly.
5th sternum with short, widely separated knob-like processes ad-
jacent to leg coxae; 6th sternum with broader, shorter, paramedial
projections, caudal ones longer. Postgonopodal sterna with nar-
row, rounded, widely segregated lobes between 9th legs; remaining
sterna with widely separated subconical lobes on segment 8 and
flattened, elevated areas on 9, becoming progressively flatter
posteriorly with variable transverse impressions between leg coxae,
and shallow, faint to moderate central impressions rendering
somewhat bicruciform appearance, all except 18th sterna with
tufts of setae on ventral surfaces adjacent to leg coxae. 2nd coxa
with large, rounded anterior, and narrower subconical posterior
lobes. Prefemoral lobes extending through segment 16.
Gonopodal aperture (Figs. 169-171, not this specimen) ovoid,
widest along anterior margin, without caudolateral pouches, with
slight anterior indentation, 1.9 mm long and 0.8 mm wide at max-
ima; anterior indentation short, broadly rounded; sides curving
mediad, not divided, rising slowly and progressively to caudo-
lateral corner, rims slightly irregular; caudolateral corner in-
distinct, continuing curvature and rise of sides; caudal margin con-
tinuous through caudolateral corner with sides, lowly elevated for
short distance then extending greatly ventrad into an enormous,
clavate, lobe extending ventrad well beyond level of coxae, drop-
ping to level of marginal elevation and curving across midline,
lobes paramedian and narrowly segregated in midline, aligned just
outside sternal processes between 9th legs; margins heavily hirsute
with hairs arising regularly from rim and overhanging opening,
thicker along anterior indentation, also arising from outer surfaces
laterally and caudally, caudal lobes densely pilose, hairs beginning
basally and becoming denser apically, hairs on lobes much shorter
and denser than those from aperture rim. Gonopods in situ (Fig.
169, not this specimen) with telopodites touching or nearly so in
midline, extending between, and terminating just beyond aperture
lobes, apices directed sublaterad. Gonopod structure as follows
(Figs. 172-173): Telopodite terminating well before level of distal
extremities of hairs. Prefemur long, about 7/8 of telopodite
length, configuration linear, curving abruptly distad, sides parallel
for most of length, tapering distad near level of curve, with two
rows of continuous, regularly spaced hairs along inner margin
culminating in slightly thicker tuft of six hairs, and distomedial tuft
ROWLAND M. SHELLEY
Figs. 168-175. Eurymerodesmus mundus. 168, projection of left mandible of holotype, lateral view. 169, aperture and gonopods in
situ of male from Comanche Co., OK, ventral view. 170, the same, lateral view. 171, the same, caudal view. 172, left gonopod of
holotype, lateral view. 173, telopodite of the same, medial view. 174, left cyphopod of female from Canadian Co., OK, caudal view. 175,
the same, dorsal view. Scale line for figs. 169-171 = 1.00 mm. Line for other figs. = 0.30 mm for 168, 0.40 mm for 172, and 0.50 mm
for 173-175.
MEM. AMER. ENT. SOC., 37
79
80 EURYMERODESMID MILLIPEDS
of about 6 hairs. Acropodite short and broadly terminal, about
1/8 of telopodite length, smoothly continuous with, and poorly
demarcated from latter, bending sharply sublaterad, slightly
bisinuate, sides tapering rapidly to acuminate tip.
Male paratype. — Agreeing closely with holotype in all
particulars.
Female from Canadian Co., OK. — Length about 19.6 mm,
maximum width 3.3 mm, W/L ratio 16.8%. Agreeing essentially
with holotype in somatic features, with following exceptions:
Subantennal setae 1-1, frontal setae 1-1. Mandibular stipes with
minute rounded process. 3rd and 4th segments with moderate
flanges lateral to legs. Sterna relatively flat and unmodified,
without projections or tufts of setae.
Cyphopodal aperture subrectangular, caudal margin slightly
elevated above metazonal surface. Cyphopods in situ with valves
directed mediad, edges and side of receptacle visible in aperture,
open side facing dorsad. Valves (Figs. 174-175) moderate-size,
slightly unequal, anterior one larger and rounded, distal corners
slightly produced. Receptacle moderately large, glabrous, located
laterad to valves, sides curving partly around operculum. Latter
relatively large, located at bases of valves on dorsal side.
Variation. — Size varies markedly, with larger
specimens occurring in the center of the range from
Oklahoma to Arkansas, and smaller ones occurring
peripherally. Piecing together a few of the less
fragmented males, with no more than two pieces, the
following measurements were made. For most
specimens merely straightening for measurement
resulted in breakage.
Table 6. Size Variation of E. mundus Males.
Length Width W/L
State County (earra) ca) (0)
NE Jefferson 21.9 3.2 14.6%
KS Riley 22.4 2.9 12.9%
OK Cleveland 31.1 5.0 16.1%
OK Comanche 32.4 4.6 14.2%
AR Sevier 33.9 5.6 16.5%
TX Johnson 21.4 2.6 12.1%
The pregonopodal legs are more densely hirsute
than those in the midbody region on some males,
and in these individuals the legs become progressive-
ly less hirsute caudad. In some males the lateral
margins of the gonopodal aperture are slightly ex-
cavated on the anterolateral corners. The distance
between the caudal lobes varies, some being more
widely segregated than others. In most individuals,
the projections extend essentially directly ventrad,
but in a few they diverge slightly laterad. These
features occur randomly throughout the range. The
lobes are virtually identical in length and configura-
tion on all males except for the southernmost from
Johnson County, Texas, where they are noticeably
shorter and less clavate.
Ecology. — The holotype was collected in a
“‘black jack forest’? (Chamberlin 1931), presumably
blackjack oak litter, on or very near the University
of Oklahoma campus. Vial labels with habitat nota-
tions are as follows: under a litter and under rocks in
the Wichita Mountains (Comanche Co., OK), on the
floor of a house (Tulsa Co., OK), under a log (Noble
Co., OK), in a storm cellar (Grayson Co., TX), and
under a rock in a pasture (Ellsworth Co., KS).
Distribution. — Eurymerodesmus mundus is the
northernmost species with presumably reliable
literature records from northeastern Nebraska and
one with a voucher specimen from near Omaha. The
known distribution covers the Central Lowland
Physiographic Province from northeastern
Nebraska to the Brazos River in northern Texas, ex-
tending eastward into the Ouachita and Coastal
Plain Provinces in eastern Oklahoma and
southwestern Arkansas (Figs. 147, 211-212). Neither
mundus nor the family is known from north of the
Missouri River. Discovery in western Missouri south
of this watercourse is anticipated. Specimens were
examined as follows:
NEBRASKA: Cass Co., South Bend, 2M, date
unknown, F. C. Kenyon (NMNH). Jefferson Co.,
Reynolds, M, 7 October 1981, W. F. Rapp (MCZ)
and M, 9 September 1987, W. F. Rapp (RLH).
KANSAS: Riley Co., locality unknown, 2M, 2
April 1952, L. Warren (FSCA) and Manhatten, M, 2
May 1940, H. H. Walker (RLH). Ellsworth Co., 6
mi. SW Ellsworth, M, 2 May 1976, C. L. Starr
(UGA). Butler Co., 8 mi. SE Augusta, M, 14 Oc-
tober 1935, Burt (NMNH).
OKLAHOMA: Craig Co., 1 mi. S Kansas state
line, F, 7 April 1973, J. H. Black (FSCA). Tulsa
Co., Bixby, M, 1 December 1980, J. M. Nelson
(NCSM). Noble Co., Perry, M, 11 May 1967, D. C.
Arnold (WAS). Logan Co., 5 mi. NE Guthrie, M,
28 October 1959, R. C. Harrel (FSCA). Canadian
Co., locality unknown, 5M, F, 1930, N. M. Newport
(SMUO) and Yukon, M, 1930, N. M. Newport
(SMUO). Caddo Co., locality unknown, 3M, F,
February 1934, Carpenter (SMUO). Grady Co.,
Chickasha, F, 4 June 1944, L. Hook (FSCA). Com-
anche CO., Wichita Mts., locality unknown, 2M, 16
October 1971, collector unknown (MSU), Mt. Scott,
MM, FF, 25 October 1974, G. Beck (MSU); Sunset
Rec. Area, 2M, 29 October 1977, J. C. Coken-
dolpher and D. Parmley (MSU); Camp Boulder, F,
2 juvs., 8 June 1928, collector unknown (NMNH);
and Panther Cr. Rec. Area, M, 6 November 1977,
D. Parmley and K. Brice (MSU). Cleveland Co.,
locality unknown, M, 5 April 1929, R. D. Bird
(NMNH) and 2M, 20 February 1931, R. D. Bird
(SMUO); and Norman, University, M, 10 March
ROWLAND M. SHELLEY 81
1929, R. D. Bird (SMUO), M, 20 February 1937, D.
Zeigler (NMNH) and M, March 1976, W. D. But-
cher (SMUO) TYPE LOCALITY. McClain Co.,
locality unknown, M, F, February 1933, Rosenmal
(SMUO). Pittsburg Co., McAlester, 2M, 2F, 10
February 1952, O. Murray (FSCA); and 3 mi. E
Krebs, M, 2F, 22 October 1961, R. C. Harrel
(FSCA). Latimer Co., locality unknown, M, F, 25
April and 27 June 1931, R. D. Bird (SMUO) and
2M, F, 9 December 1933, Carpenter (SMUO); 5 mi.
NW Wilburton, Robbers Cave St. Pk., M, F, 17
May 1960, R. C. Harrel (FSCA) and M, 25 October
1976, J. C. Cokendolpher (MSU); and Wilburton,
M, 3F, 17 November 1934, Carpenter (ILNHS,
SMUO). LeFlore Co., 3.5 mi. N Howe, M, F, 26
May 1966, L. Hubricht (RLH).
ARKANSAS: Polk Co., Rich Mt., Queen
Wilhemina St. Pk., 2M, 8 November 1966, collector
unknown (FSCA). Sevier Co., Horatio, 4M, 8F,
11-15 April 1955 and 5 April 1956, C. N. Griffith
(FSCA).
TEXAS: Cooke Co., nr. old Townsite of Lea,
M, 11F, 7 May 1968, collector unknown (FSCA).
Dallas Co., Dallas, 2M, 2F, June 1955, O. Sanders
(FSCA); and between Duncanville and Cedar Hill,
M, 3 November 1927, O. F. Cook (NMNH).
Johnson Co., Cleburne St. Pk., M, 26 December
1979, G. Spicer (NCSM).
The following additional literature records are
believed to refer to mundus and are so indicated in
figure 147.
NEBRASKA (Kenyon 1893, cited as Lep-
todesmus floridus, a synonym of Semionellus
placidus (Wood) (Chamberlin and Hoffman 1958) ):
Cuming Co., West Point. Lancaster Co., Roca.
Richardson Co., Rulo. Unknown county, La Platte.
KANSAS (Gunthorp 1913, cited as Leptodesmus
hispidipes): Cowley Co., no further data.
Remarks. — Labels in the holotype and paratype
vials read “‘Erymnogon’’ mundus rather than
Eurymerodesmus, indicating that Chamberlin
originally thought this species required a new genus.
The lobes probably created this impression, but he
must have changed his mind, and fortunately
“Erymnogon”’ was never validated, although it
would have been available as a subgeneric designa-
tion for the birdi lineage. However, Chamberlin did
incorrectly designate a male topotype in the SMUO
as a paratype of birdi.
The sample from South Bend, Nebraska, collected
by F. C. Kenyon, the northernmost known for the
family, is identified as “‘Leptodesmus floridus.’’
This was not among the Nebraska localities Kenyon
MEM. AMER. ENT. SOC., 37
(1893) reported, but those he did — West Point,
Rulo, Roca, and La Platte — probably also refer to
mundus. Thus, I include his usage of L. floridus in
the synonymy. The first three sites are indicated by
open symbols in figure 147; the last is not on the of-
ficial Nebraska map. Hence, the range for mundus
probably extends north of the Platte River. I also in-
clude under mundus the Kansas records of Lep-
todesmus hispidipes by Gunthorp (1913), since they
are either in or near the known range and likely refer
to this species.
Chamberlin (1931) characterizes the lobes as aris-
ing from the sternum behind the gonopods, but his
description of laminas extending ectad and cephalad
from the structures implies that they are part of the
aperture margin.
In addition to being the northernmost congener,
mundus also forms the western boundary of the
family in the northern part of the range (Fig. 147).
The Missouri River seems to be a boundary, not only
for mundus but also for the family. No eurymero-
desmids have been taken to the north of this river in
Iowa and northern Missouri. Consequently, I doubt
if mundus occurs in the southeastern corner of South
Dakota.
The Melacis Lineage
Located primarily in south and central Texas, on
the known southern and western peripheries of the
generic and family range, the melacis lineage is a
nondescript assemblage with few distinguishing
features. It is best characterized negatively, as the
assemblage in Texas with short acropodites that
lacks aperture lobes and pouches, and whose lateral
margins are undivided. Common traits are small
body size and short telopodites, which are either
wholly enclosed by the aperture or just overlap the
caudal margin, unless the specimen is exceptionally
compressed. The apertures are comparatively small
in three species, but dactylocyphus has propor-
tionally the largest openings in the family.
Eurymerodesmus digitatus, with intermediate-size
Openings, bridges the anatomical gap. Aperture
margins are the most unmodified in the family; the
sides are entire, and there are no lobes, pouches, or
flares. Again dactylocyphus is an exception, with
cupped, flange-like elevations at the caudolateral
corners and adjoining parts of the sides and caudal
margin, and again, digitatus spans the anatomical
gap, at least in some populations. Although the
telopodites are short, barely extending beyond the
caudal aperture margin, the relative proportions of
the prefemur and acropodite are as in the Dirdi
82 EURYMERODESMID MILLIPEDS
Fig. 176. Distribution of the melacis lineage. Dots, melacis;
triangles, dactylocyphus; squares, sanbernardiensis; stars,
digitatus; asterisk, clavatus. Open symbols denote literature
records considered reliable.
lineage and varius group, with the prefemur con-
stituting from 2/3 to 7/8 of the telopodite length.
Acropodites are broadly terminal and continuous
with the prefemur except for clavatus, where it is
Narrowly terminal, discontinuous, and _ sharply
demarcated, arising from the outer prefemoral
margin. This condition, accompanied by a clavate
prefemur with a shoulder on the inner distal margin,
is the obverse of that found in simplex, where the
shoulder is on the outer surface, and the acropodite
arises from the inner surface. Females do not display
distinguishing features enabling them to be reliably
identified, except again for dactylocyphus and to a
lesser extent digitatus. The former possesses variable
valvular projections ranging from small, nubbinlike
papillae in eastern populations to long, flexible dac-
tyliform structures in western forms that extend
caudad and overhang two to three segments. This
dactyliform condition is convergent with that in
compressus, with the important distinction that the
projections are flexible instead of being stiff and
rigid. Thus the melacis group is characterized by
small body size (all species); short telopodites (all
species); terminal, continuous acropodites (excepting
clavatus); small, unadorned apertures (excepting
dactylocyphus); and rather nondescript, unadorned
cyphopod valves (excepting dactylocyphus).
Located in south Texas, south of and parapatric
to most of the varius group, with an extension into
southwestern Oklahoma, the melacis lineage is the
only eurymerodesmid component that may extend
into Mexico. Since it occurs practically to the north
bank of the Rio Grande from Laredo to Del Rio,
melacis itself may inhabit the northern fringes of
Coahuila, Nuevo Leon, and/or Tamaulipas states in
a manner similar to Abacion texense (Loomis) in the
order Callipodida (Shelley 1984a). However, this
watercourse could be a barrier, with a new species on
the southern side. Since the family crosses several
large US rivers, including the Mississippi, it may also
traverse the Rio Grande, thereby occurring in north-
eastern Mexico and possibly farther south. Indeed, a
new species group or lineage with several com-
ponents may occur in Mexico, which may provide
clues to the relationships of the family.
The melacis lineage is comprised of one species
group, the melacis group, and five species.
Components. — melacis Chamberlin and
Mulaik; sanbernardiensis Causey; digitatus Loomis;
dactylocyphus, new species; clavatus, new species.
Eurymerodesmus melacis Chamberlin and Mulaik
Figs. 176-182, 212
Eurymerodesmus melacis Chamberlin and Mulaik, 1941:59-60.
Chamberlin, 1952:573. Chamberlin and Hoffman, 1958:80.
Loomis, 1959:161.
Type specimens. — Male holotype and 57 male,
20 female, and 12 juvenile paratypes (NMNH) col-
lected by S. and D. Mulaik in December 1939 at
Edinburg, Hidalgo Co., TX. Other paratypes, all
taken by same collectors from localities in Texas and
in the NMNH, as follows: Kerr Co., Raven Ranch,
ca. 12 mi. S Kerrville, 4F, August 1939, and 22M,
2F, 3 juvs., December 1939; Kendall Co., locality
unknown, 9M, 15F, December 1939, and Boerne,
6M, 2F, December 1939; and Bandera Co., locality
unknown, 9M, F, December 1939, and 7 mi. N
Medina, M, date unknown. The original description
does not mention the holotype, and Chamberlin and
Hoffman (1958) cited Raven Ranch, Kerr County,
as the type locality. In the 11 vials of types, both a
male from Edinburg and a female from Raven
Ranch are labeled ‘‘holotype.’’ Since the former is a
male, I regard it as the holotype. A large series was
taken from Edinburg at the same time as the
ROWLAND M. SHELLEY 83
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Figs. 177-182. Eurymerodesmus melacis. 177, projection of left mandible of holotype, lateral view. 178, aperture and gonopods in
situ of paratype from Kerr Co., TX, ventral view. 179, left gonopod of holotype, lateral view. 180, telopodite of the same, medial view.
181, left cyphopod of female paratype from Hidalgo Co., TX, caudal view. 182, the same, medial view. Scale lines for fig. 178 =
1.00 mm. Line for other figs. = 0.25 mm for 177, and 0.50 mm for 179-182.
holotype, and the other specimens are properly con-
sidered paratypes. Chamberlin and Mulaik (1941)
gave no reason for designating so many additional
paratypes from other localities, which contrasts
markedly with other species of Eurymerodesmus
authored by Chamberlin where few paratypes were
designated.
Diagnosis. — A small species; males character-
ized by moderate-size mandibular projection; aper-
ture moderate-size, much narrower than ventral
segmental width; sides not divided, without
caudolateral pouches, with or without slight but con-
tinuous elevation with slight suggestion of lobe at
caudolateral corners, descending to metazonal sur-
face on caudal margin; telopodite short, usually only
slightly overlapping caudal margin; acropodite,
MEM. AMER. ENT. SOC., 37
short, about 1/4 of telopodite length, broadly term-
inal, poorly demarcated from prefemur, bending
strongly laterad; prefemur with many hairs arranged
continuously along inner surface; cyphopod valves
of females not modified, distal corners not pro-
duced; receptacle moderate-size, sides not curving
round operculum.
Color in life. — Peritremata brick red; dorsum
black, metaterga with variably broad, brick red to
orange stripes along caudal margins connecting con-
colorous peritrematal markings; collum with brick
red to orange stripes along anterior and caudal
margins. These are the colors I noted while collecting
melacis in 1986. They correspond to Chamberlin and
Mulaik’s description (1941), who noted that the
stripe expands along the midline.
84 EURYMERODESMID MILLIPEDS
Holotype. — Body highly fragmented, length un-
measurable, maximum width 3.2 mm.
Somatic features similar to hispidipes, with following ex-
ceptions:
Width across genal apices 2.4 mm, interantennal isthmus
0.8 mm. Antennae reaching back to just beyond 2nd tergite,
relative lengths of antennomeres 4>2>3>5>6>1>7. Genae
with faint central impressions. Facial setae as follows: epicranial,
interantennal, and genal absent, clypeal about 10-10, labral about
14-14. Process of mandibular stipes moderate-size, curving slightly
inward, narrowly rounded apically (Fig. 177).
Collum extending slightly below ends of following tergite.
Paranota depressed, continuing slope of dorsum; posterior corners
rounded through segment 7, blunt on 8-12, becoming progressively
more acute caudally.
5th sternum with low, rounded lobes between both leg pairs,
caudal ones higher; 6th sternum with slightly longer lobes between
anterior legs and strong, densely hirsute projections between 7th
legs. Postgonopodal sterna with rounded, widely separated knobs
between 9th legs; remaining sterna becoming progressively flatter
and more plate-like caudally, with variable transverse grooves
originating between leg pairs and hair patches adjacent to coxae,
latter becoming progressively sparser caudally. 2nd coxa with
minute anterior projection, caudal lobe absent. Prefemoral lobes
extending through segment 15.
Gonopodal aperture (Fig. 178, not this specimen), broadly
trapezoidal, without caudolateral pouches, with moderate-size
anterior indentation, 1.6 mm wide and 0.8 mm long at maxima;
anterior indentation broadly triangular, apically rounded; sides not
divided, nearly straight, leaning slightly over opening, becoming
slightly but progressively more elevated caudad, rims essentially
smooth; caudolateral corner sharp, distinct, obtuse; caudal margin
continuous with sides through caudolateral corner, angling strong-
ly caudomediad, level for part of length with slight suggestion of
rounded elevation, then tapering to metazonal surface and con-
tinuing flush with latter across midline; margins irregularly hirsute
with long hairs arising from rim or short distance down outer sur-
faces of sides and overhanging opening, with tufts on sides of
anterior indentation, lateral hairs becoming thicker caudally,
densest on rounded part beyond caudolateral corner, diminishing
medially. Gonopods in situ (Fig. 178, not this specimen) with
telopodites crossing or overlying each other in midline, extending
slightly beyond caudal margin of aperture and terminating above
sternum between 8th legs, apices bent laterad. Gonopod structure
as follows (Figs. 179-180): Telopodite short, terminating well
before level of distal extremities of hairs. Prefemur proportionate-
ly long, about 3/4 of telopodite length, leaning slightly over coxa,
with a continuous row of widely but regularly spaced hairs along
inner surface and distomedial tuft of about 5 hairs. Acropodite
short and broadly terminal, about 1/4 of telopodite length,
smoothly continuous with, and poorly demarcated from prefemur,
bending strongly laterad, sides tapering continuously to blunt tip.
Male paratypes. — The male paratypes are all tightly coiled,
and it is impossible to unroll and straighten most for measurement
without breakage. Many shatter, rendering even length approx-
imations impossible. The smaller specimens are more flexible and
give the best measurements, but even they usually split between
two pregonopodal segments. Approximate measurements on three
specimens are length 22.5 mm long, 3.2 mm wide, W/L ratio
14.2%; 18.0 mm long, 3.5 mm wide, W/L ratio 19.4%; and 14.0
mm long, 2.1 mm wide, W/L ratio 1.50%.
The elevation of the sides of the Edinburg males varies, some
being nearly flush with the metazonal surface. The caudolateral
comer varies, some being nearly flush with the metazonal surface.
The caudolateral corner varies from obtuse to squared, or about a
right angle, and the most significant differences concern the length
of the elevation along the caudal margin and the abruptness of the
slope to the metazonal surface. In a few males the elevation stops
abruptly at the caudolateral corner, whereas it extends nearly to
the midline with a long, gradual slope in others. The rim of the
elevation is darkened in most males, and the opening extends
significantly caudad in the midline in a few. In all specimens, the
elevated part of the caudal margin is level with that of the sides,
although males with longer elevations display a suggestion of a
slightly higher rounded area just behind the caudolateral corners.
Paratypes from Kerr, Bandera, and Kendall counties closely
resemble those from Edinburg. The sides are more linear in some
and do not lean over the opening, and the rim is flared outward in
a few.
Female paratypes. — An unbroken specimen from Edinburg
measures 18.2 mm long, 3.0 mm maximum width, W/L ratio
16.5%. Agreeing essentially with males in somatic features, with
following exceptions: interantennal setae 1-1, frontal 1-1. Man-
dibular stipes with minute projection, acuminate. Sterna flat and
unmodified, glabrous.
Cyphopodal aperture elliptical, margins flush with metazonal
surface. Cyphopods in situ with edges of valves visible in aperture,
receptacle hidden under lateral margin. Valves (Figs. 181-182)
oriented transversely in aperture, anterior valve slightly larger with
rounded corner, moderate-size, extending dorsad on open side,
distal margins not produced. Receptacle moderate-size, glabrous,
located laterad to valves, sides not curving around operculum. Lat-
ter relatively large, clearly visible below open, dorsal, side of
valves.
Variation. — Eurymerodesmus melacis is a
brightly colored species with brick-red peritremata
and metatergal stripes through most of the range and
a mottled black base color. As noted by Chamberlin
and Mulaik (1941) the stripes may be of uniform
width or broader in the midline. In the western range
periphery, colors assume more of an orange hue,
becoming reddish-orange in Kinney County and
orange in Tom Green County. The specimens I col-
lected in the latter were so broadly striped and the
pigment was so intense that they appeared entirely
orange from a distance; actually the metaterga were
almost entirely orange with the prozonum mottled
black. Chamberlin and Mulaik (1941) described the
antennae as pale with the ultimate and distal half of
the penultimate articles black. I did not observe this
pattern, and the antennae, as well as the genae and
clypeus of the Kinney County specimens, were
uniformly pale. These individuals exhibited mottled
brown pigmentation on the frons, epicranium, and
interartennal regions, as opposed to black on the
segments, and the mandibular stipes were a lighter
mottled brown. Chamberlin and Mulaik (1941)
described the venter, sides, and legs as yellowish in
the type specimens. The venter was white on the Kin-
ney County individuals, and the legs possessed
ROWLAND M. SHELLEY 85
brown flecks on the dorsal surfaces, extending onto
the sides.
Because they were so tightly coiled and therefore
broke or shattered with handling, measurements
were impossible on most specimens of both sexes. A
few recent molts had not fully hardened and were
pliable.
Table 7. Size variation in E. melacis.
Males Females
County Length Width W/L Length Width W/L
San Patricio 26.7 3.6 13.5%
Terrell 17.1 2.6 15.2%
Medina 25.1 3.7 14.7% 23.1 3.8 16.4%
Medina 21.2 3.0 14.1% 22.6 3.6 15.9%
Medina 21.8 3.3 15.1% 21.9 2.8 12.7%
Medina 23.2 3.8 16.4% 20.5 2.7 13.2%
Medina 21.8 2.9 143% 20.3 2.9 14.3%
Aperture variation is as discussed for paratypes.
The telopodites appear slightly longer and narrower
in scattered individuals, and the distances they ex-
tend caudad also vary depending upon how tightly
the animal is coiled and how tightly the segments are
compressed. In some the anterior half of the aper-
ture is covered by the 6th segment. The lobes be-
tween the caudal legs on segment 6 are sharply
conical in many individuals from Bexar and Jim
Wells counties, being either subequal to or longer
than the widths of the adjacent coxae. The process
of the mandibular stipes is also quite broad in
specimens from Karnes and Frio counties.
The Terrell County male, collected in a cave at the
western periphery of the species and family, is by far
the smallest adult male of melacis and the only one
totally pallid, with no evidence of pigmentation. All
other cave specimens show evidence of the striped
color pattern even after years in preservative. The
telopodites are also much shorter in this specimens
and the acropodites are more linear and not bent as
strongly laterad. Additionally, this male was flexible
and easily unrolled for measurement; perhaps it is
somewhat cave adapted.
The cyphopod valves are mostly uniform, being
squared in profile with rounded corners. In the
female from Helotes, Bexar County, the distal cor-
ners curve into short rounded lobes, and the corners
are also slightly produced and rounded in one from
Tom Green County.
Ecology. — In February 1986 melacis was com-
mon under small branches and shallow leaf layers in
small oak thickets at the Welder Wildlife Refuge,
San Patricio County. In Tom Green County, I
found three specimens under large logs beside the
MEM. AMER. ENT. SOC., 37
Concho River. Perhaps because of the unseasonable
drought, I did not find melacis in the lush and osten-
sively desirable bottomland forests along the Rio
Grande in Cameron and Hidalgo counties, but it
may be a desertine species actually preferring arid
conditions. Thus, these forests may be unsuitable for
melacis, but it should occur away from this habitat
in drier, ‘‘inland’’ areas of Cameron, Hidalgo, and
other south Texas counties. Notations on labels with
preserved samples are as follows: under rocks (Frio,
Kerr, and Uvalde cos.), under cow dung (Bee and
Live Oak cos.), under logs (Wilson and Atascosa
cos.), and under cardboard in ‘‘barditch’’ (Zavala
Co.).
Eurymerodesmus melacis has been taken from 11
caves and can probably be anticipated in any cave in
the range, where they are accidental inhabitants
found primarily in the open on silt in entrance areas.
Except for the specimens from Longley Cave, Terrell
county, none vary significantly from epigean
populations. As suggested by the anatomical dif-
ferences, these specimens may be true troglobites,
since the opening to this cave is situated so as to vir-
tually preclude the accidental entrance of epigean
forms (J. Reddell, pers. comm.).
Distribution. — The Gulf Coastal Plain and
eastern Edwards Plateau of central and southern
Texas as far west as eastern Terrell County, extend-
ing northward from the Rio Grande along the Gulf
of Mexico to about the San Antonio River, and in-
land as far north as southern Hays and western
Taylor counties (Fig. 176). Collections are con-
centrated along an arc running from Corpus Christi
to San Antonio to Del Rio, and melacis appears rare
in south Texas, perhaps an artifact reflecting sam-
pling along major highways before the interstate era.
Thus, I shade or outline all of south Texas in figures
1 and 213-214. Specimens were examined as follows:
TEXAS: Taylor Co., 25 mi. SW Abilene, M,
March 1944, H. S. Dybas (FMNH). Tom Green
Co., 17 mi. NW San Angelo, Wardlaw Ranch, M,
F, 26 April 1967 and F, 27 November 1967, G.
Brown (FSCA); and San Angelo, along Rio Concho,
2M, F, 27 February 1986, R. M. Shelley (NCSM).
Concho Co., 10 mi. W Eden, F, 15 December 1939,
collector unknown (NMNH). Kimble Co., Junction,
M, 8 March 1986, R. W. Sites (NCSM); 4 mi. SW
Junction, M, 24 March 1978, O. F. Francke, T. B.
Hall, J. V. Moody (NCSM); and 13 mi. SW Junc-
tion, M, 2F, 24 March 1978, O. F. Francke, T. B.
Hall, and J. V. Moody (NCSM). Terrell Co.,
Longley Cave W of Pandale (Val Verde Co.), M, F,
27 June 1963, J. Reddell (FSCA). Val Verde Co.,
86 EURYMERODESMID MILLIPEDS
near Eliott Spider Cv., 8 mi. N Comstock, Marshall
Ranch, M, 1 January 1965, J. Reddell (FSCA); and
Del Rio, F. 9 October 1932, O. F. Cook (FSCA).
Edwards Co., 11 mi. NE Rock Springs, M, 24
March 1978, O. F. Francke, T. B. Hall, and J. V.
Moody (NCSM). Kinney Co., 21 mi. N Brackett-
ville, 6M, 12 January 1985, S. A. Stockwell and J.
M. Steller (NCSM). Real Co., 13 mi. N Leakey, M,
23 March 1978, O. F. Francke, T. B. Hall, and J. V.
Moody (NCSM); and Rio Frio, Prade Ranch, M, F,
21 April 1962, R. O. Albert (FSCA). Uvalde Co., 26
mi. NW Uvalde, F. 12 September 1940, S. and D.
Mulaik (NMNH); Laguna, 2M, 25 December 1941,
collector unknown (AMNH); 12 mi. NW Uvalde,
5M, F, 23 March 1978, O. F. Francke, T. B. Hall,
and J.V. Moody (NCSM); Knippa, M, F, 7 April
1937, K. P. Schmidt (FMNH); Sabinal, 2M, 2F, 4
juvs., 11 January 1933, H. C. McNamara (FSCA);
Secret Valley Cv., exact location unknown, M,
February 1984, R. M. Waters (NCSM); Tampke
Ranch Cv., 0.8 mi. NE Uvalde, 2M, 11 February
1966, J. Reddell and D. McKenzie (FSCA); and
Cedar Brake Cv., 25 mi. NE Uvalde, M, 25 October
1965, J. Reddell (FSCA). Zavala Co., 1 mi. E
Batesville, 16M, F, 15 March 1982, J. C. Coken-
dolpher (FSCA). Kerr Co., Raven Ranch, ca. 12 mi.
S Kerrville, off TX hwy. 173 north of Camp Verde,
4F, August 1939, 25M, 2F, 3 juvs., December 1939;
4F, juv. June-August 1940; and juv., June 1941, all
by S. and D. Mulaik (NMNH); 14 mi. S Kerrvilie,
4M, 2F, 9 October 1976. J. C. Cokendolpher
(MSU); and Seven Room Cv., 8 mi. SW Kerrville, 2
juvs., January 1966, J. Fish and J. Reddell (FSCA).
Bandera Co., locality unknown, 9M, F, December
1939, S. and D. Mulaik (NMNH); 7 mi. N Medina,
M, date unknown, S. and D. Mulaik (NMNH); and
Bandera, Shaw Ranch, 4M, 2 February 1962, G.
Marion (FSCA). Medina Co., 3 mi. ENE Mico, F.
19 March, year unknown, V. Roth (NCSM);
D’Hanis, 13M, 16F, 3 juvs., 18 October 1932, O. F.
Cook (NMNH); Windmill Cv., exact location
unknown, M, 10 November 1984, S. J. Hardin and
J. L. Ivy (NCSM); Koch Cv., 9 mi. N Hondo, M, 7
October 1963, J. Reddell and D. McKenzie (FSCA);
and Boehme’s Cvy., exact location unknown, 19
December 1984, J. L. Ivy (TMM). Frio Co., Pear-
sall, 2M, 4 February 1961, D. R. Whitehead (RLH).
Kendall Co., locality unknown, 9M, 15F, December
1939, S. and D. Mulaik (NMNH); and Boerne, 2M,
3F, 9 April 1911, W. P. Carr (NMNH) and 6M, 2F,
December 1939, S. and D. Mulaik (NMNH). Hays
Co., Fern Bank Springs nr. Wimberly, F, 1 April
1961, R. Highton (RLH); San Marcos, F, 22 April
1941, K. P. Schmidt (FMNH); and Swift’s Cv., ex-
act location unknown, 26 June 1982, D. L. Pate
(TMM). Comal Co., between Startzville and Tom
Cr., ca. 12 mi. NW New Braunfels, 2F, 15 March
1983, C. S. Lieb (NCSM); Brehmmer Cv., ca. 5 mi.
W New Braunfels, F, 19 March 1960, W. J. Gertsch
(AMNH); and New Braunfels, F, juv., 29 November
1942, E. S. Ross (CAS), M, 18 December 1954, L.
Hubright (RLH), and Landa Park, 4F, 19 June
1964, N. B. Causey (FSCA); and 2 mi. N Bergheim,
2M, 22 February 1986, S. J. Hardin (TMM). Bexar
Co., 20 mi. N San Antonio, 5M, 5F, 13 December
1942, E. S. Ross (CAS); Helotes, F, 6 April 1966, D.
A. Rossman (FSCA); San Antonio, 2M, F,
September, day, year, and collector unknown
(MCZ), M, March 1925, A. H. Welolet (NMNH),
Yturri Est., 15M, 10F, December 1905, O. F. Cook
(NMNH), San Antonio R. at Roosevelt Ave., 16M,
12F, 23 October 1954, L. Hubricht (RLH), and
Jessamine St., 11M, 3F, 31 October 1959, J. C.
Loomis (RLH); 6 mi. S San Antonio, 4M, 2F, 7
December 1965, T. Stewart (FSCA); Camp Bullis,
15 mi. NW San Antonio, 2M, F, 27 January 1952,
H. R. Bullock (AMNH) and 2M, F, 24 April 1966,
B. Russell and D. McKenzie (FSCA); Black Cat Cv.,
15 mi. NNE San Antonio, M, 2 December 1984, S.
J. Harden and J. L. Ivy (NCSM); and Intermittant
Spgs. nr. Leon Spgs., F, 3 November 1986, S. J.
Harden and C. Lindblom (TMM). Wilson Co., 4
mi. SE Poth, along Marcelinas Cr., M, 2F, 1 April
1961, J. F. Quinlan (FSCA); and 3 mi. W Falls City
(in Karnes Co.), F, date and collector unknown
(FSCA). Atascosa Co., Campbellton, M, FF, 15
May 1961, J. F. Quinlan (FSCA). Karnes Co., 1 mi.
NW Falls City, M, 3F, 9 April 1961, J. F. Quinlan
(FSCA); Falls City, M, F, 29 October 1961, J. F.
Quinlan (FSCA); and Karnes City, 2M, 4F, 14
November 1959, N. B. Causey (FSCA). Golida Co..,
Goliad St. Pk., 3F, 17 April 1955, L. Hubright
(RLH). Bee Co., Beeville, F, 11 November 1895,
collector unknown (NMNH) and M, 28 December
1941, R. Perita (AMNH). Live Oak Co., Largato,
2M, 2F, 18 February 1961, R. O. Albert (FSCA); 6
mi. SE Whitsett, 3M, 11 December 1984, R. S. Zack
(WASU); and George West, 2M, 6F, 3 December
1960, collector unknown (FSCA). Refugio Co.,
along Aransas R., F, May 1952, collector unknown
(FSCA). San Patricio Co., 7 mi. N Sinton, Welder
Wildlife Ref., 40M, 26F, 8 juvs., February and
December 1961, December 1965, and November
1970, R. O. Albert (FSCA), F, 18 October 1967, C.
Parrish (CAS), and 8M, 5 juvs., 21 February 1986,
R. M. Shelley (NCSM); and 4-5 mi. SW Mathis, 9M,
ROWLAND M. SHELLEY 87
OF, 3 juvs., March 1961 and January 1962, R. O.
Albert (FSCA). Jim Wells Co., along Nueces R.,
10M, 6F, 1-3 April 1962, R. O. Albert (FSCA).
Duval Co., unknown locality in NE corner, 2M, 2F,
4 April 1962, R. O. Albert (FSCA); San Diego, 9M,
F, 8 January 1961, R. O. Albert (FSCA); and 5 mi. S
San Diego, M, 2F, 17 February 1961, R. O. Albert
(FSCA). Hidalgo Co., Edinburg, 57M, 21F, 12
juvs., December 1939, S. and D. Mulaik (NMNH)
TYPE LOCALITY.
Additional literature records for melacis deemed
valid include McQueeney and Schertz, Guadalupe
County (Loomis 1959).
Remarks. — Hoffman (1981) noted that some
published Edinburg spider records are patently mis-
labeled, and since that for melacis is around 100
miles south of the closest known locality, in Duval
County, the question as to whether it too may be er-
roneous assumes considerable importance, since
Edinburg is now the type locality. The afore-
mentioned absence of melacis from the forests of the
Rio Grande Valley in contrast to its abundance in
small thickets in San Patricio County is not troubling
because melacis does not seem to be a forest-inhabit-
ing species. The site where I collected it along the Rio
Concho in San Angelo, Tom Green County, was
grassland with a few widely scattered trees and no
leaf litter, which I think reflects its habitat
preferences. Since Edinburg is away from these
woodlands, it is an ecologically plausible site for
melacis. That no specimens have been collected in
other counties in the tip of Texas, particularly inter-
vening ones like Brooks and Jim Hoff, I think
reflects their relative isolation. There are few towns
or places to look for millipeds, and collectors passing
through may have set their sights on the Rio Grande
forests instead of the arid grasslands away from the
river. Edinburg is not so far removed from other
known sites as to be implausible, and there could
also be an allopatric population there analogous to
those of b. birdi in San Patricio, Wilson, Bastrop,
and Bell counties. Thus, although confirmation with
fresh material is desirable, Edinburg is a logical site
for melacis.
As the southernmost species known essentially to
the Rio Grande in Webb, Kinney, and Val Verde
counties, melacis may be analogous to Abacion tex-
ense and occur south of the border in northern parts
of Tamaulipas, Nuevo Leon, and/or Coahuila
provinces, Mexico (Shelley 1984a). Mexican and
American biologists working in these areas should be
alert for this small, bright red milliped.
Eurymerodesmus melacis is also the westernmost
MEM. AMER. ENT. SOC., 37
eurymerodesmid, and its occurrence on the eastern
Edwards Plateau in central Texas suggests occur-
rence farther west in the Davis Mountains, the
Permian Basin, the panhandle around Lubbock, and
possibly even the southeastern corner of New Mex-
ico. I therefore spent a week in February 1986
searching for it around Lubbock and in the Davis
Mountains, with no success. I searched habitats
similar to those in which I found it at San Angelo
and the Welder Wildlife Refuge, but the only
millipeds encountered were parajulids. Future
discoveries are always possible, but I think the cur-
rent western limits are close to the actual ones.
Eurymerodesmus dactylocyphus, new species
Figs. 177, 183-189, 212
Type specimens. — Male holotype and two male
and three female paratypes (FSCA) collected by R.
O. Albert, 1 December 1963, at Maxwell, Caldwell
Co., TX. One male and one female paratypes
deposited in NCSM.
Diagnosis. — A small to moderate-size species;
males characterized by long mandibular projection;
aperture very large, occupying entire breadth of seg-
ment in ventral view; sides not divided, without
caudolateral pouches, with variable caudal elevation
continuing around caudolateral corners ranging
from slight rise to long flange; caudal margin
descending rapidly to metazonal surface; telopodite
short, wholly enclosed within aperture; acropodite
short, about 1/4 of telopodite length, broadly term-
inal, poorly demarcated from prefemur, bending
directly dorsad; cyphopod valves of females with
distal corners of valves produced into variable pro-
jections, ranging from small nubbinlike tubercles to
long, flexible dactyliform projections, extending well
beyond cypopodal aperture and overlapping suc-
ceeding two segments in situ; receptacle small,
submerged in valves, sides not curving around
operculum.
Color in Life. — Unknown.
Holotype. — Length 19.7 mm, maximum width
3.0 mm, W/L Ratio 15.2%.
Somatic features similar to hispidipes, with following excep-
tions:
Width across genal apices 1.8 mm, interantennal isthmus 0.6
mm. Antennae reaching back to caudal margin of 2nd tergite,
relative lengths of antennomeres 2>3>6>4>5>1>7. Genae
without impressions. Facial setae as follows: epicranial, inter-
antennal, and genal absent, subantennal 1-1, frontal 1-1, clypeal
about 10-10, labral about 14-14. Process of mandibular stipes
relatively long, subtriangular, narrowly rounded apically (Fig.
183).
88 EURYMERODESMID MILLIPEDS
Figs. 183-189. Eurymerodesmus dactylocyphus. 183, projection of right mandible of holotype, lateral view. 184, aperture and
gonopods in situ of holotype, ventral view. 185, left gonopod of the same, lateral view. 186, telopodite of the same, medial view. 187,
cyphopods in situ, ventral view of female paratype. 188, left cyphopod of the same, medial view. 189, the same, anterior view. Scale lines
for figs. 184 and 187 = 1.00 mm. Line for other figs. = 0.30 mm for 183 and 188-189, and 0.50 mm for 185-186.
ROWLAND M. SHELLEY 89
Collum extending slightly below ends of following tergite.
Paranota moderately depressed, continuing slope of dorsum,
peritremata somewhat flattened; posterior corners rounded
through segment 9, blunt on 10-12, becoming progressively more
acute posteriorly.
5th sternum with small paramedian lobes between leg pairs; 6th
sternum with broader, higher elevated areas. Postgonopodal sterna
with low paramedian knobs between 9th legs; remaining sterna
becoming progressively flatter, more plate-like, and less hirsute
caudally, with variably distinct transverse grooves between pairs
and broad central depressions, hairs diminishing after segment 9.
2nd coxa with large, rounded anterior and posterior lobes.
Prefemoral lobes extending through segment 10.
Gonopodal aperture (Fig. 184) very large, nearly circular but
broadly ovoid, without caudolateral pouches, with strong anterior
indentation, 1.9 mm wide and 1.0 mm long at maxima, extending
nearly to anterior edge of segment; anterior margin slightly
elevated, in form of narrow rim continuing around anterolateral
comers onto sides, anterior indentation broad, apically rounded;
sides not divided, indented slightly and elevated at anterolateral
corner, then linear for short distance, caudal half angling mediad,
anterior half continuous with elevated anterior margin, elevation
increasing rapidly and dramatically just beyond midlength, extend-
ing into long, broad, ventrally directed flange at caudolateral cor-
ners, rims mostly smooth, becoming slightly irregular on flange;
caudolateral corners slightly angular, greatly elevated; caudal
margin continuous with caudal elevation of sides, flange narrow,
inner surface curved or cupulate, very long, extending nearly to
midlength of podomeres, terminating abruptly a short distance
beyond corner, margin low medially, curving across midline and
merging with lobes between 9th legs; margins irregularly hirsute,
with tufts of three to four hairs on anterior indentation, scattered
hairs on rim of anterolateral corners and sides, a few arising from
inner surfaces of flanges, and three to four on low, central part of
caudal margin. Gonopods in situ (Fig. 184) with telopodites ex-
tending nearly directly caudad, angling slightly mediad, ter-
minating short of caudal margin, wholly enclosed by aperture,
apices directed dorsad. Gonopod structure as follows (Figs.
185-186): Telopodite relatively short, terminating well before level
of distal extremities of hairs. Prefemur proportionally long, about
3/4 of telopodite length, leaning over coxa, broad basally, tapering
slightly distad, with three rows of continuous, dense, regularly
spaced hairs along inner margin and distomedial tuft of 8 hairs.
Acropodite short and broadly terminal, about 1/4 of telopodite
length, smoothly continuous with, and poorly demarcated from
prefemur, continuing distal curvature of latter, curving dorsad,
sides narrowing rapidly to subacuminate tip.
Male paratypes. — The male paratypes agree with the holotype
in all particulars.
Female paratype. — Length 22.1 mm, maximum width 2.8
mm, W/L ratio 12.7%. Agreeing essentially with males in somatic
features, with following exceptions: Mandibular stipes with corner
slightly produced, blunt. Sterna flat and unmodified, glabrous.
Cyphopodal aperture large, rectangular, caudal margin slightly
elevated above metazonal surface, indented slightly anteriad in
midline. Cyphopods in situ (Fig. 187) with valves oriented along
body axis, protruding slightly through opening, dactyliform pro-
jections extending well beyond aperture and overlapping legs of
segments 3 and 4, directed either ventrad or laterad, appearing like
supernumerary legs. Valves (Figs. 188-189) large, unequal, lateral
one larger, distal corners extending into long, flexible, widely
separated dactyliform projections, subequal to or longer than
valves proper, projecting directly ventrad basally, curving caudad
MEM. AMER. ENT. SOC., 37
at midlength, apically blunt. Receptacle small and inconspicuous,
located on caudal side of, and submerged in, valves. Operculum
small, located under anterior corner of valves.
Variation. — As described for the holotype,
males from Hays County exhibit large, cupped aper-
ture flanges, which drop nearly perpendicularly to
the metazonal surface on the caudal margin just
beyond the caudolateral corner. However, in Bell,
Bosque, and Victoria counties the elevation is only
slightly higher than the metazonal surface. The
paramedian sternal projections between the 9th legs,
very large and coalesced in the Victoria males, are
subequal to the widths of the adjacent coxae, which
have hirsute ventrodistal lobes. In Bell County, the
9th sternal knobs are separate. On the gonopods, the
prefemur tapers distad in the Hays County males
and possesses a slight shoulder on the outer margin;
thus, the acropodite is more strongly demarcated
from the prefemur. In Bell County, the acropodite
curves strongly dorsad.
The longest dactyliform cyphopodal projections
occur in females from Hays and Caldwell counties.
They are shorter, more variable, and more unequal
in those from Bosque and Victoria counties. In the
latter, the projections vary from small nubbinlike
tubercles or papillae to digitiform elongations about
half as long as the valves. Both conditions can occur
on a single cyphopod.
Ecology. — Some of the Victoria specimens were
encountered in sedge-grass roots in a sandy region
and on a shaded river bank. A male from Bell Coun-
ty was collected along a fence.
Distribution. — The eastern fringe of the Central
Plains to the Gulf of Mexico between the Brazos and
Guadalupe Rivers (Figs. 176, 212). Specimens were
examined as follows:
TEXAS: Bosque Co., Meridian St. Pk., 5M, 26
December 1979, G. Spicer (NCSM) and 2M, 2F, 5
October 1980, G. Spicer (NCSM). Bell Co., Belton,
M, January 1931, collector unknown (NMNH); and
1 mi. W Belton, 2M, 24 December 1939 and 1941,
collector unknown (AMNH). Williamson Co.,
Three Mile Cv., ca. 3 mi. W Round Rock, M, 1
December 1962, R. O. Albert (TMM). Hays Co.,
Fern Bank Springs nr. Wimberly, 2M, 1 April 1961,
R. Highton (RLH); and San Marcos, F, 22 April
1941, K. P. Schmidt (FMNH). Caldwell Co., Max-
well, 4M, 4F, 1 December 1963, R. O. Albert
(FSCA) TYPE LOCALITY. Victoria Co., Victoria,
50 M, 35 F, 7 juvs., December 1905, August 1906,
23 November 1911, 24 November 1912, 10 March
1913, and 11 December 1915, J. D. Mitchell
(NMNBH).
90 EURYMERODESMID MILLIPEDS
Remarks. — A female of melacis was found in
the samples from Wimberly and San Marcos, sug-
gesting that their ranges abut in southern Hays
County. The male from Belton was retrieved from a
sample of b. birdi, suggesting that the two species are
sympatric in Bell County.
The gonopods of dactylocyphus appear
ridiculously small in contrast to the enormous aper-
ture. In some males the coxa is tucked under the
anterior margin and appears swallowed by the aper-
ture, which is mostly empty and thus filled with
debris. The species has by far the broadest aperture
in the genus, filling essentially the breadth of the
metazonum in ventral view and extending laterad
nearly to the level of the paranota. In some males, it
compresses the prozonum into a remnant, an ex-
tremely narrow sliver, along the anterior segmental
margin, the minimum necessary to give form to the
aperture and hold it together. This is a striking
feature of males, but I found the cyphopod projec-
tions equally sensational and decided to name the
species for this distinctive aspect of females. These
projections vary greatly, as described above, and the
long ones are convergent with those in compressus.
However, the projections in dactylocyphus differ in
being flexible, as opposed to stiff and rigid. The pro-
jections are foreshortened in figure 187, appearing
shorter than they really are, fully as long as the next
two segments combined. In some females they pro-
ject outward between the appendages of the follow-
ing segment, thus appearing at first glance as addi-
tional legs, when seen in lateral view.
Eurymerodesmus sanbernardiensis Causey
Figs. 176, 190-195, 212
Eurymerodesmus sanbernardiensis Causey, 1952a:174, figs. 6-7.
Chamberlin and Hoffman, 1958:81.
Type specimen. — Female holotype (ANSP) col-
lected by N. B. Causey, 3 January 1952, along the
San Bernard River near the crossing of US highway
59, Fort Bend Co., TX. The cyphopods and segment
3 are missing from the holotype and their separate
vial at the ANSP, and are lost. My attempt to collect
topotypes in February 1986 was unsuccessful. Since
its cyphopods roughly resemble those in Causey’s il-
lustrations (1952a, figs. 6-7), I assign sanbernardien-
sis to the form occurring at Wharton, Wharton
County, approximately 12 miles south of the type
locality.
Diagnosis. — A small to moderate-size species;
males characterized by short, triangular mandibular
projection; aperture moderate-size, much narrower
/
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a NG A
Y D>
/,
Figs. 190-195. Eurymerodesmus sanbernardiensis. 190, pro-
jection of left mandible of male from Colorado Co., TX, lateral
view. 191, aperture and gonopods in situ of the same, ventral view.
192, left gonopod of the same, lateral view. 193, telopodite of the
same, medial view. 194, left cyphopod of female from Colorado
Co., lateral view. 195, the same, anterolateral view. Scale line for
fig. 191 = 1.00 mm. Line for other figs. = 0.40 mm for 190 and
195, and 0.50 mm for 192-194.
than ventral segmental width; sides not divided,
without caudolateral pouches, elevating slightly
around caudolateral corner, descending to
metazonal surface along caudal margin; telopodite
short, at most barely overlapping caudal margin of
aperture; acropodite short, about 1/4 of telopodite
length, broadly terminal, poorly demarcated from
prefemur, curving broadly dorsad; prefemur with
many hairs arranged continuously along inner
margin; cyphopod valves of females with distal
corners produced and rounded; receptacle small,
partly submerged in valves, sides not curving around
operculum.
Color in life. — Paranota orange; rest of color
unknown. Causey (1952a) stated that the dorsum
was mottled with orange, but that the color was “‘not
completely developed.”’
Male from Wharton Co. — Length 22.7 mm,
maximum 3.1 mm, W/L ratio 13.7%.
Somatic features similar to hispidipes, with following excep-
tions:
Width across genal apices 1.8 mm, interantennal isthmus 0.8
mm. Antennae reaching back to midlength of 3rd tergite, relative
lengths of antennomeres 2>3>4>6>5>1>7. Genae without
impressions. Facial setae as follows: epicranial, interantennal,
genal, and frontal absent, clypeal about 11-11, labral about 15-15.
Projection of mandibular stipes short, triangular (Fig. 190).
ROWLAND M. SHELLEY 91
Collum extending slightly below ends of following tergite.
Paranota moderately depressed, mostly continuing slope of dor-
sum; posterior corners rounded through segment 8, blunt on 9-13,
becoming progressively more acute posteriorly.
5th and 6th sterna with hirsute elevated areas between leg pairs,
without lobes. Postgonopodal sterna flattened, plate-like, with
variably complete transverse grooves originating between leg pairs
on all segments and variably broad central depressions, with hair
patches adjacent to leg coxae becoming sparser caudally. 2nd coxa
with minute anterior swelling, without caudal lobe or swelling.
Prefemoral lobes terminating on 12th legs.
Gonopodal aperture (Fig. 191, not this specimen) broadly
ovoid, without caudolateral pouches, with broad anterior indenta-
tion, 1.2 mm wide and 0.8 mm long at maxima; anterior indenta-
tion moderately long, broadly triangular, apically acuminate; sides
curving broadly, undivided, not leaning over opening, anterior
half flush with metazonal surface, elevating at midlength and be-
coming slightly and progressively higher to caudolateral corner,
rims essentially smooth; caudolateral corner obtuse, indistinct;
caudal margin continuous with sides at caudolateral corner, taper-
ing gradually to metazonal surface, extending slightly caudad in
midline; margins glabrous. Gonopods in situ (Fig. 191, not this
specimen) with telopodites angling caudomediad, terminating
before level of caudal aperture margin, apices directed dorsad.
Gonopod structure as follows (Figs. 192-193): Telopodite short,
terminating well before level of distal extremities of hairs.
Prefemur proportionally long, about 3/4 of telopodite length,
curving over coxa, curve increasing distad, moderately broad
basally, sides tapering distad, with two rows of continuous,
regularly spaced hairs on inner surface and distomedial tuft of 5-6
hairs. Acropodite short and broadly terminal, about 1/4 of
telopodite length, smoothly continuous with, and poorly demar-
cated from prefemur, continuing broad dorsal curvature of
prefemur, sides tapering to subacuminate tip.
Female from Wharton Co. — Length 21.9 mm, maximum
width 2.9 mm, W/L ratio 13.2%. Agreeing closely with males in
somatic features, with following excepticns: frontal setae 1-1,
genal 4-4. Mandibular stipes with short, rounded projection.
Sterna flat and unmodified, glabrous.
Cyphopodal aperture elliptical, caudal margin slightly elevated
above metazonal surface, indented slightly anteriad at midline.
Cyphopods in situ with valves oriented obliquely, distal corner
protruding through opening. Valves (Figs. 194-195) small, un-
equal, caudal valve slightly larger, distal corners slightly produced
and rounded. Receptacle small, inconspicuous, partly submerged
in valves, located below latter on lateral side. Operculum small,
located at bases of valves on medial side.
Variation. — The sides of the aperture are a little
higher on two of the four males from Wharton
County. Depending upon how submerged the coxae
are in the aperture, the telopodites may or may not
extend beyond the caudal margin and overhang the
sternum between the 9th legs in situ. Specimens from
Brazoria and Colorado counties agree closely with
those from Wharton except for a slight bisinuate
curvature on the acropodites of those from Col-
orado County. In these males the structure resembles
a tarsal claw, since it is moderately long, similarly
curved, and arises from a hirsute prefemur. Females
are closely similar throughout the range.
MEM. AMER. ENT. SOC., 37
Ecology. — The type locality, which I visited in
February 1986 in an unsuccessful search for
topotypes, is a narrow bottomland forest that
merges into grassland a short distance away from the
river. There were plenty of logs, debris, and other
potential milliped habitat, but no eurymerodesmids.
The female that I collected at Richmond, Fort Bend
County, was found under leaves at a rest area near
the Brazos River.
Distribution. — A small area of the Gulf Coastal
Plain of southeastern Texas between the Brazos and
Colorado Rivers, extending from the Gulf of Mexico
to about 94 miles inland (Figs. 176, 212). Specimens
were examined as follows:
TEXAS: Austin Co., New Ulm, 9M, 6F, 3
February 1963, R. O. Albert (FSCA). Colorado Co.,
Columbus, 3M, 23 December 1961, C. L. Redus
(FSCA). Fort Bend Co., ca. 2 mi. S Kendleton,
along US hwy. 59 at San Bernard R., F, 3 January
1952, N. B. Causey (ANSP) TYPE LOCALITY;
and Richmond, F, 20 February 1986, R. M. Shelley
(NCSM). Wharton Co., Wharton, Bank of Col-
orado R., 4M, 10F, 4 juvs., December 1905, collec-
tor unknown (NMNH). Brazoria Co., Old Ocean
and 2.4 mi. S Old Ocean, 2M, juv., 23 December
1961, R. O. Albert (FSCA).
Remarks. — In his key to Texas species of
Eurymerodesmus, Loomis (1976) stated that the
valves of female cyphopods of sanbernardiensis are
of unequal length. This was probably based on
Causey’s description (1952a), which I think may
have been of an immature female as judged by the
size of the holotype versus Wharton females. As re-
defined here, the valves of sanbernardiensis are so
nearly equivalent in size that this feature would not
distinguish it from Texas congeners.
Eurymerodesmus digitatus Loomis
Figs. 176, 197-203
Eurymerodesmus digitatus Loomis, 1976:288-289, figs. 1-3.
Type specimens. — Male holotype and female
allotype (FSCA), and one male and one juvenile
paratype (NMNH), collected by J. C. Loomis, 15
February 1975, 5 mi. ESE Marble Falls, Burnet Co.,
TX.
Diagnosis. — A small to moderate-size species;
males characterized by long mandibular projection;
aperture moderate-size, much narrower than ventral
segmental width; sides not divided, without
caudolateral pouches, elevating to various heights
around caudolateral corner, descending to
metazonal surface at varying distances along caudal
92 EURYMERODESMID MILLIPEDS
Figs. 197-203. Eurymerodesmus digitatus. 197, projection of
right mandible of holotype, lateral view. 198, aperture and
gonopods in situ of paratype, ventral view. 199, left gonopod of
the same, lateral view. 200, telopodite of the same, medial view.
201, cyphopods in situ, ventral view of female from Llano Co.,
TX. 202, left cyphopod of allotype, oblique caudal view. 203, the
same, oblique anterior view. Scale lines for figs. 198 and 201 =
1.00 mm. Line for other figs. = 0.35 for 1987, 0.40 mm for
202-203, and 0.50 mm for 199-200.
margin; telopodite usually short, usually barely
overlapping caudal margin of aperture; acropodite
short, about 1/8 of telopodite length, broadly ter-
minal, poorly demarcated from prefemur, curving
variably dorsad; prefemur with many hairs arranged
continuously along inner margin; cyphopod valves
of females usually with distal corners extended,
broadly curved; receptacle small, indistinct,
submerged in valves, sides not curving around
operculum.
Color in Life. — Not known with certainty.
Loomis (1976) reported that both the color and pat-
tern were “‘apparently much as in melacis,”’ but did
not describe them per se. Thus, digitatus probably
exhibits red paranota, red caudal metatergal stripes,
and red borders on the collum.
Holotype. — Specimen highly fragmented, length
unmeasurable, maximum width 2.7 mm.
Somatic features similar to hispidipes, with following excep-
tions:
Width across genal apices 2.0 mm, interantennal isthmus 0.9
mm. Antennae reaching back beyond caudal margin of 2nd
tergite, relative lengths of antennomeres 2>3>4>5=6>1>7.
Genae without impressions. Facial setae as follows: epicranial, in-
terantennal, subantennal, and genal absent, frontal 1-1, clypeal
about 7-7, labral about 13-13. Process of mandibular stipes long,
tip broadly rounded (Fig. 197).
Collum not extending below ends of following tergite. Paranota
moderately depressed, continuing slope of dorsum, peritremata
flattened; posterior corners rounded through segment 10, blunt on
11-14, becoming progressively more acute posteriorly.
Sth sternum with low, rounded elevated areas between leg pairs,
caudal ones larger and more widely separated; 6th sternum with
knob-like processes between leg pairs, caudal ones longer and
more widely separated. Postgonopodal sterna with distinct knob-
like projections between 9th legs, narrowly segregated in midline;
remaining sterna becoming progressively flatter and more plate-
like posteriorly with progressively sparser hair patches adjacent to
leg coxae. 2nd coxa with distinct, subconical anterior lobe and
small caudal projection. Prefemoral lobes extending through seg-
ment 9.
Gonopodal aperture (Fig. 198, not this specimen) broadly
ovoid, without caudolateral pouches, with moderate anterior in-
dentation, 1.7 mm wide and 0.7 mm long at maxima; anterior in-
dentation moderately long, subtriangular, apically rounded; sides
not divided, gently curved, leaning slightly over opening, becoming
slightly and progressively more elevated caudad, rims essentially
smooth; caudolateral corner broad, slightly angled; caudal margin
continuous with sides at caudolateral corner, elevation sloping
rapidly to metazonal surface and terminating just beyond
caudolateral corner, curving gently across midline; margins with
only a few hairs arising from rims of elevations at caudolateral cor-
ners and adjacent parts of sides and caudal margin. Gonopods in
situ (Fig. 198, not this specimen) with telopodites extending caudad
in parallel arrangement along midline, extending just beyond
caudal margin of aperture, tips directed dorsad. Gonopod struc-
ture as follows (Figs. 199-200): Telopodite short, terminating well
before level of distal extremity of hairs. Prefemur short but pro-
portionally long, about 7/8 of telopodite length, upright for most
of length, trace of curvature distad, broad basally, tapering slightly
distad, with three rows of continuous, regularly spaced hairs on in-
ner surface and distomedial tuft of about 6 hairs. Acropodite short
and broadly terminal, about 1/8 of telopodite length, continuous
with, and poorly demarcated from, prefemur, continuing slight
distal curvature of latter and directed dorsad, sides tapering rapidly
to acuminate tip.
Male paratype. — Agreeing closely with holotype in all par-
ticulars.
Female allotype. — Fragmented, length unmeasurable, max-
imum width 2.4 mm. Agreeing essentially with males in somatic
features, with following exceptions: Mandibular stipes with short,
broadly rounded process. Sterna flat, with progressively sparser
hair tufts adjacent to leg coxae on segments 4-7 and an occasional
scattered hair thereafter.
Cyphopodal aperture elliptical, sides slightly elevated above
metazonal surface. Cyphopods in situ (Fig. 201) with valves ex-
tending caudad along body axis, distal corners curved, protruding
through opening. Valves (Figs. 202-203) moderate-size, subequal,
distal corners extended, long, curved, and rounded, sides ex-
tending slightly around receptacle. Latter small, located on medial
side of valves, submerged in and obscured by latter, invisible from
caudal and anterior perspectives. Operculum small, tucked under
free, dorsal side of valves.
Variation. — Eurymerodesmus digitatus under-
goes considerable variation, and forms from Blanco
ROWLAND M. SHELLEY 93
and Llano counties, adjacent to Burnet County,
closely resemble the types. In these individuals, the
aperture sides are higher, and in the Blanco County
males, the elevation continues around the
caudolateral corner for about 1/3 of the length of
the caudal margin. The gonopods are very close to
the types, but the corners of the cyphopod valves of
Llano County females are not as elongate as those of
the allotype.
The Travis County males display much wider
apertures, and that from Elbow Cave (FSCA) is also
much longer, extending nearly to the anterior
segmental margin, thereby resembling the condition
in dactylocyphus. The elevation is higher caudally on
the sides, and in that from Elbow Cave, it begins
abruptly at midlength on the sides, tapering nearly to
the metazonal surface at the caudolateral corner. In
one male from along Waller Creek, the sides lean
noticeably over the opening, whereas in others from
that site, the rim is flared laterad at the caudolateral
corner. Except for the Elbow Cave male, the gono-
pods are very close to the holotype. In this individual
the telopodites are longer, the prefemur curves more
noticeably dorsad distally, and the acropodite does
so even more strongly. Females are unknown from
Travis County.
The Tarrant County males also display longer
telopodites, extending to the level of the distal ex-
tremity of the hairs and farther from the aperture,
terminating between the 8th legs. The mandibular
process is shorter, broader, and subtriangular, and
the cyphopod valves of females are not extended.
The Oklahoma males, found mixed in a sample
with mundus, display a broader aperture with only
slight elevation at the caudolateral corner. The man-
dibular projection resembles that in Tarrant County
males.
Ecology. — Unknown. The vial labels lack in-
dications of habitat, and nothing was mentioned in
the original description.
Distribution. — A band along the boundary be-
tween the Central and Coastal Plains, extending
from southwestern Oklahoma to central Texas near
the Pedernales River (Figs. 176, 212). Specimens
were examined as follows:
OKLAHOMA: Comanche Co., Mt. Scott, 3M,
25 October 1974, G. Beck (MSU).
TEXAS: Tarrant Co., Fort Worth, Normandy
Dr., 20M, 24F, 4 juvs., 3 November 1979, G. Spicer
(NCSM). Burnet Co., 5 mi. ESE Marble Falls, 2M,
F, juv., 15 February 1975, J. C. Loomis (FSCA,
NMNH) TYPE LOCALITY. Llano Co., 13 mi. SE
Llano, 4M, 3F, 11 March 1961, J. F. Quinlan
MEM. AMER. ENT. SOC., 37
(FSCA); and 27.8 mi. N Fredericksburg (Gillespie
Co.), ject. TX hwys. 16 and 965, 10M, 13 March
1982, J. C. Cokendolpher (NCSM). Blanco Co., 2
mi. N Johnson City, 2M, 13 March 1982, J. C.
Cokendolpher (FSCA). Travis Co., Austin, 4M, 5F,
date and collector unknown (NMNH), and 4M, 19
November 1945, collector unknown (NMNH); along
Waller Cr., 7M, juv., November 1963, J. Reddell
(FSCA) and F, 9 April 1983, J. Reddell (TMM); and
Elbow Cave, 3 mi. N Austin, M, February 1963, B.
Russell (FSCA).
Remarks. — Chamberlin recognized that this
form represented a new species and inserted a label
stating ‘‘E. austini Chamb., Types’’ with the first
sample from Austin, Travis County. However, this
name was never validated and Loomis (1976)
receives authorship credit.
Eurymerodesmus digitatus is similar to melacis,
but differs in the genitalia of both sexes. The
cyphopod valves are not elongate in melacis,
whereas they are in the most proximate populations
of digitatus with females. The gonopodal apertures
are similar in general configuration, but that of
digitatus is larger and the lateral elevation is more
variable. The telopodites of melacis are uniform,
and are longer and narrower than those of digitatus.
The chief difference is the configuration and direc-
tion of the acropodite, which bends sharply laterad
in melacis and curves gently dorsad in digitatus. This
structure’s curvature is slight in all males except that
from Elbow Cave, Travis County, where it is
stronger.
In contrast to melacis, there is little difference be-
tween digitatus and sanbernardiensis, and future
collecting in Lee, Bastrop, Fayette, and Williamson
counties may show them to be subspecifically
related. For now I retain both names and distinguish
them by the length and curvature of the acropodite,
short and gently curved in digitatus, and longer and
more strongly curved in sanbernardiensis.
Though differing in details of the gonopods, aper-
ture, mandibular projection, and female cyphopods,
I do not think that the differences between the
populations of digitatus are of sufficient magnitude
to warrant taxonomic recognition for forms in
Travis and Tarrant counties, Texas, and Comanche
County, Oklahoma. The last two sites are so widely
separated from the type locality that differences
could easily be bridged by forms from intervening
areas.
As stated previously, the telopodites of digitatus
are convergent with those of dubius as shown by
comparing figures 47-48 and 199-200.
94 EURYMERODESMID MILLIPEDS
Figs. 204-209. Eurymerodesmus clavatus. 204, projection of
left mandible of paratype, lateral view. 205, aperture and
gonopods in situ of holotype, ventral view. 206, telopodite of left
gonopod of the same, lateral view. 207, the same, medial view.
208, left cyphopod of female paratype, caudal view. 209, the same,
medial view. Scale line for fig. 205 = 1.00 mm. Line for other figs.
= 0.5 mm for each.
Eurymerodesmus clavatus, new species
Figs. 176, 204-209, 212
Type specimens. — Male holotype and 7 male
and 5 female paratypes (FSCA) collected by R. O.
Albert, 3 February 1963, at New Ulm, Austin Co.,
TX. One male and one female paratypes deposited
in NCSM.
Diagnosis. — A small species; males character-
ized by short mandibular projection; aperture
moderate-size, much narrower than ventral segmen-
tal width; sides not divided, without caudolateral
pouches, elevating slightly to caudolateral corners,
descending to metazonal surface along caudal
margin; telopodite short, barely overlapping caudal
margin of aperture; acropodite moderately long,
about 1/3 of telopodite length, narrowly terminal,
arising solely from outer prefemoral margin;
prefemur with many hairs arranged continuously
along inner margin, expanding distad into clavate
shoulder on inner margin; cyphopod valves of
females with distal corners slightly extended; recep-
tacle small, partly submerged in valves, sides not
curving around operculum.
Color in Life. — Unknown.
Holotype. — Length 14.1 mm, maximum width
1.6 mm, W/L ratio 11.3%.
Somatic features similar to hispidipes, with following excep-
tions:
Width across genal apices 1.2 mm, interantennal isthmus 0.6
mm. Antennae reaching back to anterior half of 3rd tergite,
relative lengths of antennomeres 2>3>6>5>4>1>7. Genae
without impressions. Facial setae as follows: epicranial 1-1, inter-
antennal, subantennal, and genal absent, frontal 4-4, clypeal about
9-9, labral about 14-14. Process of mandibular stipes short, sub-
triangular, apically acute (Fig. 204).
Collum not extending below ends of following tergite. Paranota
moderately depressed basally, continuing slope of dorsum, peritre-
mata somewhat flattened; posterior corners blunt on segments 1-3,
rounded on 4-7, blunt on 8-13, becoming progressively more acute
posteriorly.
5th and 6th sterna with small hirsute areas adjacent to leg coxae
but without lobes or projections. Postgonopodal sterna with flat-
tened hirsute areas adjacent to 9th coxae; remaining sterna becom-
ing flatter, more plate-like, and less hirsute caudally. 2nd coxa
without lobes, gonapophyses relatively long, extending to near
midlength of 4th sternum. Prefemoral lobes small, not pro-
nounced, extending through segment 8.
Gonopodal aperture (Fig. 205) small, heart-shaped, without
caudolateral pouches, with moderate anterior indentation, 0.7 mm
wide and 0.3 mm long at maxima; anterior indentation moderate-
size, broadly triangular, apically rounded; sides not divided,
angling strongly mediad, becoming slightly elevated caudad, rims
smooth; caudolateral corner poorly defined, indistinct, continuous
with lateral curvature; caudal margin continuous with sides,
angling to point in midline, slightly elevated and thickened at ap-
proximate location of caudolateral corner, flush with segmental
surface in midline; margins nearly glabrous, with only 6 or so hairs
on thickening at caudolateral corner. Gonopods in situ (Fig. 205)
with telopodites crossing in midline and extending slightly over op-
posite side of aperture. Gonopod structure as follows (Figs.
206-207): Telopodite terminating just before level of distal extremi-
ties of hairs. Prefemur proportionally short, about 2/3 of telopo-
dite length, upright but with slight bisinuate curve, relatively nar-
row basally, expanding near midlength and continuing distad into
broad shoulder on inner margin, with two rows of continuous,
regularly spaced hairs on inner surface and distomedial tuft of
about 6 hairs. Acropodite proportionally long and narrowly ter-
minal, about 1/3 of telopodite length, arising from outer pre-
femoral margin, discontinuous with and sharply demarcated from
prefemur, curving broadly dorsad, sides tapering smoothly and
continuously, expanding slightly apically, tip rounded.
Male paratypes. — The only variation of the male paratypes is
that the telopodites in situ sometimes extend well beyond the
caudal aperture margin, terminating over the sternum between the
9th legs. The structure and configuration of the aperture and
gonopods are as described for the holotype.
Female paratype. — Length 15.8 mm, maximum width 2.6
mm, W/L ratio 16.5%. Agreeing essentially with holotype in
somatic features, with following exceptions: Subantennal setae 1-1,
frontal 2-2, genal 2-2, clypeal and labral series merging and con-
tinuing for short distances along genal borders, about 4 setae per
side. Mandibular stipes without projection, corner not produced.
Sterna flat, glabrous. Prefemora without lobes.
Cyphopodal aperture relatively broad, ovoid, sides elevated
above metazonal surface, caudal margin elevated laterally, more so
in midline, flared caudad. Cyphopods in situ with valves oriented
transversely in aperture, tips visible. Valves (Figs. 208-209)
moderately large, subequal, corners slightly extended, with lobe on
ventral surface. Receptacle small, located laterad to valves at
bases, partly submerged in latter. Operculum proportionally large,
clearly visible on medial side of valves.
Ecology. — Unknown.
Distribution. — Known only from the type local-
ity (Figs. 176, 212).
ROWLAND M. SHELLEY 95
Remarks. — The narrowly terminal, discon-
tinuous acropodite, sharply demarcated from the
prefemur, resembles the condition in simplex, except
that it arises from the outer prefemoral margin
rather than the inner. Accordingly, the distal
prefemoral shoulder is on the inner surface instead
of the outer. Thus, there is no question that the nar-
rowly terminal condition in clavatus is convergent
with that in simplex.
EcoLoGy
With so little field experience with eurymerodes-
mids, I cannot generalize about habitat preferences
and can only report in the individual accounts my
observations of the few species I have collected.
However, information on vial labels and in the
literature suggest that eurymerodesmids may seek
shelter under almost any object in a wide array of
biotopes ranging from sandy, predominantly pine
Atlantic coastal habitats, to deciduous forests in the
southeast and the Ouachita, Ozark, and Central
Lowland Provinces, to grasslands of the Great
Plains, and to xeric desertine environments of the
Edwards Plateau. They often seek shelter in caves.
I was struck by the difficulty I had in finding them
and by how spotty their occurrences can be. With all
the winter trips that I have made to Florida, I have
only encountered v. varius four times, and the only
place that I have found it farther north is the North
Carolina site. The availabie records suggest that it is
common in the southeast, yet I have collected exten-
sively in Alabama, Georgia, and the Carolinas at the
proper time of year without even one accidental
discovery. Conversely, local populations can be very
large; for example, populations of v. varius can be in
the hundreds if not thousands of individuals. The
populations I discovered in Alachua and Duval
counties, Florida, and Robeson County, North
Carolina, were easily this large, and I could have
taken far more specimens. Sizeable pitfall samples
were also obtained from Leon and Jefferson coun-
ties, Florida, and there is a very large sample of v.
christianus from Washington Parish, Louisiana.
These aggregates suggest that varius would be a
suitable subject for population studies.
Likewise in south Texas, melacis can be locally
abundant. I could have taken many more individuals
at the Welder Wildlife Refuge, San Patricio County,
and two samples from Hidalgo and Kerr counties are
quite large. The patchy distribution was evident at
the Refuge, where I found melacis in several small
thickets around the office buildings but not in
MEM. AMER. ENT. SOC., 37
similar thickets on the range. Nor did I encounter
additional specimens anywhere in south Texas
despite searching out similar environments and
working in general areas where I knew it occurs.
North of Texas and Louisiana I have only collected
one eurymerodesmid, oliphantus from Jackson
County, Illinois, despite considerable work in forests
in the Ozarks and Mississippi Valley, where they are
abundant judging from the extensive collections at
the FSCA. In summary, I have had little success in
finding eurymerodesmids anywhere, so if there is a
secret to collecting them, as there is with American
platyrhacids and cove-dwelling Appalachian species
of Sigmoria (Shelley 1981, 1982; Shelley and White-
head 1986), I have not deduced it.
According to Wood (1867), the type specimens of
impurus were discovered under old cow dung, and
Causey (1952a) remarked that two or three milliped
species may be found under a single piece of dung
since there is little other cover. This statement seems
plausible, but I did not find a single eurymerodesmid
under dung during two weeks in Texas in February
1986. I turned scores of piles in all stages of decay
and dessication, but Desmonus (Sphaeriodesmidae)
was the only polydesmoid that I encountered. The
63
1
Jan Feb Mar Apr May Jun Jul
210
Fig. 210. Seasonality of Eurymerodesmus; the number of
samples with adults per month.
Aug Sep Oct Nov Dec
96 EURYMERODESMID MILLIPEDS
few eurymerodesmids that I discovered in Texas
were all under fallen logs and branches.
Throughout their distribution, eurymerodesmids
demonstrate marked seasonality, showing a distinct
preference for the cooler months of spring, autumn,
and winter. Thus, they probably will not be en-
countered in the summer, which explains why I only
found exoskeletal fragments of one male in Texas
and the Ozarks in July 1980. To illustrate this
phenomenon, I combined the species and plotted the
samples from each month with at least one adult
(Fig. 210). Although this probably reflects field ac-
tivity to some degree, I think it portrays their avail-
ability reasonably accurately. Many other millipeds
have been collected in the eurymerodesmid range in
the summer, so many more samples would be ex-
pected from May-September if the millipeds were
present. December was the most productive month
for eurymerodesmids, followed by April, February
and March, November, and October. Eleven fewer
samples were taken in January than in October, but
24 more than in May. Thus, the seven coolest
months of the year provided 90.9% of the preserved
eurymerodesmid samples with adults. Future field
investigations should therefore be concentrated in
these months.
DISTRIBUTION
Family and genus. — Except for the Ozark and
Ouachita Provinces in Arkansas and Missouri, the
Eurymerodesmidae is a lowland milliped family oc-
curring in the Coastal Plain, Central Lowlands, and
Great Plains Physiographic Provinces (Fig. 1), the
regions of the central, south-central, and south-
eastern United States with the least topographical
variation. The center of diversity lies in the Coastal
Plain of southern Arkansas and northern Louisiana,
extending northward into the southern fringe of the
Ouachita Province. Fourteen of the 25 species,
56.0% of the total known fauna, are represented
there as are three of the four lineages and six of the
seven species groups, the mel/acis lineage and group
of Texas and southwestern Oklahoma being the sole
exceptions (Figs. 211, 213, 214). The range spans
such major rivers as the Pee Dee, Santee, Savannah,
Chattahoochee, Alabama, Pearl, Mississippi,
Sabine, Brazos, Colorado, San Antonio, and
Nueces. Inland it traverses the Red and Arkan-
sas/Canadian Rivers, two major tributaries of the
Mississippi, and the southwesternmost record, the
Terrell County, Texas, locality of melacis, is just
west of the Pecos River near its confluence with the
Rio Grande. In figure 1 a smooth curve is drawn
around range extremes in all directions, except for
the area north of the Missouri River in northern
Missouri and southern Iowa. No authentic records
are known from north of this watercourse, which
appears to be a boundary in this part of the range.
Consequently, Eurymerodesmus probably does not
protrude into southeastern South Dakota. The ir-
regular northern boundary indicates three separate
penetrations that extend progressively farther north:
one along the Atlantic Coastal Plain into south-
eastern North Carolina and two in the Central
Lowlands Province into central Illinois and north-
Fig. 211. Distributions of species of Eurymerodesmus in
Arkansas and Louisiana. Star in square, hispidipes; stars,
angularis; diamonds, compressus; vertical half-shaded dots, good;
dots/circle in square, dubius; half-shaded triangles, varius chris-
tianus; triangles, varius louisianae; dots, amplus; triangles in
squares, mewtonus; dots in diamond, oliphantus; asterisks,
simplex; horizontal half-shaded dots, polkensis; hexagons,
paroicus; star in dot, crassatus; dot/circle with line, pulaski; ovals,
serratus; squares, birdi birdi; diagonal half-shaded squares, birdi
planus; vertical half-shaded squares, mundus. Open symbols
denote literature records considered reliable.
ROWLAND M. SHELLEY 97
eastern Nebraska. The southern boundaries are the
Rio Grande, since no authentic records are known
from adjoining parts of Mexico, the Gulf of Mexico,
and the southern limit of the mesic deciduous
woodlands in Florida, as discussed in the v. varius
account. However, little sampling has taken place in
potential sections of Coahuila, Nuevo Leon, and
Tamaulipas states, so there is a reasonable probabil-
ity of future Mexican discoveries.
Boundaries in other areas are less precise, but
from my own field investigations, I doubt if
Eurymerodesmus extends much farther west in
Texas, or north in North Carolina. As discussed in
the melacis account, I unsuccessfully turned
countless logs and sifted through considerable litter
and debris around Lubbock, Texas, and in the Davis
Mountains in February 1986, so if eurymerodesmids
occur in the panhandle or southwestern Texas, it is
difficult to imagine where they could be other than
in caves or remote, secluded ravines. Southeastern
North Carolina, the area of the northernmost record
along the Atlantic Coast, has been sampled exten-
sively in all seasons of the year by biologists at the
NCSM with only the one discovery of v. varius. I
have personally searched for this species many times
in the southeastern quadrant of the state, and ex-
haustive field work has been conducted by NCSM
personnel throughout the Coastal Plain and well into
southeastern Virginia. Consequently, the presently
known limit, about eight miles north of the South
Carolina border, is probably very close to the actual
one.
Fig. 212. Distributions of species of Eurymerodesmus in Texas. Stars, impurus; dots, amplus; diamonds, birdi birdi; inverted
triangles, mundus; squares, melacis; upright triangles, dactylocyphus; verticle half-shaded dots, sanbernardiensis; horizontal half-shaded
dots, digitatus; asterisk, clavatus. Open symbols denote literature records considered reliable.
MEM. AMER. ENT. SOC., 37
98 EURYMERODESMID MILLIPEDS
Other peripheries have not been thoroughly in-
vestigated, but the western boundary in the plains
states and the inland periphery in the southeast are
considered reasonably accurate. The limit in Kansas
approximates that of the xystodesmid Pleuroloma
flavipes Rafinesque (Shelley 1980), which in turn
correlates roughly with the western terminus of the
Central Lowland Physiographic Province. The in-
terior boundary in the southeast is generally the Fall
Zone region between the Coastal Plain and Pied-
mont Plateau Physiographic Provinces. Such inland
records as Macon, Georgia, and Auburn, Alabama,
are in the Fall Zone, and there are no definite
localities from the Piedmont Plateau in any state.
The only regions where the boundaries are nebulous
are northern Illinois, and Kentucky and Tennessee.
Field work in Illinois has focused on the Shawnee
National Forest in the south, but the record of
hispidipes from Coles County, in east-central II-
linois, suggests occurrence into northern Illinois,
perhaps even to metropolitan Chicago if the range
extends to the same latitude as mundus in Nebraska.
Eastward expansion into western Indiana is not im-
plausible, although the Wabash River may restrict it
from the southwestern corner of this state. Thus,
collecting in central Illinois and adjacent Indiana
could significantly expand the family’s range. No
specimens have been taken from western Kentucky
and Tennessee, so the line in these states is hypo-
thetical, connecting records from Illinois and
Mississippi; the family is not known to traverse the
lower Ohio River as figure 1 shows. However,
eurymerodesmids may be logically anticipated in the
shaded areas of Kentucky and Tennessee, and
perhaps as far east as the western loop of the Ten-
nessee River.
Species, species groups, and lineages. — The
hispidipes lineage, comprised of the single species
and group, is known definitely from only two sites
some 470 miles apart, one in east-central Illinois and
the other in southern Arkansas (Figs. 9, 213, 214).
This distance is so great as to virtually preclude ex-
tant connection. The impurus group (Figs. 9, 213)
covers most of Arkansas except the northwest cor-
ner. It swings broadly through northern Louisiana
with outlying records from eastern Mississippi, west-
central Missouri, and east-central Texas. The
Missouri and Mississippi forms are judged con-
specific with those in Arkansas, and I connect these
areas by a single line in figure 213. However, the
species in Texas, impurus itself, has a different aper-
ture with false pouches, and is shown as isolated
(Fig. 213). The intervening area in eastern and north-
eastern Texas should be sampled in detail to deter-
mine if the areas connect geographically and whether
anatomical connection can be demonstrated between
the true pouch of angularis and that of impurus.
The varius group is the easternmost and most
widely ranging species group, although much of the
area is occupied by a single species, varius (Figs. 54,
213). South of the Ohio River, varius is the only
component east of the Mississippi, and it also ex-
tends into western Louisiana and southern Arkan-
sas. The northern part of the group’s range, in Il-
linois, Missouri, and Arkansas, contains sizeable
lacunae that need to be explored, although I show it
as continuous in figure 213. Gonopodal similarities
between amplus, newtonus, and elevatus suggest a
continuous fauna from southern Louisiana and
eastern Texas through central Missouri, and the
close resemblance between all males of oliphantus
suggests continuity between its four known areas.
However, I show the westernmost record of amplus,
from Mason County, Texas, as a small, separate
area in central Texas (Fig. 213) because of the size of
the intervening gap, about 190 miles, and the
absence of similar forms to the north and south. The
varius group should be expected in eastern
Oklahoma, where it is currently unknown; the prox-
imity of newtonus to northeastern Oklahoma sug-
gests occurrence in Delaware, Adair, and possibly
Cherokee counties.
The simplex group contains seven localized
clusters of records, six of which comprise a
reasonably compact unit in southern Arkansas,
northern Louisiana, and southwestern Mississippi,
although much more sampling is needed (Figs. 54,
213). The seventh area, representing an allopatric
population of serratus, is far to the southeast in
Alachua County, Florida. This site and the type
locality of caesariatus are the only records of the
simplex group from east of the Mississippi River,
and they suggest a substantial fauna with numerous
undiagnosed forms along the Gulf Coast in the
Florida panhandle and southern Mississippi and
Alabama.
The dubius group, the fourth and only monobasic
group in the Kewanius lineage, inhabits a broad,
contiguous area in southern Arkansas, where it is
known from many counties (Figs. 211, 213). It may
occur in northwestern Louisiana and, conceivably,
even the northeast corner of Texas.
Because it is the most diverse assemblage in
Eurymerodesmus, combining the impurus, dubius,
varius, and simplex groups, the Kewanius lineage ex-
pectedly occupies the greatest area, around 2/3 of
ROWLAND
M. SHELLEY
99
213
Fig. 213.
Comparative distributions of the species groups of Eurymerodesmus. 1, hispidipes group; 2, impurus group; 3, dubius
group; 4, varius group; 5, simplex group; 6, birdi group; 7, melacis group.
the generic range (Fig. 214). It is the easternmost
lineage, covering all the area in the southeast, and it
expands northward along the Mississippi Valley into
Illinois and Missouri, and westward through Arkan-
sas and Louisiana into eastern Texas and, by im-
plication, eastern Oklahoma. It overlaps the eastern
half of the birdi lineage and the Arkansas site of
hispidipes, and its allopatric population in central
Texas is sympatric with the melacis lineage. No over-
lap is otherwise known between the Kewanius and
melacis lineages, which appear to be parapatric in
southeastern Texas.
The birdi lineage is the northernmost ensemble,
occupying the western periphery in the northern sec-
tor of the range (Figs. 147, 213, 214). It extends ina
wide band from Nebraska and western Missouri into
MEM. AMER. ENT. SOC., 37
northeastern Texas, where it divides into a broad
branch that penetrates across the Mississippi River
into southeastern Louisiana and southwestern
Mississippi, and a narrow branch that continues into
south Texas. This is the most continuous assemblage
in Eurymerodesmus, but populations of b. birdi in
south Texas are some 214 miles from the closest
records in northeastern Texas. The gap is filled by
mundus, lending continuity to the lineage’s dis-
tribution. Aside from this hiatus in b. birdi, there are
no areas where collecting is particularly needed in the
birdi lineage, although pinpointing the western and
northern extremities would be useful.
The melacis lineage, the southernmost and
westernmost assemblage, occupies the southwestern
periphery of the genus (Figs. 176, 213, 214), over-
100 EURYMERODESMID MILLIPEDS
lapping the southern extension of the birdi lineage
and the westernmost site of Kewanius. It extends
from the Gulf of Mexico onto the Edwards Plateau
in south-central Texas, with the inland or western
border pushing northward into Oklahoma, while the
coastal boundary lies south of Houston and is
parapatric to the Kewanius lineage. Records are con-
centrated east of highway I-35, north of I-37, and
between San Antonio and Del Rio. Little material is
available from south Texas, and there is a large gap
in the north-central area, where the Fort Worth
record of digitatus is the only one between Coman-
che county, Oklahoma and the type locality in
Burnet County, near the latitude of Austin. Because
of the Fort Worth record I show a unified area for
the melacis lineage in (Figs. 213-214), but the record
in the Wichita Mountains of Oklahoma may be
detached from the rest of the range. Considerable
field work is therefore needed in the area bounded
by the Wichita Mountains on the north, I-35 on the
east, Burnet and Llano counties, Texas, in the south,
and Abilene on the west.
Although the range of the Eurymerodesmidae is
far from thoroughly collected and many distribu-
tional gaps remain, some lacunae between con-
specific populations are so large as to virtually
guarantee allopatry. For example, gaps of over 150
miles occur in hispidipes, angularis, varius, amplus,
serratus, birdi, and digitatus. The largest hiatus is in
serratus, where the Pulaski County, Arkansas, and
Alachua County, Florida, populations are
segregated by over 700 miles. Because of the vastness
214
Fig. 214. Comparative distributions of the lineages of Eurymerodesmus. 1, melacis; 2, hispidipes; 3, birdi; 4, Kewanius.
ROWLAND M. SHELLEY
of the intervening region, which covers more than
three states, it is hard to imagine that these popula-
tions connect today. Likewise in hispidipes, there is a
hiatus of around 470 miles between the known
populations, a sizeable gap suggesting absence of ex-
tant connection. The allopatric population of
amplus in grasslands of the Edward’s Plateau in
Mason County, Texas, is isolated from the bulk of
the range in forested eastern Texas by some 190
miles. In contrast to the preceding species, there are
five allopatric records of b. birdi from four non-
contiguous counties in south-central and southern
Texas, the northernmost of which is about 215 miles
south-southwest of the main area of the race in
Oklahoma, northeast Texas, and Louisiana. These
four counties are at least 70 miles apart and their
localities may or may not interconnect, but together
they appear detached from the rest of the range.
Consequently, b. birdi seems to consist of two popu-
lations — a relatively continuous one extending
from eastern Missouri and eastern Kansas through
Oklahoma, western Arkansas, and northeastern
Texas to the vicinity of New Orleans, and a smaller,
subcontinuous one in southern and south-central
Texas, ranging at least from Bell to San Patricio
counties. The dimensions of the latter cannot be ap-
proximated from present knowledge, but it could be
relatively large, encompassing much of the area
from Corpus Christi and Victoria on the coast to San
Antonio, Austin, and Belton inland.
The widely allopatric records of angularis, v.
varius, and digitatus are more likely to be joined by
future collecting, either because the lacunae are
smaller, or the habitat is more uniform, or both. My
reasoning is largely intuitive, but I have considerable
field experience in the 275 mile gap in vy. varius
between Macon, Georgia, and Robeson County,
North Carolina. I do not think that there are nearly
enough differences between coastal environments in
Georgia and the Carolinas for the latter populations
to be isolated. This hiatus is logically a collecting ar-
tifact that will eventually be filled with material from
South Carolina.
Future work. — 1 conclude this discussion of dis-
tribution with a review of future collecting needs.
Beginning in the southeast, efforts are needed to
connect the Florida and Arkansas populations of
serratus, but it is impossible to predict where addi-
tional discoveries may be made. They are more likely
in Mississippi since the eurymerodesmid fauna there
is more diverse than in states to the east, where v.
varius is the only known form aside from the one
serratus record. Otherwise, the only future field
MEM. AMER. ENT. SOC., 37
101
work that is really needed east of Mississippi is
material of v. varius from the void in South Carolina
and preferably also eastern Georgia.
Meticulous field sampling, perhaps to the extent
of spending two to three days in each county, is
needed from central Mississippi through Louisiana
and Arkansas, the region of greatest species diver-
sity. Many more forms undoubtedly occur there,
particularly in the simplex group. The record of
angularis from Oktibbeha County, Mississippi,
needs to be linked with the rest of the range in
Arkansas and Louisiana, which should be
thoroughly collected to determine the relative
statuses of the various forms with long acropodites.
Emphasis should be placed on Union and Columbia
counties, Arkansas, to determine boundaries of
compressus, and topotypes of Paresmus columbus
are needed from Magnolia, in the latter county, to
confirm or disprove its synonymy with dubius.
Careful sampling is also needed in Pulaski County
and environs to ascertain distributions of pulaski
and serratus, which were mixed in Bollman’s
material (NMNH), and forms between Montgomery
and Pulaski counties may tie serratus and polkensis
together. Collection needs dwindle to the south and
north, except for more material of simplex around
Evangeline Parish, Louisiana, and newfonus in
northwestern Arkansas. Specimens from central
Missouri will delineate the range of elevatus and
possibly connect it with newtonus, as well as close
the hiatus of 216 miles in angularis.
In the northern part of the range east of the
Mississippi River, efforts are sorely needed in central
Illinois to secure more material of hispidipes, deter-
mine its distribution, and pinpoint the northern
periphery of the family in this area. South of the
Ohio River, Eurymerodesmus should be docu-
mented from western Kentucky and Tennessee,
where new forms may be encountered.
The northern and western peripheries of the fam-
ily in Nebraska and Kansas need refinement, but the
only specific collecting needs are corroboration of
early literature records of b. birdi and mundus as ex-
plained in their accounts. A few literature records of
these species also need confirmation in Oklahoma,
but the principle requirement is in the eastern,
Ouachita section. Eurymerodesmus newtonus,
goodi, and polkensis all occur just across the state
line in Arkansas and can be anticipated in eastern
Oklahoma, where only b. birdi and mundus are cur-
rently known. In southwestern Oklahoma, more
material is needed of digitatus in and near the
Wichita Mountains and southward to the Red River.
102
Considerable field work is needed in Texas, with
emphasis on the void between Abilene and Fort
Worth, and Wichita Falls and Llano/Burnet. No
specimens, not even juveniles, have been taken from
this substantial area where eurymerodesmids can be
expected to be abundant and digitatus, melacis, and
mundus may occur. East Texas has been com-
paratively well collected, based on the number of
samples of amplus, but more work is needed to try
to link impurus in Brazos County with the rest of its
group in northwestern Louisiana. Marion and Harri-
son counties should harbor angularis, based on its
occurrence in Caddo Parish, Louisiana, and addi-
tional representatives of the impurus group may exist
around Nacogdoches, Huntsville, and the Big
Thicket Area, where only amplus is currently
known. South of Houston, male topotypes are still
desired for sanbernardiensis, to eliminate doubt of
its identity. In addition to more individuals of im-
purus, detailed collecting in neighboring Colorado,
Waller, Austin, Washington, and Fayette counties
may provide more individuals of clavatus and clarify
the relationship between its unique gonopod and the
rest of the melacis group. Farther south, material is
needed from the slight gaps in the range of dac-
tylocyphus in Gonzales and Dewitt counties, in addi-
tion to those farther north in Travis and Corzell
counties. The range of melacis in south Texas is the
best collected area of the state, but additional
specimens are needed along the southern Rio Grande
in the vicinities of Eagle Pass, Laredo, Hebbronville,
Zapata, and Rio Grande City.
For the benefit of future workers and to comple-
ment the group and lineage range maps, I append
county distribution maps for the states with the most
diverse faunas, Arkansas, Louisiana, and Texas
(Figs. 211-212). Except for north-central and north-
eastern Arkansas and the southernmost part of
Louisiana, the first two states are reasonably covered
with records, but larger voids exist in Texas, which,
because of its greater area, could alone take several
years to adequately sample. Add another two or so
years for systematic field investigations in Arkansas
and Louisiana, and perhaps five or more years can
be spent in just these three states. When the addi-
tional time for studies in other areas is considered, it
becomes obvious that research on the Eurymero-
desmidae has just begun. Many questions remain to
be answered, and the present system of species
groups and lineages may need to be completely
revamped when significantly more material is avail-
able.
EURYMERODESMID MILLIPEDS
RELATIONSHIPS
Family and genus. — The great enigma surround-
ing the Eurymerodesmidae is its phylogenetic affini-
ties. There are suggestions that it is a recent
derivative of the Xystodesmidae (Hoffman 1978,
1979), an appealing idea because of the close
similarity in body form. Possible synapomorphies
with the Xystodesmidae include the short, cylindri-
cal, truncate gonapophyses and the gently curved to
bisinuate tarsal claws, which appear to be slightly
specialized because of the basal swellings. There are
also similarities with the Holistophallidae, some
forms of which have elaborate gonopodal apertures
and strong anterior indentations. However, this
family is poorly known, and the aperture traits are
not manifested by all components. The constricted
aperture is also associated with its modified
gonopods, and thus is not homologous to that in the
Eurymerodesmidae. Until family relationships are
resolved, I regard the Xystodesmidae and Eury-
merodesmidae as sister taxa within the Xystodes-
moidea.
Apparently on an empirical basis, Hoffman
(1979) arranged the Chelodesmidea as follows:
Superfamily Chelodesmoidea: Family
Chelodesmidae.
Superfamily Xystodesmoidea: Families
Xystodesmidae, Eurymerodesmidae,
Gomphodesmidae, Campodesmidae,
and Oxydesmidae. "
Superfamily Sphaeriodesmoidea: Families
Sphaeriodesmidae, Holistophallidae.
Superfamily Platyrhacoidea: Family
Platyrhacidae.
Superfamily Rhachodesmoidea: Families
Rhachodesmidae, Tridontomidae.
The Campodesmidae, Holistophallidae, and Tridon-
tomidae were said to have around 8, 11, and 3
known species, respectively, while the Eury-
merodesmidae, with 25 known species, is the
smallest of the more speciose families. Most of these
taxa have now been subjected to reasonably inten-
sive studies, and the least investigated, the
Sphaeriodesmidae, Holistophallidae, and Rhacho-
desmidae, may hold keys to subordinal affinities as
may the western Nearctic xystodesmids in the tribes
Harpaphini and Chonaphini, which lack the
characteristic prefemoral spines found in Central
American and eastern Nearctic tribes.
Species, species groups, and lineages. — Discus-
sion of relationships within Eurymerodesmus is ten-
tative because of the large number of unknown
ROWLAND
forms that obviously await discovery. Since it will
probably be decades before substantially more
material is available, it is appropriate to draw con-
clusions from the present data. Accordingly, I pre-
sent (Fig. 215) a preliminary diagram of relation-
ships among the lineages. Unique features can be
suggested only for the melacis branch — short
telopodites, either wholly enclosed within the aper-
ture or only slightly overlapping the caudal margin,
and relatively nondescript apertures with margins
usually flat or only slightly elevated — and I judge
them to be plesiomorphies, although determination
of polarity is hampered by incomplete knowledge of
potential outgroups. Nondescript apertures are also
contradicted by dactylocyphus, which has a broad,
distinctive opening with substantial flange-like eleva-
tions at the caudolateral corners in some forms.
However, neither of these traits appears in other
lineages, in contrast to the characteristic lobes of the
birdi branch, also displayed by polkensis, and to
lesser extents pulaski and serratus, all of the
Kewanius line. The lobes of the birdi lineage also are
not shared by all of its components. They are absent
from birdi planus, whose type population has a
nondescript aperture like those in the medacis
branch, a similarity reflecting convergence. Despite
these difficulties, I think that Eurymerodesmus can
be divided into these four lineages or major com-
ponents, three of which, like the subgenera of
Sigmoria (Shelley and Whitehead 1986), display
geographic cohesiveness. They are undeniable
geographic entities and defensible from the stand-
point of geography. The exception is the hispidipes
branch, comprised of a unique species from two
allopatric populations, which does not fit in another
line.
I placed polkensis, pulaski, serratus, and caesaria-
tus in the simplex group, Kewanius lineage, because
all have narrowly terminal or subterminal
acropodites. However, they could equally well go in
the birdi lineage if greater weight were placed on the
aperture. The first three species possess undivided
apertures with variable elevations on the caudal
margins (Figs. 109, 114, 134, 137, 142, 146), whereas
simplex, paroicus, and crassatus exhibit divided
apertures with caudolateral pouches (Figs. 103, 123,
127). In particular, polkensis has strong sub-
triangular lobes (Figs. 109-110), very much like those
in the sympatric form of b. birdi (Figs. 157-158).
Likewise in Mississippi, the largely unadorned aper-
ture of caesariatus, with only very slight, indistinct
elevations at the caudolateral corners (Fig. 117), is
shared with proximate forms of birdi planus (Figs.
MEM. AMER. ENT. SOC., 37
M. SHELLEY
103
melacis-1 birdi-3 Kewanius-4
hispidipes2
As
= WA
: 215
Fig. 215. Relationships of lineages in Eurymerodesmus. The
numbers refer to figure 214.
162, 165). Thus, these four species combine features
of the simplex and birdi groups in the gonopods and
aperture, respectively, and the undivided apertures
of goodi and dubius, of the impurus and dubius
groups, may also reflect genetic influence of birdi.
Thus as with the subgenera of Sigmoria, the birdi
lineage is a distinct geographic entity, but it is in-
completely detached from the Kewanius lineage.
These forms in the area of overlap exhibit features of
both lines, whereas other forms in this area, for ex-
ample amplus, angularis, paroicus, crassatus, and
simplex, do not show characteristics of birdi. Conse-
quently, the birdi and Kewanius lineages
demonstrate the phenomenon labeled ‘‘incomplete
synapomorphy”’ by Shelley and Whitehead (1986).
Another problematical species is clavatus, placed
in the melacis lineage because of its small size and
relatively unadorned aperture (Fig. 205). Geo-
graphically, the only other possibilities are the varius
and impurus groups, Kewanius lineage, all of whose
species possess either long acropodites or divided
apertures with pouches and broadly terminal
acropodites, respectively. Consequently, clavatus
would disrupt the harmonies of these ensembles, but
its narrowly terminal acropodites are also divergent
from other forms of the melacis lineage, not to men-
tion unique to the genus, since they arise from the
outer, rather than the inner, prefemoral margins.
The acropodite of clavatus, proportionately slightly
longer than those of the other species of the melacis
branch, may reflect genetic influence of the
parapatric species, impurus, and may be an example
104 EURYMERODESMID MILLIPEDS
of ‘incomplete synapomorphy”’ between the melacis
and Kewanius lineages.
Notwithstanding these questionable species, I
think that the impurus, varius, simplex, and dubius
groups unite to form a coherent ensemble, which is
incompletely detached from the birdi branch. As
stated earlier, these four groups are distinguished on
the basis of acropodal features for convenience and
to facilitate comprehension of the genus, but they
are closely related and contain all forms with divided
apertures and pouches, or with gonopods or aper-
tures similar to such forms. For example, goodi
lacks pouches, but its gonopods possess long acropo-
dites similar to those of angularis, which has
pouches. Conversely, dubius, parapatric to goodi,
has upright telopodites with short, broadly terminal
acropodites that are only very slightly curved, but its
aperture is shared with goodi. I assigned goodi to the
impurus group because of the long acropodite,
which excludes dubius. Nor does dubius fit in the
varius group, whose components display divided
apertures with pouches, and variably curved to lean-
ing telopodites. The only alternative for dubius is a
separate species group, which is part of the parent
lineage because of the shared aperture configuration
with goodi.
Elsewhere, the simplex and varius groups blend
together through varius itself, since v. christianus,
with a distally lobed prefemur and a broadly ter-
minal but sharply discontinuous acropodite, bridges
the anatomical gap between the subterminal or nar-
rowly terminal, and therefore discontinuous,
acropodites of the former, and the smoothly con-
tinuous, poorly demarcated ones of the latter. Fur-
thermore, angularis, with divided apertures and
pouches, connects the impurus with the simplex and
varius groups. The pouches alone suffice for this
purpose, but their variability is an important addi-
tional factor. In some forms of angularis, the
pouches are broad and open, thereby resembling the
conditions in paroicus and crassatus of the simplex
group (compare figs. 18 and 23 with figs. 123 and
127), whereas the pouches of other forms are narrow
and closed, and thus more like the conditions in
amplus and varius of the varius group (compare fig.
21 with figs. 64 and 73). Finally, compressus, with a
moderately long acropodite, 3/8 of the telopodite
length, blends the impurus and varius groups by
bridging the anatomical gap between short and long
acropodites, <1/4 and =1/2 of the telopodite
length, respectively. Thus, differences between the
impurus, dubius, varius, and simplex groups enable
them to be distinguished for convenience, to simplify
Eurymerodesmus, but there are many unifying fac-
tors that merge them into a single lineage covering
much of the generic range (Fig. 214, number 4). For
ease of recognition this lineage is referenced by the
older available genus-group name, Kewanius, since
there is no single, representative species.
As stated previously, the birdi group connects
with this ensemble through polkensis, caesariatus,
pulaski, and serratus, and possibly also goodi and
dubius, but otherwise it is detached and maintains a
consistent and independent set of features in parts of
Louisiana and Arkansas where they are sympatric.
Consequently, I regard the birdi group as a separate
lineage, loosely attached to, and sister to, the
Kewanius line. With only one component species
group and no available genus-group name, it is
referenced as the birdi lineage.
The melacis group occupies a peripheral geo-
graphical position along the southern and western
generic boundaries and is overlapped by the
southern, finger-like extension of the birdi lineage
and the allopatric populations of amplus, varius
group, Kewanius lineage, to which it is otherwise
parapatric (Fig. 213). There are no discernible
anatomical similarities and no clear connections
between the melacis group and the previous
assemblages, the only possible exception being the
slightly longer acropodite of clavatus that may
reflect influence of impurus. The generally upright,
sublinear telopodites of digitatus are phenotypically
similar to, and convergent with, those of dubius,
some 240 miles to the east. Likewise, the
phenotypically similar, dactyliform cyphopodal pro-
jections of dactylocyphus and compressus are struc-
turally different and reflect convergence. Conse-
quently, the melacis group clearly represents a
separate assemblage, which is referenced as the
melacis lineage since there is no available genus-
group name. Because of its generally peripheral posi-
tion (Fig. 214, number 1) and the absences of both
pouches and true lobes on the apertures, I consider
the melacis lineage to be sister to the rest of the genus
and family (Fig. 215). It is the least differentiated
and apparently most plesiomorphic ensemble.
With only one species and two widely allopatric
samples, the hispidipes group cannot be treated in
detail. The Coles County, Illinois, site is peripheral
and allopatric to the rest of the generic range,
whereas the southern Arkansas locality is in the area
of greatest species diversity and sympatric with both
the impurus and varius groups, Kewanius lineage
(Fig. 213). There are no unequivocal anatomical
similarities with other lineages, but I detect a slight
ROWLAND M. SHELLEY
suggestion of a distal prefemoral swelling, possibly
indicating affinity with the simplex group, Kewanius
lineage. Because it cannot be linked to another
species, Hispidipes comprises a separate species
group and lineage whose phylogenetic position is
determined by default, since the positions of the
other lineages can be logically inferred (Fig. 215).
However, there are also geographical and
anatomical grounds for this placement, since its
sympatry and the aforementioned anatomical
similarity suggest closer affinity to the birdi and
Kewanius lineages than is shown by the melacis
branch. The genus-group name Eurymerodesmus,
the oldest one in the family, is available for this line,
but I hesitate to employ this name below the generic
level without being certain of its usage. Future
material may provide the necessary evidence to
merge hispidipes with either the birdi or Kewanius
lineages, thus clearly comprising a Eurymerodesmus
branch. This is currently impossible, and hence, it is
referenced as the hispidipes line.
Like Sigmoria in the east, Eurymerodesmus is a
giant mosaic complex that blankets much of the
southeastern and south-central United States (Figs.
1, 213-214). We are seeing this assemblage near its
zenith, near maximal expansion with minimal extinc-
tion, before major anatomical and geographical
discontinuities develop. It consists of four lineages or
major components that overlie each other to varying
degrees (Fig. 214), but since the hispidipes branch
contains only one species, for practical purposes
there are only three such ensembles. The Kewanius
branch covers the southeastern and south-central
areas and is overlapped in the western third of its
range by the incompletely detached birdi lineage; the
melacis line occupies the southwestern corner of the
range, where it is largely independent from, and only
minimally overlapped by, the other two com-
ponents. Thus, the birdi branch is partly superim-
posed on both the Kewanius and melacis lineages,
which are essentially parapatric (Fig. 214). Previous
workers have merely extracted forms from various
parts of these lineages and, because they display dif-
ferent combinations of the features of the apertures
and gonopods, assumed that they represent distinct
species and assigned new names at this taxonomic
level. Not having personally collected Eury-
merodesmus to the extent that I did Sigmoria, I can-
not personally attest to the degree of partitioning,
nor can I detail the extent to which clinal gradients
traverse the range without much more material, par-
ticularly in the Kewanius lineage. However, the aper-
tures and gonopods are independent of each other in
MEM. AMER. ENT. SOC., 37
105
the simplex group, reflecting influence of the in-
completely detached birdi lineage. As the acropodite
progresses from narrowly terminal to subterminal,
the aperture changes from divided with pouches in
simplex, to undivided with sizeable caudal lobes in
polkensis and indistinct elevations in caesariatus,
returns to divided with pouches in paroicus and
crassatus, and is undivided with moderate lobes in
pulaski and serratus. Trends in these two features
therefore run counter to each other, but the full ex-
tent of this phenomenon is unknown.
In Sigmoria, Shelley and Whitehead (1986) de-
fined species as taxa that appear segregated from
geographically proximate taxa and are not connected
by intergrades, hence seemingly reproductively
isolated. Evidence for isolation included syntopy
between phenotypically distinct forms, range dis-
junctions between taxa that could otherwise be con-
sidered subspecies, and abrupt range discontinuities
between proximate forms, resulting in tightly fitted
parapatry patterns. Subspecies were defined as
reasonably homogeneous taxa that are continuous
with other such taxa through intergrade or in-
termediate forms. I have attempted to apply these
definitions to Eurymerodesmus, but the nature of
the interfaces between forms can rarely be deter-
mined because of incomplete material. Thus, forms
that appear to be parapatric and hence are con-
sidered species in this first revision may eventually
need to be reevaluated. Beyond this procedural dif-
ficulty, there are interpretation problems in the
Kewanius lineage that hinder taxonomic decisions.
For example in the impurus group, I lumped
dissimilar variants together under angularis but re-
tained compressus, in the middle of this area, as a
valid species. Though seemingly inconsistent, I think
that the anatomical differences in both sexes of com-
pressus (shorter acropodite, longer and dissimilar
cyphopodal projections) warrant taxonomic recogni-
tion. Other interpretations of this situation are possi-
ble, and the question arises as to whether compressus
really is reproductively isolated or a highly localized,
point-form variant of angularis. Perhaps compressus
actually represents a combination of stages in clinal
trends in the aperture and gonopods that produce a
variant of such phenotypic distinctiveness as to ap-
pear reproductively isolated when it actually is not.
Similarly in the varius group, I combined variants of
amplus but maintained taxonomic distinction from
varius, although some forms of amplus and v. loui-
sianae with intermediate densities of prefemoral
hairs appear to bridge the anatomical gap between
the species.
106 EURYMERODESMID MILLIPEDS
The picture is clearer in the melacis lineage
because there is less interference from overlapping
branches and because of less inherent intricacy.
Relatively extensive collecting has also taken place
along the interfaces between melacis, dactylocyphus,
and digitatus, disclosing tightly fitted parapatric
spatial relationships with minimal overlap (Fig. 176).
Consequently, this situation exhibits abrupt range
discontinuities, which are prevalent in Sigmoria and
interpreted as evidence for reproductive isolation.
Slight differences between digitatus and sanber-
nardiensis, chiefly involving the orientation of the
acropodite, suggest that they may be forms of a
single species, but more material is needed from in-
tervening areas in Fayette, Bastrop, Lee, and
Williamson counties. As in the Kewanius lineage, the
question also arises as to whether the parapatric
forms really are reproductively isolated and hence
full species.
Thus as stated in the introduction, a question that
must be posed at the conclusion of this study is
whether the parapatric ‘‘species’’ of Eury-
merodesmus and Sigmoria really are true species,
reproductively isolated from each other. They
facilitate comprehension of the mosaics but may be
more on the order of semispecies. Perhaps the parent
major components in Sigmoria, which demonstrate
properties of species (Shelley and Whitehead 1986),
and the lineages in Eurymerodesmus should be
viewed as superspecies and hence the functional
species. However, semispecies and superspecies lack
nomenclatorial status and cannot be recognized tax-
onomically, so it is convenient to continue the prac-
tice of recognizing the localized, primarily parapatric
entities at the species level, particularly since many
have already been named by past authors.
The present contribution therefore applies the
species category to two markedly dissimilar entities
that logically should be discriminated. Eurymero-
desmus birdi and mundus are broadly sympatric
over a large area, unquestionably reproductively
isolated, and hence unequivocal species. Eurymero-
desmus hispidipes, with two small, widely allopatric
populations, may also be such but is too poorly
known to determine now. The “‘species’’ in the
Kewanius and melacis lineages, however, are chiefly
localized with varying degrees of differentiation.
Sympatry, when it occurs, is mostly restricted to nar-
row zones at interfaces. Hence, the latter entities are
not equivalent to birdi and mundus, so perhaps the
true species in the Kewanius and melacis lineages are
the lineages themselves. Consequently, a more
theoretically sound view of Eurymerodesmus may be
that it contains only five component species. This
decision and the companion theoretical considera-
tions are left to future workers.
CONCLUSION
Diplopodological research lags far behind that on
other animals, even other arthropods, despite the
impressive research potential of millipeds. Hoffman
(1969, 1985) equated the state of the art with those of
entomology and ornithology in the early 1800’s and
estimated (1985) that only about 20% of the global
fauna had been described. This percentage ex-
trapolates to around 50,000 “‘species’’ on the planet,
quite possibly a conservative estimate. The prospects
for the future become gloomy when it is noted that
less than a dozen systematic specialists exist to in-
vestigate this large class (Hoffman 1985), with only
three in the entire Western Hemisphere, all in the
eastern United States. Moreover, several of the more
productive and broadly-based students are at or near
the ends of their careers, and few newcomers are
entering the field. As currently practiced,
diplopodology competes poorly with more
glamorous disciplines of modern biology and may
become even more neglected in the next 25-50 years,
when the youngest present investigators retire. I
therefore think that prudence requires a different
slant to taxonomic research, that it should be
directed more towards integrating knowledge
throughout the class than towards the intricacies of
single taxa. There are far more nomenclatorial
tangles and unrevised genera than the few of us can
ever clarify, and we can describe new forms in-
definitely without beginning to impact on the global
fauna. A major objective of scientific research is the
drawing of generalities from cumulative individual
observations, and if this is ever to happen with
millipeds, it must be now while a number of
knowledgeable investigators is still alive and produc-
tive. Consequently, it is timely to expand beyond
purely descriptive and revisionistic taxonomy by
evaluating the knowledge from such studies for
possible insights into the Diplopoda as a whole.
Common patterns, shared by many taxa, may hold
broad evolutionary implications for the class, but
these patterns must first be identified.
With revisions of two speciose mosaic complexes
now complete, those of Sigmoria (Shelley 1981,
Shelley and Whitehead 1986) and Eurymerodesmus,
the questions arise as to how widespread the
phenomenon is and whether it holds general prin-
ciples for these poorly vagile arthropods. Shelley and
ROWLAND M. SHELLEY 107
Whitehead (1986) noted that problem groups, in
which species and genera are difficult to delineate,
exist throughout the class and suggested the follow-
ing as examples of potential mosaics: Rhysodesmus,
Nannaria, and the California representatives of the
tribe Xystocheirini (Xystocheir, Paimokia, Amplo-
cheir, and Motyxia) (Polydesmida: Xystodesmidae);
Amplinus and Pycnotropis (Polydesmida: Platy-
rhacidae); Chondrodesmus (Polydesmida: Chelo-
desmidae); and the family Rhinocricidae (Spiro-
bolida). I have subsequently scoured the modern
literature for evidence of additional complexes and
suggested (1989) unspecified genera in the following
families: Glomeridesmidae (Glomeridesmida); Spi-
robolellidae and Pachybolidae (Spirobolida); Spiro-
streptidae, Cambalopsidae, and Pseudonanno-
lenidae (Spirostreptida); Conotylidae and Clei-
dogonidae (Chordeumatida); and Paradoxoso-
matidae, Chelodesmidae, Platyrhacidae, Oxydes-
midae, Xystodesmidae, Sphaeriodesmidae, Polydes-
midae, Fuhrmannodesmidae, Pyrgodesmidae, and
Dalodesmidae (Polydesmida). I now augment this
roster by listing (Table 8) all milliped genera with 10
or more known or suspected species according to
Hoffman (1979); those with 20 or more congeners
are denoted by asterisks. Most of these taxa have not
received comprehensive or even cursory modern
study and cannot be reviewed. However, the
literature of many that have been investigated con-
tains evidence of similarities to Sigmoria and
Eurymerodesmus, suggesting comparable underly-
ing phenomena. Consequently, these and unstudied
genera in table 8 may be additional, but currently
undetected, mosaics.
Table 8. Speciose Milliped Genera (10 or more species) (Hoffman 1979)
PENICILLATA
POLYXENIDA
Polyxenidae
Polyxenus - about 10 species
Lophoproctidae
Lophoturus - about 15
PENTAZONIA
GLOMERIDESMIDA
Glomeridesmidae
*Glomeridesmus - 23
SPHAEROTHERIIDA
Sphaerotheriidae
*Sphaerotherium - about 50
Cyliosoma - about 10
Sphaeropoeidae
Sphaeropoeus - about 12
*Castanotherium - about 20
GLOMERIDA
Glomeridae
*Glomeris - about 80
Trachysphaeridae
*Trachysphaera - about 30
HELMINTHOMORPHA
POLYZONIIDA
Polyzoniida
Polyzonium - about 11
STEMMIULIDA
Stemmiulidae
* Diopsiulus - 37
*Prostemmiulus - about 42
*Stemmiulus - 20
SPIROBOLIDA
Messicobolidae
Messicobolus - 14
Atopetholidae
Arinolus - 11
Spirobolellidae
* Spirobolellus - about 80
MEM. AMER. ENT. SOC., 37
*taxa with = 20 species
Rhinocricidae
*FEurhinocricus - numerous
*Salpidobolus - numerous
Pachybolidae
Aulacobolus - about 12
*Centrobolus - about 44
Eucarlia - about 10
SPIROSTREPTIDA
Cambalidae
*Nannolene - about 22
Cambalopsidae
*Trachyjulus - about 23
Hypocambala - about 10
Iulomorphidae
Iulomorphia - about 14
Pseudonannnolenidae
*Pseudonannolene - about 30
*Epinannolene - about 25
Spirostreptidae
Alloporus - about 17
Archispirostreptus — about 15
Gymnostreptus - about 17
*Orthoporus - about 40
Plusioporus - about 13
Urostreptus - 11
Triaenostreptus - 10
Harpagophoridae
Zinophora - about 16
Gonoplectus - 19
Thyropygus - about 18
Remulopygus - about 10
Spissustreptus - 10
Odontopygidae
*Peridontopyge - about 30
Chaleponcus - 19
Helicochetus - 13
Kompsoprium - 12
Patinatius - 13
*Prionopetalum - 21
Rhamphidaropoides - 15
* Spinotarsus - about 96
Syndesmogenus - 13
Tibiomus - 10
JULIDA
Julidae
Julus - 13
Ophyiulus - 18
*Leptoiulus - numerous
Xestoiulus - about 16
* Allajulus - numerous
Typloiulus - 17
*Amblyiulus - numerous
Dolichoiulus - 19
Mesoiulus - about 10
Pachyiulus - about 15
* Anaulaciulus - about 30
*Megaphyllum - numerous
*Ommatoiulus - numerous
Parajulidae
Aniulus - about 17
Bollmaniulus - about 12
Oriulus - about 10
Parajulus - about 15
Mongoliulidae
Skleroprotopus - 14
Paeromopodidae
Californiulus - 10
Blaniulidae
Blaniulus - about 10
Nopoiulus - 10
SIPHONOPHORIDA
Siphonophoridae
* Siphonophora - uncertain
PLATYDESMIDA
Platydesmidae
*Platydesmus - 27
108 EURYMERODESMID MILLIPEDS
To document the evidence for mosaics for specific
genera in table 8 would be prohibitively long (general
clues to mosaics are detailed briefly in the appendix),
but the undeniable literature pattern must be
acknowledged. To varying degrees throughout the
Diplopoda there are statements and characteriza-
tions in publications on speciose genera resembling
ones made years ago about Sigmoria and Eury-
merodesmus, suggesting that these genera are other
mosaic assemblages. The cumulative weight of this
evidence implies that mosaic complexes are a major
evolutionary phenomenon warranting serious con-
sideration by diplopodologists. I focus here on large,
speciose mosaics because their literature clues are
easier to detect, but smaller, less diverse taxa can also
constitute mosaics, for example the xystodesmid
genus Dicellarius, with five species and five
subspecies (Shelley 1984c). Since they are less in-
tricate and their literature is comparatively straight-
forward, small mosaics are easier to grasp. They do
not generate the massive confusion of large mosaics
covering sizeable parts of one or more continents, so
for the purposes of this contribution, table 8 is
restricted to genera with 10 or more species accord-
ing to Hoffman (1979). There are undoubtedly other
speciose genera currently with less than 10 known
species, for example Striaria (Chordeumatida:
Striarlidae), widespread in eastern and western
North America. My purpose here is not to state
absolutes but to alert diplopodologists to a wide-
spread phenomenon and to encourage searching
beyond parochial taxonomic and faunal interests for
general underlying principles while there are still
enough broadly-based workers to do so. We will
never know the full intricacies of such widely-
ranging genera as Barydesmus, Chondrodesmus,
Mestosoma, Orthoporus, Eurhinocricus, and
Spirobolellus; the necessary levels of field sampling
are not currently possible, and many forms are
probably becoming extinct through deforestation
and habitat loss. However, diplopodologists can still
attempt to infer common properties and
Table 8. (Continued)
CALLIPODIDA
Schizopetalidae
Acanthopetalum - 15
Eurygyrus - about 14
Apfelbeckia - about 10
CHORDEUMATIDA
Conotylidae
Conotyla - 13
Taiyutyla - 14
Mastigophorophyllidae
Mastigophorophyllon - about 13
Haaseidae
Haasea - about 14
Cleidogonidae
*Pseudotremia - about 25
*Cleidogona - about 74
Trichopetalidae
Trichopetalum - about 10
Craspedosomatidae
Craspedosoma - about 10
*Pyrgocyphosoma - about 28
*Ochogona - uncertain
Chordeumatidae
Chordeuma - about 12
Anthroleucosomatidae
Anamastigona - about 12
Brachychaeteuma - 10
Caseyidae
*Caseya - 21
Opiona - 13
POLYDESMIDA
Paradoxosomatidae
Aulacoporus - 10
Aschistodesmus - 15
Eviulisoma - 15
*Scolodesmus - 20 _
Eustrongylosma - 15
Antheromorpha - 12
Orthomorpha - 17
Tectoporus - 14
Xanthodesmus - 14
Stosatea - about 14
Catharosoma - 13
*Mestosoma - about 50
Chelodesmidae
Eurydesmus - about 17
*Leptodesmus - 21
Biporodesmus - 10
*Trichomorpha - about 35
*Chondrodesmus - about 40
Brasilodesmus - 11
Caraibodesmus - 10
Strongylomorpha - about 10
Paracordyloporus - about 15
* Amphelictogon - 22
Sandalodesmus - 13
Xystodesmidae
* Xystocheir (Paimokia,
Amplocheir, Motyxia) - uncertain
*Riukiaria - about 20
*Rhysodesmus - about 70
Stenodesmus - 10
*Brachoria - about 28
*Sigmoria - 65
*Nannaria - about 28
Parafontaria - about 16
Eurymerodesmidae
*Eurymerodesmus - 25
Gomphodesmidae
*Ulodesmus - about 20
Oxydesmidae
*Coromus - numerous
Lyodesmus - about 13
Sphaeriodesmidae
* Sphaeriodesmus - 21
Platyrhacidae
*Platyrhacus - numerous
* Barydesmus - numerous
* Psammodesmus - numerous
* Aphelidesmus - about 28
*Amplinus - about 28
Pycnotropis - about 15
Rhacodesmidae
Aceratophallus - 12
Polydesmidae
*Epanerchodus - numerous
*Polydesmus - numerous
Pseudopolydesmus - about 13
Scytonotus - about 11
Cryptodesmidae
Apomus - 12
Peridontodesmus - about 11
* Aporodesmus - about 25
Nearctodesmidae
Nearctodesmus - about 13
Pyrgodesmidae
Calymmodesmus - about 16
Docodesmus - about 14
Cyrtodesmidae
*Cyrtodesmus - about 25
Dalodesmidae
Anaulacodesmus - 16
*Icosidesmus - about 38
Dityloura - 12
*Vanhoeffenia - about 70
ROWLAND M. SHELLEY
mechanisms, and search with open minds for basic
truths applicable to the broad spectrum of the class.
Interpretation of mosaic complexes is beyond the
scope of this work, but the phenomenon raises many
intriguing questions. Why do so many millipeds
form these assemblages, and how does this process
occur? Is the phenomenon applicable to any genus?
When we name and describe “‘species’’ in mosaics,
are we really characterizing species or lesser entities,
perhaps semispecies? As may be the case with the
Kewanius and melacis lineages of Eurymerodesmus,
do some of our proposed species groups and possibly
even genera instead constitute species, thus implying
that higher hierarchical levels may need to be
lowered? Do mosaic complexes hold clues or ex-
planations for other phenomena? Since partitioning
coupled with extinctions isolated the highly atypical
forms in the southern periphery of Sigmoria, could
this general process explain the origins of such
enigmatic geographical oddities as Choctella
(Spirostreptida: Pseudonannolenidae) (Shelley
1989)? Could it be the former northern component
of an ancient pseudonannolenid mosaic that is so
anatomically dissimilar as to have appeared to re-
quire a monotypic family (Chamberlin and Hoff-
man 1950; Hoffman 1969, 1979) and is now so
isolated as to be the only remaining Nearctic
representative? Does much of today’s global
diplopod picture reflect stages in the formation and
breakdown of mosaics, with some genera in expan-
sion phases, others, like Sigmoria and Eurymerodes-
mus, near their zeniths, and still others in periods of
decline, with significant anatomical and
geographical discontinuities resulting from extinc-
tions? As suggested by Shelley and Whitehead (1986)
does this phenomenon in millipeds mirror ones in
other poorly vagile organisms, for example certain
soil insects, arachnids, and possibly even other
myriapods, and could diplopods become something
of a model for investigating it? Perhaps the road to
significantly advancing diplopodology and attracting
more students is not through classical revisions and
faunistic treatments, though these are necessary, but
through the entirely different route of investigations
on shared phenomena, common to many taxa.
Perhaps the few current diplopodologists, in an ef-
fort to salvage the field and heighten its appeal to
outside specialists and potential new students, should
concentrate more on dynamic processes and less on
static descriptions, in essence focusing on what
millipeds do instead of what they are. The dynamics
of mosaic complexes seems a promising arena for
future investigations that may significantly enhance
MEM. AMER. ENT. SOC., 37
109
our understanding of milliped evolution. With a
diminishing number of specialists and a correspon-
dingly bleak outlook, the time to pursue it is now.
ACKNOWLEDGMENTS
I am indebted to the following colleagues for loaning the in-
dicated holotype and paratype specimens, in many cases twice, and
other material under their care: Jonathan Coddington, NMNH,
Polydesmus hispidipes, P. varius, Eurymerodesmus birdi, E.
booneus, E. christianus, E. creolus, E. digitatus, E. hamatilis, E.
louisianae, E. melacis, E. mundus, E. newtonus, E. oliphantus,
and E. simplex; the late Selwyn S. Roback, ANSP, Polydesmus
hispidipes, P. impurus, Eurymerodesmus angularis, E. bentonus,
E. planus, E. plishneri, E. sanbernardiensis, E. spectabilis, and E.
wellesleybentoni; Norman I. Platnick, AMNH, Eurymerodesmus
amplus, E. goodi, and Paresmus polkensis; James S. Ashe,
FMNH, Eurymerodesmus dubius, and E. schmidti; Herbert W.
Levi, MCZ, Eurymerodesmus minimus, and E. simplex; and G. B.
Edwards, FSCA, Eurymerodesmus digitatus, and E. hamatilis.
The loan by Dr. Edwards of the FSCA collection resulted in
designations of the following holotypes and neotypes: Eurymero-
desmus caesariatus, E. clavatus, E. compressus, E. crassatus, E.
dactylocyphus, E. elevatus, E. serratus, and Paresmus paroicus;
that by Dr. Coddington from the NMNH resulted in designation
of the neotype of Paresmus pulaski. My colleagues in
diplopodology, Richard L. Hoffman and William A. Shear,
donated their personal collections, and Dr. Hoffman provided
valuable review comments on a draft of the manuscript and advice
on family relationships in the Chelodesmidea. The following
curators kindly loaned material from the indicated institutional
holdings: Wojciech J. Pulawski (CAS), Richard C. Funk (EIU),
Donald W. Webb (ILNHS), Richard L. Brown (MEM), Jean-Paul
Mauries (MNHP), Norman V. Horner (MWSU), William A.
Drew (OKSU), William W. Gibson (SFAU), James E. Gillaspy
(TAI), James R. Reddell (TMM), Cecil L. Smith (UGA), Robert
L. Blinn (UMO), Harley O. Brown (UOK), and Richard S. Zack
(WASU). Dr. Funk generously agreed to transferring the impor-
tant Coles County, Illinois, male of hispidipes from the EIU to the
NCSM holdings, and the following colleagues also donated eury-
merodesmids to this collection: James C. Cokendolpher, James E.
Gardner, John M. Nelson, Henry W. Robison, Greg Spicer, and
the late William H. Cross. Donald R. Whitehead constructively
reviewed the concluding section and provided numerous ideas
about mosaics and analytical aspects. Cathy Wood typed and
retyped the manuscript, and Renaldo G. Kuhler, NCSM scientific
illustrator, prepared figures 5-6, 11, 14, 18, 21, 23, 25, 27, 30, 32,
35, 39, 42, 46, 49-51, 56, 59, 62, 64, 68, 73, 76-78, 80, 84, 89, 91,
97, 103, 109-110, 114, 117, 123, 127, 130, 134, 137, 142, 146,
149-151, 154-158, 162, 165, 169-171, 178, 184, 187, 191, 198, 201,
and 205, and inked the dissected gonopod drawings. Permission to
collect on the Rob and Bessie Welder Wildlife Refuge, Texas, was
granted by the Foundation office; that to sample in Marjorie Kin-
nan Rawlings State Historic Site, Florida, was courtesy of the
Florida Department of Natural Resources, Division of Recreation
and Parks. Short-term visitor awards from the Smithsonian In-
stitution from 1986-1988 enabled the author to search for unla-
beled eurymerodesmid samples at the NMNH. Field work in south
Texas and Coles County, Illinois, was supported in part by grants
3203-85 and 3871-88 from the National Geographic Society; that
in northeast Texas, Arkansas, Missouri, southern Illinois, and
Florida was funded by grants DEB-7702596, DEB-8200556, and
BSR-8706727 from the National Science Foundation. The last
110
award also defrayed publication costs. Finally, I would be remiss
not to acknowledge the late Dr. Nell B. Causey, whose substantial
collection of eurymerodesmids, the largest in existence and now at
the FSCA, made this study feasible. Without her invaluable
material, laborious years of field work would be necessary before
anyone could begin to clarify these fascinating arthropods.
LITERATURE CITED
ATTEMS, Cart G. 1938. Polydesmoidea II, Fams.
todesmidae, Platyrhacidae, Oxydesmidae,
phodesmidae. Das Tierreich, Lief, 69:1-487.
1940. Polydesmoidea III, Fams. Polydesmidae,
Vanhoeffenidae, Cryptodesmidae, Oniscodesmidae,
Sphaerotrichopidae, Peridontodesmidae, Rhachidesmidae,
Macellolophidae, Pandirodesmidae. Das Tierreich, Lief.
70:1-576.
BOLLMAN, CHARLES H. 1888a. Preliminary list of the
Myriapods of Arkansas, with descriptions of new species.
Entomol. Amer. 4:1-8.
1888b. Notes upon some myriapods belonging to
the U. S. National Museum. Proc. U. S. Natl. Mus.
11:343-350.
1893. The Myriapoda of North America. Bull.
No. 46, U. S. Natl. Mus. 210 pp.
BRIMLEY, CLEMENT S. 1938. Insects of North Carolina. North
Carolina Department of Agriculture, Division of En-
tomology, Raleigh, 560 pp.
BROLEMANN, HENRY W. 1895. Liste de myriapodes des Etats-
Unis, et principalement de la Caroline du Nord, faisant par-
tie des collections de M. Eugene Simon. Ann. Soc. Entomol.
France 65:43-70.
1900. Myriapodes d’Amerique. Mem. Soc. Zool.
France 13:89-131.
1915. Essai de classification des Polydesmiens
(Myriapodes). Ann. Soc. Entomol. France 84:523-609.
Causey, Nett B. 1940. Ecological and systematic studies on
North Carolina myriapods. Unpublished Ph.D. Thesis,
Zoology Department, Duke University, Durham, North
Carolina, 181 pp.
1950a. Five new Arkansas millipeds of the genera
Eurymerodesmus and Paresmus (Xystodesmidae). Ohio J.
Sci. 50:267-272.
1950b. On four new polydesmoid millipeds. En-
tomol. News 61:193-198.
1951. On Eurymerodesmidae, a new family of
Diplopoda (Strongylosomidae), and a new Arkansas species
of Eurymerodesmus. Proc. Arkansas Acad. Sci. 4:69-71.
Lep-
Gom-
1952a. On three eurymerodesmid millipeds and
notes on Paresmus impurus (Wood). Entomol. News
63:169-176.
1952b. Some records and descriptions of
polydesmoid millipeds from the United States. Chicago
Acad. Sci. Nat. Hist. Misc. No. 106, 11 pp.
1954. Three new species and new records of
southern millipeds. Tulane Stud. Zool. 2:63-68.
1963. Additional records of Louisiana millipeds.
Proc. Louisiana Acad. Sci. 26:76-79.
CHAMBERLIN, RALPH V. 1918. Myriopods from Okefenokee
Swamp, GA, and from Natchitoches Par., Louisiana. Ann.
Entomol. Soc. Amer. 11:369-380.
1920. A new leptodesmid diplopod from Louisi-
ana. Proc. Biol. Soc. Wash. 33:97-100.
EURYMERODESMID MILLIPEDS
1931. Onacollection of chilopods and diplopods
from Oklahoma. Entomol. News 42:97-104.
1938. New diplopods. Proc. Biol. Soc. Wash.
51:205-208.
1942a. New southern millipeds. Bull. Univ. Utah
32[Biol. Ser. 6(8)]:1-19.
1942b. Ona collection of myriapods from Iowa.
Can. Entomol. 74:15-17.
1943. On nine North American polydesmoid
millipeds. Proc. Biol. Soc. Wash. 56:35-40.
1946. On four millipeds from Georgia and Mis-
sissippi. Proc. Biol. Soc. Wash. 59:139-142.
1952. Some American polydesmid millipeds in
the collection of the Chicago Museum of Natural History.
Ann. Entomol. Soc. Amer. 45:553-554.
AND RICHARD L. HOFFMAN. 1950. On some
genera and families of North American diplopods. Chicago
Acad Sci. Nat. Hist. Misc. No. 71, 7 pp.
AND 1958. Checklist of the
millipeds of North America. Bull. No. 212, U. S. Natl.
Mus., 236 pp.
AND STANLEY Mutaik. 1941. Ona collection of
millipeds from Texas and New Mexico. J. New York En-
tomol. Soc. 49-57-64.
EBERHARD, WILLIAM G. 1985. Sexual Selection and Animal
Genitalia. Harvard University Press, Cambridge, MA, 244
pp.
GuntTHorP, Horace. 1913. Annotated list of the Diplopoda
and Chilopoda, with a key to the Myriapoda of Kansas.
Kansas Univ. Sci. Bull. 7:161-182.
HOFFMAN, RICHARD L. 1969. The origins and affinities of the
southern Appalachian diplopod fauna. pp. 221-246, In:
Perry C. Holt, ed., 1969. The Distributional History of the
Biota of the Southern Appalachians, Part I: Invertebrates.
Research Division Monograph 1, Virginia Polytechnic In-
stitute, Blacksburg, VA, 295 pp + i-vi.
1978. On the classification and phylogeny of
chelodesmoid Diplopoda. Abh. Verh. Naturwiss. Ver.
Hamburg, (NF)21/22:21-31.
1979. Classification of the Diplopoda. Museum
d’Histoire Naturelle, Geneva, Switzerland, 237 pp.
1981. A note on some supposed Texan localities
for some Araneus species (Araneae, Araneidae). J.
Arachnol. 10:93-95.
1985. Biological and systematic problems in-
volving soil dwelling arthropods. Quaes. Entomol.
21:543-557.
JEEKEL, C. A. W. 1971. Nomenclator generum et familiarum
Diplopodorum: A list of the genus and family-group names
in the Class Diplopoda from the 10th edition of Linnaeus,
1758, to the end of 1957. Monog. Nederl. Entom. Vereng.,
no. 5, 412 pp.
KENYON, FREDERICK C. 1893. A preliminary list of the Myria-
poda of Nebraska with descriptions of new species. Pubs.
Nebraska Acad. Sci. 3:14-18.
Loomis, HaRotpD F. 1943. A new genus of Virginia millipeds
related to Scytonotus and a new species from Florida. J.
Wash. Acad. Sci. 33:318-320.
1959. Millipeds collected enroute from Florida to
San Antonio, Texas, and vicinity. J. Wash. Acad. Sci.
49:157-163.
1969. New and known millipeds from four
southern states. Florida Entomol. 52:245-251.
ROWLAND M. SHELLEY
1976. Two new species of diplopods from Texas
and one from Mexico. Florida Entomol., 59:287-292.
McNeil, JEROME. 1887. List of the myriapods found in
Escambia County, Florida, with descriptions of six new
species. Proc. U. S. Natl. Mus. 10:323-327.
REDDELL, JAMES R. 1970. A checklist of the cave fauna of
Texas. IV. Additional records of Invertebrata (exclusive of
Insecta). Texas J. Sci. 21:389-415.
SHELLEY, ROWLAND M. 1977. Appendicular abnormalities in
the milliped family Xystodesmidae (Polydesmida). Can. J.
Zool. 55:1014-1018.
1978. Millipeds of the eastern Piedmont region
of North Carolina, U.S.A. (Diplopoda). J. Nat. Hist.
12:37-79.
1980. Revision of the milliped genus Pleuroloma
(Polydesmida:Xystodesmidae). Can. J. Zool. 58:129-168.
1981. Revision of the milliped genus Sigmoria
(Polydesmida:Xystodesmidae). Mem. Amer. Entomol. Soc.
No. 33, 140 pp.
1982. Revision of the milliped genus Auturus
(Polydesmida:Platyrhacidae). Can. J. Zool. 60:3249-3267.
1984a. A synopsis of the milliped genus Abacion
Rafinesque (Callipodida: Caspiopetalidae). Can. J. Zool.
62:980-988.
1984b. Revision of the milliped genus Cheiropus
(Polydesmida: Xystodesmidae). Proc. Biol. Soc. Wash.
97:263-284.
1984c. A revision of the milliped genus Dicellar-
ius, with a_ revalidation of the genus Thrinaxoria
(Polydesmida: Xystodesmidae). Proc. Biol. Soc. Wash.
97:473-512.
1989. Are allopatric/parapatric mosaic com-
plexes widespread in the Diplopoda? p. 23. In. A Minelli,
ed., Proceedings of the 7th International Congress of
Myriapodology. E. J. Brill, Leiden.
AND DONALD R. WHITEHEAD. 1986. A recon-
sideration of the milliped genus Sigmoria, with a revision of
Deltotaria and an analysis of the genera in the tribe
Apheloriini. Mem. Amer. Entomol. Soc. No. 35, 223 pp.
STEWART, THOMASC. 1969. Records of millipeds in twenty five
northeast Texas counties. Texas J. Sci. 20:383-385.
Woop, Horatio C. 1864. Descriptions of new species of
North American Polydesmidae. Proc. Acad. Nat. Sci. Phila.
16:6-10.
1865. The Myriapoda of North America. Trans.
Amer. Philosoph. Soc. 13:137-248.
itp
1867. Descriptions of new species of Texas
Myriapoda. Proc. Acad. Nat. Sci. Phila. 19:42-44.
Wray, Davip L. 1967. Insects of North Carolina, Third Sup-
plement. North Carolina Department of Agriculture, Divi-
sion of Entomology, Raleigh, 181 pp.
APPENDIX
Literature Clues to Speciose Mosaic Complexes in the Diplopoda
1. Restricted component distributions.
2. Parapatry of relatively localized components as revealed by
distribution maps like that of Eurymerodesmus species in Texas
(Fig. 212).
3. Statements about small or minor differences between species.
4. Uncertainty, confusion, and/or contradictions and disagree-
ments among authors, particularly concerning the composition,
generic positions, or taxonomic statuses of species groups.
Variations of general anatomical (usually gonopodal) patterns,
like those shown by the subgenera of Sigmoria and the lineages of
Eurymerodesmus, are subject to different interpretations by dif-
ferent authors, or the same author at different stages, which in
turn lead to conflicting published opinions about generic validities,
synonymies, and group compositions. When several such opinions
are published, as with genera in the tribe Amplinini (Polydesmida:
Platyrhacidae), a highly confused literature results, with contradic-
tions and disagreements similar to those in Eurymerodesmus and
Sigmoria before they were revised. Thus, muddled literature, a
reflection of inherent intricacy, is a particularly telling clue to a
potential mosaic.
A common thread in the literature of speciose genera are state-
ments about small or minor differences between species, for exam-
ple as between medlacis/sanbernardiensis/digitatus and
elevatus/newtonus/amplus/varius in Eurymerodesmus and be-
tween congeners in all subgenera of Sigmoria (Shelley 1981,
Shelley and Whitehead 1986). In the latter genus, contiguous
populations of adjacent species are often phenotypically similar
because of residual ancestral character clines traversing the bound-
aries. Hence it was difficult to formulate definitions of many
species that excluded neighboring populations of others, and
initially, virtually every sample in parts of the range appeared to
represent a separate species. Likewise in Eurymerodesmus, species
definitions are difficult in the Kewanius and melacis lineages. Con-
sequently, statements about slight differences between species in
other genera probably reflect the effects of a mosaic, which should
be expected when a genus appears to contain an inordinate number
of species.
112 EURYMERODESMID MILLIPEDS
INDEX TO TAXA
Synonyms in italics, new species in boldface
amplus, Eurymerodesmus — 45
angularis, Eurymerodesmus — 31
bentonus, Eurymerodesmus — 53
birdi, Eurymerodesmus — 72, 74
birdi birdi, Eurymerodesmus — 74
birdi planus, Eurymerodesmus — 77
booneus, Eurymerodesmus — 7
caesariatus, Eurymerodesmus — 64
christianus, Eurymerodesmus — 46
clavatus, Eurymerodesmus — 94
columbus, Paresmus — 38
compressus, Eurymerodesmus — 35
crassatus, Eurymerodesmus — 66
creolus, Eurymerodesmus — 74
dactylocyphus, Eurymerodesmus — 87
digitatus, Eurymerodesmus — 91
dubius, Eurymerodesmus — 38
elevatus, Eurymerodesmus — 59
Eurymerodesmidae — 18
Eurymerodesmus — 19
evides, Auturus — 7
floridus, Leptodesmus — 78
goodi, Eurymerodesmus — 36
hamatilis, Eurymerodesmus — 49
hispidipes, Eurymerodesmus — 25, 49, 55
Kewanius — 19
louisianae, Eurymerodesmus — 47
melacis, Eurymerodesmus — 82
minimus, Eurymerodesmus — 43
mundus, Eurymerodesmus — 78
newtonus, Eurymerodesmus — 53
oliphantus, Eurymerodesmus — 55
Paresmus — 19
paroicus, Eurymerodesmus — 65
paroicus, Paresmus — 65
planus, Eurymerodesmus — 77
plishneri, Eurymerodesmus — 74
polkensis, Eurymerodesmus — 62
polkensis, Paresmus — 62
pulaski, Eurymerodesmus — 68
pulaski, Paresmus — 68
sanbernardiensis, Eurymerodesmus — 90
schmidti], Eurymerodesmus — 74
serratus, Kurymerodesmus — 69
simplex, Eurymerodesmus — 60
simplex, Kewanius — 60
spectabilis, Eurymerodesmus — 47
varius, Eurymerodesmus — 41, 43
varius, Leptodesmus — 43
varius, Polydesmus — 43
hispidipes, Leptodesmus — 25, 49, 68, 69, 74, 78 varius christianus, Eurymerodesmus — 46
hispidipes, Polydesmus (Paradesmus) — 25 varius louisianae, Eurymerodesmus — 47
hispidipes, Polydesmus (Polydesmus) — 25 varius varius, Eurymerodesmus — 43
wellesleybentoni, Eurymerodesmus — 31
wellesleybentonus, Eurymerodesmus — 31
impurus, Eurymerodesmus — 29
impurus, Leptodesmus — 29
impurus, Paresmus — 29
impurus, Polydesmus — 29
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; Memoirs OF THe AMERICAN eroacchecis 6 ‘Society
‘The Cresson Types of Hymenoptera, Ezra T, Cresson, 1916, $12.00.
4
A Venational Study of the Suborder Zygoptera (Odonata), with Keys Ls the ‘aeeeienion of Genera, Philip: A
Munz, 1919, $10.00, —
. The Blattidae of Panama. Morgan Hebard. 1920. $8.00.
The Types of Hymenoptera in the Academy of eos i Sciences of Philadelphia other than those of Ezra T. Crewon, Ezra
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~The Eumastacinae of Southern Mexico and Central ‘America, James A.G, Rehn and John WH, Rehn. 1934, $8.00.
The Generic Names of the Sphecoid Wasps and Their Type Species. W.S.L. Pate, 1937. $8.00,
A Revision of the North American ‘Species Belonging to the Genuz Pegomyia. H.C, ‘Huckett, 1941, $6.00,
Catalogue and Reclassification of the Nearctic Ichneumonidae, Henry K. Townes, Jr. 1944, ‘$20. 00,
. Elachistidae of North America (Microlepidoptera). — Annette F. Braun, 1948. $1000. :
~ Classification of the Blattaria as Indicated by their Wings (Orthoptera), John W.H, Rehn. 1951, $10.00, -
_ A Taxonomic Study of the North American Licinini with Notes on the Old World Species of the Genus Diplocheila Brulle
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20. A Revision of the Mexican and Central American Spider ‘Wasps of the Subfamily Pompilinae (Hymenoptera: Pane,
Howard: E. Evans, 1966, $12.50, f be
‘A Taxonomic and Zoogeographic Survey of the Scarabacinae of the Antilles (Coleoptera: Scarabaeidae), Eric G.
“Matthews. 1966, $5.00.
A Monograph of the Ithomiidae (Lepidoptera) Part Wn. The Tribe Mechanitini Fox. Richard M. Fox. 1967, $9.00.
. A List of New North American Spiders, 1940-1966. _ Beatrice | R. Vogel. 1967. $9.00.
The Schizopteridae (Hemiptera: Heteroptera) with the Descr Description of New Species from Trinidad. Michael G. ‘Emsley, 1969,
$6.50.
_ATasonomic Revision of the Aquatic Bete Genus Laccophi (Dyce) of Noth Americ James R, Zimmerman, 1970. oY
$12.00, —
A Revision of Md ‘Nearctic Species of the Tribe Parydrini (Diptera: Ephydridae) Philip J, Clausen and Edwin F.
Cook. 4971, $7.00.
-‘Tischeriidae of America North of Mexico (Microlepidoptera). Annette F. Braun, 1972. $7.00,
. The Shield-backed Katydids of the Genus /diostatus. David C. Rentz, 1973, $9.50. ; ;
_ The Systematic and Morphology of the Nearctic Species of Diamesa Meigen, 1835 (Diptera: Chironomidae). Dean Cyrus
Hansen and Edwin F. Cook. 1976, $10.00.
The Stoneflies (Plecoptera) of the Rocky Mountains, Richard W, Baumann, Arden R, Gaufin and Rebecca F. Surdick. ‘197
$10.50,
_ The Genus /soperia (Plecoptera) of Western North America; Holomorphology and Systematics, and a New Stonefly Genus:
Cascadoperla. Stanley W. Szczytko and Kenneth W. Stewart, 1979, $7.50.
Revision of the Milliped Genus Sigmoria (Polydesmida; Xystodesmidac). Rowland M. Shelley. 1981. $11.00
. Proceedings of the 8th International Symposium on Chironomidae, Jacksonville, Florida, July 25-28, 1982, 1983. $25,00,
A reconsideration of the Milliped Genus Sigmoria, with a revision of Delrotaria and an analysis of the genera in the tribe
Apheloriini (Polydesmida: Xystodesmidae), Rowland M, Shelley and David R, Whitehead, 1986, $16.00
~ Biosystematics ace Dicrotendipes Kieffer, 1913 (Diptera: Chironomidae: Chironominae) of the World. John H.
Epler, 1988,
j Revision of the ae Family Eurymerodesmidae (Polydesmida; eter Rowland M. Shelley. 1989, $14.00
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