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TULANE STUDIES 
IN ZOOLOGY 


Volume 11, No. 1 August 23, 1963 


A STUDY OF THE PARASITES OF THE FLORIDA MOUSE, 
PEROMYSCUS FLORIDANUS, IN RELATION TO 
HOST AND ENVIRONMENTAL FACTORS 


JAMES N. LAYNE, 


DEPARTMENT OF CONSERVATION, CORNELL UNIVERSITY, 
ITHACA, NEW YORE 


| Marine Biological Laboratory 
LIBRARY 


SEP 1°3963 
WOODS HOLE, MASS. 


TULANE UNIVERSITY 
NEW ORLEANS 


TULANE STUDIES IN ZOOLOGY is devoted primarily to the zoology of the waters 
and adjacent land areas of the Gulf of Mexico and the Caribbean Sea. Each number is 
issued separately and contains an individual monographic study, or several minor studies. As 
volumes are completed, title pages and tables of contents are distributed to institutions ex- 
changing the entire series. 


Manuscripts submitted for publication are evaluated by the editor or associate editor 
and by an editorial committee selected for each paper. Contributors need not be mem- 
bers of the Tulane University faculty. 


The editors of Tulane Studies in Zoology recommend conformance with the principles 
stated in chapters I and II (only) of the Style Manual for Biological Journals published in 


1960 by the American Institute of Biological Sciences, Washington, D. C. 


The editors also recognize the policy adopted by the Federal Council for Science and Tech- 
nology, and endorsed by the Conference of Biological Editors, that page charges for publica- 
tion of scientific research results in scientific journals will be budgeted for and paid as a 
necessary part of research costs under Federal grants and contracts. Accordingly, writers 
crediting research grant support in their contributions will be requested to defray publica- 
tion costs if allowable under the terms of their specific awards. 


Illustrations and tabular matter in excess of 20 percent of the total number of pages may 
be charged to the author, the levy applied being the excess above 10-point typesetting costs. 


Manuscripts should be submitted on good paper, as original typewritten copy, double- 
spaced, and carefully corrected. Two carbon copies in addition to the original will help 
expedite editing and assure more rapid publication. 


An abstract not exceeding three percent of the length of the original article must accom- 
pany each manuscript submitted. This will be transmitted to Biological Abstracts and any 
other abstracting journal specified by the writer. 


Separate numbers or volumes may be purchased by individuals, but subscriptions are not 
accepted. Remittance should accompany orders from individuals. Authors may obtain 
copies for personal use at cost. 


Address all communications concerning manuscripts and editorial matters to the editor; 
communications concerning exchanges, and orders for individual numbers to the Director, 
Meade Natural History Library. 


When citing this series authors are requested to use the following abbreviations: Tulane 
Stud. Zool. 


Price for this number: $0.60 


Harold A. Dundee, Editor 

Gerald E. Gunning, Associate Editor 
Department of Zoology, 

Tulane University, 

New Orleans, Louisiana 70118, U.S.A. 


Royal D. Suttkus, Director 

Meade Natural History Library, Francis L. Rose, 
Tulane University, . Assistant to the Editors 
New Orleans, Louisiana 70118, U.S.A. 


ete “46 nee. 


a ee 


TULANE STUDIES 
Iw ZOOLOGY 


VOLUME il 
1963-1964 


TULANE UNIVERSITY 
NEW ORLEANS 


TULANE STUDIES IN ZOOLOGY is devoted primarily to the zoology of the waters 
and adjacent land areas of the Gulf of Mexico and the Caribbean Sea. Each number is 
issued separately and contains an individual monographic study, or several minor studies. 
As volumes are completed, title pages and tables of contents are distributed to institu- 
tions exchanging the entire series. 


Manuscripts submitted for publication are evaluated by the editor and by an editorial 
committee selected for each paper. Contributors need not be members of the Tulane 
University faculty. 


MEMBERS OF THE EDITORIAL COMMITTEES FOR 
PAPERS PUBLISHED IN THIS VOLUME 


R. Tucker Abbott, Academy of Natural Sciences, Philadelphia, Pennsylvania 
M. W. Boesel, Miami University, Oxford, Ohio 

Frank A. Brown, Jr., Northwestern University, Evanston, Illinois 

J. S. Dendy, Auburn University, Auburn, Alabama 

Milton Fingerman, Tulane University, New Orleans, Louisiana 

Willard D. Hartman, Yale University, New Haven, Connecticut 

Sewell H. Hopkins, Texas A. & M. University, College Station, Texas 

Allen McIntosh, U. S. Department of Agriculture, Beltsville, Maryland 
Robert W. Menzel, Florida State University, Tallahassee, Florida 

Tametake Nagano, Tohoku University, Sendai, Japan 

George H. Penn,* Tulane University, New Orleans, Louisiana 

Mary Hanson Pritchard, University of Nebraska, Lincoln, Nebraska 

Harald A. Rehder, United States National Museum, Washington, D. C. 
Muriel I. Sandeen,* Duke University, Durham, North Carolina 

Franklin Sogandares-Bernal, Tulane University, New Orleans, Louisiana 
Horace W. Stunkard, American Museum of Natural History, New York, New York 
Kiyoshi Takewaki, Tokyo University, Tokyo, Japan 

Henry Townes, University of Michigan, Ann Arbor, Michigan 

Emily H. Vokes, Tulane University, New Orleans, Louisiana 


* deceased 


NUMBER 
ile 


QO 


CONTENTS OF VOLUME 11 


A STUDY OF THE PARASITES OF THE FLORIDA MOUSE, PEROMYS- 
CUS FLORIDANUS, IN RELATION TO HOST AND ENVIRONMENTAL 
HACTORS: 2 = sh Et UP BE ae ree ool 


James N. Layne 
THE SPONGE FAUNA OF THE ST. GEORGE’S SOUND, APALACHEE 


BAY, AND PANAMA CITY REGIONS OF THE FLORIDA GULF COAST 
Frank J. Little, Jr. 


HORMONAL CONTROL OF THE REFLECTING RETINAL PIGMENT IN 
ib TSOPOD LIGIA COLES! BRANDTs.— = =~ 


Milton Fingerman and Chitaru Oguro 

HORMONAL AND ENVIRONMENTAL REGULATION OF THE MOLTING 
CYCLE IN THE CRAYFISH FAXONELLA CLYPEHATA____ ponies te 
William C. Mobberly, Jr. 

MURICIDAE (GASTROPODA) FROM THE NORTHEAST COAST OF 


SOUTH AMERICA, WITH DESCRIPTIONS OF FOUR NEW SPECIES. 
Harvey R. Bullis, Jr. 


CHIRONOMIDAE (DIPTERA) OF LOUISIANA. I. SYSTEMATICS AND 
IMMATURE STAGES OF SOME LENTIC CHIRONOMIDS OF WEST- 
CENTRAL LOUISIANA______ Se Di dane LO Rt Nell ceo Te, CNA le NaS WIN tetas. VE Poe 


PAGE 


ou) 


99 


James E. Sublette 109 


CHIRONOMIDAE (DIPTERA) OF LOUISIANA. II. THE LIMNOLOGY OF 
THE UPPER PART OF CANE RIVER LAKE, NATCHITOCHES PARISH, 
LOUISIANA, WITH PARTICULAR REFERENCE TO THE EMERGENCE 
OF CHIRONOMIDAB 2... A eS a ee ey De bet eee 


DIGENETIC AND ASPIDOGASTRID TREMATODES FROM MARINE 
BSS. OF) CUMNCAOLAN DY JA MAT CAS coe 8 ee ee ee 


51 


Printed in the U.S.A. 
at New Orleans, by 
HAUSER PRINTING Co., ING. 


TULANE STUDIES IN ZOOLOGY 


Volume 11, No. 1 


August 23, 1963 


CONTENTS 


-. HENCE LOO OF Gs 01 (6 1 fee a ile od ea eee 
REBATE RIAUE: S AUNTIE INLET EG DD Si sooner reer eee i se 
BRIDESGRIPTTON (OB! DAB IAT Gece me orate ree ese Se 
MURESUL TS: 2. Rico) FEE sere eee re ae ee 8 ee 


PMMA EG t-sralee oa». 2 Oy I es wi eee NS EE eas 
ie Species: Recordsjand Relacionsttips tO n1Oste= = 
des Bttect of Host DensingoumeyilenCes= = eS 
peehitect Of Habitatontbreuemcen sen ee ae Te 
AM AE Etect Of Season onubneyalencome ae en ee 


lie. Species Records andehclanonsmips tOnOSt: 2) = ee 
2. Eitect or Hose Denstsonvbrevalences rie 5 2 ee 
pemiefiece of Habicatombteyalencen sere. ann oe ee 
eee bitect of Seasonion ereyalence: see Se eect hao etre Be 


eB 
rj 
on 
Bb 


Jee Species Recotdsand Relationships to Ost == = = Se 
Pep ELecnor ilost Dens ityonse tev alence 25s 
5. Eifect Of Elabitat om etevalence 2) es Be a, ee eee 
An Effect Of Seasom Onskteyalences = et.) BEN eee ee 
MODOERIESS..«.<. bec Serer yer Res elo a EEE a ON rene Oe 
ike Species Record and) Relationshipsto Host... == ee 
Pee Biicce Of Host Densityiambrevalence.. = =e eee 
Se trect:or Habitae onjbtevalence. eee se 2 a a ee 
4a wtrechiol Season One Ptevalences.- = = ee 
NEE EOS PEGE ak Eh Ne tate ee ee Wn a ard cB Sneed Oe 


_ PARC CY 40 Ve Minders ae OR ee. SPO OE DA et ee a bee. EIB ty hi ee 


ae; 1) es 


2. 


merIN GI atOd eS. =. ses ee ed ee ae oe ee, 
LCHIEASEOGNIOS: =o ole eo ae ee SE OL 5 ee 


RBISBREINCES CATED c= tees Samia a Wohl se Mele ay RS ee ae Mae. ee 
IABSURAGIa oN Bie og PN ae hs So A ae 0 Sy Cr SS ae Pe 


EDITORIAL COMMITTEE: 


— 
US) 


— 
2 WY Ov Ov ! 


13 


GEORGE H. PENN,* Professor of Zoology, Tulane University, New Orleans, Louisiana 


FRANKLIN SOGANDARES, Associate Professor of Zoology, Tulane University, New 


Orleans, Louisiana 


ALLEN MCINTOSH, Leader, Parasite Classification and Distribution Investigations, 


U. S. Department of Agriculture, Beltsville, Maryland 


* deceased 


A STUDY OF THE PARASITES OF THE FLORIDA MOUSE, 
PEROMYSCUS FLORIDANUS, IN RELATION TO 
HOST AND ENVIRONMENTAL FACTORS! 


JAMES N. LAYNE, 


Department of Conservation, Cornell University, 
Ithaca, New York 


I. INTRODUCTION 


The kinds and abundance of parasites as- 
sociated with a particular mammalian host 
depend upon a complex interaction of many 
factors. Knowledge of these factors and 
their relative importance is essential to a 
full understanding of the significance of 
parasitism in terms of the biology of the 
host. In the present study an attempt has 
been made to obtain as complete a represen- 
tation as possible of the parasitic fauna of 
the Florida mouse, Peromyscus floridanus 
(Chapman), and to examine the relation- 
ships between the occurrence and prevalence 
of particular parasite species or larger groups 
and such factors as age, sex, and density of 
host, habitat, and season. 


Peromyscus floridanus occurs only in Flor- 
ida, where it is further restricted in its dis- 
tribution to a narrow range of habitats in 
certain parts of the state. It is taxonomically 
distinct from other species of the abundant 
and widespread genus Peromyscus and is re- 
ferred to the monotypic subgenus Podomys. 
The Florida mouse apparently has affinities 
with western species. Its presently restricted 
range probably represents the survival in a 
Floridian refuge of a formerly more wide- 
spread population eliminated in other parts 
of its range by environmental changes dur- 
ing the Pleistocene. Information on the 
parasites of this rodent is therefore of inter- 
est both from the standpoint of its contribu- 
tion to a knowledge of the population dy- 
namics and environmental relationships of 
the species in its present range and habitats 
as well as perhaps providing some indication 
of the ecological factors that might have 
played a role in its evolutionary and distri- 
butional history. 


II. MATERIALS AND METHODS 
The data on parasites included in this 
paper were obtained from over 800 mice 
collected between February, 1957, and Oc- 


1 This study was supported by Grants No. 
G-3215 and No. G-13240 from the National 
Science Foundation. 


tober, 1960. Collections were made at 35 
localities. Thirty-three of these were in six 
counties (Alachua, Clay, Gilchrist, Levy, 
Putnam, and St. Johns) in the northern half 
of peninsular Florida and two were in High- 
lands County in the southern part of the 
State (Fig. 1). All of the known habitats 
of P. floridanus were sampled, in approxi- 
mate proportion to their importance, and an 


SAND PINE SCRUB 

LONGLEAF PINE / TURKEY OAK 
UPLAND HAMMOCK 

PINE FLATWOODS 

SLASH PINE /TURKEY OAK 


XPD +O 


35. x“ 
ae 


L J 


‘ 


Figure 1. Map of peninsular Florida show- 
ing location of habitat types sampled. 


attempt was made to procure adequate sea- 
sonal representation from each habitat type. 
However, low population levels in some 
habitats prevented obtaining sufficient num- 
bers of mice in all seasons. 


Mice were collected with small (2 x 2% 
x 614 in.) Sherman live traps baited with 
rolled oats. The majority of specimens ex- 
amined for parasites were trapped specific- 
ally for laboratory data. A smaller number 
of animals from which parasite data were 


4 Tulane Studies in Zoology 


obtained consisted of specimens that had 
died in traps during mark-and-release studies. 

Traplines of varying numbers of stations 
spaced at approximately 50-ft. intervals were 
used in general collecting. Two traps were 
set at each station. Traplines ordinarily were 
run for periods of one to three days, and the 
number of mice taken per 100 trapnights 
was used as a general index of population 
density. Traps on mark-and-release study 
plots were set in a grid pattern of 50- or 
100-ft. intervals. 

Samples of from 542 to 610 mice were 
examined for different types of ectoparasites. 
Most of the specimens were brought alive 
to the laboratory in traps. Each mouse was 
then transferred directly to a waxed sand- 
wich bag containing a pledget of ether- 
soaked cotton. After the mouse had died, 
the bag was torn open along the sides and 
the specimen, cotton, and bag carefully 
searched for parasites. After the ectopara- 
sites that had dropped off the specimen into 
the bag or become entangled in the cotton 
were collected, the mouse was carefully 
combed and examined, usually with the aid 
of a dissecting microscope, to recover those 
still attached or lodged in the pelage. In 
some cases, specimens were quick frozen in 
the waxed bags after death and examined 
at a later time. Mice found dead in traps on 
mark-and-release study areas usually were 
kept in plastic bags until examined. 

The majority of the mice obtained in the 
course of general collecting or from live- 
trapping study plots were routinely surveyed 
for the more conspicuous types of endopara- 
sites at the time of necropsy. The liver, 
stomach, and intestines of most of the mice 
were preserved in 10% formalin. The livers 
of 698 specimens were subjected to a forma- 
lin-ether concentration technic (Ritchie, 
1948) for examination for certain helminth 
eggs, and the alimentary tracts of 186 speci- 
mens from different habitats were intensive- 
ly searched for parasites. In addition, a 
detailed survey for endoparasites was made 
in a series of 38 freshly-killed mice. Urine 
and kidney tissue of four mice from a single 
locality were cultured and examined for 
leptospires, and blood samples from the same 
individuals were tested for leptospire anti- 
bodies. Blood smears from a number of 
specimens taken during the course of the 
study were checked for parasites after stain- 
ing with Wright's. 


Vol. 11 


Fleas, lice, ticks, and botfly larvae occur- 
ring on individual hosts were counted. The 
presence of mites was recorded. Although 
actual counts of individuals on infested hosts 
were not made, relative abundance was 
noted. Because of limitations of time, not 
every ectoparasite collected was identified. 
The percentages of infested hosts from 
which parasites in a particular group were 
identified are as follows: trombiculid mites, 
5%: non-trombiculid mites, 85%; ticks, 
70%; lice, 100% (only 1 collected); fleas, 
88%; and cuterebrids, 100%. All endopara- 
sites collected were identified to the lowest 
taxonomic level possible, and numbers and 
sites of occurrence in the host were recorded 
for helminths and pentastomids. The abun- 
dance of intestinal protozoans was estimated 
from microslide preparations of fecal smears 
stained with hematoxylin. 

For the purposes of this study, two age 
classes of mice were recognized: young and 
adult. The former category included mice 
in juvenal pelage or undergoing the post- 
juvenal molt, and the latter, specimens in 
which the postjuvenal molt was complete. 
These age classes correspond to chronological 
ages of approximately 4 to 16 weeks and 17 
weeks or older, respectively. 


Ill. DESCRIPTION OF HABITATS 


Mice were collected from several major 
habitat types that are essentially equivalent 
to ones recognized by Rogers (1933), Carr 
(1940), and Laessle (1942, 1958). These 
habitats, together with the number of sta- 
tions of each type sampled, include: sand 
pine scrub—10, longleaf pine/turkey oak— 
16, slash pine/turkey oak—1, upland ham- 
mock—6, and pine flatwoods—2. The up- 
land hammock category employed here cor- 
responds to that termed “ecotonal” in an 
earlier paper (Layne and Griffo, 1961). 

The Florida mouse typically is associated 
with sand pine scrub and longleaf pine/ 
turkey oak habitats. Sand pine scrub is found 
on fine, white, and excessively well-drained 
sandy soils. Sand pine (Pinus clausa) and 
three species of oaks of shrubby habit (Qzer- 
cus myrtifolia, QO. virginiana, and Q. chap- 
mannt) are diagnostic woody plants of this 
association. In most of the scrubs sampled 
in this study, the sand pines were widely 
spaced, and the oaks formed dense, low 
clumps with extensive: patches of bare sand 
between them. Forbs were sparse, and the 


No. | 


litter layer was largely restricted to the area 
beneath clumps of shrubs. Two of the sta- 
tions included in the scrub category were 
somewhat atypical in that slash pine (Pinus 
elliottw) replaced sand pine. In all other 
respects, however, these areas were similar 
to true scrub. 


The longleaf pine/turkey oak association 
also occurs on well-drained sandy soils, usu- 
ally on low ridges or hills. Longleaf pine 
(Pinus australis) and turkey oak (Q. lae- 
vis) are ordinarily the only common tree 
species present. The pine is often sparse 
and in some instances may be absent. At 
some of the stations studied the oaks were 
so closely spaced as to produce nearly a 
closed canopy, but in most cases the stand 
was more open. The shrub and sapling 
understory varied from dense and brushy to 
nearly absent. Forbs were generally more 
abundant than in sand pine scrub habitats, 
and the ground cover was ordinarily better 
developed. Wire grass (Aristida stricta) is 
a characteristic component of the ground 
vegetation in longleaf pine/turkey oak habi- 
tats and may be present as scattered clumps 
or form a nearly continuous cover. The 
former condition, in which open patches of 
ground are frequent, is preferred by the 
Florida mouse. As the turkey oak is decidu- 
ous, the seasonal change in the aspect of the 
vegetation is more pronounced in the long- 
leaf pine/turkey oak habitats than in the 
others studied. 


The slash pine/turkey oak association has 
a limited distribution in the southern part 
of the state. It is intermediate between 
sctub and typical longleaf pine/turkey oak 
in species composition and general aspect. 


Upland hammocks are open woodlands 
with sandy soils that contain more organic 
matter than those of the foregoing habitats 
and which, though still quite dry and well 
drained, tend toward a moister state than 
either scrub or pine/oak associations. South- 
ern red oak (Q. falcata) is a characteristic 
tree in upland hammocks and a number of 
other species, including persimmon. (Dzo- 
spyros virginiana), mockernut hickory (Ca- 
rya tomentosa), loblolly pine (Pinus taeda), 
and laurel oak (Q. laurifolia) may also 
occur. The understory and ground cover 
ranged from heavy to sparse in the examples 
included in this study. Those stations that 
produced the best catches of mice usually 


Layne: Parasttes of Florida Mouse 5 


had numerous stumps, logs, and bare patches 
of exposed soil. Also included in the upland 
hammock category are several stations repre- 
senting ecotones between longleaf pine/tur- 
key oak and mesic hammocks (broad-leaved 
woodlands containing such evergreen species 
as American holly and magnolia) and live 
oak hammocks. Both of these types environ- 
mentally were similar to typical upland ham- 
mocks. The ecotone sites had essentially the 
same species composition and aspect, dif- 
fering chiefly in being more limited in ex- 
tent and thus possibly more influenced by 
contiguous habitats. Live oak hammocks 
differed from typical upland hammocks in 
having the live oak as the dominant tree and 
a lesser variety of other woody plants. The 
understory was usually more open than in 
most of the upland hammocks with red oak 
as the principal tree as the result of closer 
spacing of the live oaks. 


Pine flatwoods occupy level, poorly drained 
soils that are relatively rich in organic mat- 
ter. Pines are the dominant trees, with the 
particular species present depending largely 
upon the edaphic conditions of the site. 
The understory may be variously developed 
and may include young pines and a variety of 
hardwood species. The lower strata are typt- 
cally dense and may range from mostly 
shrubs or palmetto, or an intermixture of 
both, to principally grasses and forbs. As is 
also true of the other habitats mentioned, 
fire is an important factor in determining 
the particular vegetative characteristics of 
flatwoods stands. Peromyscus floridanus ap- 
parently only rarely inhabits pine flatwoods, 
and then only the better drained types dur- 
ing dry seasons or years. In the present 
study, mice were collected in only two flat- 
woods sites, both with longleaf pine and 
grassy ground cover. 


Sand pine scrub and longleaf pine/turkey 
oak are relatively xeric environments. Stud- 
ies by Rogers (1933) of evaporation rates 
in six woodland habitats in northern Florida 
showed scrub and turkey oak to have the 
highest overall evaporation rates, with the 
former having the greatest increment due 
to insolation (Fig. 2). Cooper et al, (1959) 
recorded temperatures in the litter layer of 
a scrub stand in central Florida. Monthly 
maximum temperatures were over 100° F 
every month of the year and above 125° in 
8 months. The highest values occurred from 


6 Tulane Studies in Zoology 


25 

20 
q 
ra 
~~ 
E is 
WwW 
= 
< 
rr 
z 
So 10 
= 
< 
c 
ro) 
a 
$ 
W 5 

SCRUB TURKEY PINE MESIC 
OAK FLATWOODS HAMMOCK 

Figure 2. Average evaporation rates in 


four habitats in northern Florida. Based 
on weekly measurements given by Rogers 
(1933) for the period from 26 February 
through 23 July. White and black bulb 
atmometer readings are shown by solid 
and open portions of bars, respectively. 
The latter represent the increment due to 
insolation. 


May through August, exceeding 60° in 
June. 

Slash pine/turkey oak habitats on the 
basis of general observations appear to agree 
closely with scrub and typical turkey oak in 
their environmental conditions. 

Upland hammocks, though still compara- 
tively dry, are less so than the preceding 
habitats. They occupy an intermediate posi- 
tion on the moisture scale between sand pine 
scrub and mesophytic hammocks, which are 
considered to represent the climax com- 
munity in northern Florida (Laessle, 1942). 


The environment of pine flatwoods is 
subject to rather drastic fluctuations; there- 
fore this habitat type does not fit as neatly 
into a sequence based on a moisture gradient 
as do the preceding. During droughty peri- 
ods, flatwoods may be exceedingly dry, only 
to become quickly flooded after a_ brief 
period of rain. Under normal weather con- 
ditions, however, flatwoods are probably ap- 
preciably more mesic than either scrub or 
turkey oak, and possibly upland hammocks 
as well. 


Vol. 11 


IV. RESULTS 
A. Mites 


1. Species records and relationships to 
host—Seven species of mites in four fami- 
lies were recorded. These include Ewschon- 
gastia peromysct (Ewing), Gahrlepia ( Wal- 
chia) americana Ewing, and Trombicula 
crossleyt Loomis of the family Trombiculi- 
dae; Eulaelaps stabularis (C. L. Koch) and 
Haemogamasus liponyssoides Ewing of the 
family Haemogamasidae; Ornithonyssus ba- 
coti (Hirst) of the family Dermanyssidae; 
and Haemolaelaps glasgowi (Ewing) of the 
family Laelaptidae. 


Trombiculid mites were collected from 
within the external auditory meatus, on the 
margin of the pinnae, and from the under- 
side of the tail base, edge of the anus, and 
external genitalia. The ear canal was the 
most frequent site of attachment of chiggers, 
and ear-infesting chiggers were the most fre- 
quent ectoparasites recorded. The prevalence 
of chiggers in the ears of 610 specimens ex- 
amined was 69.3%, as compared to 4.3% 
for infestations on other parts of the body. 
Trombiculids occurred in the ears of 70.5% 
of 519 adults and in 62.6% of 91 young, 
whereas infestations on other parts of the 
body were observed in 3.8% of the adults 
and 6.6% of the young. In neither case 
were the age differences significant when 
tested by chi-square (P>.05). 


Samples of trombiculids for identification 
were collected from 22 mice selected at ran- 
dom from different habitats. In this series, 
T. crossleyi occurred on 10 hosts, E. pero- 
mysct on 7, Trombicula sp. on 6, and G. 
americana on 4. T. crossleyi was found in 
the ear canal of all specimens except 1, 
which had a single mite of this species be- 
neath the base of the tail. Clusters of nu- 
merous individuals ordinarily occurred in 
the ear canal. Trombicula sp. was taken once 
on the margin of the pinna and 5 times on 
the underside of the tail base or on the 
perineal areas. E. peromysci was recorded 
only from the ear canal, and G. americana 
only from the ventral tail base and perin- 
eum. These limited data suggest species dif- 
ferences in the selection of attachment sites. 


Trombicula sp. and G. americana occurred 
together on 2 of the 22 hosts from which 
chiggers were identified, Trombicula sp. 
and T. crossleyi on 1, and T. crossleyi and 
E. peromysci on 2. In view of the similarity 


No. 1 


of attachment site and the high frequency 
of single infestations of the last two species, 
the seemingly low number of joint occur- 
rences may indicate competition as the re- 
sult of similar niche requirements, as was 
suggested for other trombiculid species by 
Jameson and Brennan (1957). 


Five-hundred and ninety mice were sur- 
veyed for mites other than trombiculids, and 
25.9% of this sample was infested. The in- 
cidence of non-trombiculid mites on 497 
adults was 28.0%, as compared to 15.0% 
on 93 young. The difference is highly sig- 
nificant on the basis of the chi-square test 
(B01). 

Samples of body mites from 130 mice 
were identified. H. glasgow: was the com- 
monest species, occurring on 118 (90.8% ) 
of the specimens. E. stabularis was recorded 
from 10 (7.7%) of the mice, H. lzponys- 
soides from 6 (4.6%), and O. bacoti from 
2 (15%). The number of individuals of 
each species in the total sample indicates 
the same order of abundance as does preva- 
lence. Of the total of 389 randomly collected 
mites identified, 89.2% (299 adults, 148 
nymphs) were H, glasgowi; 6.2% (24 
adults), E. stabularis; 3.9% (3 adults, 12 
nymphs), H. liponyssoides; and 1.0% (4 
nymphs), O. bacoti. Mixed species infesta- 
tions occurred in only 8 of the 130 samples. 
E. stabularis, H. glasgowt, and H. liponys- 
soides occurred together in 1 sample; E. 
stabularis and H. glasgow2 in 3; and H. glas- 
gowr and H. ltponyssoides in 4. 

The adult sex ratio in H. glasgow, the 
only species represented by a sufficient num- 
ber of specimens to warrant this calculation, 
was markedly unequal. Only 15 (5.2%) of 
286 sexed specimens were males, as com- 
pared to 271 (94.8%) females. 

2. Effect of host density on prevalence— 
To analyze the effect of host density on the 


Layne: Parasites of Florida Mouse ii 


incidence of various groups of ectoparasites, 
three population levels were recognized. 
Catches of less than 5, 6-10, and 11 or more 
mice per 100 trapnights were designated as 
“low,” “moderate,” and “high” populations, 
respectively. The prevalence of parasitism 
at each density level was calculated by habi- 
tat and season, then comparisons of levels 
of prevalence in pairs of populations repre- 
senting different density levels, e.g., low- 
moderate, moderate-high, and low-high, 
were made. In those instances where a 
change of prevalence occurred with increase 
in population density, the average change in 
prevalence as well as the direction was de- 
termined. Because of seasonal effects on 
prevalence in a number of ectoparasites, 
actual comparisons of pairs of values were 
made within the same season. With the 
sizes of the samples involved, no difference 
could be seen in the direction or magnitudes 
of change associated with low-moderate, 
moderate-high, and low-high pairs, thus the 
data for these separate comparisons were 
combined into two density categories, “low- 
er” and “higher.” 


The relationships between host density 
and prevalence of mites are shown in Tables 
1 and 2. In the case of trombiculids, the dif- 
ferences in frequencies of the three possible 
categories of response of prevalence to 
higher host density fall far short of statistical 
significance when tested by chi-square 
(.80>P>.70). The differences between ob- 
served and expected frequencies in the three 
categories approach more closely to sig- 
nificance (.10>>P>.05) in the case of non- 
trombiculid mites. Considering only the 
cases of increase or decrease of prevalence 
corresponding to higher population density, 
ear-infesting trombiculids exhibit nearly 
twice as many increases than decreases, with 
a correspondingly greater change in preva- 


TABLE 1. 
Influence of increased host density on the prevalence of trombiculid mites 
attached within the ear canal 


Number of No 

Habitat Comparisons Change 
N 
Sand pine scrub 4 1 
Longleaf pine/turkey oak 5 0 
Upland hammock 7 4 
Pine flatwoods 1 ~ 
Combined Habitats 7 5 


Decrease Increase 
X change in X change in 
N prevalence N prevalence 
il 64.1 2, 62.5 
4 14.4 il 6.1 
0 = 3 46.4 
= = i 52.6 
5 24.3 Ti 46.1 


8 Tulane Studies in Zoology 


Vol. 11 


TABLE 2, 


Influence of increased host density on the prevalence of 
non-trombiculid mites 


Number of No 


Habitat Comparisons Change 
N 
Sand pine scrub 4 1 
Longleaf pine/turkey oak 5 0 
Upland hammock 5 0 
Pine flatwoods i 0 
Combined Habitats 15 1 


Decrease Increase 
X change in xX change in 
N prevalence N prevalence 
0 - 3 16.0 
2 20.1 3 23.7 
3 29.4 2 37.6 
1 25.8 0 - 
6 25.7 8 24.3 


lence in the increase than in the decrease 
category. A similar but less pronounced 
trend is suggested by the data for non- 
trombiculid mites. In neither group of para- 
sites, however, is the evidence for a correla- 
tion between host and parasite abundance 
particularly convincing. 


3. Effect of habitat on prevalence—None 
of the trombiculid mite species was recorded 
from all of the habitats studied. T, crossleyi 
occurred in sand pine scrub, longleaf pine/ 
turkey oak, and upland hammock; Trombz- 
cula sp. in longleaf pine/turkey oak and 
upland hammock; G. americana in scrub 
and longleaf pine/turkey oak; and E. pero- 
mysct in scrub, upland hammock, and flat- 
woods. Since the sample of identified trom- 
biculids was small, there is some question 
as to how accurately the above data reflect 
the actual habitat tolerances of the several 
species represented. Most likely further 
sampling would have increased the known 
habitat distribution of at least some of the 
forms. 


Table 3 gives the seasonal and overall 
prevalence of trombiculid mites in the four 
major habitat types studied. The incidence 
of trombiculids in the ear canal, presum- 
ably mostly or entirely T. crossleyi and E. 
peromysct, varies markedly with habitat. 
The prevalence ranges from a low of 41.4% 
in scrub to a high of 83.0% in upland ham- 
mock, with intermediate values for turkey 
oak (70.6%) and flatwoods (65.5%). 
When an R x C test for independence 
(Snedecor, 1946) is applied to these data, 
with the calculation of expected values for 
individual habitats being based on the inci- 
dence in all habitats combined, the observed 
differences are highly significant (P<.01). 
The incidence of trombiculids occurring. on 
other parts of the body is low in all habitats, 


and the differences between habitats are not 
significant (.20>P>.10). 

The trend in prevalence of ear-infesting 
trombiculids in relation to habitat probably 
is primarily a reflection of the environ- 
mental requirements of the free-living 
nymphal and adult stages. An influence of 
habitat on the prevalence of chiggers has 
been noted by a number of workers (e.g. 
Ewing, 1944; Jameson and Brennan, 1957; 
Michener, 1946; Mohr, 1947; Pearse, 1929; 
Worth, 1950), and in many instances abun- 
dance of chiggers has been correlated with 
habitats having a well-developed litter layer 
or heavy ground vegetation and relatively 
moist conditions. Jenkins (1947) found 
that Ewtrombtcula batatas laboratory colonies 
were most successful when maintained on a 
nearly saturated medium at temperatures 
ranging from 25 to 35° C. At relative 
humidities above 85%, optimum egg-laying 
took place between 27 and 34°, a decline 
occurring when temperature exceeded 35°. 
Apparently, therefore, temperature, humid- 
ity, and the development of litter or ground 
cover are among the critical factors deter- 
mining the suitability of a given habitat for 
chiggers, a combination of relatively moist 
conditions, limited temperature fluctuation, 
and abundance of litter or ground cover 
tending to favor high populations. 


Upland hammocks and pine flatwoods ap- 
proach these conditions more closely than 
the other habitats represented in this study. 
Sand pine scrub apparently provides the 
poorest habitat for trombiculids, while typi- 
cal longleaf pine/turkey oak woodlands 
rank only slightly better. The lower preva- 
lence of ear trombiculids in these habitats 
as compared to upland hammock bears out 
these assumptions. On the basis of general 
environmental conditions, flatwoods would 


Layne: Parasttes of Florida Mouse 


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10 Tulane Studies in Zoology 


Vol. 11 


TABLE 4. 
Habitat and seasonal prevalence of non-trombiculid mites 


Longleaf pine/ 


Period Sand pine scrub turkey oak Upland hammock Pine flatwoods 
No. Percent No. Percent No. Percent No. Percent 
exam. infested exam. infested exam. infested exam. infested 
Jan.-Mar. = = 61 26.2 40 BieD 28 s2nill 
Apr.-June 81 19.8 20 40.0 54 44.4 31 25.8 
July-Sent. 29 13.8 61 14.8 ie 16.7 — — 
Oct.-Dec. 1 0.0 50 24.0 62 19.4 = = 
Total maki 18.0 192 23.4 228 Sih sil 59 28.8 


be expected to have a relatively higher prev- 
alence than that observed. The apparently 
low value for this habitat may be explainable, 
at least in part, on the basis of a sample 
bias. Data for other habitats indicate a 
strong interaction between habitat and sea- 
son, with a marked increase in the abun- 
dance of chiggers in the period from July 
through December. This interval is not rep- 
resented in the sample from _ flatwoods. 
Thus, if the same seasonal trend in chigger 
prevalence occurs in this habitat as in others, 
the overall prevalence should equal or even 
exceed that for upland hammock. The alter- 
nation between wet and dry conditions may 
also influence trombiculid populations in 
flatwoods. 

Infestation rates of non-trombiculid mites 
are slightly higher in the moister habitats 
(Table 4). However, the observed fre- 
quencies in different habitats do not differ 
significantly from expected values based on 
the overall prevalence (.02>P>.05). The 
occurrence and relative abundance of the in- 
dividual non-trombiculid species identified 
from 130 mice taken in different habitats 
(Table 5) indicate that H. glasgowz is the 
most common species on P. floridanus in 
every habitat. It does not exhibit any pro- 
nounced habitat specificity, although Worth 
(1950) reported this species to be more 
prevalent in moist habitats in south Florida. 
H. liponyssoides is the only other species 
occurring in all habitat types, appearing to 


be most abundant in flatwoods. E. stabularis 
seems to be more abundant in the moister 
of the two environments, longleaf pine/ 
turkey oak and upland hammock, from 
which it was recorded. 

The apparent low degree of correlation 
between habitat type and prevalence of non- 
trombiculid mites as compared to chiggers 
may be a reflection of the greater influence 
of physical environmental factors on the 
latter as a result of their free-living nymphal 
and adult stages. Since non-trombiculid spe- 
cies are parasitic in both nymphal and adult 
stages and require the host’s nest for their 
life cycle, they may be more independent 
of the direct effects of environment than 
chiggers. Thus, over the environmental 
range encountered in this study, their popu- 
lations may depend to a greater extent on 
the kinds and abundance of mammalian 
hosts in a given habitat. 

4. Effect of season on prevalence—When 
the data on prevalence of trombiculid mites 
in all habitats are combined into 3-month 
periods (January-March, April-June, July- 
September, October-December ) , pronounced 
seasonal trends in abundance are indicated. 
The prevalence of trombiculids in ears is 
low during the first six months of the year 
(43.7% for the January-March period and 
41.9% for the April-June interval) and high 
in the last half of the year (97.3% in the 
July-September period and 90.4% in the 
October-December quarter). The differences 


TABLE 5. 


Percentages of mice carrying different non-trombiculid mites in 
four habitats 


Species of Mite 
Eulaelaps stabularis 


Haemogamasus liponyssoides 5.6 
Ornithonyssus bacoti = 
Haemolaelaps glasgowt 94.6 


Sand pine scrub turkey oak 


Longleaf pine/ Upland Pine 
hammock flatwoods 
5.6 11.3 ~ 
2.8 3.2 21.4 
5.6 - _ 
86.1 90.3 100.0 


No. 1 


in prevalence in the quarterly periods are 
highly significant (P<.01) when tested for 
independence against expected values based 
on the overall incidence. Monthly infesta- 
tion rates for all habitats give a more precise 
indication of seasonal trends in abundance 
of ear chiggers. The prevalence exceeded 
90% from June through November, de- 
clined in December (79.6%) and January 
(82.1%), reached its lowest levels in Feb- 
ruary (4.5%), March (18.2%) and April 
(4.6%), then increased again in May 
(37.5% ). These data indicate a yearly cycle 
in prevalence, with peak abundance occur- 
ring during the summer and fall months. 


In the case of individual habitats, com- 
plete seasonal data on prevalence is avail- 
able for longleaf pine/turkey oak and up- 
land hammock, and ia both habitats the sea- 
sonal trends are similar to those of the com- 
bined data. Although certain periods are 
not represented in the data for sand pine 
scrub and flatwoods, the trends suggested by 
the partial information also indicate agree- 
ment with the overall pattern. However, 
while the four habitats appear to exhibit the 
same seasonal patterns in prevalence, infesta- 
tion rates in individual habitats during com- 
parable seasons show considerable variation. 
Thus, upland hammock exhibits a higher 
incidence in the January-March period than 
either turkey oak or flatwoods (data not 
available from scrub for this interval ), while 
flatwoods has a much higher prevalence in 
the April-June period than any of the other 
habitats. These differences probably repre- 
sent the interaction between habitat and sea- 
sonal factors. For example, the generally 
higher prevalence in upland hammock and 
flatwoods during the early part of the year 
as compared to scrub and turkey oak may 
indicate a higher survival of tromb‘culids in 
these habitats during the winter as a result 
of more favorable cover, moisture, and tem- 
perature conditions. 

Although the differences are less pro- 
nounced than those of ear-infesting trombi- 
culids, the prevalence of chiggers on other 
parts of the body exhibits seasonal variation. 
The differences are highly significant 
(P<.01). The prevalence in the January- 
March and October-December periods was 
0.8%, as compared to 4.5% in the April-June 
interval and 8.6% in the July-September 
quarter. The general trend is similar to that 


Layne: Parasites of Florida Mouse ia 


of ear trombiculids except that the peak in 
abundance may occur somewhat earlier. 

The seasonal changes in abundance of 
chiggers probably reflect a response to cli- 
matic conditions, particularly temperature 
and humidity. The period of greatest abun- 
dance occurs when temperatures are high 
and rainfall is abundant. Strong seasonal 
trends in abundance of various species of 
trombiculids on mammalian hosts have been 
reported by other workers (e.g., Elton e¢ al., 
1931; Jameson and Brennan, 1957; Mich- 
ener, 1946; Worth, 1950), with peak abun- 
dance often coinciding with increased moist- 
ness of the habitat. 

Seasonal differences in the prevalence of 
non-trombiculid mites are also pronounced. 
The departure of the observed frequencies 
for quarterly periods from expected values 
calculated from overall incidence is highly 
significant (P<.01). In contrast to chig- 
gers, however, the increased prevalence of 
other mites occurred during the January- 
June period. The months of greatest abun- 
dance, 53.3% and 58.9%, respectively, were 
March and April. Insofar as data are avail- 
ab'e, these trends for individual habitats 
agree generally with that for all habitats 
combined. As in the case of trombiculids, 
d'fferences in prevalence between habitats 
in comparable seasons probably reflect the 
interaction between habitat and season. 

The observed trends in seasonal abun- 
dance of non-trombiculid mites are chiefly 
due to Haemolaelaps glasgowz, which is the 
most abundant species in the sample. The 
increased prevalence of this species during 
late winter and early spring agrees generally 
with the findings of Morlan (1952) and 
Smith and Love (1958). The latter also 
found H. glasgowz to be more abundant in 
wet years. 

The increased occurrence of non-trombi- 
culid mites on hosts in the cooler months 
may reflect a peak in an annual population 
cycle. This trend may also be the result, at 
least in part, of a greater tendency of mites 
to remain on the host rather than in the nest 
in cool weather. Paralleling the situation 
in the case of habitat, the seasonal changes 
in apparent abundance of non-trombiculids 
are of lesser magnitude than those of chig- 
ger mites. Again, this may indicate that the 
former, being more closely associated with 
the nest, are less subject to outside environ- 
mental conditions. A similar relationship 


12 Tulane Studies in Zoology Vola 
TABLE 6. 
Influence of increased host density on tick prevalence 
Number of No 
Habitat Comparisons Change Decrease Increase 
x change in X change in 
N N prevalence N prevalence 
Sand pine scrub 4 0 3 10.6 1 5.3 
Longleaf pine/turkey oak 5 D, 0 ~ 3 8.1 
Upland hammock 9 1 5 10.9 3 10.8 
Pine flatwoods 1 0 1 itilal 0 — 
Combined Habitats 19 3 9 10.8 7 8.9 


in the seasonal prevalence of trombiculids 
and other mites on small mammals in Cali- 
fornia was observed by Jameson and Bren- 
nan (1957). 
Be faces 

1. Species records and relationships to 
host—Four species of ticks collected on P. 
floridanus included Dermacentor variabilis 
(Say), Amblyomma maculatum Koch; Am- 
blyomma americanum (Linn.), and Ixodes 
minor Neumann (=I. bishoppi Smith and 
Gouck ).* The prevalence of ticks on a total 
of 600 mice examined for these parasites 
was 7.8%. The prevalence on adults (7.2%) 
was not significantly different (.50>P>.30) 
from that on young (11.2%). D. vartabilis, 
the only tick commonly represented, oc- 
curred on 29 of 33 hosts from which ticks 
were identified. All were nymphs or larvae. 
A, americanum (2 larvae) was collected on 
only two mice, and A. maculatum (1 nymph) 
and I. mimor (1 adult), were each taken on 
a single host. The average number of ticks 
on infested hosts was 1.3. The number of 
D. vartabilis larvae or nymphs on mice 
ranged from 1 to 8, with a mean of 1.6. 


*Smith and Gouck (1947) described Jao- 
des bishoppi from Georgia. However, ac- 
cording to G. M. Kohls (personal communi- 
cation) J. bishoppi is probably a synonym 
of /. minor earlier described from Guate- 
mala, and on this basis the latter name is 
employed in this paper. 


In a series of 39 infested mice, ticks were 
attached to the pinna of 31 (79.5%), to 
the shoulder of 6 (15.4%), and within the 
ear canal of 3 (7.7%). The one specimen 
of I. minor was attached to the shoulder. 

2. Effect of host density on prevalence — 
The frequencies of no change, decrease in 
prevalence, or increase in prevalence of 
ticks in comparisons between lower and 
higher host population levels (Table 6) do 
not differ significantly from those expected 
on a random basis (.20>P>.10). In con- 
trast to mites there is slight indication of a 
negative correlation between tick prevalence 
and host density. 

3. Effect of habitat on prevalence—D. 
variabilis was recorded in each of the major 
habitats studied, whereas the other species 
were each taken in one habitat type: J. 
minor in sctub, A. maculatum in upland 
hammock; and A. americanum in longleaf 
pine/turkey oak. Because of the few records 
involved, the extent to which the data reflect 
actual habitat specificity of these ticks is 
questionable. 

Table 7 presents the habitat and seasonal 
distribution of 47 tick infestations on 600 
mice. The differences between observed and 
expected habitat-specific frequencies is high- 
ly significant (P.<.01), with the greatest 
number of infestations occurring in upland 
hammock. The increased prevalence of ticks 
in upland hammock habitats probably is due 


TABLE 7. 
Habitat and seasonal prevalence of ticks 


Longleaf pine/ 


Period Sand pine scrub turkey oak Upland hammock Pine flatwoods 
No. Percent No. Percent No. Percent No. Percent 
exam. infested exam. infested exam. infested exam. infested 
Jan.-Mar — - 58 6.9 40 15.0 28 3.6 
Apr.-June 81 2.5 16 0.0 54 16.7 31 0.0 
July-Sept 30 10.0 68 1.5 76 6.6 - = 
Oct.-Dec. 1 100.0 52 5.8 65 16.9 - - 
Total ial 5.4 194 4.1 235 13.6 59 Uee/ 


No. 1 


mainly to the existence of favorable environ- 
mental conditions. The relatively high hu- 
midity of this habitat type, absence of ex- 
tremely high temperatures, and abundance 
of brush, debris, and litter probably offer 
more suitable conditions for tick survival 
and reproduction than the other vegetative 
associations represented. Since the adult 
stages of the ticks recorded, except possibly 
I. minor, are found typically on larger mam- 
mals, the population levels of raccoons, opos- 
sums, bobcats, and foxes presumably would 
interact with habitat factors in influencing 
tick abundance. Evidence indicates that 
larger mammals were not abundant in the 
flatwoods habitats represented in the present 
study, and this might be at least partly re- 
sponsible for the lower prevalence of ticks 
in this environment than would be predicted 
on the basis of vegetative characteristics 
alone. 


4. Effect of season on prevalence—rThe 
prevalence of ticks in 3-month intervals 
through the year was as follows: January- 
March, 8.7%; April-June, 6.4%; July- 
September, 5.2%; and October-December, 
12.7%. Although the observed seasonal fre- 
quencies do not differ significantly (.10> 
P>.05) from expected values, the limited 
data suggest, however, that tick abundance 
may be highest in the fall and early winter 
months. 


C. Lice 


One specimen of Hoplopleura hirsuta Fer- 
ris was collected in a sample of 574 mice 
examined, a prevalence of 0.2%. The louse 
was taken on a Florida mouse collected in 
pine flatwoods in March. The cotton rat, 
Sigmodon hispidus, was abundant at this 
locality, and in all probability the louse was 
a straggler from this host. 


D. Fleas 


1. Species records and relationships to 
host—Four species of fleas recorded in- 
cluded Polygenis floridanus Johnson and 
Layne, Polygenis gwynt (Fox), Ctenophthal- 
mus pseudagyrtes Baker, and Hoplopsyllus 
affinis (Baker). 

One hundred and eighty (33.2%) of the 
542 mice examined carried fleas. The preva- 
lence of fleas on adults was 33.6% as com- 
pared to 30.8% on young, the difference 
not being statistically significant (.90> 
P>.80). The numbers of adult males and 


Layne: Parasttes of Florida Mouse 13 


females infested were practically equal 
(males, 33.3%; females, 34.0%), but a 
greater number of young males (36.8% ) 
carried fleas than females (25.0%). The 
number of fleas on infested mice ranged 
from 1 to 16, averaging 2.3. The mean was 
somewhat lower in adults (2.2) than in 
young (3.0), but in neither age class did 
the flea index of the sexes differ appreciably 
(adult males, 2.3; adult females, 2.1; young 
males, 3.0; young females, 2.9). 

Three hundred and sixty-five fleas were 
identified, this number representing all of 
the fleas carried by 158 of the 180 animals 
infested. Polygenis floridanus occurred on 
94.9% (150) of the mice and comprised 
92.3% (337) of the total identified sample. 
It was the only common flea on Peromyscus 
floridanus, being known thus far only from 
this host (Johnson and Layne, 1961). Poly- 
gents gwynt, perhaps the most abundant flea 
on other small terrestrial rodents in Florida, 
occurred on only 10.1% (16) of the mice 
and constituted only 5.2% (19) of the 
sample. C. pseudagyrtes was taken on 5.1% 
(8) of the specimens and represented only 
2.2% of the fleas determined. One H. af- 
fénis was collected. Since the typical hosts of 
this flea are hares and rabbits (Kohls, 1940), 
its presence on the Florida mouse is con- 
sidered accidental. 

The majority (89.8%) of infestations 
were of single species, and in no instance 
was more than two species of fleas found 
on the same host. Polygenis floridanus was 
the only flea recorded in 85.4% of the in- 
festations, P. gwyni in 3.8%, and C. pseuda- 
gyrtes in 0.6%. P. floridanus and gwynt oc- 
curred together on 5.7% of the hosts, P. 
floridanus and C. psendagyrtes on 3.8%, and 
P. gwyni and C. pseudagyrtes on 0.6%. The 
mean numbers of specimens per infested 
host of each species were: P. floridanus, 
2.2; P. gwynt, 1.2; and C. pseudagyrtes, 1.0. 

2. Effect of host density on prevalence — 
The prevalence of fleas apparently is more 
closely related to host numbers than in the 
case of acarine parasites (Table 9); the ef- 
fect of change in host density on the flea 
index is more pronounced than would be 
expected on the basis of chance (P<.01). 
In 22 pairs of values compared, an increase 
was recorded in 15 (68.2%) and a decrease 
in 6 (27.3%). The mean percentage change 
in prevalence in going from a lower to 
higher population was approximately equal 


14 Tulane Studies in Zoology Vol. a0 
TABLE 8. 
Influence of increased host density on flea prevalence 
Number of No 
Habitat Comparisons Change Decrease Increase 
X change in X change in 
N N prevalence N prevalence 
Sand pine scrub 4 0 0 — 4 38.8 
Longleaf pine/turkey oak 5 0 3 20.7 2 31.9 
Upland hammock iby 1 2 Syile!55 9 26.1 
Pine flatwoods 1 0 1 162 0 - 
Combined Habitats 22 1 6 27.6 15s 30.3 


for both increases and decreases (Table 8) 
in infestation rates. 

3. Effect of habitat on prevalence—tThe 
differences in prevalence in different habi- 
tats are highly significant (P<.01). In- 
festations were more frequent than expected 
in sand pine scrub and longleaf pine/turkey 
oak and less so in upland hammock and pine 
flatwoods habitats (Table 9). 

C. pseudagyrtes was collected in upland 
hammock and flatwoods only, its prevalence 
being 3.1 and 1.7%, respectively in the two 
habitats. H. affimis occurred in slash pine/ 
turkey oak, from which numerous Polygenis 
floridanus were also obtained. P. floridanus 
and gwyni were each taken in the four 
major habitats studied, but the relative abun- 
dance of these species varied from one habi- 
tat to another. P. gwyni occurred most fre- 
quently in sand pine scrub (7.4%) and 
flatwoods (10.3%) and had a low preva- 
lence in longleaf pine/turkey oak (1.3%) 
and upland hammock (0.9%). P. flort- 
danus, on the other hand, was most abundant 
in sand pine scrub (34.6%) and longleaf 
pine-turkey oak (35.5%), and least so in 
upland hammock (23.8%) and flatwoods 
(27.6%). 

A relationship between the host’s environ- 
ment and the occurrence and abundance of 
particular flea species has been shown by a 
number of workers, including Gabbutt 
(1961), Jameson (1947), Jameson and 


Brennan (1957), Pearse (1929), and Worth 
(1950). The habitat distribution of two of 
the flea species associated with the Florida 
mouse may be related mainly to the eco- 
logical distribution of their typical small 
mammal hosts, while that of the third may 
be due in greater measure to the direct ef- 
fect of environmental factors. C. pseuda- 
gyrtes commonly occurs on moles and 
shrews, and the greater abundance of these 
mammals in upland hammocks and _ flat- 
woods than in either scrub or turkey oak 
probably accounts for the appearance of this 
flea on the Florida mouse only when it oc- 
curs in the first two habitats. Polygenis 
gwyni is a characteristic flea of the cotton 
rat. In this study cotton rats occurred com- 
monly only in flatwoods and in certain scrub 
habitats, and it is in these vegetation types 
that P. gwynz is encountered most frequently 
on the Florida mouse. Within scrub habi- 
tats, the prevalence of Polygenis gwyni on 
the Florida mouse showed a close positive 
correlation with the size of the cotton rat 
population. 

The greater abundance of Polygenzis flori- 
danus in scrub and turkey oak habitats than 
in upland hammock and flatwoods may be 
indicative of preference for more xeric con- 
ditions, since host predilection can be ruled 
out. The environmental distribution of flea 
and host show a close correspondence. Pres- 
ent data indicate that sand pine scrub and 


TABLE 9. 
Habitat and seasonal prevalence of fleas 


Longleaf pine/ 


Period Sand pine scrub turkey oak Upland hammock Pine flatwoods 
No. Percent No. Percent No. Percent No. Percent 
exam. infested exam. infested exam. infested exam. infested 
Jan.-Mar. - ~_ 50 52.0 39 28.2 28 21.4 
Apr.-June 50 40.0 14 92.8 68 30.9 Sill 41.9 
July-Sept. 32 43.8 61 21.3 56 1225 - - 
Oct.-Dec. 1 0.0 52 38.5 60 26.7 — - 
Total 83 41.0 eT 40.7 223 24.7 59 Bye 


No. 1 


longleaf pine/turkey oak are the primary 
habitat types of the Florida mouse. Upland 
hammock is less important, and the species 
occurs only sporadically in flatwoods, and 
then only under particular conditions. The 
low prevalence of Polygents floridanus in 
flatwoods might be the result of general 
environmental conditions or specific micro- 
climates of the nests of the host in this habi- 
tat. Also, possibly the initial flea population 
on mice dispersing from more favorable 
habitats may have been low. 

4. Effect of season on prevalence——Fleas 
were most abundant (41.0%) in the period 
from April to June, while the prevalence 
dropped to 22.8% in the July-September 
interval. In the following 3-month period 
infestations increased to 31.8% and_ rose 
still higher (36.8%) in the January-March 
interval. These values depart significantly 
(P<.01) from the assumption of equal 
prevalence in all seasons. Trends of seasonal 
abundance in upland hammock and longleaf 
pine/turkey oak agree with that of the com- 
bined data. Although data are available from 
only the January-June period in flatwoods, 
the trend in prevalence of fleas during this 
interval follows the general pattern also. In 
sand pine scrub, however, a higher preva- 
lence is indicated for the July-September 
period relative to the April-June interval 
than in other habitats. 

The one H. affenzs was collected in April. 
C. pseudagyrtes was obtained in upland ham- 
mock habitats in March, April, and June; 
and in flatwoods in May. P. gwyni occurred 
on mice only during the first six months of 
the year. Its prevalence in the January- 
March interval was 3.1% and in the April- 
June period, 7.9%. The data for this species 
suggest, therefore, an increased abundance 
on hosts during the late winter and spring. 
Similar findings were reported for this spe- 
cies on cotton rats in Georgia by Morlan 
(1952) and Smith and Love (1958). 

The overall seasonal trend in flea abun- 
dance can be attributed largely to P. flort- 
danus. The prevalence of this species in the 
January-March period was 31.6%, in the 
April-June interval, 34.8%, in the July- 
September period, 22.0%, and in the Oc- 
tober-December period, 28.4%. The mean 
number of fleas per infested host exhibited 
a similar trend, being 2.9, 2.5, 1.5, and 2.4 
for the four periods. Copulating pairs of 
fleas were observed on hosts during live- 


Layne: Parasites of Florida Mouse 15 


trapping operations in March. Where com- 
parable periods are represented, the data for 
individual habitats show about the same 
seasonal patterns. Sand pine scrub, however, 
appears to have peak flea abundance at a 
somewhat later time of the year than other 
habitats. The data indicate that the abun- 
dance of fleas on hosts is highest during the 
cooler months of the year, reaching a peak 
in the spring. The extent to which the trend 
involves actual changes in flea populations 
or seasonal differences in the activity or 
behavior of the fleas or hosts is unknown. 

Total and seasonal sex ratios were calcu- 
lated for Polygenis floridanus. In a sample 
of 378 specimens collected in all months of 
the year 48.9% were males, 51.1% females. 
The highest proportion of males (57.3% ) 
occurred in the January-March sample, fol- 
lowed by the April-June period (48.6% ). 
Males constituted 47.8% of the fleas col- 
lected in the July-September interval and 
only 42.1% of those from the October to 
December period. Although the sex ratios 
calculated for 3-month periods do not differ 
significantly (.3>P>.2) from equality, a 
trend toward a higher number of females in 
late fall and early winter is suggested. This 
may indicate a sex difference in tempera- 
ture response similar to that shown for 
Xenopsylla cheopis by Cole (1945). The 
correlation of high flea prevalence and in- 
creased proportion of males in the spring 
with observed copulation suggests the pos- 
sibility that changes in sex ratio or overall 
abundance of fleas on hosts also may be as- 
sociated with reproductive activity. 


E. Botflies 


1. Species record and relationships to 
host—The species of botfly, Cuterebra, in- 
festing the Florida mouse is questionable. 
An unsuccessful attempt was made to rear 
specimens obtained from this host. How- 
ever, adult flies were reared from larvae 
taken from cotton mice (Peromyscus gos- 
sypinus) trapped in the same habitats as 
Peromyscus floridanus. According to Dr. 
C. W. Sabrosky these forms, presumably the 
same parasitizing the Florida mouse, closely 
resemble C. angustifrons Dalmat, although 
differing slightly from the latter in some 
particulars. 

The overall prevalence of Cuterebra larvae 
in 630 mice was 3.6% (23 specimens in- 
fected). The difference in infection be- 


16 Tulane Studies in Zoology 


tween adults (3.5%) and young (4.4%) 
was not significant (.98>P>.95). The lo- 
cation of the bots was recorded for 20 mice. 
The larvae were located in the inguinal 
region of 18 specimens, in the lumbar re- 
gion of 1, and in the sacral region of 1. 
Ten of 15 mice carrying living larvae had 
a single bot. Two larvae were present in 
each of 3 mice, and 2 animals each contained 
3 larvae. 

2. Effect of host density on prevalence — 
Because of the low prevalence rate in other 
habitats, host abundance and prevalence of 
cuterebrids can be considered for upland 
hammock habitats only. In three pairs of 
values, the prevalence was decreased in two 
and increased in one case with higher popu- 
lation density. The small sample of course 
precludes drawing any definite conclusions 
about the relationship between prevalence 
of cuterebrids and host density, although, as 
suggested by the data for ticks, an increase 
of hosts might be associated under some cir- 
cumstances with a reduction in the fre- 
quency of botfly infections. Wilson (1945) 
observed an increase in the occurrence of 
cuterebrids correlated with a low population 
of Peromyscus leucopus, and a similar trend 
is indicated by the data of Scott and Snead 
(1942) and Wecker (1962). 

3. Effect of habitat on prevalence—Habi- 
tat differences in the prevalence of botfly 
infections (Table 10) were highly sig- 
nificant in terms of expected values based 
on prevalence in all habitats combined 
(P<.01). Upland hammock was the only 
habitat type in which botfly infection reached 
significant proportions. The prevalence of 
infections in longleaf pine/turkey oak and 
pine flatwoods was low, and no mice with 
cuterebrid larvae were recorded from sand 
pine scrub. The apparently strong restric- 


Vol. 11 


tion of infections to upland hammock as- 
sociations becomes even further pronounced 
when the data for longleaf pine/turkey oak 
areas are examined more closely. All of the 
records of botfly larvae in mice in this habi- 
tat type came from a single station in which 
a long term live trapping study is in prog- 
ress. The study area consists of about 26 
acres of longleaf pine/turkey oak habitat 
surrounded on two sides by mesophytic ham- 
mock, with a narrow ecotone between the 
two habitat types. The Florida mice in this 
area are confined largely to the turkey oak 
association, although some are trapped in 
the ecotone. None has ever been taken in 
the hammock. An examination of the trap- 
ping records of the infected individuals from 
this area showed that, although their home 
ranges lay largely in the turkey oak zone, 
occasionally they did range into the moister 
ecotone areas. No animals known to live 
entirely within the turkey oak were recorded 
with botfly infections. 

The restricted ecological distribution of 
cuterebrid infections probably can be at- 
tributed to the effect of physical environ- 
mental factors on the adult or larval stages 
of the parasite. The adult flies may prefer 
moist, shady situations and may occur less 
frequently in more open, drier environ- 
ments. Evidence on the life cycle of the 
forms infesting Peromyscus suggests that the 
flies lay their eggs around the homesites or 
in local areas frequented by the mice and 
that the eggs hatch into larval stages which 
are attracted by the proximity of the host 
(Dalmat, 1943; Penner and Pocius, 1956). 
A poorer survival of larval stages might 
also be a factor contributing to the low level 
of infections in the drier habitats. 

4. Effect of season on prevalence.——tLiv- 
ing bots or fresh exit sites were recorded 


TABLE 10. 
Habitat and seasonal prevalence of cuterebrid larvae 


Longleaf pine/ 


Period Sand pine serub turkey oak Upland hammock Pine flatwoods 
No. Percent No. Percent No. , Percent No. Percent 
exam. infected exam. infected’ exam. infected exam. infected 
Jan.-Mar. = — 58 0.0 <«-j; 40 4.52 28 0.0 
Apr.-June Tfe4 0.0 23 O20 ie 2 DO 5.4b 31 3.2 
July-Sept. 46 0.0 70 0.0 et 2.6 - _ 
Oct.-Dec. i 0.0 63 6.3 65 16.9¢ — - 
Total 119 0.0 214 1.9 238 7.6 59 ey ( 


* one old sear, 1 dead. 
> one with old sear. 
“two with old sears. 


No. 1 


in June (1); July (3), October (4), and 
November (7). The differences in preva- 
lence in 3-monthly periods are significant 
(P<.01) when tested for independence. 
The seasonal distribution of infections thus 
indicate an increased prevalence in the fall 
months. Similar seasonal trends in preva- 
lence of cuterebrid larvae in Peromyscus 
have been reported for more northerly re- 
gions (Burt, 1940; Test and Test, 1943; 
Dalmat, 1943; Sealander, 1961; Wecker, 
1962). The infection rate in Florida mouse 
populations is lower generally than those 
reported for Peromyscus leucopus by several 
authors (Test and Test, 1943; Dalmat, 1943; 
Scott and Snead, 1942; Hirth, 1959; Sea- 
lander, 1961, Abbott and Parsons, 1961; 
Wecker, 1962), a possible reason being that 
P, leucopus occurs more regularly in habi- 
tats favorable to Cuterebra than does P. 
floridanus. 
F. Leptospires 

No evidence of leptospire infections was 
obtained from four mice collected at one 
sand pine scrub station. This sample is far 
too small to give any indication of the true 
status of leptospires in this host. 


G. Protozoa 


Blood smears of 10 mice and fecal sam- 
ples of 38 specimens were examined for 
parasites. All blood smears were negative. 
Endamoeba ? muris (Grassi), Trichomonas 
? muris (Grassi), and Gzardia ? muris 
(Grassi) were recorded in fecal prepara- 
tions. Endamoeba occurred in 2 (7.1%) 
of 28 mice from which smears were made, 
Trichomonas in 17 (60.7%), and Giardia 
in 12 (42.8%). One young mouse had 
relatively large numbers of Trichomonas and 
Giardia, and the former was abundant in one 
adult female. According to Dr. Elliott Lesser, 
who made the examinations and determina- 
tions, the remainder of the mice had rela- 
tively light infections of these protozoans, 
with the incidence of Endamoeba being par- 
ticularly low. 

Preserved feces from 8 of 10 mice from 
scrub habitats contained parasitic protozoans. 
Two specimens had only Trichomonas, 4 
only Eimeria sp. (oocysts), and 2 both. 


H. Helminths 


1. Trematodes—The ova of an unidenti- 
fied, apparently dicrocoeliid, fluke were re- 
covered from livers subjected to the forma- 


Layne: Parasttes of Florida Mouse 17 


lin-ether concentration technic. Although 
careful dissections of numbers of both fresh 
and preserved livers were made, no adult 
flukes were observed, or were recognizable 
fragments of adults found in the egg- 
containing sediments from treated livers. 

The prevalence of this parasite, as indi- 
cated by the occurrence of ova in livers, was 
7.0% in a sample of 723 mice from all habi- 
tats combined. Considering only popula- 
tions from which infected specimens were 
recorded, the prevalence of infection was 
15.0%. In these populations, 15.9% (50) 
of 315 adults were infected, as compared to 
3.8% (1) of 26 young. Although sugges- 
tive of the existence of an age difference in 
infection rare, the difference is not sta- 
tistically significant (.10>P>.05). 

The correlation between habitat type and 
infection rate is high (P<.01). Forty-five 
(88.2%) of 51 infected mice were from up- 
land hammock or pine flatwoods habitats 
(Table 11). The evidence for the restric- 
tion of infections to the moister of the hebi- 
tat types utilized by Florida mice is strength- 
ened by the fact that the one infected mouse 
recorded from sand pine scrub was collected 
in close proximity to a mesic hammock and 
that each of five infected animals from long- 
leaf pine/turkey oak were from a live-trap- 
ping study area and had capture records 
showing that they ranged into the moister 
ecotone area between the turkey oak and 
an adjoining mesic hammock. The close 
correlation between the occurrence of the 
parasite and the moistness of the habitat is 
probably related to the ecological distribu- 
tion of an intermediate molluscan host. 

Infections were recorded in all months 
with the exception of May, August, and Sep- 
tember. In those populations in which 11- 


TABLE 11. 
Prevalence of unidentified trematode in 
different habitats based on presence 
of eggs in liver 


Number Percent 

Habitat Examined Infected 
Sand pine scrub 120 0.88 
Slash pine/ 

turkey oak 45 0.0 
Longleaf pine/ 

turkey oak 246 2.0 
Upland hammock 256 15.6 
Pine flatwoods 56 8.9 

Total (es 7.0 


* captured near mesophytic hammock 


18 Tulane Studies in Zoology 


fections were present, the mean prevalence 
was 15.9% for the January-March period, 
30.9% for April-June, 18.0% for July- 
September, and 6.7 for October-December. 
The data are, therefore, suggestive of a 
tendency toward a higher frequency of in- 
fections in the warmer months of the year. 


2. Cestodes—Larval cestodes were rare, 
only 3 cases being recorded in over 700 
specimens examined from all habitats. One 
mouse from longleaf pine/turkey oak had 
cysticerci of either Cladotaenta or Paruterina 
in the liver. The former is a parasite of 
hawks in the adult stage and the latter, a 
parasite of owls. One mouse from. slash 
pine/turkey oak and another from sand pine 
scrub habitat about 14 mile away had Taenza 
lyncis Skinker cysticerci in the liver. Only 
one specimen was present in each case. 

The only adult cestode recorded was Hy- 
menolepis ? nana (Siebold). It occurred 
in 4.8% (9) of 186 mice examined from 
all habitats combined. Although the preva- 
lence of infections was higher in adults 
(5.1%) than young (3.6%), the differ- 
ence is not significant. The mean number 
of worms per infected host was 3.0, with a 
range of from 1 to 11. Hymenolepis was 
recorded in mice from all habitats, with the 
highest number of infections being encoun- 
tered in sand pine scrub and upland ham- 
mock environments (Table 12). A_ sea- 
sonal trend in infections is suggested by the 
data. The highest prevalence during the 
year (12.0%) occurred during the January- 
March period. From April to June, 3.6% 
of the mice were infected. No infections 
were recorded in the July-September inter- 
val, and the prevalence in the October- 
December period was only 2.0%. 

3. Nematodes.—Five species of nematodes 
were recorded. These included Capzllaria 
hepatica (Bancroft), Rictularia ?coloraden- 
sis (Hall), Aspiculuris americana Erickson, 


Volante 


Syphacia peromysci Harkema, and Tricho- 
strongylus ransom Dikmans. 

The prevalence of C. hepatica infection in 
the livers of 723 mice from all habitats was 
2.9% (Layne and Griffo, 1961). Consider- 
ing only those populations in which infec- 
tions were recorded, the prevalence was 
12.7%. Aspiculuris occurred in 32.2% (60) 
of 186 mice examined from the four major 
habitats. The mean number of worms in 
infected hosts was 4.2 with a range of from 
1 to 31. Rictularia occurred in 9.1% of the 
mice. The average number of worms in 
infected mice was 5.7, with extremes of 2 
and 14. Harkema (1936) observed a higher 
prevalence rate for this parasite in Pero- 
myscus leucopus in North Carolina, but usu- 
ally found only single worms, rarely two, in 
infected hosts. Syphacia was present in 2.2% 
of the animals, averaging 2.2 worms per 
host. Only one mouse had more than one 
worm (6). One infection of Trichostrongy- 
lus was recorded, a prevalence of 0.5%, and 
only one individual was collected. 


Seven of 84 mice infected with either 
Hymenolepis, Rictularia, Aspiculuris, Sypha- 
cia, or Trichostrongylus possessed more than 
one form. Three had both Hymenolepis and 
Aspiculuris, while single cases each were re- 
corded for the following combinations: Ric- 
tularia/ Aspiculuris, Syphacia/Trichostrongy- 
lus, and Aspiculuris/Syphactia. 

A general trend of higher infection rates 
in adults as compared to young was noted 
in all nematodes for which adequate sam- 
ples were available. C. hepatica occurred in 
15.5% of the adults in infected populations 
and in only 2.8% of the young (Layne and 
Griffo, 1961). The prevalence of Rictularia 
in adults was 10.1% as compared to 3.6% 
in young. Corresponding values for Aspz- 
culuris were 34.2 and 21.4%. These dif- 
ferences are statistically significant (P<.05) 
only in the case of C. hepatica. 


TABLE 12. 
Prevalence of five helminths in different habitat types 
Total No. Percent Hymeno- Rictu- Aspicu- Sypha-  Trichos- 
Habitat Examined Infected lepis laria luris cia trongylus 
Percent Percent Percent Percent Percent 
infected infected infected infected infected 
Sand pine scrub 45 48.9 8.9 35.6 Tals 0.0 0.0 
Longleaf pine/ 
turkey oak 59 42.4 We Hert 40.7 0.0 0.0 
Upland hammock 52 48.1 5.7 0.0 44.2 0.0 0.0 
Pine flatwoods 31 38.7 3.2 0.0 25.8 12.9 3.2 


No. 1 


C. hepatica exhibited a marked habitat 
restriction, occurring only in mice from sand 
pine scrub and slash pine/turkey oak habi- 
tats (Layne and Griffo, 1961). The com- 
bined prevalence of five helminths of the 
alimentary tract was highest in sand pine 
scrub and upland hammock, lowest in pine 
flatwoods, and intermediate in longleaf 
pine/turkey oak (Table 12). When tested 
for independence, however, these differences 
are not statistically significant (.80>P> 
70). Of the other four nematode species, 
Aspiculuris had the greatest habitat range, 
showing greater abundance in mice from 
longleaf pine/turkey oak and upland ham- 
mock and less in sand pine scrub and pine 
flatwoods. Rictularia was the most prevalent 
nematode in sand pine scrub, occurring out- 
side of this habitat only in longleaf pine/ 
turkey oak, where it was infrequent. Sypha- 
cia and Trichostrongylus were recorded only 
in pine flatwoods. 

Some suggestion of seasonal trends in 
prevalence is evident in the data for three 
of the nematodes. The prevalence of C. 
hepatica reached its highest level (9.8% ) 
in the April-June period, declining to 3.0 
and 2.4%, respectively, in the July-Septem- 
ber and October-December samples. No in- 
fections were recorded in the January-March 
interval. In contrast, Rictularia and Aspi- 
culuris were most prevalent in the January- 
March period. The former declined from 
14.0% to 10.9% in the April-June interval. 
The lowest infection rate occurred in the 
July-September sample (3.2%), that for the 
October-December period being somewhat 
higher (6.0%). Aspiculuris infections 
dropped from 42.0% in the January-March 
sample to 30.9% in the April-June period. 
The prevalence of this parasite was 38.7% 
in the July-September interval and 20.0% 
in the October-December period. 


I. Pentastomids 


Nymphal stages of the pentastomid Poro- 
cephalus crotali (Humboldt) were found in 
7 (0.8%) of 840 Florida mice examined. 
Apparently the occurrence of P. crotali 
nymphs in the Florida mouse constitutes the 
third record of this parasite in North Ameri- 
can mammals. Penn (1942) reported its 
presence in the muskrat (Ondatra ztbetht- 
cus) in Louisiana, and Self and McMurry 
(1948) recorded it from Peromyscus leu- 
copus in Oklahoma. 


Layne: Parasites of Florida Mouse 19 


All infections were in adult mice. In 
two specimens the parasites were partially 
embedded in the liver. One female had a 
nymph in the mesovarium and one in the 
bladder. A male had nymphs in the mesen- 
teries of the epididymides, and a female had 
them in the mesenteries of the abdominal 
viscera and attached to the lung and liver. 
In the above instances the number of nymphs 
present was small, ranging from one to five 
in the cases where counts were made. Two 
mice were heavily infected. A female had 
numerous nymphs located on the liver, in- 
testine, lungs, body wall, kidney, and bladder 
and in the mesenteries of the abdominal 
organs. A male had the nymphs beneath the 
tunica albuginea of the testes, around the 
cauda epididymis, in the trunk mesenteries, 
on the surface of the liver, and in the mesen- 
tery near the lung. None of the mice in- 
fected with Porocephalus exhibited any evi- 
dence of weakness or debilitation when alive 
nor any gross effects of the parasites when 
necropsied. Esslinger (1962) found that 
the tissue responses of the liver of the rat 
to immature stages of Porocephalus re- 
sembled those produced by other agents of 
visceral larva migrans and metazoan para- 
sites generally. 

Of the 840 mice examined for Poro- 
cephalus, 202 were from sand pine scrub 
habitats, 49 from slash pine/turkey oak, 
300 from longleaf pine/turkey oak, 230 
from upland hammock, and 60 from pine 
flatwoods. Infections occurred only at a 
single scrub station. Here, 103 adults and 
11 young mice were collected, giving a 
prevalence of Porocephalus in this popula- 
tion of 6.1% for both age groups combined 
or 6.8% for adults. The difference in in- 
fection rate of adult males and females was 
not significant on the basis of the small 
sample involved. 

The scrub station from which pentasto- 
mids were recorded was a slight ridge sur- 
rounded by low pine flatwoods, marshes, 
and cypress swamps. There was evidence 
of a high snake population at this locality, 
and such semi-aquatic forms as the cotton- 
mouth (Agkistrodon piscivorus) and water 
snake (Natrix fasciata) were recorded. Pos- 
sibly, therefore, the occurrence of Poro- 
cephalus at this station was in part due to 
an abundance of snakes. However, addi- 
tional factors probably were involved, since 
certain other scrub stations as well as those 


20 Tulane Studies in Zoology 


in other habitat types appeared to support 
high snake populations while Porocephalus 
was apparently absent. 

Twelve mice collected in February were 
free of infections. Three of 59 specimens 
taken in April (11), and May (48) were 
infected, as were 3 of 30 specimens from 
July (5) and August (25). Of 11 speci- 
mens examined in October (6) and No- 
vember (5), 1 had a Porocephalus infection. 
Thus, although the data are not conclusive, 
a slight trend toward a higher frequency of 
infections in late summer and fall is indi- 
cated. Whether this trend is related to the 
life cycle of the parasite or to seasonal dif- 
ferences in the movements or behavior pat- 
terns of mice or snakes is uncertain. 


V. DISCUSSION 


Thirty-two parasites, including 18 ecto- 
parasites and 14 endoparasites, were recorded 
from the Florida mouse. Although further 
collecting may yield additional forms, par- 
ticularly microorganisms and helminths, the 
present list probably is a relatively complete 
representation of the parasitic fauna of this 
rodent. Comparative data for other small 
mammals are few. However, Elton ef al. 
(1931) found 41 species of parasites asso- 
ciated with the wood mouse (Apodemus 
sylvaticus) in England. 

Based on combined data for all localities, 
habitats, and seasons, the overall level of 
parasitism on the Florida mouse is low. The 
prevalence of ectoparasites as a group con- 
siderably exceeds that of endoparasites. The 
difference is especially pronounced if Pro- 
tozoa are excluded from consideration. A 
number of the parasites recorded were rare 
in all habitats or occurred with any fre- 
quency only in particular habitat types. Of 
the seven categories of ectoparasites recog- 
nized for the purpose of analysis, only 
non-infesting trombiculids, non-trombiculid 
mites, ticks, and fleas occurred on more than 
5% of the mice in the total sample. In each 
of these groups, only one or two species 
dominated in the identified samples. These 
included the mites Trombicula crossleyi, 
Euschongastia peromysct, and Haemolaelaps 
glasgowt, the tick Dermacentor variabilis, 
and the flea Polygenis floridanus. Two of 
the ectoparasitic species collected, Hoplo- 
pleura hirsuta and Hoplopsyllus  affinis, 
were clearly accidental. In the case of endo- 
parasites, Endamoeba, Giardia, Trichomonas, 


Vol. 1 


Eimeria, an unidentified trematode, Hyme- 
nolepis ?nana, and Aspiculuris americana 
approached or exceeded the 5% prevalence 
level. Thus, even with the rather liberal 
criterion of commonness accepted here, only 
12 of the 32 parasites of the Florida mouse 
would be considered as generally common. 
Roughly the same proportion, 13 out of 41 
species, of the parasites of the wood mouse 
was considered to be of possible importance 
in the study by Elton et al. (1931). 

Parasitism apparently plays a relatively 
minor role in the population dynamics of 
the Florida mouse. There is no evidence to 
indicate that parasites Constitute an impor- 
tant source of direct mortality. In addition 
to the generally low prevalence rates, the 
numbers of ecto- or endoparasites carried 
by individual hosts were seldom high for 
the particular parasite involved. Nor was 
any animal encountered during the course 
of the study in a sick or weakened state that 
could be attributed to its parasite load, al- 
though the possibility that sick or weak mice 
may be less responsive to traps must be con- 
sidered. However, in some cases even rela- 
tively low parasite burdens probably may 
produce effects on the host’s physiology or 
behavior that render the animal more sus- 
ceptible to predation or various environ- 
mental stresses. At the general levels of 
abundance of parasites found in this study, 
endoparasites would probably be more im- 
portant in this regard than ectoparasites. 
Certain ectoparasites without an obvious di- 
rect effect on the host also may be involved 
in inter- and intra-specific transmission of 
bacterial or viral diseases that may produce 
debilitation or death, although no evidence 
for this is as yet available for Peromyscus 
floridanus. 

Although the overall parasite level of the 
Florida mouse is low, the actual combina- 
tions of parasitic species and prevalence 
rates in different host populations are far 
from uniform. Differences in patterns of 
parasitism may be related to such factors 
as age of host, host density, habitat, and sea- 
son and are of interest from the standpoint 
of their causation as well as their signifi- 
cance in terms of the biology of the host. 
Adult mice appear to carry more parasites 
than young. However, a marked difference 
in the effect of host age on prevalence exists 
between ecto- and endoparasites. The aver- 
age prevalence of ectoparasites was 24.4% 


No. 1 


on adult mice and 21.8% on young. For 
those endoparasites (unidentified trema- 
tode, Hymenolepis, Capillaria, Rictularia, 
Aspiculuris, and Porocephalus) for which 
adequate data for a comparison of age- 
specific infection rates were available, mean 
prevalence was 14.6% in adult and 5.9% in 
young mice. 

Several factors might contribute to the 
greater difference in age-specific prevalence 
of endoparasites compared to ectoparasites. 
One of these is the relative mobility of the 
infective stages of the two kinds of para- 
sites. Most of the ectoparasites have a rela- 
tively mobile, free-living infective stage, in 
contrast to that of endoparasites which is 
typically an ovum, cyst, or essentially seden- 
tary larval stage. A young mouse outside its 
burrow thus would have a higher probability 
of encountering an ectoparasitic organism 
than an endoparasitic one, consequently ac- 
quiring a population of the former more 
rapidly than the latter. 

The extent to which parasites are acquired 
in the nest might also have a bearing on 
the problem of the different age relation- 
ships in prevalence of ecto- and endopara- 
sites. Infestation in the natal nest might be 
expected to contribute significantly to a 
faster build up on the young mouse of those 
ectoparasites (non-trombiculid mites and 
fleas) which require the host’s nest for at 
least part of the life cycle. The present data, 
however, do not indicate any trend toward 
higher prevalence of these forms than of 
those ectoparasites in which infestations are 
probably ordinarily acquired outside the nest. 
This may be due to the fact that most of 
the young in the samples were of post- 
weaning age and might, therefore, have al- 
ready moved from the natal burrow and 
established their own nests, in which mite 
and flea populations had not yet had suf- 
ficient time to build up. Drummond (1957) 
showed that the nesting activity of white- 
footed mice (Peromyscus leucopus) was a 
particularly important influence on the 
fluctuations in numbers of species and indi- 
viduals of mites in nests. 

Circumstantial evidence suggests that nests 
may not be an important source of endo- 
parasitic infections in the Florida mouse. 
The low prevalence of endoparasites in 
young might in itself be cited in support of 
this. Some endoparasites also exhibit dis- 
tinct trends in prevalence associated with 


Layne: Parasites of Florida Mouse vail 


habitat type, which suggests that physical 
environmental factors outside the nest may 
be influencing the distribution of the para- 
sites in question. Furthermore, if fecal con- 
tamination is assumed to be a principal 
route of nest infections, then the fact that 
the Florida mouse appears to practice good 
nest sanitation would perhaps also serve to 
limit the incidence of nest-acquired infec- 
tions. No accumulations of feces were found 
in two nests excavated, and captive mice 
generally defecated in parts of the cages re- 
moved from the nest. Another factor that 
might contribute to a lower rate of endo- 
parasitic infections in young mice in the 
natal nest is the existence of some degree 
of immunity. 

A final possibility is that the difference 
in age-specific prevalence rates of the two 
major groups of parasites is actually partly 
or entirely due to the age classification used. 
In ectoparasites, excepting botflies, an in- 
festation presumably is recognizable as soon 
as it occurs, whereas there may be a variable 
period between the acquisition of the in- 
fective stage of an endoparasite and the 
time at which the infection would be appar- 
ent at necropsy. Since the average interval 
a mouse spends in the young age class is 
considerably shorter than that spent in the 
adult category, it follows that a significant 
proportion of infections acquired by young 
mice may not reach a stage at which they 
would be detected at routine autopsy until 
the animals have passed over into the adult 
age group. Therefore, the actual numbers of 
endoparasites carried by young mice may 
have been appreciably greater than observed. 
Unfortunately, the present data do not per- 
mit any conclusions as to the relative im- 
portance of the above-mentioned factors, 
either singly or in combination, in explain- 
ing the differences noted. Whatever the 
cause, the lighter parasite load, particularly 
of endoparasites, of young mice is of sig- 
nificance from the standpoint of population 
dynamics in that it can be assumed to con- 
tribute to better survival of this age class. 

Comparisons of prevalence rates of five 
groups of ectoparasites at different host 
population levels showed no overall effect 
of population density on the proportion of 
mice infested. In going from a lower.to a 
high population level, decreases in preva- 
lence averaged 42.2% and increases, 46.6% 
The number of instances in which no change 


22 Tulane Studies in Zoology 


occurred averaged 10.4%. The mean de- 
crease in prevalence was 22.1% and the 
average increase, 27.4%. Because of the 
small number of comparisons available, 
trends in individual habitats cannot be con- 
sidered conclusive. However, sand pine 
scrub differed from other habitat types in 
showing a stronger correlation between ecto- 
parasite prevalence and host abundance. 

The relationship between host population 
level and parasite prevalence varies between 
different types of ectoparasites. The data 
for fleas and mites indicate a positive corre- 
lation between host abundance and parasite 
levels, while an inverse relationship is sug- 
gested in the case of ticks and botflies. The 
prevalence of fleas shows the strongest asso- 
ciation with host numbers of any group of 
ectoparasites. This trend is largely due to 
the species Polygenis floridanus, which thus 
far is known only from the Florida mouse. 
The ultimate factor responsible for this nar- 
row host restriction may be microclimate 
of the nest of Peromyscus floridanus, al- 
though several proximate factors also may 
Operate to prevent host transfer (Johnson 
and Layne, 1961). 


In view of this host specificity, the num- 
bers of Polygenis floridanus would be ex- 
pected to conform more closely to the popu- 
lation level of the Florida mouse than those 
of other ectoparasites which have a broader 
host range. The relationship between preva- 
lence of mites and host population level was 
considerably less pronounced than in the 
case of Siphonaptera. The prevalence of non- 
trombiculid mites on a particular small mam- 
mal species appears to depend at least in part 
on the abundance of other possible hosts in 
the same habitat. The Florida mouse was 
the principal small mammal species at many 
of the stations sampled during this study, 
which might explain the apparent correla- 
tion between mouse and mite population 
trends. As chiggers are parasitic only in the 
larval stage and do not require the nest en- 
vironment to complete their life cycle, the 
correlation, though slight, between their 
prevalence and mouse density may be due to 
a certain degree of host specificity in the 
commoner species such as Trombicula cros- 
sleyi. The particular habits or relative abun- 
dance of the Florida mouse in the habitats 
studied may also be factors influencing the 


Vol. 11 


host density correlated abundance of chig- 
gers. 

Botflies and ticks have a low infestation 
rate on the Florida mouse, usually occurring 
as one or only a few individuals. They do 
not require the nest for their life cycle, al- 
though ticks may seek refuge in the nest 
(Drummond, 1957). The inverse relation- 
ship between the prevalence of these para- 
sites and mouse abundance suggested by the 
data might be taken to indicate an effect of 
the parasite on host population. An alter- 
native explanation which assumes a limited 
number of parasites so that an increase in 
host density results in reduced frequency of 
infestations seems more likely. Two factors 
might operate to prevent an increase of 
parasites, after an appropriate lag, with an 
expanding mouse population: (1) a limita- 
tion on the numbers of parasites in certain 
habitats by environmental factors (physical 
or biotic) independent of the number of 
mice present; (2) the existence of marked, 
short-term fluctuations in host populations 
that even in an environment with an other- 
wise high carrying capacity for the particular 
parasites would inhibit a build-up of para- 
site numbers. Elton et al. (1931) advanced 
a similar explanation for the inverse rela- 
tionship between wood mouse population 
density and certain endoparasites. 

Of the factors considered in this study, 
habitat is probably the most important in 
determining the patterns of parasitism in 
particular host populations. The average 
prevalence of ectoparasites was lowest in 
sand pine scrub (22.2% ), slightly higher in 
flatwoods (22.5%) and longleaf pine/turkey 
oak (24.4%), and highest in upland ham- 
mock (27.1%). The data on prevalence of 
ectoparasites in flatwoods are probably bi- 
ased by the fact that samples were available 
for this habitat type only in the early part 
of the year, while all seasons are represented 
in the remaining habitats. Since there is 
evidence that in such groups as trombiculid 
mites, ticks, and botflies greatest prevalence 
occurs in late summer, fall, and early winter, 
it is possible that the overall ectoparasite 
prevalence in flatwoods is actually higher 
than indicated by the available data. On the 
other hand, the marked fluctuations of soil 
moisture in this habitat may be detrimental 
to some types of parasites. 

The abundance of ectoparasites as a group 
apparently is correlated with the moistness 


No. 1 


of the habitat and development of ground 
cover and litter. This trend is particularly 
clear for ear-infesting chiggers and non- 
trombiculid mites and present, though less 
obvious, in ticks and botflies. Fleas are the 
only group of ectoparasites exhibiting a re- 
verse trend, being more abundant in sand 
pine scrub and longleaf pine/turkey oak 
habitats. This habitat distribution pattern 
primarily reflects that of Polygenis flori- 
danus. 

No obvious correlation exists between the 
prevalence of endoparasites as a group and 
common environmental factors such as 
moisture or ground cover. Sand pine scrub 
had the highest average infection rate 
(9.7% ), longleaf pine/turkey oak the low- 
est (5.8%), and upland hammock and flat- 
woods occupied intermediate positions with 
rates of 8.2 and 6.8%, respectively. Indi- 
vidual parasite species, however, did show 
distinct trends in abundance correlated with 
habitat type. This suggests that the various 
kinds of endoparasites are more variable in 
their environmental requirements than the 
ectoparasitic forms and may thus exhibit 
greater habitat restriction. Other evidence 
points to the existence of narrower habitat 
specificity of endoparasites as compared to 
ectoparasites. Only 4 (24%) of 17 species 
of ectoparasites were collected from a single 
habitat, whereas 6 (60%) of 10 macro- 
endoparasites are in this category. If the 
similarities (presence or absence of species) 
in total parasite faunas are analyzed, sand 
pine scrub agrees with other habitat types 
in an average of 54% of the parasites. 
Longleaf pine/turkey oak exhibits a 58% 
agreement, upland hammock 66%, and flat- 
woods, 57%. The differences, though slight, 
suggest that variation in species composition 
as well as in prevalence of large groups is 
associated with habitat differences. When 
the habitat similarities are determined sepa- 
rately for ecto- and endoparasites, a differ- 
ence in habitat tolerance is indicated for the 
two groups. The average similarity in spe- 
cies composition of ectoparasites between 
different habitat types is slightly higher 
(61%) and the range of values for indi- 
vidual habitats less variable (57-65%) 
than for endoparasites, for which corre- 
sponding figures are 57% and 47-67%, 
respectively. On the basis of its endopara- 
sitic fauna, scrub is again more distinctive 
than other habitats. Sand pine scrub is also 


Layne: Parasites of Florida Mouse 


i) 
1S) 


the only habitat type in which the preva- 
lence of parasites with an egg as the infec- 
tive stage greatly exceeds those with free- 
living larvae. This suggests that high soil 
temperatures and dry conditions may limit 
the success of parasites with free-living lar- 
val stages in the scrub habitat. 


The extent to which new species of ecto- 
and endoparasites are added as the habitat 
range is increased is shown in Figure 3. The 
data for this graph were obtained by system- 
atically varying the sequence of the four 
principal habitats involved in the study and 
determining the mean percentage of the total 
species of parasites added with the addition 
of each habitat. The results seem to indicate 
that for any single habitat a higher propor- 
tion of the ectoparasite fauna is represented 
than of endoparasites and as a consequence 
the rate at which new species are added per 
habitat is higher in the latter. This again 
supports the supposition that the ectopara- 
sites of the Florida mouse tend to show less 
habitat restriction than endoparasites. 


The above data demonstrate clearly that 
populations of the Florida mouse living in 
different habitats are subjected to different 
regimens of parasite pressure, and that cer- 
tain parasites of negligible significance over 


100 


90 


80 


70 


% OF TOTAL SPECIES 


60 


sok ———— ECTOPARASITES 
rik esa? Ae ie ENDOPARASITES 


| 2 3 4 
NUMBER OF HABITATS 


Figure 3. A comparison of the increase of 
ecto- and endoparasite species on the Flor- 
ida mouse with increase in habitat types. 


A 


the entire geographic and ecologic range of 
the species as a whole may be of some local 
importance. This provides an example of 
the variation in selective forces that may 
Operate on populations of a species in dif- 
ferent habitats and influence its ability to 
invade new environments. 

Both the overall prevalence and abundance 
of particular groups of ecto- and endopara- 
sites are influenced by season. The highest 
prevalence of ectoparasites as a group occurs 
in the late fall and early winter (27.8%) 
and declines to the lowest level in the April- 
June interval (20.9%). Individual groups 
show varying degrees of departure from this 
general pattern. The peak prevalence cf 
endoparasite infections falls in the January- 
March interval (14.0% ) and exhibits a gen- 
eral decline through the year, reaching the 
lowest point in the October-December peri- 
od (7.7%). The causes of these trends may 
be different in the two groups of parasites. 
The general pattern in ectoparasites is prob- 
ably attributable mainly to the direct effects 
of weather on the life cycles of the parasites, 
particularly in the case of chiggers, ticks, and 
botflies. In other groups, such as non-trom- 
biculid mites and fleas, seasonal trends may 
be influenced by variation in the nesting 
habits or activity of the hosts. Two factors 
may be important in explaining the seasonal 
patterns of general endoparasite abundance. 
The peak levels occur in that part of the 
year when the food supply of the mice ap- 
pears to be most limited. The mice may 
forage more intensively and do more digging 
for food at this time and as a consequence 
have a higher probability of acquiring an 
infection than at other seasons. The other 
factor concerns a change in the age com- 
position of populations during the year. 
Much of the breeding of the Florida mouse 
is concentrated in the late fall and early 
winter. Thus when reproduction has been 
successful, young age groups predominate in 
the population. The observed differential in 
adult and young infections noted earlier 
could therefore account for at least a part 
of the fall and winter decline in infection 
rate. Elton et al. (1931) attributed seasonal 
changes in the frequency of the nematode 
Nematospiroideas dubius in the wood mouse 
to changes in host age composition. Addi- 
tional factors may be involved in the trends 
shown by particular parasites, for example, 
the unidentified trematode and Porocephal- 


24 Tulane Studies in Zoology 


Vol. 11 


ws, in which seasonal changes in the habits 
or activity of intermediate or definitive hosts 
might in turn influence the prevalence of 
infections in mice. 


The influence of seasonal changes in para- 
site prevalence on the host probably is 
greater in the case of endoparasites than 
ectoparasites. As noted previously, at the 
general levels of abundance shown by the 
two groups, endoparasites probably would 
be expected to have more effect on the 
health of the host than ectoparasites, assum- 
ing that none of the ectoparasites recorded 
transmits some as yet unknown viral or 
bacterial disease that causes significant mor- 
tality. In addition, the annual peak in endo- 
parasite burden coincides with the time of 
year in which populations appear to be 
under greatest stress from other environ- 
mental factors, particularly low temperatures 
and food shortage. 


VI. SUMMARY 


Data on the parasites of the Florida mouse, 
Peromyscus flortidanus, were obtained from 
35 localities during the period February, 
1957, to October, 1960. Samples of from 
542 to 610 mice were surveyed for different 
kinds of ectoparasites, while the numbers of 
specimens examined for various groups of 
endoparasites ranged from 4 to 840. Where 
adequate data were available, analyses were 
made to determine the influence of sex, age, 
and density of host, habitat, and season on 
parasite composition and prevalence. 

Thirty-two species of parasites were re- 
corded, only 12 of which were of relatively 
common occurrence from the standpoint of 
the species as a whole. Ectoparasites as a 
group were more abundant than endopara- 
sites. Overall infestation rates for the former 
ranged from 69.3% for trombiculid mites 
in the ear canal to 0.2% for lice. Proto- 
zoans were the most frequent endoparasites 
(42.0%), larval cestodes the rarest (>.4%). 
In addition to generally low prevalence rates 
for various kinds of ecto- and endoparasites, 
the numbers of parasites carried by indi- 
vidual hosts were seldom high for the par- 
ticular parasite involved. No evidence of 
mortality or weakening of mice attributable 
to parasites was obtained, and it is concluded 
that parasites probably play a_ relatively 
minor role in the ecology of the Florida 
mouse, although in some populations para- 


No. 1 


sitism may be of more significance than 
in others. 

The mean prevalence of six groups of 
ectoparasites on adult mice was 24.4% as 
compared to 21.8% on young. Age differ- 
ences in prevalence were more pronounced 
in the case of endoparasites. Six species had 
a mean prevalence of 14.6% in adults and 
5.9% in young. The lighter parasite load, 
particularly of endoparasites, of young mice 
probably favors a higher survival rate in this 
age class and is therefore of significance 
from the standpoint of the population dy- 
namics of the host. 

No general correlation between overall 
ectoparasite abundance and host numbers 
was demonstrated, certain groups appearing 
to exhibit a rendency to increase in preva- 
lence with increase in host density and others 
showing a reverse trend. The most marked 
positive correlation between prevalence of 
a parasite and host numbers was shown by 
fleas. This relationship is presumed to re- 
flect the intimate association between the 
Florida mouse and the flea Polygenis flori- 
danus, which appears to be restricted to this 
host. The prevalence of ticks and botflies 
may vary inversely with host abundance. If 
actual, this trend might indicate that in the 
habitats studied these parasites are held at 
relatively low levels of abundance by factors 
not directly related to the Florida mouse or 
that short-term fluctuations in the small 
mammal populations in these habitats pre- 
vent an increase in their numbers. 

Of the host and environmental factors 
considered in this study, habitat exerted the 
strongest effect on parasite composition and 
abundance. Among ectoparasites, mites, 
ticks, and botflies tended toward greater 
abundance in moister woodlands with greater 
development of ground cover and litter lay- 
er, while fleas were the only group showing 
greatest prevalence in drier habitat types. 
No overall correlation between endopara- 
sites as a group and habitat type was appar- 
ent, although individual species exhibited 
strong variation in habitat-specific preva- 
lence. 

Prevalence of ectoparasites as a group was 
highest in late fall and early winter. Indi- 
vidual kinds departed from this overall trend 
to varying degrees. Endoparasites as a group 
were most abundant from January through 
March. The trends in ectoparasite abundance 
are probably mainly correlated with seasonal 


Layne: Parasites of Florida Mouse 25 


changes in physical environmental factors. 
The peak in endoparasite abundance may be 
related to greater foraging activity of mice 
during a time of food scarcity or to variation 
in age composition of populations. The in- 
fluence of seasonal changes in parasite pres- 
sure on the host is probably greater in the 
case of endoparasites than ectoparasites, since 
the heaviest endoparasite burden corresponds 
with the period of the year in which mouse 
populations may be under greatest stress. 


VII. ACKNOWLEDGMENTS 


I am grateful to the following persons 
who kindly aided in the identification of 
parasites: Dr. Phyllis T. Johnson, Gorgas 
Memorial Laboratory, Panama (fleas, in 


part, and lice); Dr. Glen M. Kohls, Na- 
tional Microbiological Institute, Rocky 
Mountain Laboratory (ticks); Dr. Elliott 


Lesser (intestinal protozoa), Dr. Allen Mc- 
Intosh, Mrs. M. B. Chitwood, and Mr. W. W. 
Becklund (helminths and pentastomids), 
U.S.D.A., Animal Disease and Parasite Re- 
search Division, Beltsville, Md.; Dr. Curtis 
W. Sabrosky, U. S. National Museum (cu- 
terebrids; and Dr. R. W. Strandtman, Texas 
Technological College (mites). Dr. Frank- 
lin H. White, University of Florida, tested 
a small series of mice for leptospires. Dr. 
James V. Griffo, Jr., Fairleigh Dickinson 
University, aided in several phases of the 
field and laboratory work, provided identi- 
fications for some of the helminths collected, 
and kindly reviewed the manuscript of this 
paper. Dr. Carl O. Mohr, University of 
California (Berkeley), and Dr. William L. 
Jennings, Florida State Board of Health, 
also have read and criticized the manuscript. 
Grateful acknowledgment also is made to 
the following persons who have also par- 
ticipated in various phases of this study: 
Dr. D. E. Birkenholz, W. O. Wirtz, II, Dr. 
G. E. Woolfenden, R. McFarlane, and C. R. 
Myers. 


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murine typhus antibodies in cotton rats 
of southwest Georgia during wet and dry 


periods. Trans. Am. Microscop. Soc. 77: 
48-56. 

SNEDECOR, GEORGE W. 1946 Statistical 
methods. Iowa State Coll. Press, Ames. 
485 pp. 


TEST, FREDERICK H., and Avery R. TEST 
1943 Incidence of dipteran parasitosis in 
populations of small mammals. J. Mam- 
mal, 24: 506-508. 


WECKER, STANLEY C. 1962 The effects of 


No. 1 


betfly parasitism on a local population 
cf the white-footed mouse. Ecology 43: 
561-565. 


WILSON, L. WAYNE 1945 Parasites col- 


lected from wood mouse in West Virignia. 
J. Mammal. 26: 200. 


WortH, C. B. 1950 Observations on ecto- 
parasites of small mammals in Ever- 
glades National Park and Hillsborough 
County, Florida. J. Parasitol. 36: 326-335. 


ABSTRACT 
Thirty-two species of parasites, in- 
cluding 18 ectoparasites and 14 endo- 
parasites, were recorded from the Flor- 
ida mouse, Peromyscus  floridanus. 


Layne: Parasites of Florida Mouse 27 


Only 12 species had a prevalence equal- 
ling or exceeding 5% in the total sam- 
ples of mice examined. The present data 
include that, for the species as a whole, 
parasitism is not a major factor in the 
ecology of the Florida mouse and prob- 
ably has little direct role in the regu- 
lation of population size. Although the 
overall level of parasitism is relative- 
ly low, conspicuous differences in the 
kinds and abundance of parasites may 
occur between populations. Age com- 
position and density of the host popula- 
tions, habitat, and season are shown 
to be among the factors influencing 
patterns of parasite distribution and 
abundance on P. floridanus. 


TULANE STUDIES IN ZOOLOGY 


Volume 11, Number 2 


October 18, 1963 


. INTRODUCTION 


PG OMMEGIING Sit AIONG eee 


. SYSTEMATICS 


. ARTIFICIAL KEY TO THE SPECIES 


. ACKNOWLEDGMENTS________ So EY et eal) Or os 


. REFERENCES 


| ABSTRACT ___ 


CONTENTS 


. METHODS ___ 


EDITORIAL COMMITTEE: 


Dr. WILLARD D. HARTMAN, Curator of Division of Invertebrate Zoology, Peabody 
Museum of Natural History, Yale University, New Haven, Connecticut 


Dr. ROBERT W. MENZEL, Associate Professor of Marine Sciences, Oceanographic 


Institute, Florida State University, Tallahassee, Florida 


Dr. SEWELL H. Hopkins, Professor of Zoology, Department of Biology, Agricul- 
tural and Mechanical College of Texas, College Station, Texas 


THE SPONGE FAUNA OF THE ST. GEORGE'S SOUND, APALACHEE BAY, 
AND PANAMA CITY REGIONS OF THE FLORIDA GULF COAST! 
FRANK J. LITTLE, JR., 


Institute of Marine Science, 
Port Aransas, Texas 


I. INTRODUCTION 

The sponge populations of the Gulf of 
Mexico have been little investigated although 
the western coast of Florida harbors a com- 
mercial sponge fauna second only to that of 
the eastern Mediterranean (de Laubenfels, 
1948). The commercial sponge fisheries of 
this country are centered in the Gulf. 


Although sponges have been exploited for 
years, only two studies on the general sponge 
fauna of this area have been published. Car- 
ter (1884) listed tentative genera and a few 
species collected along the West Coast of 
Florida. These names were based on dry, 
fragmental specimens. De Laubenfels (1953a) 
reported on a collection of sponges made by 
the staff of the University of Miami Marine 
Laboratory in the eastern Gulf of Mexico in 
1948. In his study, data from twenty-two 
stations were reported from the western 
coast of Florida in the area between the Dry 
Tortugas and Dog Island, to the west of 
Apalachee Bay. Three stations were occu- 
pied in the Apalachee Bay region and the 
results from these stations (de Laubenfels, 
1953a) have been included in this paper. 

The purpose of the present investigation 
was to survey the sponge fauna of the Apa- 
lachee Bay Region. This involved extensive 
collecting from stations in the area over a 
period of two years. In addition, specimens 
were obtained from Dr. John Morrill who 
made collections in the St. Mark’s Light area 
i 1955. 

The initiation of a detailed faunal investi- 
gation of the Panama City, Florida, area by 
the staff of the Oceanographic Institute, 
Florida State University, made possible the 
addition of several specimens from that vi- 
cinity for comparison. Some notes on the 
ecology of the sponge fauna are included. 


1 Submitted at Florida State University 
in partial fulfillment of the requirements 
for the degree of Master of Science. Con- 
tribution No. 177 from the Oceanographic 
Institute, Florida State University. 


II. COLLECTING STATIONS 


In Fig. 1 are located the collecting sta- 
tions in the Apalachee Bay -St. George’s 
Sound area including 14 occupied by the 
author and associates in 1956 and 1957 and 
three occupied by the University of Miami 
in 1948 (de Laubenfels, 1953a). Station 
depths are indicated in Fig. 1. Station 15, 
Panama City, was under investigation by 
Dr. Meredith Jones of the Oceanographic 
Institute, Florida State University, in 1959. 
A large part of his collections was obtained 
by dredging, the remainder by hand. De- 
scriptions of the stations follow: 

Station 1—29°55'36” N., 84°26'30” W. 
Depth: 1.5-3.5 meters. Substrate: primarily 
outcrops of Tampa limestome; much open 
sandy bottom as well. 

Station 2.—29°54'18”" N., 84°26’ W. 
Depth: 1 meter; intertidal in places. Sub- 
strate: chiefly sand; broken shells or other 
invertebrates serve as substrates for some 
species of sponges. 

Station 3—29°54'30" N., 84°23’-84°24’ 
W. Depth: 1 meter; intertidal in places. 
Substrate: chiefly oyster bars in the area; 
some clear areas of sand and mud. 

Station 4.—29°47'06"-29°48’ N., 84°19’ 
30” W. Depth: 10-14 meters. Substrate: 
rock and sand. 

Station 5.—29°49’44” N., 84°16'18” W. 
Depth: 12 meters. Substrate: sand. 

Statton 6.—29°51’ N., 84°11'24" W. 
Depth: 8 meters. Substrate: sand. 

Station 7.—29°49’30’-29°49’48” N., 84° 
07'31"-84°08’ W. Depth: 9.5 meters. Sub- 
strate: rock and sand. 

Station 8—30°05' N., 84°11'30" W. 
Depth: intertidal on bar, 2-3 meters in chan- 
nel south of bar. Substrate: oyster bar, mud 
and sand in channel south of bar. 

Station 9—30°04'30" N., 84°11’ W. 
Depth: 1 meter. Substrate: sand and Tha- 
lassia testudinum Konig. 

Station 10.—30°03’ 
Depth: 2.5-9.5" meters: 


N., 84°05’ W. 
Substrate: Tampa 


OS) 
Le) 


Z St George. 2:: 


a 
14 


84°30! 
Figure 1. 


Tulane Studies in Zoology 


Volt 


Map of the Apalachee Bay region, showing stations (drawn by Dr. Stuart 


Grossman, Institute of Marine Science, Port Aransas, Texas). 


limestone outcrops in the sandy Thalassia 
grass flat. 

Station 11—29°57'30"-29°58’ N., 83° 
55’-83°56’ W. Depth: 2.5 meters. Sub- 
strate: primarily sand and Thalassia grass 
flat. Some limestone outcrops occur. 

Station 12.—29°53'56” N., 84°20’-84°21’ 
W. Depth: 0 to 1 meter. Substrate: Beach. 
Note: Specimens from this station probably 
come from a sponge bed located approxi- 
mately 29°56’ N., 84°15’ W., according to 
local residents. 

Station 13—29°55'18” 
W. Depth: 5.5-6.5 meters. 
dominantly muddy. 

Station 14—29°46'45"” N., 84°42'12” 
W., Depth: 6 meters. Substrate: sand and/ 
or mud. 

Station 15.—This includes the entire Pan- 
ama City area of the north Florida Gulf 
Coast, both offshore and estuarine areas. 

University of Miami Stations —October, 
1948 (de Laubenfels, 1953a). 

Station 20.—29°50’ N., 84°32’ W. 
Depth: 12.5 meters. Substrate: not indi- 
cated. 


N., 84°14'12” 
Substrate: pre- 


Station 21—29°59' N., 84°05’ W. 
Depth: 6.5 meters. Substrate: not indicated. 

Stations 18, 22.—29°39’ N., 83°56’ W. 
Depth: 14-14.5 meters. Substrate: not in- 
dicated. 


Ill. METHODS 


Wading, skin-diving with face-mask and 
swim-fins, dredging, and beachcombing were 
employed in the collection of specimens 
from the areas investigated. 


Upon collection, fresh specimens were 
fixed immediately in ninety-five per cent 
isopropyl alcohol, since delay causes physio- 
logical and physical distortion, especially of 
the flagellate chambers (de Laubenfels, per- 
sonal communication). The original alcohol 
was decanted and replaced with seventy per 
cent isopropyl alcohol after an hour. 

Fixation and storage in formalin were 
avoided since these procedures eventually 
reduce the sponge to a gummy mass. Neu- 
tral formalin fixation (for histological pur- 
poses) may be used providing the specimen 
is soon placed in at least two changes of 
seventy per cent alcohol to remove any traces 


No. 2 


of the formalin (de Laubenfels, personal 
communication ). 


Hand sections were cut and mounted after 
the method of de Laubenfels (1953b). Par- 
affin mounts and microtome sections were 
made on most specimens. After the paraffin 
was removed by xylene, the section on the 
slide was removed from the xylene, blotted, 
and treated in the same manner as hand-cut 
sections. Spicule mounts of specimens with 
siliceous spicules were prepared according 
to the method of de Laubenfels (1953b). 
Sponges with a cortex, a special dermal 
skeleton, or a dermal membrane required 
spicule mounts from both this outer area 
and the endosome. Differences in the spic- 
ule populations of ectosome and endosome 
often are significant taxonomic characters. 
Mounts of boring sponge spicules were made 
by the method of Old (1941). Spicule 
mounts of calcareous sponges were prepared 
in much the same manner as those of bor- 
ing sponges, except that concentrated po- 
tassium hydroxide solution was substituted 
for the nitric acid. This more tedious meth- 
od was used since the normal spicule mount- 
ing procedure of de Laubenfels (using 
KOH instead of HNO:) always leaves a 
thick coating of white substance on the 
slide which renders observation difficult. 

Measurements of mean size and range of 
spicules were based on not less than 10 
(usually 20) spicules of each category. Some 
data are expressed in a formula: 7.e. Lower 
limit-mean-upper limit (length) X lower 
limit-mean-upper limit (diameter ). 


IV. SYSTEMATICS 

Sixty-four species, in 48 genera, are in- 
cluded here. Of these, 56 species, in 41 
genera, were found during the course of 
this investigation. The remainder were re- 
corded only by de Laubenfels (1953a). None 
belong to the class Hexactinellida; but the 
Calcarea and Demospongiae are well repre- 
sented. 


The classification of de Laubenfels (1936a) 
is followed. All specimens have been as- 
signed Oceanographic Institute, Florida State 
University, numbers designated by “OIL”. 
Duplicate specimens have been deposited 
with the United States National Museum and 
are referred to by USNM numbers. 


Little: The Sponge Fauna 


)) 
Ov 


CLASS DEMOSPONGIAE 
ORDER KERATOSA Bowerbank 
Family SPONGIDAE Gray 


Spongia barbara Duchassaing and Miche- 
lotti, 1864—-The commercial “yellow 
sponge” was taken at Station 21 in October, 
1948 and reported as S. zimmocca barbara 
by de Laubenfels (1953a). It was later re- 
stored to specific rank in a paper published 
shortly after his death (de Laubenfels and 
Storr, 1958). 

Spongia graminea Hyatt, 1877.—The well 
known species known as the “Key grass 
sponge” (de Laubenfels and Storr, 1958) 
was taken at Station 21 and reported by de 
Laubenfels (1953a). 

Spongia sp. (?)—OI 1052, USNM 23553, 
USNM 23558 (figs. 2-4). Specimens fit- 
ting closely the published description of the 
“Gulf grass sponge,’ Spongia graminea tam- 
pa de Laubenfels and Storr 1958 (page 110), 
were abundant at Stations 1 and 10. Depth 
was between 1.5 and 3.5 meters, and the 
sponges were found on a rock substrate. 


These specimens also fit the written de- 
scription of the “glove or finger sponge,” 
S. cheiris reported by de Laubenfels and 
Storr 1958 (page 112), who found S. 
cheiris at Alligator Harbor, Florida. This 
report stems from specimens taken at Sta- 
tion | in the presence of de Laubenfels and 
later macerated, dried and sent to him by 
the author. Dr. de Laubenfels identified the 
sponge 2s the “g'ove sponge” in the field 
at the time of collection. 

All this would lead to the conclusion 
that these specimens were indeed repre- 
sentative of S. cheiris. Unfortunately the 
specimens resemble the type specimens of 
S. graminea tampa, while the type specimen 
of S. che’r?s seems close to, if not identical 
with, Hyatt’s type specimen of S. graminea, 

In addition there is the matter of color. 
S. graminea tampa is reported to be drab to 
pale taupe in color, whereas S. cheirts is 
black as is S. graminea. The author's speci- 
mens in I'fe were white with faint lavender 
t nts. 

Dr. Willard Hartman of the Yale Pea- 
body Museum feels that S. chezrts de Lau- 
benfels and Storr is identical with S. gra- 
minea Hyatt and that S. graminea tampa de 
Laubenfels and Storr may indeed be a sepa- 
rate species (personal communication ). 


Figures 2-4. 2 (top). Spongia sp. (USNM 


23553). 3 (middle). Spongia sp. section. 
4 (bottom). Spongia sp. section. 


Hippiospongia lachne de Laubenfels, 
1936.—The “sheepswool sponge” of com- 
merce was taken at Stations 18 and 22, and 
reported by de Laubenfels (19532). 

Hippiospongia gossypina  (Duchassaing 
and Michelotti, 1864)—The “velvet sponge” 
is recorded only at Station 22 (de Lauben- 
fels, 1953a). 

Aulena columbia de Laubenfels, 1937— 
The third report and the only record for the 


34 Tulane Studies in Zoology 


Vol. 14 


area is by de Laubenfels (1953a) from Sta- 
tion 20. 

Ircinia fasciculata (Pallas, 1766)—Ol 
1000 and USNM 23556. The “stinker or 
garlic sponge” has the peculiar sulfurous 
odor characteristic of all species of Ircznia. 
The filaments, which are characteristic of 
the genus, had a mean diameter of 3.7 p 
and ranged from 2 to 5.5 p in diameter. 
The bulbs at the terminal ends of the fila- 
ments averaged 9.8 p» in diameter, with a 
range from 8.6 to 12.1 p. 


This sponge may be distinguished from 
all other members of the genus, except I. 
ramosa, by its brownish-white color. Also, 
its conules are much closer together than 
those of any other Floridian species in the 
genus except I, ramosa. The shape is vari- 
able, from massive to lobate and even oc- 
casionally ramose. In ramose specimens the 
branch ends tend to be pointed rather than 
bluntly rounded as in I. ramosa (de Lau- 
benfels, 1950a). 

The “garlic sponge” was extremely abun- 
dant throughout the year, usually on rock 
substrates at a depth of 2 to 15 meters. It 
was taken at Stations 1, 2, 10, 11, 12, 20, 
and 22. In addition it was found at Panama 
City, Florida, z.e. Station 15, on buoys. 


Ircinia ramosa (Keller, 1889) de Lauben- 
fels, 1948—USNM 23689. One beachworn, 
macerated specimen was found at Station 12 
on September 25, 1956, shortly after a heavy 
storm. It was preserved in dry condition. 

The specimen was quite ramose, and had 
the characteristic bluntly rounded branch 
ends of the species, rather than pointed 
branch ends as in I. fasciculata (de Lauben- 
fels, 1950a; Hartman, 1955). The branches 
were relatively broad, though flattened. At 
its widest point one branch measured 4.3 x 
1.2 cm. Another branch was more rounded 
but still appeared slightly flattened; it meas- 
ured) 2 x, 1.Gvem; 

The surface was conulose with conules | 
to 2 mm high and averaging 2.2 mm apart, 
with a range of 1 to 4 mm. Oscules were 
scattered at random over the surface and 
were between 0.5 and 4 mm in diameter. 


Filaments characteristic of the genus ap- 
peared abundantly. Although the mean sizes 
of the filaments and their tylote ends were 
not different from those of I. fasciculata, 
the size range in I. ramosa is distinctly small- 
er. Filaments of this specimen ranged from 


No. 2 


2 to 4.4 » in diameter, with a mean of 3.6 p. 
The knobbed ends ranged from 4.4 to 9.9 p, 
with a mean of 8.3 p. The top figures of 
these ranges are distinctly smaller than 
those of the local specimens of I. fasciculata 
recorded here. 

Ircinia campana (Lamarck, 1814) de Lau- 
benfels, 1948—OI 1006 and USNM 23579. 
This is the vase-shaped Ircinia with conules 
of medium size, 4 to 8 mm apart (de Lau- 
benfels, 1936a, 1953a). It is reported to have 
a somewhat reddish color. 

A specimen was taken at Station 20 and 
reported by de Laubenfels (19532). 

Specimens were taken at Stations 1, 4, 10, 
and 12 during the course of this investiga- 
tion. Depth, in Apalachee Bay, ranged be- 
tween 1.5 and 12.5 meters, and substrate 
was rock or sand. Specimens were basically 
white in color with overtones of pink, giv- 
ing the flesh almost the color of Caucasian 
skin. The conules were about 1 mm high 
and only 2 to 4 mm apart. Flagellate cham- 
bers were hemispherical and small, with a 
mean diameter of 38 » and a range from 
27 to 46 p. 

The filaments of this species have been 
reported as 3 to 4 w in diameter (de Lau- 
benfels, 1936a) or 10 to 14 p» in diameter 
(Lendenfeld, 1888). The Apalachee Bay 
area specimens fall close to de Laubenfels’ 
measurements. Filaments proper averaged 4 
p in diameter, range 2.2 to 4.8 p, while the 
bulbous endings averaged 9.7 » in diameter 
with a range from 8.6 to 11.4 p». The fila- 
ments became distinctly narrower close to 
the bulbous ending. These narrower areas 
averaged 2.6 » and ranged from 1.8 to 4 p 
in diameter. 

Ircinia strobilina (Lamarck, 1816) de 
Laubenfels, 1948—OI 1040, USNM 23573. 
This is a cake-shaped Ircinia. Its coloration 
is reported to vary from dark grey to black 
and its conules are 6 to 12 mm apart (de 
Laubenfels 1936a, 1948). It was taken from 
a sunken ship off Panama City, Florida, 
depth 12.5 meters, substrate iron. The speci- 
men is a flat cake 9 cm in diameter and 2 
cm high. 

Verongia longissima (Carter, 1882) de 
Laubenfels, 1936—This is a long thin, ra- 
mose Verongia. Its color in life is reported 
to be gray, drab, or dull yellow (Carter 
1882; de Laubenfels, 1936a, 1948), slowly 
turning carmine or grey upon death (de 


Laubenfels, 1936a, 1948). © 


Little: The Sponge Fauna 5D 


The species was not found during the 
course of the present investigation and was 
reported from Station 20 only in the area 
by de Laubenfels (1953a, page 515). 

Verongia sp—OI 998, USNM 23552. 
This is also a long thin, ramose Verongza, 
which is persistently light brown on the 
upper surface and dull yellow on the lower 
one. On dying in air, or in alcohol, it quickly 
turns to dark purple and in alcohol remains 
thus indefinitely. It was found common at 
Stations 7, 10, and 11 and _ beachworn, 
macerated specimens were seen at Station 12. 

Depth ranged from 2 to about 13 meters. 
Of significance, on every occasion when 
the author viewed this species underwater 
it was not attached, but merely lying on the 
bottom, generally on the sandy substrate of 
a grass flat or other somewhat protected 
area. The abundance of beachworn, macer- 
ated specimens seems to support the hypo- 
thesis of unattached habit. 

Diameter of the individual branches of 
the sponge was about 1 cm while the length 
of some observed specimens exceeded 30 
cm, the branches often intertwining to some 
extent. 

Consistency in life is softly spongy. 

The surface was minutely conulose, the 
conules being 0.5 to 1.5 mm high and 1 to 
2.2 mm apart. The oscules were 3 to 6 mm 
in diameter and 0.9 to 2.8 cm apart; they 
were scattered over the surface of the sponge 
in an irregular fashion though a majority 
were located on the upper surface. A dermis 
about 15 pw thick covers the sponge. 

The skeleton consisted of an irregular 
meshwork of spongin fibers averaging in 
size about 560 x 670 p (range: 400 to 1050 
uw). The concentrically laminated spongin 
fibers averaged 105 p (range: 48 to 230 p) 
in diameter, each with a central pith zone 
constituting 30 to 60 percent of its overall 
diameter. 

The small ovate flagellate chambers aver- 
aged 20.2 p» in diameter (range: 14 to 33 4). 

A comparison of the data from these 
specimens with those from V. fistwlarts, 
aurea, longissima, and fulva (aurea per de- 
Laubenfels 1948) yields the impression that 
this sponge may indeed fall within the scope 
of V. aurea as recognized by de Laubenfels 
(1948: 85, 87), especially in view of the 
rapid color change noted above, but its liv- 
ing coloration, conule arrangement and spac- 
ing, dermal thickness, flagellate chamber 


36 Tulane Studies in Zoolog) 


size, and oscular location resemble more 
closely those of V. longissima and therefore 
its final allocation is deferred to some future 
date. 

Family DySIDEIDAE Gray 

Dysidea etheria de Laubenfels, 1936.—OI 
1019, USNM 23557. This lamellate sponge 
is characterized by beautiful sky blue colora- 
tion, primary and secondary fibers that are 
both heavily cored with coarse debris, and 
a conulated surface. The bright blue color 
distinguishes it in the field. 

Specimens were found at Station 8, No- 
vember 17, 1956, and station 10 in the sum- 
mer of 1957. The species appears to be 
fairly common, at least seasonally. 


Dysidea crawshayi de Laubenfels, 1936.— 
OI 1047, USNM 23586. This is the third 
record of this sponge. It was redescribed 
briefly by de Laubenfels (1948: 145), and 
later redescribed by him in detail (1950a: 
26-28). 

One specimen was taken from the grass 
flat at Station 11, at 2.5 meters, by J. Bran- 
ham, R. Hathaway and R. Bhatnagar on 
October 31, 1957. This specimen tended to 
be amorphous but had some low, broad lobes. 
Its color was not quite characteristic of the 
species but was a pinkish red, instead of the 
orange color previously recorded. Primary 
fibers were heavily cored with detritus and 
secondary fibers less so, which corresponds 
well with the original description. 

Size was 6 cm in diameter and 3 cm in 
height. The mean sample size of the eury- 
pyllous flagellate chambers was 69 p (range: 
53 to 84 p). 

Euryspongia rosea de Laubenfels, 1936.— 
OI 1044, USNM 23574. One specimen was 
taken on the grass flat at Station 11 on Oc- 
tober 31, 1957, by J. Branham, R. Hathaway 
and R. Bhatnagar. 

Shape was lobate to ramose, total height 
18 cm, total diameter 9 cm. The diameter 
of each branch was about 2 cm. Color was 
light to medium brown. This differs from 
the recorded rosy red color and possibly may 
be accounted for by the several hours the 
specimens spent in air before reaching the 
laboratory. 

Flagellate chambers ranged between 50 x 
30 pw, and 80 x 50 p. 

lanthella ardis de Laubenfels, 1950.—OI 
1045, USNM 23576. This amorphous sponge 
was primarily a dark plum color externally 


Volyad 


and pink internally during life. It was taken 
at Station 7 on rock and sand, November 3, 
1957. Upon its surface there appeared to 
be a yellow slime or sheen similar to that on 
lotrochota birotulata (Higgin). Apparently 
it is not uncommon since it was found in two 
of the dredge hauls made in the area. 


The consistency of my specimens was that 
of soft cork. The surface was covered with 
conules about 1 mm high and 2 to 5 mm 
apart and there was a definite, dense, dermis 
30 to 45 p» thick covering the sponge. The 
endosome, in sections, appeared quite fleshy 
containing ovate, sacklike flagellate cham- 
bers in profusion. These were 22-31.9-49 pu 
in diameter. The laminated spongin fibers 
were 96-210.8-345 m in diameter and ap- 
peared dendritic in arrangement, that is, they 
branched but seldom, if ever, anastomosed. 
This is attributed to the fact that the fibers 
were generally 1 to 2 mm apart in the sec- 
tions and that their anastomoses were not 
seen because of the thinness of the sections. 
The fibers contained a large central pith 
area constituting about one-third of the di- 
ameter in smaller fibers to two-thirds of 
the diameter in larger ones. The small cells 
within the fibers, which set this genus apart 
(de Laubenfels 1948: 157), were also noted. 


These specimens most closely match those 
that de Laubenfels (1936a: 31-32) originally 
reported as I. basta and which he later con- 
sidered conspecific with ardis (1950a: 33). 
The morphology of my specimens and of 
de Laubenfels’ basta specimen resembles 
closely that of the type specimen of I. ardis 
with the exception of color, flagellate cham- 
ber size, and dermal thickness. 

Color in both my specimens and de Lau- 
benfels’ basta may be said to be purple while 
that in the ardis type specimen is reported 
yellow to emerald green (de Laubenfels 
1950a: 31). Flagellate chamber size in de 
Laubenfels’ basta specimen is 25 to 45 p 
which matches the data from my specimens 
well, while the size in the ardis type speci- 
men is about 30 to 60 p. Dermal thickness 
in both my and the basta specimens is gen- 
erally between 30 and 45 » while in the 
ardis type specimen it is reported to be 15 
(de Laubenfels 1950a: 32). 


In spite of the differences noted above, 
I am reluctant to designate this as a new 
species at present because of the overall 
morphological similarity exhibited among 


No. 2 


the specimens. Further specimens are needed 
to give an estimate of the range of variation 
in each population. 

I am indebted to Drs. Willard D. Hart- 
man and Patricia R. Bergquist for indicat- 
ing the proper generic assignment of these 
specimens. 


Family APLYSILLIDAE Vosmaer 

Darwinella joyeuxi Topsent, 1889.—OI 
1007, USNM 23550. The genus is peculiar 
for having triaxon horny spicules and is set 
apart for that reason. 

During the course of the present investi- 
gation specimens of this species were taken 
at Stations 4, 8, and 13 between 2.5 and 14 
meters on rock and sand bottoms. Shape 
was massive to amorphous. De Laubenfels 
(1953a: 517-518) previously reported a 
specimen from Station 20 as D, mulleri. As 
discussed below, this specimen is considered 
identical with those taken during this in- 
vestigation. 

This dull red, softly spongy, conulose 
sponge reached a maximum height of 10 cm 
and diameter of 15 cm. Its principal lami- 
nated spongin fibers averaged 50 p (range 
32 to 61 mw) in diameter and rarely were 
cored with siliceous spicular detritus and 
other material. The secondary, or connect- 
ing, fibers which had a mean diameter of 
19.74 (range 10 to 30 pw) were not so 
cored. The arrangement of these fibers ap- 
peared to be quite ordinary for the genus, 
as was the general architecture. 

The horny, almost equi-rayed, triaxon 
spicules had rays averaging 10.4 x 559 yp 
(range 7 x 437 to 16 x 650 p»). 

The ovate flagellate chambers had mean 
dimensions of 26.2 x 58.7 w (range 14 x 39 
to 39 x 61 pee 

Were it not for the slight amount of 
detrital coring of the principal fibers, these 
specimens would fall to D. australiensis 
Carter as recognized by Topsent (1905: 
CIXXVI, CLXXXIT) and Levi (1952: 
-38-39), or to D. mulleri Schultze if de Lau- 
benfels’ (1948: 168-170) broad concept of 
that species is considered valid. 


Checking of de Laubenfels’ (1953a) 
specimen slide from Station 20 failed to 
yield any evidence of detrital coring which, 
however, was rare even in my specimens. 
In all other respects, however, his speci- 
men compared favorably to mine. De Lau- 


Little: The Sponge Fauna 37 


benfels’ statement (1948: 171) that some 
of the horny spicules in D. joyeuxi anasto- 
mose to form a reticulation independent of 
the principal keratose fibers was not veri- 
fied by reference to the original description 
or to Topsent’s later reference to this species 
(1905: CLXXXIV, CLXXXVII-CLXXXIX) 
and therefore is to be disregarded. 

Probably the specimens of Darwinella 
heretofore reported by de Laubenfels (1950a: 
38-39; 1953a: 517-518) are all D. joyeuxi, 
in view of the overall agreement between 
them and the specimens reported here and 
the relative rarity of coring material in these 
specimens. On the other hand, it may also 
be true that de Laubenfels’ view is the cor- 
rect one and that they represent D. mulleri. 
If this is true then D. joyeuxi also falls to 
muller’ on the basis of the evidence pre- 
sented above. At this time I prefer to main- 
tain the distinction between the two, at 
least until new specimens and data are forth- 
coming which may clarify the issue. 


Family HALISARCIDAE Vosmaer 

HALISARCA PURPURA, sp. nov— 
(figs. 5-9). OI 1038, USNM 23589. 

USNM 23589 is designated as the holo- 
type; April 14, 1957 by Mr. J. Branham. 

Locality and abundance—This species was 
reported abundant and encrusting on the 
turtle-grass, Thalassia testudinum Konig, at 
Station 9 on the date of collection. Subse- 
quent visits by the author failed to yield any 
new specimens. Depth was less than 1 meter. 

Shape and size—Basically encrusting, but 
the surface was somewhat lobate and there- 
fore appeared almost wrinkled. The largest 
specimen was 6 cm long and 1-1.8 cm in 
diameter. A smaller specimen was 3.8 x 0.4 
x 0.8 cm. 

Color—Color in life was a striking pur- 
plish-red, brighter than maroon. This color 
was found throughout the fresh organism by 
Mr. Branham but the color in alcohol was 
drab grey throughout. 

Consistency.—Soft, almost colloidal. 

Surface —Smooth. 

Oscules—Not observed; they are pre- 
sumed to be very small. 

Ectosomal anatomy.—Thete was a proto- 
plasmic dermis, over a 600 to 700 p thick 
alveolar zone of small subdermal cavities 
which ranged in size from 2 x 2 w to 20 x 


38 Tulane Studies in Zoology Vol. 11 


‘Figures 5-7. Read with page turned sideways. 5 (top). Halisarca purpura, sp. nov., Sec- — 
tion. 6 (lower left). Halisarca purpura, sp. nov., section of outer alveolar zone. 7 (low- — 
er right). Halisarca purpura, sp. nev., section of choanosome. ; 


No. 2 


37 p. This alveolar zone contained many 8 
to 32 cell developmental stages. 

Endosomal anatomy—This consisted of 
flagellate chambers, canals, and hyaline jelly 
between them. The flagellate chambers had 
a mean size of 21 x 38 m (range in length: 
coero ss) uw; diameter 12 to. 30 4). They 
were long and_ sack-like and sometimes 
branched. There were a great many develop- 
ing spermatocytes and oocytes in the endo- 
some, indicating sexual maturity at the time 
of collection. 

Skeleton —None; only the colloidal ground 
substance was present. 

Discusston: De Laubenfels (1948: 175) 
stated that the flagellate chambers of Halz- 
sarca dujardini Johnston are “commonly 25 
microns in diameter by 60 microns to 150 
microns long.” The color is a dull yellow- 
ish brown. The diameter of the flagellate 
chambers of H. magellanica Topsent (1901b: 
44) are reported to range between 70 and 
100 pw. H. magellanica is purple in color. 

Specimens taken at Station 9 by Mr. Bran- 
ham have flagellate chambers which are in 
the range of djardini, while their color re- 
sembles that of magellanica. Since there are 
few anatomical characters to go by in this 
genus, we might have an intermediate be- 
tween the two species. Indeed, some author- 
ities regard other members of the genus as 
conspecific with dwjardint, but considering 
the wide range of the genus and the isolated 
occurrence of the forms reported, as well as 
the few distinct morphological characters, at 
the present I regard them as separate species 
as does de Laubenfels (1932, 1948). 

A detailed study of the embryology in the 
manner of Levi (1956a) may further clarify 
the situation. 


ORDER HAPLOSCLERINA Topsent 


Family HALICLONIDAE de Laubenfels, 1932 

Haliclona rubens (Pallas, 1766) de Lau- 
benfels, 1932—OI 1037, USNM 23554. 
This dull red, ramose sponge was plentiful 
throughout the year along the beach at Sta- 
tion 12. Most specimens, however, were 
badly beachworn and macerated. 

The oxeas which make up the skeleton in 
this species averaged 136 x 4.1 p im size 
(range in length: 115 to 157 p»; width 1.8 
to 7.3 »). This agrees well with the range 
of Hartman’s 1955 data from the Gulf of 
Campeche, and does not vary greatly from 
the de Laubenfels (1936a, 1949a) values 


Little: The Sponge Fauna ay) 


Figures 8-9. 8 (top). Halisarca purpura, 
sp. nov., section of spermatocysts. 9 (bot- 
tom). Halisarca purpura, sp. nov., section 
of oocytes. 


from Florida and the Bahamas, though Car- 
ter (1882) and Wilson (1902) report larger 
spicules. Carter found 230 p oxeas while 
Wilson’s are reported to be 3 p longer than 
the largest spicules in my specimens, or 160 
x 4 ». Hartman (1955) tabulated all of 
these data. 

Haliclona rubens was reported in an an- 
notated checklist for the study area (Men- 
zel, 1956). 

Haliclona viridis (Duchassaing and Mi- 
chelotti, 1864) de Laubenfels, 1936—OI 
1014, USNM 23587. The coloration of this 


40 Tulane Studies in Zoology 


sponge varied from light green to grey- 
brown. Its skeleton consisted eniirely of 
oxeas, plus a few stylote and strongylote 
spicules that were clearly derived from the 
oxeas. 

De Laubenfels (1953a) reported the spe- 
cies from Station 21 (USNM 23396). 
Measurements made on a slide of this speci- 
men indicate that the oxeas had an average 
size of 165.9 x 5.7 » (range in length: 144 
to 201 p; width: 2 to 10 »). 

Specimens on rock at Stations 4, 7, and 
13, between 6.5 and 14 meters, were taken 
during the course of the present investiga- 
tion. Spicule size was much nearer the “3 
by 120 microns” size given by de Lauben- 
fels (1950a) for Bermuda. The mean size 
found was 120.2 x 3.3 » (length range: 96 
to 153 »; width range: 1 to 7 p). 

Haliclona permollis (Bowerbank, 1866) 
de Laubenfels, 1936—OI 1036, USNM 
23585. This is a brownish-grey or lavender 
Haliclona. Its shape varies from thickly en- 
crusting to massive and amorphous. Its skel- 
eton is reported to be comprised of an iso- 
dictyal reticulation of oxeas. Sometimes a 
few of the oxeas are modified to styles, but 
this is not uncommon in the genus. 


This species is cosmopolitan and variable. 
De Laubenfels found spicule ranges of 3 x 
90 to 5 x 100 » from Plymouth (Note in 
de Laubenfels’ Card Index of Porifera), 6 to 
8 x 150 » from California. (1932; 121), 4 
x 105 to 5 x 110 » from Bermuda (1950a: 
47). The skeleton of the present specimen, 
taken from Panama City, had oxeas averag- 
ine 5 x 150 fe Geanve] Ux 109 tony x 
LS.) 

Because of the pronounced isodictyal re- 
ticulation of the skeleton, and the lack of 
any dermal or cortical specialization, this 
specimen is placed here. However, the length 
of the spicules may indicate that it is a 
separate species. 

Haliclona sp. (?)—USNM 23686, 23687 
(figs. 10, 12). On both November 4 and 
November 16, 1956, specimens were taken 
at pace 9 by Mr. R. Hathaway and the 


author. 


One specimen consisted of a number of 
ramose arms, 0.5 to 1 cm in diameter, ex- 
tending from a base of 1 x 2 cm. The maxi- 
mum overall length was about 6 cm. An- 
other specimen appeared to be one of the 
ramose arms plus a few pieces of such an 


Vol. 11 


arm. The piece was 0.5 to 1 x 6.5 cm. One 
specimen in alcohol was light greenish tan 
and the other was white. Consistency was 
softly compressible and both specimens 
were easily torn. 


The surface of both specimens was typi- 
cally haliclonid and the endosome of both 
was isodictyally recticulate in places. There 
were also vague tracts containing 3 to 6 
spicule rows. The principal spicule was a 
sharp-pointed oxea 96-124-139 x 1.8-6.1- 
10.6 » in the tan specimen and 103-131- 
167 x 2-6.7-11 pw in the white one. There 
seems to be a tendency for the larger spi- 
cules to become strongylote or stylote; for 
example, in the tan specimen the strongy- 
lote type was 77-104.2-123 x 7-88-13 p 
and the stylotes were 105-118.6-125 x 7-8.6- 
11 p». Only the thicker spicules were so 
modified; there seemed to be no juvenile 
forms of these types. In addition the overall 
length seemed to diminish as this rounding- 
up occurred, as indicated by the fact that 
the mature oxeas seemed to be longer than 
the styles of comparable width, and the 
styles in turn were longer than the strongyles. 
There was no localization of these types 
within the specimen. This, with the reticu- 
late nature of the sponge and the ramose 
branches from the main body, seem to in- 
dicate the genus Pellina Schmidt but the 
lack of any dermal specialization indicates 
Haliclona., 


Haliclona erina de Laubenfels (1936b: 
457) seems extremely close to the present 
specimens, and it may be that they are con- 
specific with it. De Laubenfels listed the 
oxeas as being 3 x 120 to 10 x 200 m but 
made no mention of a pronounced strongy- 
lote and stylote modification. Indeed, his 
slide shows relatively few such forms. For 
this reason, I feel that the present conserva- 
tive Course 1s wisest. 


Family DESMACIDONIDAE Gray 


Xytopsene sigmatum de Laubenfels, 1949. 
—OI 1017, USNM 23548. This bright or- 
ange, amorphous sponge with conical ele- 
vations was found at Station 1, on Tampa 
limestone between 1 and 3 meters deep, 
throughout the year. It reached a height of 
6 cm and a base diameter of up to 10 cm. 

The spiculation is distinctive, containing 
tylotes, two sizes of sigmas, and also two 
sizes of isochelas that are primarily arcuate 


No. 


i) 


Little: The Sponge Fauna 41 


1 a2 3 4 5 » 6 ? 


8 9 10 1 12 13 14 AS. 


Figures 10-11. 10 (top). Haliclona sp. (USNM 23686, 23687). 11 (bottom). Callyspon- 


gia repens, sp. nov. (USNM 23551). 


but verge towards palmate. In his original 
description de Laubenfels listed only one 
type of isochela, but Dr. Willard Hartman 
found two in slides of specimens from Sta- 
tion 1. De Laubenfels’ slide also shows two 
sizes of isochelas. 


The sizes of the various spicule types are 
as follows: in the ectosome, tylotes 262- 
280.3-314 x 3.0-4.36-5.5 p, chelas 33-39.8- 
44 and 11-14.5-15 » in chord length, sigmas 
40-45.2-51 and 11-13.6-15 p» in chord length, 
in the endosome, tylotes 249-270.5-301 x 


2.9-4.24-5.7 p, chelas 22-38.9-44 and 13- 
15.2-18 p, and sigmas 40-43.9-53 and 11- 
(ABS 

This is the second record of the sponge. 
This specimen was identified by the late Dr. 
de Laubenfels, who originally described the 
species from the Western Bahamas. 


Family CALLYSPONGIIDAE de Laubenfels 

Callyspongia vaginalis (Lamarck, 1814) 
de Laubenfels, 1936—OI 996, USNM 
23565. This, the common tube sponge, was 
found previously in the area at Station 20. 


42 Tulane Studies in Zoology 


Living specimens of the sponge were taken 
at Station 4, depth 11 meters, from a rock 
substrate. Beachworn specimens were found 
throughout the year at Station 12. 


The color in life was buff brown. Hollow 
cylindrical tubes 3 cm in diameter and 20 
cm in height were found. Spiculation was 
entirely of oxeas, and the structure was 
typically callyspongiid, z.e., the dermal spe- 
cialization consisted of a secondary reticu- 
lation of small fibers with the coarser pri- 
mary meshwork, thereby giving an overall 
appearance of distinctly smaller mesh size 
at the surface. 

The spicules of area individuals averaged 
92) x3) (tanse: eax Z-ro 101 x A. a), 
which is fully 20 » longer than the largest 
thus far recorded for the species. In view 
of the overall agreement with the published 
description (de Laubenfels, 1936a: 56) I do 
not feel that this difference constitutes suf- 
ficient evidence for designation as a new 
species, and therefore it is placed here. 

The surface of many of the tubes was cov- 
ered with small bright sky-blue spots 1 to 
2 mm in diameter and 3 to 4 mm apart. 
These apparently represent a species of 
Parazoanthus, presumably P. parasiticus 
(Duchassaing and Michelotti) Verrill as 
described by Duerden (1903: 495). My 
specimens however were blue whereas Duer- 
den’s were brown, 2.¢., clear with pale brown 
tentacles. Thus possibly these represent a 
new species. Evidently these organisms are 
common on Callyspongia for most speci- 
mens observed were seen to have them, 
either actually present or represented by pits 
in beachworn macerated specimens. 


CALLYSPONGIA REPENS, sp. nov.— 
OI 1008, USNM 23551 (figs. 11, 13). 

The holotype is designated as USNM 
Zia55 1; 

Locality and abundance—One specimen 
was taken during September, 1955, by Dr. 
John Morrill in the vicinity of Station 10. 
It was also taken in abundance on September 
26, 1956, as beachworn specimens, at Station 
12 shortly after a storm. The substrate was 
rock. 

Shape-——Repent ramose; it has branches 
1 to 2 cm in diameter which intermingle 
and coalesce as they cross each other making 
the sponge seem almost flabellate. It may 
be hollow but is not the conspicuous tube 


Vol. 11 


that C. vaginalis is; rather its branches may 
or may not be hollow depending upon cir- 
cumstances and thickness, the thicker branch- 
es more commonly being hollow. 

Size—The largest specimens reached a 
total length of 18 cm and a diameter of 9 
cm. 


23687 


Figure 12. Spicules of Haliclona sp. 


Color—Y ellow-green to cream-colored in 
life, pale yellowish tan. 

Oscules—Widely scattered, generally ter- 
minal or on the upper side; diameter about 
6 mm. 

Consistency—Somewhat spongy and elas- 
tic but fragile. 

Ectosomal anatomy—Typically callyspon- 
giid; there is a primary meshwork of fibers 
averaging about 390 x 520 pw (range: 250 
to 775 »), which encloses a finer secondary 
meshwork of smaller fibers about 94 x 
135 mw (range: 57 to 210 »). Bothysetsser 
fibers generally are heavily cored with spi- 
cules; the primary fibers contain 1 to 11 
spicule rows and range in diameter from 19 
to 58 », while the secondaries contain only 
1 to 4 spicule rows and are between 10 and 
29 pw across. 

Endosomal anatomy.—Fibro-reticulate, the 
endosomal portions of the primary surface 
fibers form a meshwork averaging about 280 
x 400 » and range in diameter from 110 to 
640 pu. Very little sponge tissue was seen, 
for the most part it seems confined to the 
areas adjacent to the fibers. 

Skeleton.—The spiculation resembles that 
of C. procumbens (Carter) as described by 
de Laubenfels (1936a: 57) under the name 
Patuloscula plictfera (Lamarck) and later 
corrected by him (1950a: 61; 1953a: 523). 
There are oxeas, some verging to strongyles, 
75-100.3-107 x 1-3.9-7 pw, as well as mi- 


Little: The Sponge Fauna 43 


a 
SOw 


Figure 13. Spicules of Callyspongia repens, sp. nov. 


croxeas of which many are bent to resemble 
toxas so closely as to be mistaken for them. 
Three classes of microxea were found. There 
were raphides, 18-33.8-55 y, and two sizes 
of bent microxeas many of which went so 
far as to become pseudo-toxas. The larger 
ones were 23-25.1-32 x 1.8-2.2-2.7 » while 
the smaller were 7-14.4-20 » long. There 
were also small siliceous objects 3-4.1-6 x 
0.8-1.2-1.6 » which looked like little bright 
kidney beans in the field. 


Discussion —This species most resembles 
C. procumbens (Carter 1882: 365) from 
which it differs chiefly in color, somewhat 
smaller mesh size than that reported by de 
Laubenfels (1936a: 57-8), and in the 
densely packed fibers as opposed to the 
sparsely cored fibers of procumbens origin- 
ally reported and verified in de Laubenfels’ 
preparations. Indeed, de Laubenfels’ ma- 
terial matches the literature data (Carter 
1882: 365; Dendy 1890: 355-56) with the 
one exception that de Laubenfels’ slides con- 
tain thinner spicules than the material re- 
ported on by previous workers. Burton's 
(1934: 539) report of toxas in the type 
specimen of procumbens is also matched by 
de Laubenfels’ data. 


Though the spicule size of this species 


better approximates the data presented by 
Carter and Dendy than does that of de Lau- 
benfels, the other morphological differences 
cited above preclude the designation of my 
specimens as procumbens both on a quanti- 
tative basis and on the purely qualitative 
impression resulting from the comparative 
material examined. 


ORDER POECILOSCLERINA Topsent 
Family ADOCIIDAE de Laubenfels 


Adocia neens (Topsent, 1918) de Lau- 
benfels, 1936—OI 1031, USNM 23601. 
This species was found as an encrustation 
3 to 4 mm thick and 3 to 4 cm in diameter 
on Geodia gibberosa Lamarck at Station 11, 
depth 2.5 meters, on October 31, 1957. Color 
was white in life and is the same in alcohol. 

The presence of a neatly reticulate skele- 
ton of oxeas verging toward strongyles, and 
a detachable reticulate dermal skeleton, place 
the specimens in this species. There seem 
to be two sizes of spicules. One averaged 
126 x 6 mw (tange: 110 to 134 »), the 
other averaged 110 x 3 p (range: 98 to 
116 »). The small ones are undoubtedly 
immature. De Laubenfels (1936a: 58) in- 
dicated that the spicules of his specimen 
were 118 x 5 p» in general and some were 


44 Tulane Studies in Zoology 


as small as 105 x 1 pw. Thus the Apalachee 
Bay specimen has slightly longer spicules, a 
difference which I feel not marked enough 
to justify designation of a new species. 


Family COELOSPHAERIDAE Hentschel 

COELOSPHAERA FISTULA, sp. nov. 
—OI 1049, USNM 23583 (figs. 14, 16). 
The holotype is designated as USNM 23583. 

The specimens were taken from the Sta- 
tion 4 area April 29, 1957, by J. Branham 
and R. Hathaway. 

Locality and abundance——This species was 
relatively common in the vicinity of Station 
4 throughout the year. It was growing on 
tests of dead sand dollars, Mellita quin- 
guiesperforata (Leske), at a depth of 12 to 
14 meters. 

Shape.—Fistulate, like a small bent finger 
standing erect upon the surface of the sand 
dollar test. No basal mass was observed. 

Size-——Up to 1 cm in height and between 
5 and 7 mm in diameter. This represents 
the largest found though many were con- 
siderably smaller. 

Color —White, both in life and in alcohol. 

Oscules—No obvious vents were found, 
as is often the case in the Coelosphaeridae. 

Ectosomal anatomy.—This consisted of a 
dermal region, 75 to 100 w thick, densely 
packed with spicules in confusion; their 
interstices were also packed with organic 
material, flesh, and perhaps spongin. 

Endosomal anatomy.—The endosome was 
wanting and was replaced by a hollow fluid- 
filled area. Presumably the fluid was sea 
water but no investigations were made con- 
cerning it. 

Skeleton—The spicules were packed in 
the ectosome in confusion and did not in- 
vade the hollow central cavity. The mega- 
scleres were tylotes 107-211-240 x 4-4-5 yp. 
The mean size of the heads at each end was 
5.7 » (range: 5 to 7 pw ). The microscleres 
were unguiferate isochelas 9-10.5-12 jp, and 
sigmas 25-40.8-53 p. The chelas generally 
bore four teeth at each end. A few were 
seen. that seemed to have three teeth but 
observation was difficult due to their posi- 
tion on the slide. Under lower magnifica- 
tions these may appear to be arcuate iso- 
chelas. 

Discussion—The sponge is placed in this 
genus due to its structure and spiculation. 
The size and range of its spicules effectively 


Vole 


exclude it from any of the existing described 
species. In C. actimioides (Hallmann, 1914) 
from Australia, the microscleres are closest 
in size to those of the Station 4 specimens, 
but the chelas are arcuate and the mega- 
scleres are over 100 pw longer. C. tunicata 
(Schmidt, 1870), the only West Indian 
member of the genus to date, does have 
broad spatulate three-toothed _ isochelas. 
However, they are too large, averaging about 
31 » (Topsent, 1920: 17) and do not re- 
semble closely those of the present speci- 
mens. 


Rhizochalina oleracea Schmidt, 1870.— 
USNM 23688 (figs. 15, 17). Specimens 
were found growing on Geodia gitbberosa at 
Station 11, October 31, 1957. It was de- 


ENS I EBB 


LL 
7 8) 9 


Figures 14-15. 14 (top) Coelosphaera fis- 
tula, sp. nov. (USNM 238588) on WMellita 


test. 15 (bottom) Rhizochalina oleracea 
Schmidt (USNM 23688). 


ey 


——— 


25y 
Figure 16. Spicules of Coelosphaera fistula, 
sp. nov. 

SSS ee 
ae | 


SO 


Figure 17. Spicules of Rhizochalina oleracea 
Schmidt. 


scribed as Phloeodictyon nodosum by George 
and Wilson (1919: 152-53) from Beau- 
fort, N. C. This was corrected by de Lau- 
benfels (1947: 35) on the basis of his 
study of “many West Indian specimens of 
oleracea,” though the results of these studies 
were never published. A search of de Lau- 
benfels’ slide collection yielded five speci- 
men slide sets from the West Indian region 
definitely identified as oleracea by him. The 
spicular and gross morphological data from 
these is compared to both the literature data 
and my own in Table 1. Wells, e¢ al. (1960: 
212) also confirm de Laubenfels’ opinion 
after examining George and Wilson’s type 
specimen. 


My specimen consisted of fingerlike cy- 
lindrical fistulae arising from an encrusting 
basal portion 2 to 5 mm thick. The fistulae 
were 2 to 4 mm in diameter and 2 to 4 cm 
long. They were hollow and the ends of 
each were closed and were not oscular sites. 
Color was white. 

The surface of the fistular wall was 
smooth, being a tangentially arranged, uni- 
spicular, triangular reticulation supported 
by a more or less perpendicular unispicular 


Little: The Sponge Fauna 45 


reticulation above vague tracts or confusedly 
arranged spicules lying parallel to the sur- 
face. The perpendicular reticulation is some- 
what vague in itself and holds the surface 
layer one spicule length above the interior 
layer. The endosome is wanting in the fistu- 
lae, being replaced by the large central 
Cavity. 

The ectosome of the basal portion 1s simi- 
lar to that of the fistulae except that the tan- 
gential dermal reticulation appears more 
polygonal due to a somewhat more dense 
arrangement of the spicules which comprise 
it. Also, here we find that the tracts just 
below the perpendicular reticulation are 
more definite. They are more closely packed 
with spicules and range from 60 to 135 p» 
in diameter. These, in turn, are supported 
above extensive subectosomal cavities 90 to 
380 » in diameter by perpendicular exten- 
sions of some of the endosomal tracts. 

Below the subectosomal cavities lies a 
loose mass of vague and distinct tracts of 
spicules 45 to 125 » in diameter forming a 
vague reticulation. There are also many 
spicules loosely scattered in vague bundles 
or in confusion throughout the flesh. 

The general morphology of this sponge is 
in close agreement with that of George and 
Wilson’s, Topsent’s (1920: 2), and de Lau- 
benfels’ specimens (Table 1). The size range 
of the oxeas which comprise the skeleton 
matches well the data of Wilson (1902: 
395), George and Wilson, Topsent, and de 
Laubenfels smaller spiculed specimens 
though his larger spiculed specimens, 2. 
USNM 22390, 22388, and BNMH TW 17 
VIII, may require reallocation on further 
study (see Table 1). The color difference 
from brown (Topsent) to brownish white 
(George and Wilson) is considered insig- 
nificant, especially since de Laubenfels’ speci- 
mens are recorded as ranging between yel- 
lowish drab and pale grey in life. 

That this sponge has not been found here 
before is probably due to its small size and 
inconspicuous habitus. Probably it is far 
more common than this one collection in- 
dicates. 


Family PLOCAMIIDAE Topsent 
HOLOPLOCAMIA DELAUBEN- 
FELSI, sp. nov—OI 1039, USNM 23596 
(fig. 18). 
The holotype is designated as USNM 
25596: 


Vol. 11 


Tulane Studies in Zoology 


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Little: The Sponge Fauna 47 


FY 
3 


See eee eee 
25y 


Figure 18. Spicules of Holoplocamia delaubenfelsi, sp. nov. 


Locality and abundance—One specimen 
was found growing on an oyster shell at- 
tached to a specimen of Verongia sp. at Sta- 
tion 10, August 13, 1957 by Dr. John Mor- 
rill and the author. Depth was 2.5 meters. 


Shape.—Encrusting; the specimen was en- 
crusted completely around the oyster shell 
making the sponge appear superficially mas- 
sive and amorphous. 


Size—Diameter of the encrusted shell 
was about 3 cm and it was completely cov- 
ered. Height of the sponge was approxi- 
mately 1 to 3 mm. 

Color—Bright orange-red in life, and 
dark drab in alcohol. 

Oscules—No definite vents were seen. 

Consistency—Firm, yet slightly elastic. 

Ectosomal anatomy—No definite dermis 
was observed. The surface was somewhat 
hispid since the tufts of plumose columns 
of spicules project through the surface. 


Endosomal anatomy.—The endosome was 
made up of two regions, a confused mass of 
spicules which formed a base, and plumose 
columns of spicules which extended to the 
surface. These principal tracts contained 3 
or 4 spicule rows and were 20 to 30 p in 
diameter. They seemed almost axinellid in 
that they were so plumose. These were 
joined by secondary tracts 1 or 2 spicules 
wide, 8 to 10 » in diameter, and one spicule 
long, thus appearing like rungs of a ladder 
between two upright plumose columns. The 


resulting mesh varied in size up to 150 x 
190 », which seemed to be the modal size. 
Also observed were a few pieces of con- 
centrically laminated spongin fiber 85 to 
170 » in diameter, containing pith. These 
appear to be dendritic because some branch- 
ing was observed but no recrossing. These 
fibers were 290 to 960 mw apart in the 
sections. 


Skeleton—The megascleres consisted of 
strongyles and styles, both of which may 
be slightly acanthose at or near their 
rounded ends. The strongyles, mean size 138 
x 9 » (range: 112 to 153 »), made up the 
bulk of the basal mass while the styles, mean 
size 265) x 15a, (range: 199 ato 30672.) 
made up the bulk of the plumose columns, 
though both were found throughout the 
sponge. The strongyles characteristically had 
unequal ends, the smaller end corresponding 
to the pointed end in styles. They appeared 
like styles that had rounded up short of their 
goal. The microscleres included toxas, mean 
size 79 » (range: 48 to 103 pm), and pre- 
dominantly palmate isochelas, mean chord 
length 14 pw (range: 11 to 16 pw). The 
shovels on the isochelas averaged about 4 
across. A few arcuate chelas within this size 
range were also found. 

Discussion—Lévi (1952: 54) advocated 
dropping Holoplocamia in synonymy to 
Plocamilla Topsent (1928: 63), but Topsent 
stressed that Plocamilla was set up for 


48 Tulane Studies in Zoology 


sponges in which there was no differenti- 
ation of primary from secondary tracts such 
as are cited by him (1928: 63) for Plocamia 
and therefore also for Holoplocamia since 
this is separated from Plocamia only by the 
presence of spiny rather than smooth prin- 
cipal (diactinal) spicules (de Laubenfels 
1936a: 75; plus personal communication ). 
Therefore, I feel that the genus should be 
retained distinct from Plocamulla. 

All other species within this genus have 
diacts entirely spined except the one described 
by Sollas (1879: 44) as Plocamia plena from 
West Africa. Its diacts are described as only 
slightly spined at the ends like those of the 
present specimen. The present specimen dif- 
fers from H. penneyi de Laubenfels (1936a: 
76), a Tortugas sponge, in size and relative 
thickness of spicules, in color, which in H. 
penneyi is brownish orange, and in the fact 
that the chelas of H. penneyi have rotated 
shafts so that one shovel is at right angles 
to the one at the other end. 

The species is named in honor of the late 
world sponge authority, Dr. M. W. de Lau- 
benfels, whose interest and confidence in me 
have been major factors in sustaining my 
work. 


Family CyAMONIDAE de Laubenfels 


Cyamon vickersi (Bowerbank, 1863) Gray, 
1867.—OI 1046, USNM 23563. One speci- 
men was taken at Station 4, November 3, 
1957. Depth was 11 meters. The specimen 
was an encrustation less than 1 mm thick 
on a piece of limestone. The color was or- 
ange. 

The species may be common; its hard 
texture and thinly encrusting habitus led to 
overlooking many such pieces of orange- 
encrusted limestone in the course of dredg- 
ing. This specimen was brought back to the 
laboratory only as a check. 

The principal spicules were tetraxons, 
and a few pentactines (which are aberrant 
tetraxons) with the rays approximately 
equal, rounded and acanthose near the end 
of each ray. The rays in this specimen aver- 
aged about 41 » (range: 20 to 55 »). There 
were also a few subtylostyles about 877 x 
13 p, as well as very long, thin styles which 
echinated the surface, being perpendicular 
to it with their points out. They were 2 to 
6 » wide and up to 2 mm long, but most 
were broken. No acanthostyles, which sup- 
posedly give rise to the tetraxons (Dendy 


Viola 


1921: 117), were found. A few tetraxons 
had one acanthotylote-tipped arm very long 
in relation to the other arms, which seemed 
to be stubby in these cases. Thus, the part 
corresponding to the point in Dendy’s de- 
veloping tetraxons is acanthotylote, possibly 
indicating an intermediate stage in develop- 
ment. 

The species was redescribed in detail by 
de Laubenfels (1936a, 1950a). 


Family MyXILLIDAE Hentschel 


Merriamium tortugasensis de Laubenfels, 
1936—OI 1009, USNM 23561 (figs. 19, 
21). Several specimens of this sponge were 
taken at Station 7, depth 10 meters, Novem- 
ber 3, 1957. One other small specimen was 
taken at Station 4, depth 12.5 meters, the 
same day, thus indicating that the sponge is 
common, at least seasonally, within the area. 
The substrate was rock. 

Color, alive, was fire red. Shape was mas- 
sive. The largest specimen measured 5 x 3 
cm wide and 6 cm high. 

The spiculation of tornotes, acanthostyles, 
and arcuate isochelas is distinctive. 


Family TEDANIIDAE Ridley and Dendy 


Tedania ignis (Duchassaing and Miche- 
lotti, 1864) de Laubenfels, 1936—OI 1061, 
USNM 23560. Several specimens of this 
sponge were taken at Station 10, August 13, 
1957, and also at Station 11, October 31, 
1957. Depth in both cases was about 2.5 
meters; substrate in the first case was Sar- 
gassum and in the second was Geodia gtb- 
berosa. Habitus was encrusting to massive. 
This sponge is thought to be extremely 
abundant, at least seasonally. 

The orange to pinkish-red coloration plus 
the spiculation of tylotes, styles (rarely sub- 
tylostylote), and roughened raphides are 
distinctive. A further distinctive feature of 
the species is that the heads of the tylotes are 
faintly microspined near the apex. It was 
redescribed in detail by de Laubenfels 
(1936a, 1950a, 1950b). 

Lissodendoryx isodtctyalis (Carter, 1882) 
Topsent, 1889—OI 1011, USNM 23584. 
One specimen of this species was found at 
Station 10, August 13, 1957 permeating be- 
tween the trellis of ascon tubes of Leuco- 
solenia canariensis (Miklucho-Maclay); depth 
2.5 meters. It was subsequently found in the 
Panama City area encrusting on steel buoys 
at a depth of less than 1 meter. 


No. 2 


The lavender to brownish-green color of 
the exterior and the yellow interior are dis- 
tinctive, as is the spiculation which consists 
of tylotes, slightly bent styles, sigmas, and 
arcuate isochelas. Hartman (1958: 41) dis- 
cussed in detail whether we should separate 
the carolinensis type (Wilson, 1911) and 
the zsodictyalis type (Carter, 1882). They 
were placed together by de Laubenfels 
(1947: 35). There are distinct differences 
in the microsclere populations of the two 
groups. The carolinensis type has large sig- 
mas and small chelas while the 7sodzctyalis 
type has small sigmas and large chelas. Hart- 
man concluded that since both of these vari- 
ations occur side by side in the Mediterrane- 
an they should not be separated at present. 

The specimens collected from this area 
were of the carolinensis type having larger 
sigmas than chelas. The dimensions of the 
spicules are: tylotes, mean 166 x 4 p (range: 
146 to 176 »); styles, mean 168 x 6 yp 
(range: 157 to 183 p); sigmas, mean chord 
length 37.9 » (range: 21 to 63 pm); arcuate 
isochelas, mean chord length 24.9 » (range: 
16 to 28 ee 

The species was redescribed by de Lau- 
benfels (1936a, 1950a, 1953b). He noted 
(1953b: 21) the permeating habitus men- 
tioned above. 


Family MICROCIONIDAE Hentschel 


Microciona prolifera (Ellis and Solander, 
1786) Verrill, 1873—OI 1002, USNM 
23562. This orange-red to dull brick-red, 
encrusting to lamellate, lumpy sponge was 
abundant and was collected throughout the 
year at Stations 1, 2, and 9. Depth was be- 
tween 0 and 2 meters. Its size ranged from 
a thin encrustation to a lumpy structure up 
to 6 cm wide and 9 cm high. Shape is largely 
governed by environment. On Sargassum 
and Thalassia it was usually encrusting. On 
limestone in areas of current, it was generally 
lamellate, and on sand substrate it appeared 
as a sub-spherical mass of short lumpy 
branches. In this last area it was in a tidal 
pool and hence little disturbed by currents. 

The skeleton consisted of plumose col- 
umns of subtylostyles with small heads, 
echinated by acanthostyles. The microscleres 
were toxas and palmate isochelas, with an 
occasional arcuate isochela. There were two 
categories of subtylostyles, 165-440-529 x 
11-19-21 pw and 133-266-402 x 2-4-7 yp. 


Little: The Sponge Fauna 49 


The larger ones were often almost stylote 
in that the head was so lightly developed. 
Two categories of acanthostyles also were 
found, one entirely acanthose 71-87-103 x 
5-7-7 p, and the other with only the head 
acanthose 116-174-223 x 9-11-12 pw. The 
palmate isochelas had a mean chord length 
of 17.1 » (range: 14 to 20 pw), while the 
toxas averaged 33.5 pw (range: 16 to 45 p). 
This species was redescribed by George 
and Wilson (1919: 157) from Beaufort, 
N. C. They also described a new species, 
Esperiopsis obliqua, in the same paper (page 
148): this was said by de Laubenfels (1947: 
35) to be conspecific with M. prolifera. 
Wells et al. (1960: 218) on the other hand 
cited evidence that E, obligua is a valid spe- 
cies and restored it to specific rank as Tena- 
ciella obliqua since its spicules are enclosed 
in the spongin fibers rather than echinating 
them as in Esperiopsis. Hartman (1958: 
36) wrote an excellent discussion of M. pro- 
lifera in which the data of previous authors 
were brought together in tabular form. 


EURYPON CLAVATELLA, sp. nov. 
=O111030, USN 2457 Satis. 22). 

The holotype is designated as USNM 
2508: 

Locality and abundance—One specimen 
was taken at Station 4, depth 10 meters, on 
rock and sand bottom. 


Shape—A thin encrustation on limestone. 


Size—Less than 1 mm thick, in patches 
up to 1 cm square. It was nearly invisible 
and subsequent investigation undoubtedly 
will yield more specimens. 

Color—No positive data are available at 
present. The limestone originally was cov- 
ered with a vivid purple (lavender-red ) sub- 
stance, even in areas where there was no 
sponge. This color faded to white and was 
attributed to a coral. The sponge is drab 
in alcohol. 

Consistency.—Softly fragile. 

Surface —Hispid, which is characteristic 
of the genus. 

Oscules—None observed. 

Ectosomal anatomy.—No specialization. 

Endosomal anatomy.—Microcavernous and 
fleshy. There seems to have been a basal 
plate of spongin from which vague plumose 
tracts of spicules rose vertically to the sur- 
face, a distance of about 500 » on the aver- 
age. 

Skeleton—The vague plumose columns 


50 Tulane Studies in Zoology 


of spicules were made up primarily of tylo- 
styles and were echinated by smaller acan- 
thostyles. Many tylostyles and styles stand 
erect on the base between the columns, as 
do the acanthostyles. The tylostyles meas- 
ured 249-384-470 x 14-15-21 » with an aver- 
age head diameter of 20 p. The styles aver- 
aged 361 x 4 w (range: 351 to 392 »). The 
acanthostyles measured 75-102-145 x 5-6-9 
pw, and their head ends averaged 8.4 p in 
diameter. The styles possibly represented 
acanthostyles which lost their spines as they 
increased in size. In addition, what appeared 
to be vermiform stylote and tylote spicules 
were seen in the spicule slide. Some were 
highly contort. They ranged in mean chord 
length from 18 x 0.5 to 35 x 2 p. None 
was found in the sections, however, and I 
concluded that they were not proper spicules. 


Discussion —This species resembles most 
closely E. clavata (Bowerbank) Gray, which 
was redescribed by de Laubenfels (1950a: 
79) from Bermuda. However, the great dif- 
ference in spicule size leads to the conclu- 
sion that this is a distinct species. The tylo- 
styles in this specimen only approach one 
quarter the size of those of clavata and the 
range in size of the acanthostyles, while over- 
lapping to a considerable extent, is much 
larger in clavata. A further difference is 
the presence of the styles. 


If the presence of vermiform spicules is 
confirmed subsequently and actually they are 
proper spicules in this species, then it would 
fall to Bubarts, Their absence in pieces con- 
taining the basal plate suggests that they 
were not proper to the specimen. 


Thalyseurypon vasiformis de Laubenfels, 
1953.—This drab to black, vase-shaped 
sponge was taken at Station 21 on October 
27, 1948. Originally it was described by 
de Laubenfels (1953a: 525). It was not 
found during the course of the present in- 
vestigation. 


Family OPHLITASPONGIIDAE de Laubenfels 


Carmia macilenta (Bowerbank, 1866) 
Gray, 1867—OI 1018, USNM 23559 (fig. 
23). This species is new to the Gulf and 
Caribbean region and therefore a detailed 
description is warranted. The genus was 
incorporated into the genus Mycale by Top- 
sent (1928: 84); Burton (1956: 129) re- 
ported the species from the central eastern 
coast of Africa. 


Vol. 11 


The genus Carmia is considered here to 
be valid, after the classification of de Lau- 
benfels (1936a: 118). 


Locality and abundance—This sponge was 
common at both Stations 10 and 11, depth 
2.5 meters. It was found on Sargassum, 
Thalassia, and Ulva. On October 31, 1957, 
it was found at Station 11, and on August 
13, 1957, it was taken at Station 10. It was 
taken also in the vicinity of Station 10 in 
September, 1955, by Dr. John Morrill. 


Shape and size—Generally it was thinly 
encrusting on the available vegetation, but 
in Dr. Morrill’s specimen it was thickly en- 
crusting (on Sargassum). The largest mass 
was about 2 cm long and 0.5 cm in diameter 
with the Sargasswm stem in the middle. In 
some places thickness exceeded this, and 
there were, of course, many smaller areas of 
thin encrustation. 

Color—Orange-red to orange in life; it 
faded quickly to grey-white in alcohol. 

Surface—Superficially hispid in effect, 
probably due to numerous tracts of spicules 
which terminated in surface brushes. 


Oscules—One oscule on the 1955 speci- 
men measured 4 mm in diameter. It was 
partly closed by a thin sphincter membrane 
to an aperture 2 mm in diameter. Small 
oscular openings about 2 mm in diameter 
were about 3 mm apart in other specimens. 

Ectosomal anatomy —A protoplasmic der- 
mis, supported in large part by tangential 
tracts of spicules directly beneath it, is pres- 
ent as well as the perpendicular tracts which 
pierce it. 

Endosomal anatomy.—The endosome com- 
prised a region of many dense spicule tracts 
forming a confused mesh or trellis-work 
with the mesh size ranging from 70 x 100 
to 300 x 500 p. What little flesh there was, 
was scattered about the edges of these 
meshes along the tracts. Here the micro- 
scleres were found in profusion. The pri- 
mary spicule tracts contained 12-20 spicule 
rows and ranged from 30 to 70 » in diame- 
ter. Secondary spicule tracts containing 5-8 
spicule rows, 15 to 25 m in diameter, also 
meandered about. The haphazard crossings 
of these two types of spicule tracts formed 
the mesh. 

Skeleton—Megascleres consisted of sub- 
tylostyles with small heads, plus styles. The 
subtylostyles averaged 211 x 3 mw with a 
range from 173 to 256 p. A few immature 


No. 2 


ones measured about 171 x 2 pw. The styles 
had a mean length of 218 pw (range: 173 to 
255 m). Microscleres included palmate an- 
isochelas in three categories, plus sigmas and 
toxas. The chord length sizes of the chelas 
were 13-16.3-18, 22-24.2-31, 40-43.6-46 p. 
The sigmas were 35-84.9-96 y, while the 
toxas were 53-111-221 yp in length. 


Discussion.—Possibly this sponge might 
constitute a new species in view of some 
minor differences between it and Bower- 
bank’s description, but I concur with the 
opinion of the late Dr. M. W. de Laubenfels 
(personal communication) that these are 
not enough to separate it from macilenta at 
present. 


Family AMPHILECTIDAE de Laubenfels 


Toxemna tubulata (Dendy, 1905) Hall- 
mann, 1917.—OI 1027, USNM 23595. Sev- 
eral small specimens of this yellow sponge 
were taken from the surface of Geodta gib- 
berosa at Station 11 on October 31, 1957, at 
2.5 meters depth. 

This sponge was redescribed in detail by 
de Laubenfels (1936a: 124), but the generic 
mame was incorrectly written as Toxemma. 
He described the shape as massive to amor- 
phous in contrast to Dendy’s original de- 
scription (1905: 105) which stressed cy- 
lindrical shape. 

The specimens from this area were slender 
(1 to 2 mm diameter) cylindrical digitate 
processes rising from the substrate a dis- 
tance of about 1 cm, thus bearing out the 
original description. The spicules were 
styles 244-259.6-297 x 4-4.5-5 jm, and abun- 
dant microscleres containing raphides 29- 
103-230 x 1-1.5-2 mw, sigmas whose chord 
length measured 13-20.9-29 mw, and toxas 
which measured 26-33.8-42 pm. 


ORDER (?) HALICHONDRINA Vosmaer 


The order is not established firmly and is 
open to severe criticism. It was discussed 
by de Laubenfels (1953a: 530) who sug- 
gested that the order should be dropped and 
“most or all of its families be merged with 
those of the Poecilosclerina.” De Laubenfels, 
however, followed the established classifica- 
tion. It is followed here also. 


Family AXINELLIDAE Ridley and Dendy 

Axinella polycapella de Laubenfels, 1953. 
—Beachworn specimens were found at Sta- 
tion 12 during 1957. The species was taken 


Little: The Sponge Fauna 51 


also at Stations 18, 20, 21 and 22 in 1948 
and described by de Laubenfels (1953a: 
530). It is not now represented by a speci- 
men. 

The spiculation of this species is princi- 
pally of oxeas though a few styles may be 
found; in addition, two strongyles were 
found, one 205 x 11 p from the axis, and 
the other 168 x 10 » from the outer area. 
The spicules measured in the axis consisted 
of oxeas 230-260.8-297 x 9-10.9-15 p» and 
two styles 240 x 11 » and 249 x 12 pw. The 
oxeas of the outer areas were smaller on the 
average, measuring 182-243.3-307 x 2-8.2- 
12 p, as were the only two styles found. 
They were 192 x 14 and 187 x 17 p. 

These measurements indicate a greater 
range in spicule size than is recorded by 
de Laubenfels; also, the axis seems to have 
larger spicules than the outer portion. 
Nevertheless, because of overall morpho- 
logical agreement, both macro- and micro- 
scopic, the specimens are placed here with 
confidence. 

Homaxinella waltonsnuthi de Laubenfels, 
1953—This flabellate or palmate sponge 
was not found during the course of the pres- 
ent investigation but was taken at Station 
20, October 26, 1948. It was described 
originally from the area by de Laubenfels 
(195345555))- 


Family HALICHONDRIIDAE Gray 


Halichondria panicea (Pallas, 1766) Flem- 
ing, 1828—OI 1010, USNM 23566 (fig. 
20). This sponge superficially is much like 
Haliclona permollis (de Laubenfels 1954b: 
20), from which it is set apart by having a 
special dermal skeleton. It is basically amor- 
phous with the oscules often on volcano-like 
protrusions 0.5 to 1 cm in diameter. The 
maximum size found was 20 cm in diameter 
x 7 cm high. 

The color has been recorded as basically 
yellow (de Laubenfels, 1934), but the speci- 
mens found here ranged from light greenish- 
brown to light green in color, with one 
specimen pink. This wide range of color is 
characteristic of the species (de Laubenfels, 
1953b). A particularly distinctive character- 
istic is that the spicules are spread about in 
confusion in the cavernous endosome. How- 
ever, in the present specimens there were 
some vague tracts containing 5 to 8 spicules 
in cross section. The skeleton is made up 
exclusively of oxeas with great variation in 


Tulane Studies in Zoology 


! 4 


Figures 19-20. 19 (top) Merriamium tortugasensis de 


(bottom). Halichondria panicea (Pallas) 


size. In this case they were between 73 x 2 
and 57 aes nn 

Because of the similarity between H. pant- 
cea and H. bowerbanki Burton (1930), the 
specimens collected were examined in detail 
in regard to the dermal reticulation and 
sizes of dermal and endosomal spicules. 
Hartman (1958: 33-34) pointed out that 
H. panicea has “a very regular network of 


Laubenfels (USNM 28561). 20 


Fleming (USNM 23566). 


multispicular tracts” with occasional spicules 
lying across the oblong areas made by the 
tracts, while H. bowerbanki is different; its 
multi-spicular tracts “when present, are 
widely spaced and divide up the dermis into 
further subdivided by a 
pattern of overlapping individual spicules.” 
In addition he pointed out that the dermal 
spicules of panicea are smaller than those 


larger areas, 


No. 2 

<< => 
©> 

——— 
> 

S = —=—— 

See 

SOy 
Figure 21. Spicules of Merriamium tortu- 


gasensis de Laubenfels. 


[Pe 
50y 


Figure 22. Spicules of Hurypon clavatella, 
sp. nov. 


Little: The Sponge Fauna 53 


of the endosome while dermal spicules of 
bowerbanki are the same size or larger than 
the endosomal ones. 


Concerning Hartman’s first difference, I 
must point out that while he seems to be 
right, the judgment must be made extremely 
cautiously and after several dermal areas of 
the specimen have been observed. Individ- 
ual areas or fields of panicea may closely re- 
semble bowerbanki and probably the reverse 
is also true. 


Dermal spicule size also may be variable. 
The average size of the dermal spicules in 
various specimens was from 1.0 to 74.2 p 
smaller than the endosomal ones, the mean 
average difference being 37.7 yw. Another 
specimen collected at the same spot as the 
specimen with the least difference showed 
an average difference of 55.5 pu. The mean 
length of the endosomal spicules ranged 
from 307 to 409 p, while the dermal spicule 
average ranged from 264 to 364 up, thus 
showing the great variability of the speci- 
mens within the species. 


The species was found in abundance dur- 
ing the fall of the year at Stations 2 and 11, 
and at a depth of 5 meters in the Panama 
City area. Substrate was either sand or grass 


Figure 23. Spicules of Carmia macilenta (Bowerbank) Gray. 


54 Tulane Studies in Zoolog 


flat. Data from seven specimens are repre- 
sented. 

Halichondria melanadocia de Laubenfels, 
1936.—OI 1022, USNM 23590. This very 
dark brownish, amorphous sponge was taken 
from a steel buoy opposite the Shipyard in 
Panama City by staff of the Florida State 
University Oceanographic Institute. It had 
a few small finger-like processes extending 
from the surface. These were about 1 cm 
high and 0.5 cm in diameter. Its skeleton 
and architecture placed it in the genus Hali- 
chondria. The spicules were oxeas 144 x 2 
to 460 x 13 yw. The mean size of the en- 
dosomal ones was 308.0 x 8.7 yw and the 
mean size of the ectosomal spicules was 
320.5 x 7.4 ». In view of the great range of 
size the difference between the means is not 
considered significant; the overall mean size 
is 314.3 x 8.0 ». This greatly overlaps the 
range given by de Laubenfels (1936a: 134). 

Family SEMISUBERITIDAE de Laubenfels 

RHAPHISIA MENZEL, sp. nov.—Ol 
1042, USNM 23588 (fig. 24). 

The holotype of this species is designated 
as USNM 23588. 

Locality and abundance——One specimen, 
taken at Station 2, Bay Mouth Bar, Alligator 
Harbor, on March 4, 1957. Depth was 1.5 
meters and substrate was sand. 

Shape and size—Basically a sub-spherical 
mass 3 cm in diameter and 2 cm in height. 

Color—tThe exterior was lavender to a 
depth of about 0.5 cm. The interior was 
brown. 

Consistency.—Soft and spongy. 

Surface—The surface was covered with 
closely set lamellate projections, like flat- 
tened conules, each in turn bordered by 
minute conules less than 1 mm in height. 


heres Stee eee 


eS an 
eee 


a 


[a eee 
25y 


Figure 24. Spicules of Rhaphisia menzeli, 
sp. nov., showing two spicules centrotylote 
as is occasionally the case with each type. 


Vol. 11 


The lamellate projections generally were less 
than 1 mm apart. The overall appearance 
was fuzzy. 

Oscules—One oscule was partially hidden 
by the encircling projections or lamellate 
conules; it was 3 mm in diameter. The os- 
cules were difficult to locate due to the 
nature of the surface. 

Odor—Even in alcohol there was a dis- 
tinct fetid odor about this sponge. 


Ectosomal anatomy.—wNo specialization; 
the spicule tracts from the endosome entered 
the lamellate protrusions and branched to 
form their skeleton. 

Endosomal anatomy.—The endosome was 
microcavernous and confused. There were 
primary spicule tracts ascending to the sur- 
face. These contained 10 to 12 spicules per 
cross section and ranged from 38 to 46 p» 
in diameter. There were also secondary tracts 
containing 3 to 7 spicule rows, 15 to 23 p 
in diameter. The secondary tracts were 
sometimes parallel, and sometimes vertical, 
to the surface, very close to a primary tract 
and connected to it by a trellis of spicules. 
These trellises sometimes appeared to cre- 
ate a curious entwining effect between the 
two tracts. The random crossings of these 
tracts formed meshes 70 x 70 to 535 x 460 p. 
The flesh was confined to the region of tracts 
or interconnecting spicules. 


Skeleton —The spiculation may be re- 
garded as being comprised of a single cate- 
gory but with much individual variation. 
Some were simple, smooth, sharp-pointed 
oxeas 75-106-178 x 3-4-5 p, while others 
which might be considered immature or 
microxeas were 89-94-98 x 1-1.3-2 pw. The 
larger spicules frequently had a wide varia- 
tion of shape. Not uncommonly they were 
centrotylote, and some were also stylote and 
strongylote. 


Discusston—All the sponges now in 
Rhaphisia except the type species, R. laxa 
Topsent (1892, xvii), and R. myxa de Lau- 
benfels (1951: 263), from Hawaii, have 
much larger spicules than those of this speci- 
men. R. myxa is clearly closest to the Gulf 
form in physical characteristics, differing 
primarily in color, mesh size, and smooth 
surface. The viscous nature of both of the 
above species was not particularly noted in 
this specimen, possibly because it was re- 
ceived preserved in alcohol. 

The species is named in honor of Dr. 


No. 2 


R. Winston Menzel of the Oceanographic 
Institute, Florida State University, who in- 
spired and helped greatly in the completion 
of this work. 


Family HYMENIACIDONIDAE de Laubenfels 


Hymeniacidon heliophtla (Wilson, in: 
Parker, 1910) de Laubenfels, 1936—OI 
1013, USNM 23581. This orange-pink 
sponge was found thickly encrusting on the 
stone jetties at St. Andrews State Park, 
Panama City, Florida, throughout the year. 
The elongate, conical processes character- 
istic of the species (de Laubenfels, 1950a; 
Parker, 1910) were not in evidence except 
for slightly elevated rounded protrusions. 
Patches of the sponge 20 cm in diameter 
and 2 cm in height were found. In some 
places it was so abundant that it practically 
covered the surface of the rock. The broad 
area of attachment and the tidal currents in 
the channel undoubtedly were responsible 
for this variation of shape. 

The skeleton was composed exclusively of 
styles 128-278-345 x 2-4-5 yp. These spicules 
were somewhat smaller than usual. The na- 
ture of the conulose surface, the color, and 
the structure, identify the specimens as H. 
heliophila. 

It was redescribed by Wilson (1911), 
George and Wilson (1919), de Laubenfels 
(1936a, 1950a), and by Wells et al. (1960). 


ORDER HADROMERINA Topsent 
Family SPIRASTRELLIDAE Hentschel 

Sphectospongia vesparta (Lamarck, 1814) 
Marshall, 1892—OI 997, USNM 23547, 
USNM 23580. This is the common “logger- 
head sponge” of the Tortugas. It is massive 
and cake-shaped, growing as large as 60 cm 
in diameter and 40 cm high. Consistency 
when wet is cork-like, but after drying the 
sponge becomes woody. The color is creamy 
brown to dark brown. The spiculation con- 
sists of tylostyles averaging 386 x 9 pw (range 
214-482 »), and rare spirasters, mean length 
13 p» (range: 9-18 »). Most of the spirasters 
had three contortions or bends. 

It was found abundantly throughout the 
year at Stations 1 and 10, and was also re- 
corded from Stations 20 and 21 (de Lau- 
benfels, 1953a) thus giving a depth range 
in this area between 2 and 13 meters. Sub- 
strate is rock, generally soft limestone of 
the Tampa type. 


Little: The Sponge Fauna 55 


This species was redescribed in detail by 
de Laubenfels (1936a: 140). 

Spirastrella coccinea (Duchassaing and 
Michelotti, 1864) de Laubenfels, 1936— 
OI 1033, USNM 23598. This thinly en- 
crusting, red-brown to orange sponge fades 
to grey or white in alcohol. Its spiculation 
is of tylostyles plus abundant spirasters in 
the cortex (de Laubenfels, 1936a: 143). 

Several small patches, 1 cm in diameter 
and 1 to 2 mm in height, were found on 
pieces of limestone dredged at Station 7, 
November 3, 1957, depth 9.5 meters. The 
color was orange in life. 

The tylostyles were 134-227.5-287 x 3- 
4.2-5 p in size while the spirasters averaged 
16 » (range: 9 to 40 ») though only one 
was found of the largest size. In general the 
maximum seemed to be about 24 p. Thus 
the megascleres seem to be smaller than de 
Laubenfels’ (1936a) 6 x 360 pu record while 
the microscleres are larger than his 12 x 2 
to 20 x 4 p report. In addition the spirasters, 
while abundant in the sponge, did not “pack” 
the ectosome as he indicated. 

These differences do not seem great 
enough to justify describing it as new; 
therefore it is left here with some reserva- 
tion. 

Spirastrella coccinopsis de Laubentels, 1953. 
—This was not found during the course of 
the present investigation but was taken at 
Station 20 on October 26, 1948, and re- 
ported by de Laubenfels (1953a: 537) as 
a dubious new species. He indicated that it 
may be synonymous with S. coccinea since it 
differs primarily in color and size. 

Anthosigmella varians (Duchassaing and 
Michelotti, 1864) de Laubenfels, 1936— 
OI 1032, USNM 23594. This dingy brown 
sponge has a multitude of shapes, sometimes 
elongate and cylindrical, at other times mas- 
sive or amorphous. Its consistency was com- 
pared to that of cheese by de Laubentels 
(1949a). 

The spiculation was reported by de Lau- 
benfels (1949a: 19) as consisting of tylo- 
styles 360 x 6 p, typical spirasters, and 
spitasters that are essentially C-shaped with 
blunt, knob-like protrusions arranged only 
along the convex side. It is reported as a 
hard sponge to identify because often these 
“typical C-shaped” spirasters are rare or even 
wanting, though in some cases they do form 


the bulk of the microscleres (de Laubenfels, 
1953a: 539). 

This sponge was found at Station 7, 
depth 9.5 meters, on a rock bottom. It was 
grey and formed (encrusted) around a 
worm tube. In alcohol its color was drab 
brown or brownish grey. 

The spiculation was somewhat different 
in the endosome and ectosome. The tylo- 
styles of the endosome were 278-376.4-460 
x 4-5.7-7 » while those of the ectosome were 
smaller, measuring 240-326.2-412 x 4-5.3- 
7 p. In addition, the normal spirasters of 
the endosome, 7-19.3-31 p, were larger than 
those of the ectosome which were 4-13.0- 
31». The C-shaped spirasters were confined 
largely to the ectosome, only a few being 
found in the endosome. They were 7-9.9- 
Lyi size: 

The species also was recorded from Sta- 
tion 20 on October 26, 1948, by de Lauben- 
fels (1953a). 

Halicometes stellata (Schmidt, 1870) Top- 
sent, 1898—USNM 23571. Small portions 
of this species were collected in the Panama 
City area by Dr. Meridith Jones during the 
early part of 1958. The specimens were en- 
crustations, 0.5 to 1.5 cm in diameter and 
2 to 5 mm in height and are numbered I-II 
and 3E3 in Dr. Jones’ collection. No color 
or ecological data were available for these 
specimens. 

Because of the small size of the specimens, 
virtually all of the material was used in mak- 
ing sections and spicule mounts. 

The tylostyles, a few of which tend to be 
stylote, were 211-569.1-1015 x 3-7.1-12 p 
which is in good agreement with de Lauben- 
fels’ (1950a: 99) Bermuda report, though 
the microscleres were somewhat larger than 
he reported. The oxyspherasters averaged 
26.4 » (range: 18 to 33 »), while the chi- 
asters averaged 12.8 pw (range: 7 to 22 p). 

The minute size of the specimens and 
the difference in microsclere size lead the 
author to place this sponge here with some 
reservation. 


Family PLACOSPONGIIDAE Gray 


Placospongta carinata (Bowerbank, 1858) 
Vosmaer, 1902.—OI 1043, USNM 23582 
(figs. 25, 27). This is an encrusting species 
whose surface is hard, almost stony, and is 
divided by cracks and ridges into roughly 
polygonal areas. It was common at Station 


56 Tulane Studies in Zoology 


Volk att 


7 


10, depth 3 meters, encrusting on limestone 
in patches up to 10 cm in diameter and 3 
mm thick. It was collected August 13, 1957. 


Color in life was orange, while in alcohol 
it was brown. The surface was relatively 
smooth, but broken up by the above-men- 
tioned polygonal plates, about 12 x 20 mm 
in size. The cracks along the raised ridges 
which separated the polygonal areas were 
about 1 mm wide and were assumed to con- 
tain both oscules and pores. 


The ectosome consisted of a dense, stony 
cortex of microscleres firmly bound together 
by fibrous or protoplasmic structures. The 
endosome, what little there was, contained 
tracts of megascleres which penetrated the 
base of the cracks, thinned out, and finally 
lined the wall of the lumen which was 
formed by the edges of the crack coming 
together at its apex. These tightly packed 
megasclere tracts were about 350 p» in diam- 
eter in the endosome and thinned to about 
125 » across the narrow neck of the crack, 
before it widened in the ridge area to form 
the lumen. After following the walls of 
the lumen, the thin megasclere tracts came 
together in the opening of the crack to form 
a thick mass of megascleres and finely micro- 
spined spirasters which looked as if they 
were too large for the opening and thus 
folded back into the lumen, forming another 
smaller lumen within the spicule mass. The 
diameter of this mass in the crack mouth 
was 285 to 385 p. Sections look similar to 
Vosmaer’s figure (1902: Pl. II, fig. 5). 


The megascleres were tylostyles averaging 
729 x 10 p (range: 359 to 910) pein 
main, cortex-building microsclere was a sel- 
enaster, mean size 57 x 68 p (range in diam- 
eter: 53 to 76 w). Selenasters look similar 
to sterrasters but arise in a different fashion. 
Juvenile selenasters appear as two basic 
types of spirasters: the first is finely micro- 
spined and varies in shape from a kidney 
bean to a short and often bent microstrong- 
yle, size range 7-10-14 p; the second is a 
somewhat larger, highly contorted spiraster 
with long, 4 to 9 p, often dichotomous 
spines. Its overall measurements were 15- 
20-24 x 13-16-22 » while the shaft was be- 
tween 4 and 7 p» in diameter. In addition, 
there were all sizes of immature selenasters, 
some with long sharp spines and older ones 
with the spines shorter and seemingly more 
closely packed. 


No. 2 


De Laubenfels (1936a: 153) reported P. 
melobesotdes Gray from the Tortugas but 
since his specimen showed only a few tylo- 
style fragments and mature selenasters, his 
identification was only tentative. Possibly 
he actually found a specimen of this same 
species though this is impossible to tell from 
his slide. 

P. carinata heretofore has been reported 
only from the Indo-Pacific (see Vosmaer 
and Vernhout, 1902) and from Madagascar 
(Levi, 1956b). In view of this distribution 
possibly we might be dealing with a new 
species. However, because of close agree- 
ment with descriptions of previous workers, 
this sponge is tentatively placed here. 


Family CLIONIDAE Gray 


Cliona celata Grant, 1826—OI 999, 
USNM 23567. This is the common yellow 
boring sponge of the region, which outgrows 
its burrows to form tall, massive, cylindrical, 
papillate chimneys. Its galleries are 1 to 4 
mm in diameter. Its papillae are the same 
diameter and | to 4 mm in height. Its skele- 
ton consists entirely of tylostyles which, in 
this case, were 285-372-399 x 5-7-9 up. 

Early authors sometimes mentioned spir- 
asters in Connection with young specimens, 
but Old (1941) found none and redescribed 
the species with the megascleres only. De 
Laubenfels (personal communication) field- 
identified some of these specimens as célata 
rather than C. cartbboea Carter. He also de- 
scribed both species for field identification 
(1953). 

The species was found abundantly through- 
out Alligator Harbor and vicinity and was 
taken specifically at Stations 1, 2 and 3. 
Depth in no case was over 2 meters. 

Cliona cartbboea Carter, 1882—This spe- 
cies was not found during the present in- 
vestigation but was taken at Station 21 on 
October 27, 1948, and reported by de Lau- 
benfels (1953a). 

It is much like celata and may be mistaken 
for it. The main distinguishing character 
is burrow or papilla size, which is on the 
average much coarser. The burrows and 
papillae reach a diameter and height of 4 
to 6 mm, or about 2 mm larger than those 
of celata (de Laubenfels, 1936a, 1953b). 
The spicule size is about the same, though 
caribboea is reputed to have slightly thinner 
tylostyles (de Laubenfels, 1950a). 


Cliona truitti Old, 1941—OI 1024, 


Little: The Sponge Fauna d7 


USNM 23568. This boring sponge was col- 
lected from oyster reefs in Alligator Harbor, 
Station 3, by Dr. R. Winston Menzel. It 
was boring small holes and galleries 1 to 2 
mm in diameter in the shell of Crassostrea 
virginica Gmelin at an intertidal depth. 


The skeleton was composed of tylostyles 
142-180-212 x 4 yp, finely spined microxeas 
68-90-109 x 2-4-5 p, and finely spined spir- 
asters 7-11.2-14 x 2-2.4-4 w which often 
were distinctly angulated one to three times, 
but sometimes straight. 

Cliona vastifica Hancock, 1849 (?).—OI 
1025, USNM 235069. This ts a boring sponge 
found on an unidentifiable beachworn frag- 
ment of shell at Station 2, Bay Mouth Bar at 
Alligator Harbor, during October, 1956. It 
was found by Dr. R. Winston Menzel and 
Mr. R. T. Damian. Depth was intertidal. 

Papillae protruding from the galleries 
were 0.5 to 1 mm in diameter and less than 
1 mm high. 

The megascleres were tylostyles 153- 
245.5-278 x 3-4.8-7 p. The microscleres were 
composed of finely spined or smooth mi- 
croxeas and spirasters. The microxeas were 
64-84.2-115 x 2-2.6-4 » while the spirasters 
were 11-24.2-29 x 2-2.4-3.3 mw (shaft diam- 
eter) and were distinctly angulated one to 
four times. 

Although the tylostyle and microxea di- 
mensions neatly fit Old’s (1941: 11) tabu- 
lation, the spirasters seem slightly too large. 
In addition, the specimens I have examined 
of vastifica had fairly fine microspines on 
the spirasters, but in this specimen the 
microspines on the spirasters were so coarse 
that they almost doubled the diameter of the 
spiraster at points where they occurred. For 
this reason this sponge is only tentatively 
placed here. 


Cliona lampa de Laubenfels, 1950.—OI 
1023, USNM 23591. This is a boring sponge 
which permeates the substrate instead of 
excavating galleries as other boring sponges 
do. The color is a distinctive brick red. 


The species was reported by de Laubenfels 
(1953a) from Station 22 and was found also 
during the course of the present investiga- 
tion at Station 1, August 4, 1957, by the 
author. It was identified immediately in 
the field by the late Dr. M. W. de Laubenfels 
who was accompanying the expedition. Its 
host in this case was a large coral, Siderastrea 
siderea (Ellis & Solander), which looked 


58 Tulane Studies in Zoology 


normal except that it was red to a depth 
of about 0.5 cm instead of being its usual 
yellow-brown color. 


The skeleton consisted of tylostyles 153- 
210.7-240 x 2-3.9-6 » with heads 4-5.8-7 p 
in diameter, and abundant microscleres of 
two categories. There were finely micro- 
spined microxeas 77-92.0-105 x 2-2.4-3 
and straight streptasters 5-8.1-13 x 1-1.9- 
2.2 pw. These size ranges greatly overlap those 
given by de Laubenfels (1950a: 110) in his 
original description. They are, however, 
somewhat larger than his. In view of the 
permeating habitus, straight spirasters, and 
de Laubenfels’ previous record in the area, 
the differences are not sufficient to justify 
designation as a subspecies. 


The skeleton of this species is similar to 
that of C. vastifica and might be mistaken 
for it. The primary differences are the per- 
meating, as against excavating, habitus and 
the straight streptasters of this species. The 
spirasters of vastrfica are distinctly angulated. 

Cliona viridis (Schmidt, 1862) Gray, 1867. 
—OI 1029, USNM 23593 (fig. 28). This 
is a light yellowish-brown, thickly encrusting 
to amorphous sponge, soft in texture, whose 
base dimensions were 3 x 2 cm and height 
was | cm. It held much shell detritus, espe- 
cially near the base, which may have been 
there as substrate. The spiculation consisted 
exclusively of tylostyles 210-296-440 x 1-5- 
7 » whose heads averaged 8 p (range: 4 to 
11 » diameter). There were no microscleres. 


The specimen was found in Alligator Har- 
bor in the vicinity of Station 3, depth 1 
meter, on March 4, 1957, by Mr. Raymond 
i, Damyran: 

This sponge was described as Swberites 
undulatus by George and Wilson (1919: 
140) from Beaufort, N. C. It was placed 
incertae sedis by de Laubenfels (1947: 34), 
primarily because no specimens could be 
found during his stay at Beaufort. He noted 
also that dead specimens of Microciona pro- 
lifera that have lost their flesh spicules 
would come to a residual spiculation ap- 
proaching that of S. wndulatus. 

The most striking, and only important dif- 
ference so far noted, is that of color. George 
and Wilson’s specimen was light grey while 
this one is a light yellowish-brown. This dif- 
ference is not considered significant and the 
specimen was identified originally as S. wn- 
dulatus. 


Vol. 11 


The finding of George and Wilson's orig- 
inal type specimen of S. undulatus by Wells 
et al. (1960: 232) and their subsequent 
synonymizing it with Cliona viridis necessi- 
tates placing my specimen here. 


ORDER EPIPOLASIDA Pallas 
Family TETHYIDAE Gray 


Tethya aurantia (Pallas, 1766) Topsent, 
1900.—OI 1020, USNM 23577. This was 
an orange, subspherical, tuberculate sponge 
4 to 6 cm in diameter and the same in 
height. The endosome was olive-drab in 
color. 

The megascleres were strongyloxeas and 
small styles, some of which showed the sub- 
tylostylote modification. The strongyloxeas 
in the ectosome averaged smaller than those 
of the endosome, though their range was 
greater. The styles were confined to the 
ectosome. 

The mean size of the ectosomal strongyl- 
oxeas was 1221 x 21 mw (range: 765 to 1828 
yw), while the endosomal ones averaged 1326 
x 25 w (range: 1010 to 1775 p), though a 
few immature strongyloxeas were found 
here averaging 553 x 7 pw. The styles were 
144-253.4-452 x 4-7.9-11 up. 

The microscleres were composed of spher- 
asters, 32-76-107 » in diameter, microspined 
chiasters 9-13-18 », and microspined oxyas- 
ters 12-17-25 p. 

This species was abundant throughout the 
year at Station 10, depth 2.5 meters, on a 
limestone substrate. The species also was 
reported from Station 1 but this station is 
not now represented by a specimen. 

The specimens are placed here because of 
the small styles which characterize the ecto- 
some. Only one other species, T. extensa 
Hentschel, from Australia, has styles of the 
same order of magnitude as this species, but 
the spicules of extensa are fully 200 p 
longer than those of the Apalachee Bay 
specimens. 


ORDER CHORISTIDA Sollas 
Family ANCORINIDAE Gray 


Unimia trisphaera de Laubenfels, 1953.— 
This spherical, dark mahogany sponge was 
not found during the course of the present 
investigation. It was reported from Station 
20 only where it was taken October 26, 1948. 
It was described by de Laubenfels (1953a: 
546). 

Stelletta grubit Schmidt, 1862—OI 1004, 


No. 2 


USNM 23575, USNM 23603. This is the 
“oyster sponge” which plays host to, and 
probably protects from fouling organisms, 
the sponge oyster Ostrea permollis Sowerby. 
This relationship first was recorded by the 
author (1958). This sponge is mistaken 
easily for Geodia gtbberosa (Lamarck) which 
was erroneously reported to play host to the 
oyster (Menzel, 1957). It is easily distin- 
guished from Geodia by its lack of ster- 
rasters in the cortex. All oyster sponges col- 
lected to date have been this species. It is 
common throughout the year at Stations 1, 
2, and 4, and was collected previously at 
Station 20 (de Laubenfels, 1953a). Depth 
range is 1 to 13 meters. 

The skeleton of this species contains very 
long oxeas, plagiotriaenes and anatriaenes 
as well as a microsclere population contain- 
ing finely microspined oxyeuasters plus 
eutylasters which are finely microspined at 
the tips of the tylote-ended rays. The mega- 
scleres of the ectosome were a little longer 
than those of the endosome while the micro- 
scleres of the ectosome were smaller than 
those of the endosome. 

In the ectosome the long oxeas were 
1100-1317-1560 x 12-27.6-36 pw. The pla- 
giotrizenes were 815-984-1170 x 10-23.2- 
38 » with clads 48-79.5-105 » long. Only 
one anatriaene 1416 » long was found, while 
the shaft diameter on broken individuals 
was 10-13.2-19 w and the clad length was 
38-59.4-86 p. The fairly rare oxyeuasters 
were 4-11.5-15 , while the eutylasters were 
GERI 

In the endosome the oxeas were 885-1258- 
1450 x 9-18.1-38 » with plagiotriaenes 565- 
906-1100 x 9-15.4-34 p, clads 29-60.4-96 p, 
and anatriaenes 675-1062-1415 x 10-14.5- 
29 pw with clads 19-45-86 w long. The 
oxyeuasters were 7-12.9-22 » and the euty- 
lasters were 11-16.3-22 in diameter. 

This species was redescribed in detail by 
de Laubenfels (1950a) from Bermuda. 


Family GEODIIDAE Gray 


Geodia gibberosa Lamarck, 1815.—\OI 
1015, USNM 23564. This massive to lobate, 
basically dirty-white sponge was abundant 
at Stations 4, 7, and 11 throughout the 
spring and fall. It is often a mass of knobby 
fist-like projections up to 50 cm in diameter 
with individual knobby projections measur- 
ing up to 10 x 5 x 20 cm high. The pro- 
jections are often packed closely together 


Little: The Sponge Fauna 59 


and the effect is somewhat like a bushel of 
potatoes, closely packed, all standing on end. 

The megascleres consisted of huge oxeas 
well over 1.5 mm, most of which were 
broken; orthotriaenes or plagiotriaenes with 
rhabds over 1 mm long but also broken, 
with clads 23-46 p» long; protriaenes all over 
1 mm but broken, with clads 130-230 p 
long; and, in one specimen, tylostyles 107- 
162-290 x 1-3-4 p, and styles of mean size 
225 x 5 w (range: 205-246 p). 

The microscleres were composed of ster- 
rasters (mean diameter: 62 x 60 p; range 
53-69 »); oxyeuasters 12-18-21 pw; and oxy- 
asters 4-6-9 w in diameter. 

This species was collected also at Station 
20, October 26, 1948, and reported by de 
Laubenfels (1953a: 551). The species was 
redescribed in detail by de Laubenfels 
(193G6a, 1950a). 

We do not know at present whether the 
sponge oyster invades this species, as prev- 
iously reported. Extensive collections and 
experiments are being carried on at the 
Oceanographic Institute, Florida State Uni- 
versity, by Mr. Milton Forbes to determine 
the exact relationship between oyster and 
sponge, and the degree of specificity which 
governs it. As previously reported in this 
paper, Ostrea permollis has so far been found 
only in the sponge Stelletta grubu in this 
area. A few specimens of the sponge with- 
out the oyster also have been taken. 


Family CRANIELLIDAE de Laubenfels 

Cinachyra alloclada Uliczka, 1929.—OI 
1048, USNM 23597. The shape of this spe- 
cies is recorded as sub-spherical and _ its 
color is recorded as yellow (de Laubenfels, 
1936a). 

One specimen was taken at Station 13 
from sand and mud bottom, March 10, 1957, 
by N. Hulings and A. N. Sastry. 

The shape of this specimen was like that 
of a child’s top, with a convex upper surface. 
It appeared to be a piece ripped out of a 
spherical sponge, including the rounded sur- 
face and part of the endosome which tapered 
to a point, showing the radiate structure of 
the sponge nicely. Size was 3.5 cm in diam- 
eter and 3 cm in height (z.e., radis to apex 
of pointed part of endosome). Color was 
yellow-brown; it faded somewhat in alcohol. 

The convex upper (outer) surface was 
covered with numerous large pits 1-3 mm in 
diameter and 4 mm or more in depth. 


60 Tulane Studies in Zoology 


The skeleton consisted of huge oxeas, sinu- 
ous broken-up anatriaenes, and protriaenes 
of about the same length as the oxeas. There 
were two sizes of oxeas found; the first 
averaged 3094 x 28 p (range: 2731-4552 pn), 
and the second had a mean size of 1414 x 
9 w (range: 1155-1683 »). The anatriaenes 
had clad lengths measuring 38-61-99 p» while 
the protriaenes had clad lengths measuring 
64-84-180 py. The microscleres were com- 
posed of sigmaspires with tylote ends 9- 
13-16 p. 

This specimen is placed here since the 
sigmaspires were within the 12-18 pw range 
of the species and because some of the 
triaenes were of the “Kudu” type (de Lau- 
benfels, 1936a: 175). Had the range fallen 
into the 17-20 » category and had the clads 
of the triaenes not been of the “Kudu” type, 
the specimen would then fall to C. cavernosa 
(Lamarck ). 

Possibly 


alloclada is conspecific with 


BUSNM 23555 


4 ) 6 8 9 if Tat 


Figures 25-26. 25 (top). Placospongia ca- 
rinata (Bowerbank) Vosmaer (USNM 
23582). 26 (bottom). Cvraniella crania 


(Muller) Schmidt (USNM 23555). 


Vol. 11 


cavernosa as de Laubenfels suggested (1953a: 
552), but in view of the distinctive spicule 
type which characterizes alloclada, I suggest 
keeping them separate until better evidence 
is available. 

Craniella crania (Muller, 1776) Schmidt, 
1870.—OI 1012, USNM 23555 (figs. 26, 
29). This species tends to be globular, but 
may take other forms in shallow water; my 
specimens are subspherical to semicylindrical 
in shape. Color is usually grey-brown or 
drab; the surface is usually hispid and often 
feels and looks like coarse, heavy felt. The 
structure is usually radiate internally. 


The spiculation consisted of very long 
thin oxeas 444-881-1736 x 8-12-15 p, and 
of anatriaenes and protriaenes in the same 
size range. Most were broken in slide prepa- 
ration. In addition there was a microsclere 
population of sigmoid spirasters 11-13-16 p. 
There were also enormously long, slender 
root spicule anatriaenes at the base of the 
sponge. They were 2 to 6 p in diameter 
and had clads which measured 31-35.0-44 yp. 


This species was found commonly at Sta- 
tions 8 and 10, and one was also dredged 
at Station 14. This gives a depth range in 
this area from less than 1 to 6 meters, and 
a range in substrate from oyster shell to rock, 
and to sand. 


This species was described in detail by 
George and Wilson (1919: 142) as Tez#lla 
laminarts, which was made synonymous with 
C. crania by de Laubenfels (1947: 34-35). 
It was also redescribed under its correct 
name by de Laubenfels (1949b: 25). Wells, 
et al. (1960: 236) restored laminaris to spe- 
cific rank as Craniella laminaris, differen- 
tiated from C. crania principally on the basis 
of growth form, color, and habitat, though 
the sigmoid spirasters were reported as 
slightly smaller than those of C. crania. I 
do not feel that such action is warranted on 
the limited qualitative impressions reported 
by them. More quantitative data are needed. 
For this reason I have taken what I consider 
to be the more conservative course since 
spiraster size and habitat, as indicated by 
the presence of root spicules, might indicate 
the specimen as laminaris, while color and 
shape do not. 


CRANIELLA CINACHYRA (de Lauben- 
fels, 1936) new combination—OIl 1021, 
USNM 23572 (fig. 30). 


LS | 
SOw 


Little: The Sponge Fauna 61 


Figure 27. Spicules of Placospongia carinata (Bowerbank) Vosmaer. 


This specimen was taken off Dog Island 
in 1956, perhaps in the region of the Uni- 
versity of Miami Station 20. It was received 
in dry condition with no ecological data. It 
is an oval cake-shaped mass 10 x 8 cm in 
diameter and 3 cm high. Dry color is 
yellowish-gray. Technical difficulties and 
weather conditions prevented work off Dog 
Island during the course of the investigation 
and this species was not encountered again. 

The megascleres consist of long oxeas, 
some of which were as thin as 4 » and quite 
sinuous, and of prodiaenes and protriaenes 
1.4-2.8 » in diameter which appeared rela- 
tively rare. All the megascleres were over 
1 mm in length; most were broken in the 


Tylostyles of Cliona viridis 
(Schmidt) Gray. 


Figure 28. 


preparation of slides so that it was impos- 
sible to ascertain the total length. The micro- 
scleres are sigmaspires whose ends are slight- 
ly rounded, often tylote. Chord measure- 
ment indicated they were 7-9.4-11 p in size. 

This specimen originally was identified as 
Trachygellius cinachyra by me and rechecked 
after the contention of Wells (personal com- 
munication) that there were triaenes in it 
which had been missed. Subsequent investi- 
gation yielded what are interpreted to be 
pieces of triaenes but only a few had un- 
broken, recognizable clads. The investiga- 
tion of de Laubenfels’ original section slide 
revealed the presence of triaenes, although 
in his spicule slides only fragments of tri- 
aenes could be found. I could not get any 
good data on the size of the anatriaenes in 
de Laubenfels’ section, since they were with- 
in spicule tracts and had to be observed by 
focusing up and down; also, the section it- 
self contains only three or four which can 
be demonstrated. 

The above findings justify the transfer of 
this species to the genus Craniella, The lack 
of any anatriaenes in my slides presents 
some difficulty but the close morphological 
agreement between this specimen and my 
specimens of C. crania, with the exception 
of sigmaspire size, could easily lead one to 


62 Tulane Studies in Zoology Vol. 11 


ROOT 


[ae rae wean ea 
50u 


Figure 29. Spicules of Craniella crania (Muller) Schmidt. 


eS 
5 Oy 


Figure 30. Spicules of Craniella cinachyra (de Laubenfels), new comb. 


No. 2 


place it there. Indeed, possibly this species 
is conspecific with C. crania. This would go 
far toward explaining de Laubenfels’ ap- 
parent error in identifying cimachyra as 
crania (Wells et al., 1960: 233). At present 
however, I feel that the more conservative 
course should be followed until the problem 
is resolved clearly by further work. 


The above discussion raises some question 
as to the finding of Trachygellins cinachyra 
by Wells e¢ al. (1960: 233) off North Caro- 
lina. Those specimens should be reexamined 
for triaenes and their proper systematic 
position determined. 


ORDER CARNOSA Carter 
Family CHONDRILLIDAE Gray 


Chondrilla nucula Schmidt, 1862.—OI 
1050, USNM 23599. This is the “chicken 
liver” sponge. It is flat to lobate in shape 
and does not look or feel like the usual con- 
cept of a sponge, but is smooth, slippery, 
and shiny. It has the consistency of cooked 
egg white or raw liver. The color is usually 
pale drab. It has no spongin skeleton and 
no megascleres; its only spicules are spher- 
asters which in this case averaged about 30 
x 20 » in diameter (range: 14-36 1). These 
spherasters were not round as is shown by 
the two axial measurements. 

It was found on another sponge August 
13, 1957, at Station 10, depth 3 meters, by 
Dr. John Morrill. The species was rede- 
scribed in detail by de Laubenfels (1950a: 


153). 


CLASS CALCISPONGIAE 


ORDER SYCONOSA de Laubenfels 
Family SCYPHIDAE de Laubenfels 


SCYPHA ACANTHOXEA, sp. nov.— 
OI 1035, USNM 23602 (fig. 31). 

The syntypes of this species are designated 
as USNM 23602. 


Locality and abundance.—Relatively abun- 
dant in the Panama City region and was 
found on steel buoys at a depth of less than 
1 meter. 

Shape and size—A small cylinder or ball 
2 to 5 mm in diameter and 1 to 4 mm long. 
The average size was 2 mm in diameter and 
4 mm in length. 

Color —Greenish-white. 

Surface —Hispid. 

Oscules—Single, apical, 0.3 to 0.5 mm 
diameter. --..): 


Little: The Sponge Fauna 63 


a 
NN 
a x 
NS \ 
NS IS 
~ s a = 
~ : 
\ SS. " 
SS <4 
SS ™~S SS 
Ss SS 
ww : 
= 
100y 
Figure 31. Distinctive acanthose oxeas of 


Scypha acanthoxea, sp. nov. Other spicule 
types are not illustrated. 


Flagellate chambers —350 x 70 to 525 x 
250 p. 

Ectosomal anatomy.—No _ specialization 
except for tufts of oxeas perpendicular to 
the surface. They are located in the distal 
ends of the flagellated chambers and give 
the sponge the hispid effect. There was no 
actual ectosome. The distal part of the fla- 
gellated chambers was naked, except for the 
spicule tuft. 


Endosomal anatomy.—Triaxon _ spicules 
were strewn between the flagellate chambers 
forming a skeleton. The rays of some pro- 
jected into the central spongocoel as did the 
rays of the few tetraxons present. The struc- 
ture was typical of the genus. 


Skeleton—Six categories of spicules were 
present. There were three sizes of triaxons, 
some of which had at least one ray that was 
sinuous. Among them were a few tetraxons 
of the same size categories. The three sizes 
were measured on the basis of the length 
of their individual rays. Their ray lengths 
were: 58-144.7-220 p, 58-90.1-115 p, and 
29-44.1-86 p. In addition there were two 
classes of regular oxea, 7,e. the dermal spic- 
ules. They were 527-727.6-1169 x 8.6-9.0- 
9.6 w and 229-333.4-489 x 2-6.4-9.6 p. 

There was one further category of oxea: 
z.e., with one end normal or slightly swollen 
and the other coarsely acanthose for about 
one-fourth the length of the spicule. The 
acanthose part varied from about one-tenth 
the length in long ones to a very short one 
that was virtually completely acanthose. 
These were 345-728.6-2299 x 3-4.3+7) yu. 
They are considered primarily coronal spic- 
ules though two or three have been seen 


64 


sticking into, and even completely through, 
the spongocoel. 

Discusston—tThere do not seem to be any 
other members of the genus that have oxeas 
or axon rays that match these specimens, nor 
are there any other members of the genus 


with such acanthose-ended oxeas. I think 
= 4 
> we > 
oO a WW kK 
ee a eae 
<a > fad a 2S a) 
ud Ee = e) > = 
>< n” ep) = I= = 
‘S) (eS) Oo oO © Oo 
IB == a aS =e ac 
= = KE E pe 
za Fae, FZ Fae Fa FZ 
< < <a < at <x 
O oO Oo oO (&) O 
< § < <a =< a <a 
ae ¢ \ 
| \ 
| H | 
| [> | 
, >¢ { | 
‘ ? | | 
{ Nie | les 
} S ) >) | 2 S 
BI eal , | hss 
| | (° oN } { 7 Ss q 
\ \ \ 7 | i | } ‘ Ss 
PST ip ae y 
isi he dS >] le>{ ) \¢ 
Lu 
55 zi 
Soe) he 
Lu Fae oO = ep) 
< = So = ro) ro) 
uJ > oc ox al — 
>< t- = re) > > 
oO dp) dp) - [= i 
\ | \\ —~ /\ ii ~\ h 
[\ ' 


Ee ee: 


Tulane Studies in Zoolog) 


Vol. 11 


these traits constitute good criteria for desig-' 
nation of a new species. The species is 
named for the peculiar acanthose spicule 
which characterizes it. 

Credit is due to Dr. Willard Hartman of 
the Yale Peabody Museum who first pointed 
out these queer, relatively rare spicules to me. 


oO 
S 
Lon) 
x = = ) 
= Bereee O x 
= E = ca 
wo oc rs 
Ser a af 
Lu Lu Lu — 
ere aa a= 
= = 
no ate 2 On 
<< OO.” 7p) SS! 
| = = y 4 | 
= == < ‘ 
“ d \ 
iS SY \ 
\4 \ 
{ v 
LJ 
uJ LJ 
an 
=) in 
= Lu LJ LJ 
<a Li = z z= 
(dp) = 
oO e = a tad Lu 
= EE eae = 
>= Oo S © oc oc 
_ = 
Oo @ 2 Jeo aae 
= =i Oo oc = oc 
on oO a ro) at oO 
Ar) 


—_ 


I | | 
U a a 


Figure 32. Types of megascleres (after de Laubenfels, 1953b). 


No. 2 


ORDER ASCONOSA de Laubenfels 

Family LEUCOSOLENUDAE Minchin 
Leucosolenia canartensis (Miklucho- 
Maclay, 1868) Dendy and Row, 1913——OI 
1041, USNM 23600. This sponge’s small 
yellow trellis-work of ascon tubes, each 1 
mm in diameter, forms masses up to 3 or 4 
cm in diameter. It is identified readily by 
its lemon yellow color which is retained in 
alcohol, and by its pronounced ascon struc- 


Little: The Sponge Fauna 


65 


ture. It contains only simple triaxon spicules. 

The species was abundant at Station 10, 
depth 3 meters, on October 13, 1957, where 
it was growing on Sargassum. Identification 
was verified by de Laubenfels (personal com- 
munication ). 


This is the only record of the sponge for 
this area though de Laubenfels reported and 
described it from the Tortugas (1936a: 201) 
and Bermuda (1950a: 149). 


i) 
a E 
a a oO Ld oO 
Lu Lu uJ ap) re = 
= = = ac w oc a Ss 
op) dp) (dp) uJ Lu <q £ 
at <x at re rs E = = 
a ar =) ” ” oO a 
oc LJ id <a 2 < Lu 
a ae = =] = oc oc = 
Ee = as CE - OW = ” 
wn op) oO = oO Tp) w” aS} 
INS yas a a 
va ie i So OX » Ss 
} ep Nn Ue 
N oe: \ 72 / ; ; 
iy, a a mw Sh oat 
Z en Sj os <— GP 
S AL a oe Z~, <] ”) IS. > aS y, 
Ga ie eke 
; Wie NR _ < A S) - © / 
ae 2 Se ee | 
: = raen Sy C / 3 c 2 | 
Ses , mle 
Nu as ye yeni oN EOS 
Z < =e oe ae) 
VN Css LY SS 
ep) 
< g 
7 =i 
LJ 
= =. ae n 
© S) O <a 
wo a — a 
WW te = ts a 
— oO — 
re —t = Ss oO i 
BES) x eS) a ” Oo. 
is ie) ac a = = 
cs = < a <t <t 


= 


V 


CD <_> sims 
of. 
ie 
a 


Figure 33. Types of microscleres (after de Laubenfels, 1953b). 


66 


Tulane Studies in Zoology 


V. ARTIFICIAL KEY TO THE SPECIES 
Most spicule types encountered are illus- 


trated in Figures 32 and 33. De Laubenfels 
(1948, 1953b) and Hyman (1940) may be 
referred to for further explanation and illus- 
tration of the terms used in this key. 


ip 


Oo 


—] 


Proper spicules of silica or cal- 


cium carbonate present 19 
Proper spicules absent = oe 
Spongin fibers present 3 
Spongin fibers absent, color 


in life purplish-red, brighter 

than maroon _Halisarea purpura 
Skeleton a network of fibers _ 
Skeleton fibers tree-like; they 

branch but seldom or never 

anastomose. Color dull red, 

shape massive to amorphous, 

triaxon spicules of spongin 
oe 4 Darwinella joyeuxi 
Flagellate peeuibers spherical 

or ovate and small, general- 

ly 50 microns or less in di- 

ameter 
Flagellate chambers sack- 

shaped and large, generally 

over 50 microns in diameter 


Fibers show little evidence of 
stratification or axial spe- 
cialization, often opaque, of- 
ten spongy when dry and 
may soften when returned 
to water ___ 2 ae 

Fibers markedly — stratified, 
very evident axial speciali- 
zation, often with pith; when 
once dry they remain hard 
and brittle permanently _.- 15 


Main fibers not trellised or 

fascicular a= 
Main fibers trellised or fascic- 

ular, persistently brittle once 

dry: sharply set off by the 

presence between the skele- 

tal fibers of filaments akin 

to spongin, 3 to 10 microns 

wide and about 1 mm long; 

sponge has strong sulfur or 

Par lie Od Or eee eee ee eee 12 
Primary fibers “cored” with 

foreign material; secondary 

fibers not so cored . =o ee 
Huge ramifying subdermal cav- 

ities, great emphasis on the 

uncored secondary fibers so 

that the primary (cored) fi- 

bers are rare or wanting _.........11 
Normal structure and flesh in 

sponge; flagellate chambers 

normally placed Serres, 
Peculiar structure in that ‘the 

flagellate chambers are 

scarcely more than _ holes 

punched in thin sheets of 

tissue ¢252/0ee veh ee > Aulena columbia 


~ 


2: 


10. 


11. 


14. 


15. 


16. 


Ae 


Vol at 


Shape subspherical with wide- 
ly distributed oscules each 
on a lobate protrusion; color 
drab to black — Spongia barbara 
Shape massive to cylindrical ________.10 
Shape tends toward being an 
inverted truncated cone; col- 
or drab to black _. Spongia graminea 
Shape tends to be cylindrical to 
vase shape; color white with 
lavender tints _______......__.Spongia sp. 
Long wall-like ridges between 
subdermal canals; consider- 
able areas (over canals) that 
are relatively smooth valley- 
like plains ___ Hippiospongia lachne 
Areas between subdermal can- 
als restricted to small island- 
like areas so that the sur- 
face appears covered by tu- 
bercles ______. Hippiospongia gossypina 
Shape ramose to lobate; color 
brown to brownish white; 
conules 2 to 4 mm apart ae 13 
Vase or cake shape; conules 
4 to 12 mm apart 
Branch ends acute, or pointed 
TIreinia fasciculata 
Branch ends bluntly rounded 
its A ee Ircinia ramosa 
Vase-shaped; reddish or white 
with reddish tinge; conules 


AStors mine pals tae Ircinia campana 
Cake-shaped; grey; conules 6 
too 12) mmraparti = Ircinia strobilina 


Exceedingly ramose; predomi- 
nant color in life dull yellow 
though the upper side may 
be light brown; fibers con- 
tain a central conspicuous 
pith extending their entire 
length 

Ramose to amorphous or lam- 
ellate; color in life dark pur- 
ple (may be covered with 
yellow sheen); fibers may 
appear slightly cored with 
foreign material (they con- 
tain distinctive small cells 
especially in the central pith 
revion): Tanthella ardis 


Color in life dull yellow or 
grey, slowly turning reddish 
or carmine, or drab upon 
death or in alcohol 
_ Verongia longissima 
Color dull yellow with upper 
side light brown, quickly 
turns dark purple on death 
or in. alcoho) = es Verongia sp. 


Both primary and secondary 
fibers cored with foreign 
material 

Primary fibers so cored, but 
secondary fibers clear; shape 
lobate to ramose; color light 
brown to rosy red 
ajo Ve oA eee Euryspongia rosea 


No. 


18. 


19. 


21. 


22. 


23. 


24. 


25. 


26. 


21. 


28. 


29. 


30. 


2 Little: The Sponge Fauna 67 


Color orange to pink-red 
_Dysidea crawshayi 


Color sky blue _ _ Dysidea etheria 


Caleareous spicules present _._________.20 
Siliceous spicules present ___________-21 
Color greenish white; sycon 


with rare acanthoxeas 
Scypha acanthoxvea 
asconoid 
_Leucosolenia canariensis 
Megascleres as well as micro- 
scleres present __.__ eee... 
Only microscleres, spher asters 
present; color white, consis- 
tency cartilaginous; surface 
smooth and shiny like cooked 
ege-white Chondrilla nucula 
Tetraxons 


Color yellow; 


bt 


22 


absent as mega- 
SCLEG CS wee mes Aaa eae eae ee eee ny 23 
Tetraxons present as mega- 
SGlene cies etn. oh: Sele erg ee Deere ale 2 2) 60 
Diactine megascleres present 
Only 2s eee eee 25 
Monactines and/or diactines 
present as megascleres ___ Csi 
Monactines and diactines pres- 
eMitrals; Me MAS Cleves) sae ee 36 
Monactine megascleres present 
qialky; | CS ee es Oe ee eee 42 
Diactine megascleres only, no 
MNLCEOS ClLERCS yes tee ey salle Ee 26 
Diactine megascleres plus mi- 
CROSCLERCSymene te ee ek ees eo 34 
No dermal specialization; re- 
ticulation of endosome mere- 
ly continues to surface 27 
Dermal mesh or reticulation 
smaller, or a definite special 
dermal skeleton present 30 
Skeleton of oxeas only _...- 28 


Skeleton mainly of oxeas, but 

with a strong tendency for 

the larger ones to be stron- 

gylote or stylote _.___ Haliclona sp. 
Sponge light green to grey- 

brown; encrusting to mas- 

sive with digitate processes; 

cxeas about 120 microns long 

and some few may be stron- 

gylote or stylote __ Haliclona viridis 
Sponge lavender to brownish 

AMEN CCIE CTO tareteeeeaes Ne, eee eine 229 
Sponge brownish grey to lav- 

ender; thickly encrusting to 

massive and amorphous; ox- 

ea lergth averaging about 

150 microns; skeleton regu- 

larly reticulate — Haliclona permollis 
Sponge dull red even when 

dry; shape ramose — Haliclona rubens 


Dermal mesh smaller than 

that of the endosome; sponge 

a light brown hollow tube 

with oxeas the only spicules 

Ti ee eee Callyspongia vaginalis 
Definite special dermal skele- « 

(OVO. WEWAECSVO GE ekenpee te een OME 


dl. 


33) 


34. 


Co 
-~] 
. 


38. 


Sponge white or light brown 

fingerlike fistula rising from 

a flat basal mass; fistula 

2-4 mm diameter and 2-4 cm 

high; skeleton consists of 

oxeas only; endosome want- 

ing in the fistula giving it 

the form of a slender hollow 

tube _ _ Rhizochalina oleracea 
Sponge encrusting to massive 

or amorphous though it may 

have slender finger-like pro- 

cesses (less than 2 em high) 

standing erect on its surface ___32 
Sponge white; encrusting with 

a few finger-like processes 

rising from the surface; skel- 

eton of oxeas only, though 

some verge towards stron- 

gyles - Adocia neens 
Sponge light greenish - _ brown 

(cecasionally pink) to yel- 

low, light green or brown _ O68 


Color yellow to light green 
(occasionally pink) to light 
greenish-brown 

Halichondria panicea 

Color dark, ‘brownish 

_ Halichondria melanadocia 


Menaeciores! are tylotes only _ 3b 
Megascleres are oxeas only; 
repent ramose; color yel- 
lowish green to cream; der- 
mal mesh smaller than endo- 
somal; spiculation of oxeas 
about 100 microns long plus 
microxeas and _ ravhides, 
many of which are bent to 
simulate toxas — Callyspongia repens 


Shape a small hollow fistula 
up to 5-7 mm in diameter 
abcut 1 em high; spiculation 
of tylotes, arcuate isochelas 
and sigmas; color white 

ae Coelosphaera fistula 
Shape amorphous with conical 
elevations; spiculaticn of ty- 
lotes; arcuate verging to pal- 
mate isochelas; two sizes of 
sigmas; color bright orange 

ic eee See See Xytopsene sigmatum 
Diactine and monactine mega- 
scleres only, no microscleres; 
color bright red to orange- 
red; shape ramose; dis- 
tinct axial modification in 
branches: spiculation of 

styles and cxeas 


Avinella polycapella 
Diactine and monactine mega- 


scleres plus microscleres == = 37 
Tylotes comprise part of meg- 

asclere population —_ ee to 
Tylotes not found as part of 

the megasclere population. ..______.59 


Color orange to pink-red; shape 
encrusting to massive; spic- 
ulation of tylotes and styles 


68 


39. 


40. 


41. 


42. 


43. 


44, 


45. 


Tulane Studies in Zoology 


(which may be subtylosty- 

lote to a slight extent) plus 

roughened raphides as mi- 

croscleres Tedania ignis 
Exterior color yellow to brown- 

ish-green; interior color yel- 

low; shape permeating to en- 

crusting and sometimes mas- 

sive and amorphous; spicu- 

lation of tylotes and slightly 

bent styles plus sigmas and 

arcuate isochelas as micro- 

scleres Lissodendoryx isodictyalis 
Megascleres comprised of tor- 

notes and acanthostyles, plus 

arcuate isochelas as micro- 

scleres; color fire red in 

life; amorphous 

Merriamium tortugasensis 

No tornotes in the megasclere 

DOULA OTs eee 


Spicules all under 500 microns 
in’ length) —— seen All 
Spicules over 1 mm in n length 
common; spiculation of huge 
strongyloxeas (many over 1 
mm long’), styles about 200- 
250 microns long, plus micro- 
scleres consisting of spher- 
asters, chiasters and oxyas- 
ters; shape subspherical and 
tuberculate; color orange 
ae _Tethya aurantia 


Spieulanonr Fe oxeas, styles, 
and strongyles, plus microx- 
eas or immature oxeas; all 
spicules basically oxeas but 
with wide variation in form; 
exterior color lavender, in- 
terior color brown; shape 
amorphous) 2 Rhaphisia menzeli 

Spiculation of slightly acan- 
those, slightly bent styles and 
strongyles, plus toxas and 
predominantly palmate iso- 
chelas (a few are arcuate) ; 
color bright orange; shape 
encrusting 


..-.. 40 


Monactine megascleres only 
present, no microscleres _. 43 
Monactine megascleres ge 
microscleres present — ead 
Both tylostyles and styles pres- 


ent, with acanthostyles; 
shape thinly encrusting 
Eurypon clavatella 


Tylostyles or styles present, 


but not both “ = ale 
Styles only present 2k 45 
Tylostyles present, with or 

without acanthostyles —__ eG 


Shape flabellate to palmate 
with a special axis; color 
bright orange-red; spicules 
styles about 220 x 10 mi- 
crons in size 


Homaxinella waltonsmithi 


Holoplocamia delaubenfelsi 


46. 


48. 


49. 


Vol. 11 


Shape thickly encrusting to 
massive and amorphous; col- 
or orange-pink; spicules 
styles 130-350 microns long 
and 2-5 microns wide 
Hymeniacidon heliophila 
plus acanthostyles 
present; vase-shape; color 
black _ Thalyseurypon vasiformis 
Tylostyles only present ___.__ 47 
Sponge soft; light yellowish 
brown; shape encrusting to 
amorphous Cliona viridis 
Sponge boring or massive cy- 
lindrical; consistency corky; 
color yellow in life; surface 
papillate _ 


Surface papillae or galleries 1 

to 4 mm in diameter and 

height, > =a Cliona celata 
Surface papillae or galleries 4 

to 6 mm in diameter and 


Tylostyles 


heioht: 2. eee Cliona caribboea 
Plain styles: present ——————————— 50 
Plain stylessabsent a 52 
Plain styles present with or 

Wathout Subtylostyless= =.= 51 
Plain styles plus’ tylostyles 


present, plus spherasters and 
chiasters as microscleres 
Jha bate Halicomites stellata 


Pla styles the only mega- 

scleres, plus raphides, toxas, 

and sigmas as microscleres 
ho 22 tees Toxemna tubulata 


Plaia styles plus subtylostyles 
with very faint heads; mi- 
croscleres palmate anisoche- 
las, toxas and sigmas; shape 
encrusting; color orange to 
orange-red Carmia macilenta 
Subtylostyles with faint heads, 
plus acanthostyles, palmate 
isochelas, and toxas; shape 
encrusting to lumpy and lam- 
ellate; color dull brick red 
to orange-red. Microciona prolifera 
Tylostyles the only megascleres 
present (often there are large 


microxeas, some of mega- 

sclere size) 22.22 eee 53 
Boring sponges; microxeas of- 

ten approaching megasclere 

Size ut ee eee 54 
Non-boring sponges; no mi- 

croxeas present eee 55 


Spirasters distinctly angulat- 

ed oon ence GONE VESTER 
Spirasters not distinctly an- 

gulated but more or less rod 

shaped ioe OD 


Boring galleries in eee ne 
molluses; microxeas mostly 
70-120 ‘microns long, spi- 
rasters sometimes slightly 
aneulate ds eee Cliona truitti 


56. 


57. 


58. 


59. 


60. 


61. 


62. 


63. 


64. 


Substrate of coral or limestone 
not bored but permeated by 
a fine network of sponge 
tissue to a depth of 2-5 mm; 
color brick red; spirasters 
straight and rod shaped 

_Cliona lampa 


Onlyue one ae ue Smeg 
Spirasters, present) 22 eof 
Sterrasters abundant as ar- 
mor, plus spirasters; sur- 
face divided into polygonal 


plates _ Placospongia carinata 
Color reddish brown to orange 

or bright crimson _ ee EDS 
Color grey or light bron aa 59 
Color orange or red-brown; 


shape encrusting 
eee no ee Spirastrella coccinea 
Color bright crimson; shape 

massive and amorphous 

a _Spirastrella coccinopsis 

@oler grey; shape encrusting: 

to amorphous; generally with 

two types of spirasters, one 


normal and the other C- 
shaped — _ Anthosigmella varians 
Color light yellow-brown to 


dark suede; shape massive 

to cake-shaped; consistency 

woody when dry 

- 1 AO Spheciospongia vesparia 
Both tetraxons and large oxe- 

as, v.e. oxeas over 500 mi- 

GOMES) itoy eine oy See ee 61 
Tetraxons without oxeas, te- 

traxons equi-rayed and acan- 

those near ends, subtylo- 

styles also present and long 

(up to 2 mm); thin styles 

but no microscleres Cyamon vickersi 
Tetraxons are calthrops, also 

there are microxeas, oxyeu- 


asters, tylasters, chiasters, 

and peculiar tri -sphaeroid 

streptasters Unimia trisphaera 
Tetraxons are not calthrops 

butpirigdencs! 22 * 2 = Se een ee 62 
Only one type of microsclere 

present —_ Stoel 03) 
More than one type ‘of micro- 

sclere present —_ £165 


Surface with large’ “pits” 4-6 
mm wide and deep; color 
yellow to yellow brown ; 
some triaenes of the “Kudu” 
type; microscleres sigma- 


Splines: = ae Cinachyra alloclada 
Surface without pits; no “Ku- 

due’ type triaenes' 2s G4 
Microscleres sigmoid spiras- 


ters 11-16 microns chord 
length; color grey to brown; 
surface often felted 
ers es Te Craniella crania 
Microscleres sigmoid  spiras- 
ters 7-11 microns chord 
length; dry color yellowish- 
Ray eae ee Craniella cinachyra 


$2 Little: The Sponge Fauna 69 


65. Microscleres are  sterrasters, 
especially in stony cortex, 
and oxyeuasters._____Geodia gibberosa 
Microscleres are eutylasters 
and rare oxyeuasters; no 
sterrasters _Stelletta grubii 


VI. ACKNOWLEDGMENTS 


I am deeply indebted to both Dr. R. Win- 
ston Menzel of Florida State University, who 
introduced me to sponges as an area of re- 
search and directed the general aspects of 
this work as supervising professor, and to 
Dr. M. W. de Laubenfels, who inspired and 
largely directed the detailed aspects of this 
work until his death, February 4, 1958. The 
bequest of Dr. de Laubenfels’ library and 
materials proved invaluable to the study. 
Mrs. Beth Jones de Laubenfels has continued 
her late husband’s interest and encourage- 
ment. 

I am indebted to Drs. Charles B. Metz and 
Donn S. Gorsline, members of my graduate 
committee. Dr. Willard D. Hartman of the 
Peabody Museum of Natural History, Yale 
University, greatly improved the finished 
work with a detailed and critical analysis of 
the data. 

Dr. John Morrill, Raymond T. Damian, 
and Akella Sastry provided helpful sugges- 
tions and extensive collection of specimens. 

My wife, Jane, collected specimens, typed, 
and assisted in all phases of this work. 


VII. REFERENCES CITED 
Burton, M. 1934 Sponges. Brit. Mus. (Nat. 
Hist.) Great Barrier Reef Expedit. 1928- 
29, sci. Repts. V4: 53-614. 
pee 1956 The sponges of 
Atlantide Rept. No. 4: 111- 


West Africa. 
147. 

Carter, H. J. 1882 Some sponges from the 
West Indies and Acapulco in the Liver- 
pool Free Museum described, with general 
and classificatory remarks, Ann. Mag. 
Nat. Hist. (ser. 5) 9: 266-301, 346-368. 


; SV hoe ISSA Gatsionue of ma- 
rine “sponges, collected by Mr. Jos. Will- 
cox, on the west coast of Florida. Proc. 
Acad. Nat. Sct. Phil. 36: 202-209. 


DE LAUBENFELS, M. W. 1932 The marine 
and fresh-water sponges of California. 
Proc. U. S. Nat. Mus. 81: 1-140. 


1934 New sponges from 
the Puerto Rican Deep. Smithson. Mise. 
Coll. 91: 1-28. 

yee ett ne TY Ree A 1936a A discussion of the 
sponge fauna of the Dry Tortugas in par- 
ticular and the West Indies in general, 
with material for a revision of the fami- 


70 Tulane Studies in Zoology 


lies and orders of the Porifera. Carnegie 
Inst. Wash., Publ. 467: 1-225. 

A 1936b A comparison of 
the shallow-water sponges near the Pa- 
cific end of the Panama Canal with those 
at the Caribbean end. Proc. U. S. Nat. 
Mus. 83: 441-466. 


oe 1947 Ecology of the 
sponges of a brackish water eeoment 
at Beaufort, N. C. Ecol. Monogr. ; 31- 


46. 

_.. 1948 The Order Kerato- 
sa of the Phylum Porifera—a monograph- 
ic study. Allan Hancock Found., Publ. 3: 
1-217. 

—_ _... 1949a Sponges of the 
western Bahamas. Amer. Mus. Novitates, 
No. 1431: 1-35. 

= ae 1949b The sponges of 
Woods Hole and adjacent waters. Bull. 
Mus. Comp. Zool. 103: 1-55. 

1950a The Porifera of 

Trans. Zool. 


the Bermuda Archipelago. 
Soc. London 27: 1-154. 

16 SE Peat 1950b An ecological dis- 
cussion of the sponges of Bermuda. Ibid. 
27: 155-201. 


island of Hawaii. 


1951 The sponges of the 
Pacifie Sev. 5: 256-271. 

a _. 19583a Sponges from the 
Gulf of Mexico. Bull. Mar. Sci. Gulf and 
Carib. 2: 511-557. 

ahs _... 1958b A Guide to the 
Sponges of Eastern North America. Univ. 
Miami Press, 32 pp. 

= _and J. F. Storr 1958 The 
taxonomy of American commercial 
sponges. Bull. Mar. Sci. Gulf and Carib. 
8: 99-117. 

DENDY, A. 1890 Observations on the West 
Indian Chalinine sponges, with descrip- 
tions of new species. Trans. Zool. Soc. 
London 12: 349-368. 

ean i. IRM ee 1905 Report on the 
sponges collected by Professor Herdman, 
at Ceylon, in 1902. Herdman, Rept. Pearl 
Oyster Fish. Gulf of Manaar. Publ. Roy. 
Soc. London (suppl.) 18: 57-246. 

pe ree. ae 1921 The tetraxonid 
sponge -spicule: a study in evolution. 
Acta Zool. (Stockholm) 2: 95-152. 

DUERDEN, J. E. 1903 West Indian sponge 
encrusting actinians. Bull. Amer. Mus. 
Nat. Hist. 19: 495-503. 

GrorRGE, W. C. and H. V. WiLson 1919 
Sponges of Beaufort (N. C.) harbor and 


vicinity. Bull. U. S. Bur. Fish. 36: 129- 
179: 
HARTMAN, W. D. 1955 A collection of 


sponges from the west coast of the Yu- 
catan peninsula with descriptions of two 


new species. Bull. Mar. Sci. Gulf and 
Carib. 5: 161-189. 

: 1958 Natural history of 
the marine spenges of southern New 


Vol. 11 

England. Peabody Mus. Nat. Hist., Yale 
Univ., Bull. 12) 1-155. 

Hyatt, A. 1877 Revision of the North 


American Poriferae; with remarks upon 
foreign species, part 2. Mem. Boston Soc. 
Nat. Hist. 2: 481-554. 

HyMAN, L. H. 1940 The Invertebrates. I. 
Protozoa through Ctenophora. McGraw- 
Hill Book Co., 726 pp. 

LENDENFELD, R. VON 1888 Descriptive Cata- 
logue of the Sponges in the Australian 
Museum, Sydney. Australian Museum, 
Sydney (published in London), xvi + 
260 pp. 

Levi, C. 1952 Spongiares de la cote du Sen- 
egal. Bull. Inst. Franc. Afrique Noire. 
14: 34-59. 

Sook el, ae eee 1956a Etude des Halisarca 
de Roscoff. Embryologie et systematique 
des Demosponges. Arch. Zool. Exper. et 
Gen. 93: 1-181. 

ia _... 1956b Spongiares des 
ecotes de ‘Madagasear. Mem. Inst. Set. 
Madagascar, ser. A 10: 1-23. 


LitTLe, F. J. JR. 1958 The sponge fauna of 
the St. George’s Sound, Apalachee Bay, 
and Panama City regions of the Florida 
Gulf coast. (Unpublished M.S. thesis, 
Florida State University.) 

MENZEL, R. W. 1956 Annotated check-list 
of the marine fauna and flora of the St. 
George’s Sound- Apalachee Bay region, 
Florida Gulf coast. Fla. St. Univ., Ocean- 
ogr. Inst. Contrib. No. 61: 1-78. 

obs ae ee 1957 Marine biology of 
Alligator Harbor, Florida. Assoc. South- 
eastern Biol. Bull. 4: 51-54. 

Oxup, M. C. 1941 The taxonomy and distri- 
bution of the boring sponges (Clonidae) 
along the Atlantic coast of North Ameri- 
ca. Chesapeake Biol. Lab., Publ. 44: 1-30. 

PALLAS, P. S. 1766 Hlenchus Zoophytorum. 
Hagae-comitum apud Petrum van Cleef. 

PARKER, G. H. 1910 The reactions of sponges 
with a consideration of the origin of the 
nervous system. Jour. Exp. Zool. 8: 1-41. 


SoLtuas, W. J. 1879 On Plocamia plena, a 
new species of echinonematous sponge. 
Ann. Mag. Nat. Hist. (ser. 5) 4: 44-53. 

TOPSENT, E. 1889 Quelques spongiaires de 
Banc de Campeche et de la Pointe-a-Pitre. 
Mem. Soc. Zool. France 2: 30-52. 

ee 1892 Diagnoses d’eponges 

nouvelles de la Mediterranee et puis par- 

ticulierement de Banyuls. Arch. Zool. 

Exp. 10: xvii-xxviii. 

ae 1901 Spongiaires. Huped. 

Antarct. Belge, Resultats Voyage S. Y. 
Belgica 1897-1899, pp. 1-54. 

1905 Etude sur les Den- 

droceratida. Arch. Zool. Exp. 3: ¢elxxi- 

CXC. 

_. 1920 Spongiaires du Mu- 
see Zoologique de Strasbourg. Monaxo- 


No. 2 


nides. Bull. Inst. Oceanogr. Monaco, No. 
381: 1-36 

i 1928 Spongiaires de 1’At- 
lantique et de la Mediterranee, provenant 
des croisieres de Prince Albert Ier de 
Monaco. Resultats des Camp. Se. Albert 
I de Monaco, vol. 74, pp. 1-376. 

VOSMAER, G. C. J. and J. H. VERNHOUT 1902 
The Porifera of Siboga-Expedition. I. 
The genus Placospongia. IN: Siboga-Ex- 
peditae, 6(a): 1-17. 

WELLS, H. W., M. J. WELLS, and I. E. Gray 
1960 Marine sponges of North Carolina. 
Jour. Elisha Mitchell Sci. Soc., 76: 200- 
245. 


Witson, H. V. 1902 The sponges collected 
in Porto Rico in 1899 by the U. S. Fish. 
Commission Steamer Fish Hawk. Bull. 
U.S. Fish. Comm. 1900, 2: 375-411. 


ee 1911 Development of 


Little: The Sponge Fauna 


ral 


sponges from dissociated tissue cells. Bull. 
WU. SS) Bur: Fish, 30: 1-30 


VIII. ABSTRACT 


A survey of the sponge fauna of the 
Apalachee Bay region of the Florida 
Gulf Coast was made during 1956-57. 
Collections were included from the Pan- 
ama City, Florida, area. Sixty-five spe- 
cies in forty-seven genera were found, 
including seven hitherto undescribed 
species: Callyspongia repens, Coelo- 
sphaera fistula, Eurypon clavatella, Ha- 
lisarca purpura, Holoplocamia delau- 
benfelsi, Rhaphisia menzeli, and Scy- 
pha acanthoxea. In addition, Trachy- 
gellius cinachyra de Laubenfels 1936 
was transferred to the genus Craniella. 
A detailed key to the sponges of the 
area is included. 


TULANE STUDIES IN ZOOLOGY 


Volume 11, Number 3 


October 25, 1963 


CONTENTS 


HORMONAL CONTROL OF THE REFLECTING RETINAL PIGMENT IN THE 
ISOPOD LIGIA OLFERSI BRANDT 


Milton-FingermaniamdiCiitann Oguros. <5 te D> 


EDITORIAL COMMITTEE: 


FRANK A. BROWN, JR., Morrison Professor of Biology, Northwestern University, 
Evanston, Illinois 

TAMETAKE NAGANO, Professor of Tohoku University, The Biological Institute, 
Tohoku University, Sendai, Japan 

KTYOsHI TAKEWAKI, Professor of Zoology and Chairman of Zoology Department, 
Tokyo University, Tokyo, Japan 


HORMONAL AND ENVIRONMENTAL REGULATION OF THE MOLTING 
GYCER IN IHE CRAYFISH FAXONELLA GLYPEATA 


NWwAillitanay CMG benly sity emer br een A hee se ee ee 79 


EDITORIAL COMMITTEE: 


FRANK A. BROWN, JR., Morrison Professor of Biology, Northwestern University, 
Evanston, I[]linois 

MurikL I. SANDEEN (deceased), Associate Professor of Zoology, Duke University, 
Durham, North Carolina 

MILTON FINGERMAN, Professor of Zoology, Newcomb College, Tulane University, 
New Orleans, Louisiana 


HORMONAL CONTROL OF THE REFLECTING RETINAL PIGMENT IN THE 
ISOPOD LIGIA OLFERSI BRANDT! 
MILTON FINGERMAN 
and 
CHITARU OGURO, 
Department of Zoology, Newcomb College, Tulane University, 
New Orleans, Loutsiana, and 
Akkeshi Marine Biological Station, Akkeshi, Japan 


The light-adapted and dark-adapted posi- 
tions of the retinal pigments involved in 
photomechanical adaptation of compound 
eyes to changes in illumination have been 
described for three species of isopods. 
Whether retinal pigments of isopods are in- 
dependent effectors or are under nervous or 
endocrine control has not been determined. 

Peabody (1939) was the first investigator 
to describe migration of retinal pigments in 
isopods. She used Idotea balthica and Idotea 
metallica. In her paper she described only 
distal and proximal retinal pigment cells. 
Kleinholz (1961) stated that the fixative 
she used might have dissolved the reflecting 
pigment. 

Nagano (1949) described the structure of 
the eye in the isopod Ligia exotica including 
the positions of the retinal pigments in light- 
adapted and dark-adapted specimens. Of in- 
terest herein is his diagram showing the 
reflecting pigment proximal to the basement 
membrane in illuminated specimens and dis- 
tal to the basement membrane in specimens 
kept in darkness. The objectives of the pres- 
ent investigation were to determine (1) 
whether the light-adapted and dark-adapted 
positions of the reflecting pigment in the 
eyes of Liga olfers: are the same as in Ligza 
exotica and (2) whether migration of the 
reflecting pigment in L’gza olfersi is under 
hormonal control as in higher crustaceans. 


MATERIALS AND METHODS 


The specimens of Ligia olfersi used in this 
investigation were collected on pilings along 
the south shore of Lake Pontchartrain in 
New Orleans, Louisiana. The authors are 
indebted to Dr. Thomas E. Bowman of the 
United States National Museum for identi- 
fying this species. In the laboratory the Ligia 


1This investigation was supported in 
whole by Public Health Service Research 
Grant B-838 from the National Institute of 
Neurological Diseases and Blindness. 


were kept in covered aquariums containing 
damp pieces of paper. 


To observe the reflecting pigment, his- 
tological sections of the eyes, 10 yp thick, 
were prepared. After the appropriate ex- 
perimental treatment the animals were 
dropped into boiling water for 15-30 sec- 
onds to stop rapidly further migration of 
the pigment. The heads were then removed 
and fixed in Carnoy’s fluid. Bouin’s fluid 
was unsuitable because it dissolved the re- 
flecting pigment. Paraffin sections were 
then prepared. As a measure of the position 
of the reflecting pigment, a reflecting pig- 
ment index was devised. This index was the 
ratio of the mean width of the reflecting 
pigment in both eyes divided by the dis- 
tance from the center of the spherical lens 
to the basement membrane. The latter dis- 
tance is unaffected by light and darkness. 
Use of this ratio minimized the effect of 
size differences among the specimens. The 
distances were measured with the aid of an 
ocular micrometer and reflected illumina- 
tion. The reflecting pigment under such 
illumination appeared silvery-white. The 
center of the lens rather than the cornea 
was chosen as one of the fixed points for 
measurement because the cornea was fre- 
quently torn loose in the histological sec- 
tions. Each unit of the ocular micrometer 
at the magnification used corresponded to 
10.9 pp. Measurements were made of 20 
ommatidia, 10 in each eye. The average 
ratio for both eyes was then calculated and 
represented the index of that particular 
specimen. 

Tissue extracts were prepared in the usual 
manner. The concentrations were two-thirds 
of a sinus gland, z.e. one-third of the isopod’s 
complement, per 0.02 ml physiological saline 
and one-third of the supraesophageal gang- 
lia in the same volume. The dose injected 
into each isopod was 0.02 ml. 

Student’s ¢ test was used for determination 


76 Tulane Studies in Zoology 


of the level of significance between means. 
The 5% level was considered the maximum 
for a significant difference. 


OBSERVATIONS AND RESULTS 
Normal migration of the reflecting retinal 
pigment in Ligta olferst 

The object of the first set of experiments 
was to determine the positions of the re- 
flecting pigment in light-adapted and dark- 
adapted eyes. Specimens were placed both 
in a photographic darkroom and in white 
enameled pans under an incident illumina- 
tion of 120 ft-c for two hours, at the con- 
clusion of which the isopods were killed. 


Inspection of the sectioned eyes revealed 
that the reflecting pigment always remained 
distal to the basement membrane (Fig. 1). 
In dark-adapted eyes this pigment always 
abutted against the basement membrane, 


Se I 


DA 


LA 


Figure 1. Diagrammatic representation of 
ommatidia from dark-adapted (DA) and 
light-adapted (LA) eyes. BM, basement 
membrane; C, cornea; L, lens; RP, reflect- 
ing pigment. 


Vol. 11 


but in about one-half of the light-adapted 
eyes some of the pigment had migrated a 
short distance away, about 10 », from the 
basement membrane. Furthermore, measure- 
ments of eyes from 15 light-adapted and 11 
dark-adapted specimens revealed that the 
mean width of the reflecting pigment in the 
light-adapted eyes was 43.9 » but in the 
dark-adapted eyes was 74.5 p. The differ- 
ence between the means is statistically highly 
significant (p<0.001). The mean distances 
from the center of the spherical lens to the 
basement membrane were 167.3 p» and 
171.1 » for light-adapted and dark-adapted 
eyes respectively. The difference between 
these means is insignificant statistically. 
Consequently, the difference between the 
widths of the reflecting pigment could 
hardly have been due to size difference alone. 
The respective reflecting pigment indexes 
for light-adapted and dark-adapted eyes were 
0.262 and 0.435 (Fig. 2A, B). The differ- 
ence between these means is also highly 
significant statistically (p<0.001). Accord- 


0.5 


INDEX 


PIGMENT 


REFLECTING 


A B Cc D E le G H 


Figure 2. Mean reflecting pigment indexes 
of (A) light-adapted specimens, (B) dark- 
adapted specimens, (C) dark-adapted speci- 
mens that had received physiological saline, 
(D) dark-adapted specimens that had re- 
ceived an extract of sinus glands, (E) dark- 
adapted specimens that had received an ex- 
tract of supraesophageal ganglia, (F) light- 
adapted specimens that had received physio- 
logical saline, (G) light-adapted specimens 
that had received an extract of sinus 
glands, and (H) light-adapted specimens 
that had received an extract of supraesoph- 
ageal ganglia. 


No. 3 


ing to these reflecting pigment indexes the 
reflecting pigment is 1.66 times wider in 
dark-adapted than in light-adapted eyes. 


Effect of extracts of sinus glands and 
supraesophageal ganglia on the 
reflecting pigment 

The object of this experiment was to de- 
termine whether migration of the reflecting 
pigment might be controlled by a principle 
in the sinus glands or supraesophageal gang- 
lia. Extracts were injected into specimens 
maintained in the darkroom and into speci- 
mens in white containers under an illumi- 
nation of 120 ft-c. The injections in the 
darkroom were performed with the aid of a 
dim, red photographic lamp. Forty-five 
minutes after injection of the extracts the 
isopods were sacrificed. The experiments 
with illuminated specimens were performed 
three times, with isopods in the darkroom 
two times. Nine illuminated specimens were 
injected with extract of supraesophageal 
ganglia, 10 illuminated isopods with sinus 
gland extract. Six isopods were injected with 
each extract in the darkroom. Control speci- 
mens received saline alone. 


The extracts had no effect on the isopods 
kept in darkness (Fig. 2C, D, E). However, 
these extracts caused a dark-adaptational re- 
sponse in the light-adapted specimens (Fig. 
2F, G, H). The saline caused a slight but 
statistically insignificant dark-adaptational 
response in the light-adapted controls. How- 
ever, the dark-adaptation shown by the il- 
luminated isopods injected with tissue ex- 
tracts was equal to that of specimens kept 
in the darkroom for two hours. These re- 
sponses were statistically highly significant; 
p<0.001 for the sinus gland extracts and 
p<0.01 for the extracts of supraesophageal 
ganglia. 


DISCUSSION 


The description of the positions of the 
reflecting pigment in light-adapted and 
dark-adapted specimens of Ligia olfersi pre- 
sented above does not agree with the de- 
scription of the same phenomenon presented 
by Nagano (1949) for Ligia exotica. He 
reported that the reflecting pigment lay dis- 
tal to the basement membrane in darkness 
and migrated proximal to the basement 
membrane in light. Herein, however, we 
noted that this pigment always remained 


Fingerman and Oguro: Reflecting Retinal Pigment 1) 


distal to the basement membrane. Elonga- 
tion of the reflecting pigment in dim light 
would render this pigment more capable of 
reflecting light onto the retinula cells, there- 
by increasing the visual efficiency of the 
eye in dim light. 

Kleinholz (1936) has shown that eye- 
stalk extracts will cause light-adaptation of 
the reflecting pigment in the prawn Palae- 
monetes vulgaris. Nagano (1947) found 
the same response in the shrimp Paratya 
compressa. With Ligia olfersi, however, the 
response of the reflecting pigment to ex- 
tracts of sinus glands and supraesophageal 
ganglia was dark-adaptation. These observa- 
tions constitute the first report of (1) a 
reflecting pigment dark-adapting principle 
among crustaceans and (2) a retinal pig- 
ment activator in isopods. Perhaps further 
investigation will show that the principle 
occurs among higher crustaceans as well as 
in the isopod Ligia olfersi. 


SUMMARY AND CONCLUSIONS 


1. The reflecting retinal pigment of the 
isopod Ligia olfersi migrates in response to 
light and darkness. 


2. The reflecting pigment always remains 
distal to the basement membrane. The width 
of the reflecting pigment is greater in dark- 
adapted than in light-adapted eyes. 


5 


3. The sinus gland and supraesophageal 
ganglia contain a principle that causes dark- 
adaptation of the reflecting pigment. 


REFERENCES CITED 


KLEINHOLZ, L. H. 1936. Crustacean eye- 
stalk hormone and retinal pigment migra- 
tion. Biol. Bull., 70: 159-184. 

a ete li oh oe Bo 1961. Pigmentary Effec- 

In: The Physiology of Crustacea. 
Vol. II, Sense Organs, Integration, and 
Behavior, T. H. Waterman, ed. Academic 
Press Ine., New York. 

NAGANO, T. 1947. Physiological studies on 
the pigmentary system of Crustacea. II. 
The pigment migration in the eyes of the 
shrimps. Sci. Repts. Tohoku Univ., Ser. 
IV. (Biol.) 18: 1-16. 

pees ae ee eT 1949. Physiological stud- 
ies on the pigmentary system of Crusta- 
cea. III. The color change of an isopod 
Ligia exotica (Roux). Sci. Repts. Tohoku 
Univ., Ser. IV (Biol.) 18: 167-175. 

PEABODY, E. B. 1939. Pigmentary responses 
in the isopod, Jdothea. J. Exp. Zool., 82: 
47-83. 


78 


ABSTRACT 


The reflecting retinal pigment of the 
isopod Ligia olfersi migrates in re- 
sponse to light and darkness. The pig- 
ment, occupying a wider area in dark- 
adapted than light-adapted eyes, al- 
ways remains distal to the basement 
membrane. Extracts of the sinus glands 


Tulane Studies in Zoology 


and supraesophageal ganglia cause a 
dark-adaptational response of the re- 
flecting pigment. This paper repre- 
sents the first report of endocrine reg- 
ulation of a retinal pigment in isopods. 
The reflecting pigment is involved in 
photomechanical adaptation of the eye 
in response to changes in illumination. 


Vol. 11 


HORMONAL AND ENVIRONMENTAL REGULATION OF THE MOLTING 
CY CLESIN THE CRAYFISH FAXONELLA CLYPEAT A‘ 


WILLIAM C. MOBBERLY, JR., 


Department of Biology, Northeast Louisiana State College, 
Monroe, Louisiana 


I. INTRODUCTION 


The molting process, one of the more in- 
teresting aspects of crustacean physiology, 
has been divided by a number of authors 
(Drach, 1939; Carlisle and Dohrn, 1953; 
Travis, 1955a; and Passano, 1960b) into 
four periods: (1) premolt, a period of ac- 
tive preparation for molt, which includes a 
gradual thinning of the cuticle and storage 
in the gastroliths or hepatopancreas of the 
inorganic constituents needed for hardening 
of the new exoskeleton, (2) molt, the split- 
ting and shedding of the old exoskeleton, 
(3) postmolt, a period of rapid redeposi- 
tion of chitin and inorganic salts to produce 
a new cuticle, and (4) intermolt, a period 
in which the exoskeleton is hard and calci- 
fication is maximal. In the premolt stage 
of astacurans, specifically, gradual resorp- 
tion of the inorganic material in the exo- 
skeleton and deposition of calcium salts in 
the form of gastroliths in the antero-lateral 
walls of the cardiac stomach occur. 


Darby (1938) has stated that operative 
injury appears to hasten the next molt in 
the shrimp Crangon armillatus. R. Smith 
(1940) did not find a shortened intermolt 
period in the crayfish Procambarus clarki 
when an injury, other than eyestalk removal, 
was inflicted early in intermolt. On the 
other hand if the subtropical land crab 
Gecarcinus lateralis lacks many limbs, it 
will molt despite an unfavorable environ- 
ment (Bliss, 1956). She found that when 
six to eight limbs were missing, G. lateralis 
molted on completion of limb regeneration; 
when only one or two limbs were missing, 
only one of nine crabs molted. 

Several investigators have been concerned 
with the relationship of light and tempez- 
ature to the molting process. With G. Jat- 
eralis, Bliss (1954) obtained inhibition of 
premolt regeneration and growth in speci- 
mens maintained under constant illumina- 


1 Part of a dissertation submitted in par- 
tial fulfillment of the requirements for the 
Ph.D. degree in Zoology at Tulane Univer- 
sity, May, 1962. 


tion. She postulated that this effect was 
mediated through the eyes. Stephens (1955) 
found that Orconectes virilis responded to 
daily illumination (20 hours) by an in- 
creased tendency to molt. Hess (1941) 
demonstrated that temperature is a factor 
that influences molting in Crangon armail- 
latus. Specimens did not begin molting 
until the temperature had risen to approxi- 
mately 29° C and when the temperature fell 
below this in the afternoon, molting ceased. 
Light of 75 ft-c had very little if any effect 
on diurnal molting in this shrimp. The in- 
cidence of molting in the lined shore crab 
Pachygrapsus crassipes was also clearly dem- 
onstrated to be associated directly with 
water temperature. Hiatt (1948) found that 
exuvial frequency was highest during the 
summer months and relatively low from 
November to March. 

The molting cycle is under hormonal and 
environmental control. Brown and Cunning- 
ham (1939) found that removal of both 
eyestalks from the crayfish Orconectes im- 
munis caused an acceleration of molting. 
When the contents of eyestalks were im- 
planted into eyestalkless animals, molting 
activity was postponed. These investigators 
concluded that eyestalk tissue liberated a 
humoral substance into the blood which in- 
hibited molting. Abramowitz and Abramo- 
witz (1939, 1940) working with the fiddler 
crab Uca pugilator, R. Smith (1940) with 
the crayfish Procambarus clarki, Kyer (1942) 
with the crayfish Orconectes virtlis, and 
Scudamore (1942) with the crayfish Or- 
conectes immunis, also concluded that the 
eyestalk was a source of molt-inhibiting 
hormone. In the crayfishes Orconectes rusti- 
cus and Orconectes immunis, Stephens 
(1951) found that molt inhibition was ob- 
tained with implants of supraesophageal 
ganglia and circumesophageal connectives. 
Carlisle (1954) using the green crab, Car- 
cinus maenas, suggested that molting was 
partially inhibited during the molting season 
by some factor emanating from a source 
other than the eyestalk. 


80 Tulane Studies in Zoolog 


Scudamore (1947) suggested that the 
sinus gland had a retarding effect on gastro- 
lith formation in the crayfish whereas cen- 
tral nervous tissue extracts (supraesophageal 
ganglia-thoracic ganglia) stimulated gastro- 
lith formation. This observation indicated 
that central nervous organs outside of the 
eyestalk were a source of a molt-accelerating 
substance. In the prawn Palaemon serratus, 
Carlisle (1953) did not find a molt-inhibit- 
ing hormone in the eyestalk. Eyestalk abla- 
tion led to a significant lengthening of the 
intermolt period, which suggested that a 
molt-accelerating hormone was present in 
the eyestalk. Carlisle and Dohrn (1953) 
reported that eyestalk extracts of the shrimp 
Lysmata seticaudata contained a molt-acceler- 
ating factor. 

Gabe (1956) was the first investigator 
to show that Y-organs, paired, bilateral tis- 
sue lying beneath the external adductor 
muscles of the mandibles, were character- 
istic features of malacostracans. He sug- 
gested that the secretory activity of these 
glands was correlated with molting. Car- 
lisle (1957) reported that the immediate 
cause of cessation of molting in the crab 
Mata squinado was degeneration of the Y- 
organ, which secreted a molt-promoting 
hormone. Echalier (1959) found that bi- 
lateral extirpation of Y-organs caused a 
definite blockage of growth and of molting 
in the crab Carcinus maenas, Working with 
the crayfish Orconectes l'mosa, Durand 
(1960) demonstrated histological changes 
in the Y-organ. Activity of the Y-organ 
showed a brief period of activity extending 
from about three days before molt to four 
to five days after molt. 


STATEMENT OF THE PROBLEM 


The present investigation was undertaken 
to learn (1) the normal molting cycle of the 
crayfish Faxonella clypeata, (2) whether 
the molting cycle is influenced by photo- 
period, (3) whether temperature is directly 
associated with incidence of molting, (4) 
whether loss of limbs shortens the intermolt 
period, (5) whether a molt-accelerating fac- 
tor exists in F. clypeata, and (6) whether 
a molt-inhibiting hormone exists in Faxo- 
nella clypeata. 


Il. MATERIALS AND METHODS 


The crayfish Faxonella clypeata (Hay) 
is a small crayfish; adults are approximately 


Vol. 11 


15 mm (11.0-19.1 mm) in cephalothorax 
length. The species inhabits fresh-water 
ponds, ditches, rivers, and swamps in the 
Southeastern United States. Ovigerous fe- 
males retire to burrows. Otherwise members 
of both sexes are active throughout the year 
except when the habitat is dry. 


The experimental animals were collected 
with a dip net near Pearl River, Louisiana. 
The population studied occurred in three 
shallow pine-land roadside ditches that were 
exposed to direct sunlight part of each day 
and contained an abundant plant growth. 
Two ditches paralleled both sides of Louisi- 
ana Highway 41 for approximately 400 yards 
and a third ditch ran for approximately two 
miles alongside a logging road that branched 
off Louisiana Highway 41, thus forming a 
T-shaped collecting area. 


Drought conditions, characterized by com- 
plete absence of standing water anywhere in 
the ditches, and wet conditions, character- 
ized by the presence of standing water 
throughout the ditches, occurred periodically 
during the study period. The condition de- 
pended on the rainfall (U.S. Weather Bu- 
reau climatological data) for the area (Fig. 
1). The depth of water in the ditches fluctu- 
ated from zero to 34 inches, but an inter- 
mediate condition usually existed in which 
the deeper parts of the ditches were wet and 
the shallower parts dry. Collections were 
made only when the ditches contained water. 
Drought conditions occurred during June, 
October and November of 1960. Otherwise 
collections were made at least twice a month. 

Black (1958) suggested that a cephalo- 
thorax length of 11.5 mm was the lower 
limit for sexually mature males and femaies 
of F. clypeata. The data presented herein 
were obtained from animals with a cephalo- 
thorax length of 12.0 mm or longer to elimi- 
nate the possibility that animals undergoing 
pre-maturity molts would influence the de- 
termination of molting peaks for the adult 
population. 

Preliminary experimental observations on 
changes associated with premolt showed that 
gastrolith formation began within 24 hours 
after bilateral eyestalk ablation. The appear- 
ance of gastroliths indicated that the animals 
were undergoing premolt and was used in 
the following experiments as the basis for 
determining whether premolt had begun. 
The absence of gastrolith formation was 


INCHES 
wn N o = 
———$>—_——-—_ > —_  —_—_—_—e—_ 
O 
O 
@) 
O 


Mobberly: Molting Cycle in Crayfish 81 


eee U 


gp eg 
O N Dad 2 WwW ZS ™ 2 A Sy 20) IN 4D Oe M A M 
1959 MONTHS 19 6l 


Figure 1. Annual variation in rainfall at Pearl River, Louisiana, expressed as monthly 
mean number of inches of rain. The solid bars at the base of the figure indicate the pe- 
riod during which 60 per cent or more of the crayfish contained gastroliths. 


considered evidence that premolt had been 
inhibited or had not begun. A soft exo- 
skeleton indicated that the animal had just 
entered the postmolt phase. 

Procedures for statistical analysis of data 
used in this study were obtained from the 


book by Snedecor (1956). 


Ill. EXPERIMENTS AND RESULTS 
Normal Annual Molt Cycle 


To determine the natural molt cycle of 
F., clypeata, collections were made at least 
twice monthly, except for periods of drought, 
for 20 months beginning in October, 1959, 
and ending in May, 1961. A random sample 
of animals from each collection was dis- 
sected to determine the incidence of gastro- 
liths (Table 1). The collections consisted 
of 9,212 animals and of these, 3,426 were 
checked for gastroliths. 

The collection data (Fig. 2) indicated three 
major molting periods: fall-winter (Novem- 
ber-December-January), spring (April-May), 
and summer (July-August). Since molting 


occurred throughout the year, a gastrolith 
incidence of 60 per cent or above was taken 
as indication of a molting peak. On only 
two occasions (September 3 and September 
17, 1961) was the percentage of gastroliths 
below 20. The mean for periods between 
peaks of molting was 35 per cent with a 
range of zero to 56 per cent. 


Effect of Bilateral Eyestalk Ablation 


The precise environmental conditions ex- 
perienced by the crayfish could not be dupli- 
cated in the laboratory. Consequently, meas- 
urement of the time between two molts of 
the same crayfish could not be determined. 
An experiment was designed to determine 
the number of days between bilateral eye- 
stalk ablation and the subsequent shedding 
of the exoskeleton. With the aid of a dis- 
secting microscope, both eyestalks were re- 
moved from crayfish by severing the base 
of the eyestalk with a scalpel. The animals 
were placed in a dry pan for 10 minutes to 
allow the blood to coagulate and then they 
were placed in water. Preliminary experi- 


82 Tulane Studies in Zoology 


Vol. 11 


TABLE 1. 
Incidence of gastroliths in Faxonella clypeata 


With Without Percentage With 
Date Gastroliths Gastroliths Gastroliths 
Octoberwg 195 iaccc-ccseeee soso eee 7 18 28 
INOWieMUD eT (ec. 25 25-2ee ve oes ees 30 24 56 
November 7. 2:----- eens 23 6 79 
November 3c Ose. eee 39 39 50 
December 22a eee 69 43 62 
Sanary: (7, 96 0k eee 14 2 88 
January: 2e-. ee 62 26 70 
SATUS V sO eee eee, 36 14 72 
Hebruanyi6= peu 52 63 45 
February Ui eee 96 148 39 
Marching... eee ee ees 32 74 30 
Marchilibe es ieee es ste es 18 29 38 
Wier CpG ieee ee ae 45 102 31 
Mian cht? eres oe ee il47/ 56 27 
AnreiiGr ack See See Lit 100 52 
PAT PIG Ad See et a be ee ee ET iabal 74 60 
UY ip Le ae oe ase ae aS ee 101 iil 90 
INS 2e lage nde Se eee aR Oe Ss iapterr eee 20 8 fal 
Whey litera Beet see aes pes ee ae, 59 29 67 
ia iiiet eee ea Le od 15 21 42 
Prt bp ay (eee Ace ee ee ne ene ae 16 9 64 
iyo eee ee ee eee ee ee, 16 IL 48 
PAULO; UIS ty Obs ee eee eee ee in ee 78 27 74 
Ani UuSts0 Sees ee 13 3 81 
PAGIY OFULS trp Gee sce eee ae 5 94 
CASIO tcc eet eee sere 48 85 36 
SeHceMM eto. ees ee ee 0 5 0 
Seprenilb etre eee eee 3 26 10 
Decemberalis 2). ss ses ee 13 8 62 
December? Ges ee ee 9 4 69 
Jenene iy MOGI es ee 28 7 80 
MANNA RU) -3e eo oe ne cee 30 2 94 
AMUN ee eee ee eee ee 25 3 29 
VAMTIATN LE Se ee ee ee ce 18 3 86 
TEL TRUE eA a ae ee ewe 62 ilal 85 
Ne Dra Tay: iS eee ee es V2 98 56 
Marchac eae eee: 104 39 Ue 
Manche? beset ae eae eee es 44 144 23 
HN 0) of | Mo Ree care Ae ee eee Cn ee 20 60 25 
ADVIUG2O Sees see ee es 50 30 63 
ay Te ee eee 106 (ie 58 


ments showed that animals cooled at 10° C 
for 15 hours before their eyestalks were re- 
moved had a higher percentage of survival 
than animals that had not been cooled but 
instead had the stubs electrically cauterized 
after eyestalk ablation. Forty-six eyestalk- 
less crayfish were kept for 15 days in aquar- 
iums containing aerated tap water at room 
temperatures of 21-24° C. Postmolt crayfish 
were selected for use in this experiment and 
were not fed during the course of the ob- 
servations. The experiment was repeated 
twice. 

In the first five days of the experiment, 
83 of 138 animals died before molting and 


were not included in the results depicted in 
Fig. 3. By the tenth day all but two molted. 


They died on the 13th and 15th day, re- 
spectively, without having molted. Conse- 
quently the percentage of animals molting 
was 96.4 per cent. The first molt occurred 
on the fifth day after eyestalk ablation and 
the last one occurred on the tenth day with 
a median of 7.9 days for those animals 
molting. 


Influence of Environmental Factors 


Light. To determine the effect of day- 
length on gastrolith production in crayfish, 
four groups of animals were maintained 
under different photoperiods from August 9 
through October 10, 1960. This experiment 
was repeated two times, from January 1 to 
March 18, 1961 and from March 25 to June 


No. 3 


100 


80 


PERCENTAGE 
3 


aN 
(e) 


On NI Di) el Nee Agen NA 


1959 


MONTHS 


Mobberly: Molting Cycle in Crayfish 83 


O 
Jd nt nS} 


JA TA SS OM IN Died. Fs NAb AYE NA 


I96| 


Figure 2. The annual molt cycle of F. clypeata expressed as the percentage of animals 


with gastroliths. 


12, 1961. Animals for the first experiment 
were collected on August 6, 1960; for the 
second experiment on January 1, 1961; and 
for the third experiment on March 25, 1961. 
Males and females were present in each 
group. The crayfish were maintained in cov- 
ered rectangular stainless steel tanks, 49 cm 
long and 37 cm wide, placed side by side 
in an air-conditioned laboratory. The water 


PER CENT MOLTING 


DAYS 


Figure 3. Relationship between the total 
percentage of crayfish that molted and the 
number of days after bilateral eyestalk 
ablation. 


was approximately two inches deep and was 
changed every 15 days. The animals were 
not fed during the course of the experiment. 
Illumination was provided in each tank by 
one frosted 10-watt bulb suspended 20 cm 
above the water surface. The intensity of 
illumination at the surface of the water was 
approximately 40-45 ft-c. Illumination be- 
gan at 6 A.M. and the duration of light was 
controlled by automatic time clocks set to 
provide 6 and 12 hours of illumination daily. 
Other groups were kept in constant darkness 
and in constant illumination. 


All groups started with 90 animals. Group 
I with mortalities of 33, 61, and 24 per cent 
in the three experiments respectively was 
maintained in constant light; Group II with 
mortalities of 38, 62, and 35 per cent was 
exposed to 12 hours of illumination; Group 
III was exposed to six hours of light per day 
and had mortalities of 44, 68, and 25 per 
cent; and Group IV with mortalities of 26, 
61, and 30 per cent was maintained in con- 
stant darkness. A random sample (36, 10, 
and 24 animals) was selected on the begin- 
ning day and a sample (10, 6, and 10 ani- 


84 Tulane Studies in Zoology 


mals) at intervals of 15 days thereafter for 
75 days. The selected animals were removed 
from each tank and sacrificed to determine 
the incidence of gastroliths. Exuviae were 
found when the water was changed but the 
number of molts per tank was not deter- 
mined. 

The results are summarized in Fig. 4. 
Group I showed the least tendency toward 
gastrolith formation. A large peak of gastro- 
lith production was evident in the 15 day 
sample of the January-March animals and a 
small peak in the 45 day sample of the 
March-June animals. 

In Group II gastrolith production peaks 
were noted as follows: 60 day sample in 
August-October animals, 30 and 75 day 
samples in January-March animals, and 30 
and 60 day samples in March-June animals. 

Group III had the greatest tendency to- 
ward gastrolith formation. Peaks were as 


100 


AUG-OCT. 


PER CENT WITH GASTROLITHS 


JAN-MARCH 


Vol. fi 


follows: 30 and 75 day samples in August- 
October animals, 30 and 60 day samples in 
January-March animals, and a 45 day peak 
in the March-June animals. 

In Group IV gastrolith production peaks 
were noted as follows: 30 and 60 day sam- 
ples in the August-October animals, 30 and 
60 day samples in the January-March ani- 
mals, and 30 and 60 day samples in the 
March-June animals. 

The total percentages of gastroliths pro- 
duced by the animals of each group were: 
Group I, 41 per cent; Group II, 60 per cent; 
Group III, 64 per cent; and Group IV, 54 
per cent. The differences in behavior be- 
tween the animals maintained in constant 
light and Groups II, HI, and IV were treated 
statistically using Student’s ¢ test. The means 
of Group I and Group IV were not signifi- 
cantly different whereas Groups II and III 


MARCH -JUNE 


30 45 
DAYS 


Figure 4. Gastrolith production in Faxonella maintained at different daily light periods 
of 40-45 ft-c.: circles (constant illumination), circles with right half filled (12 hours), 
circles with left half filled (6 hours), and dots (constant darkness). A probable 40 day 
intermolt period is indicated by the vertical dashed lines. 


©. 415; 30.45.60. 75 © I5 60 75:0 IS 30° 45 390Gne 


No. 3 


were significantly different (p. 0.05 and 
0.01 respectively ) from Group I. 


Temperature. To determine the effects 
of low temperature on the length of the pre- 
molt stage, seven groups of eyestalkless cray- 
fish were placed alternatively between room 
temperature and a low temperature for vary- 
ing periods of time. Both eyestalks were 
removed from postmolt animals as previous- 
ly described and the eyestalkless crayfish 
were placed in pans. The pans were white 
enamel with a bottom diameter 14.5 cm and 
contained aerated tap water approximately 
1.5 inches deep. The room temperature (20- 
28° C) was maintained by using an air- 
conditioned laboratory and the low temper- 
ature (8-10° C) by placing the animals in 
a refrigerator. Temperatures were measured 
by means of maximum-minimum thermom- 
eters. 


Group I was maintained at room temper- 
ature for 24 hours and then placed in the 
low temperature for 24 hours and then back 
to room temperature for 24 hours. This 
alternation of exposure of animals to room 
temperature and then low temperature was 
continued until all of the animals had either 
molted or died. Group II was alternated 
every 48 hours, Group II every 72 hours, 
Group IV every 96 hours, Group V every 
120 hours, Group VI every 144 hours, and 
Group VII, serving as the low temperature 
control, was maintained at 8-10° C for the 
course of the experiment. Group VIII, serv- 
ing as the room temperature control, was 
the batch used above (Fig. 3) to determine 
the length of time between bilateral eyestalk 
ablation and shedding of exuviae. The ex- 
periment was repeated two times. Animals 
that died during the course of the experi- 
ment were not included in the results. 


Mobberly: Molting Cycle in Crayfish 85 


Table 2 summarizes the results as mean 
number of days at room temperature before 
the animals molted. Groups II and IV had 
the lowest means (7.9 and 7.1 days re- 
spectively) and Groups I and VI had the 
highest means (8.6 and 9.4 days respec- 
tively ). 

The seven experimental groups showed no 
significant differences; constant low tem- 
peratures (8-10° C), however, inhibited the 
process of premolt to such an extent that 
the animals did not molt. Analysis of vari- 
ance by means of the “F” test was used on 
the results. An F value of 0.90 was obtained 
which indicated no significant difference 
among the seven groups. 


Influence of Limb Loss on Gastrolith 
Formation 


To determine the effect of limb loss on 
the incidence of gastroliths, four groups of 
crayfish with different numbers of limbs 
missing were kept for a period of 21 days in 
glass aquariums containing aerated tap water 
approximately two inches deep. Postmolt 
crayfish were used and were not fed during 
the course of the experiment. The aquariums 
were kept in an air-conditioned laboratory 
(26-30° C) under identical light conditions. 


The chelipeds would undergo autospasy 
when pressure was exerted on the merus by 
means of forceps. Walking legs were re- 
moved at the base by clipping with fine 
scissors. The next day 15 animals with the 
proper number of limbs missing were se- 
lected for each group. Group I had one 
cheliped missing, Group II had two cheli- 
peds missing, Group HI had two chelipeds 
and the first two pairs of walking legs miss- 
ing, and Group IV having no appendages 
missing served as the control. The crayfish 


TABLE 2. 
Number of days at room temperature required for Faxonella to molt after bilateral 
eyestalk ablation 


No. of Mean No. of 
No. of Animals Alternate Days in Room 
Groups Animals that Molted Period in Temp. Required 
Low Temp. for Molting 
I 60 22 24 hours 8.6 
II 60 19 48 hours 7.9 
iil 60 18 72 hours 8.2 
IV 60 18 96 hours Foal 
V 60 18 120 hours 8.3 
VI 60 18 144 hours 9.4 
VII 30 0 continuous 0.0 
VIII 138 53 0 hours 7.9 


86 Tulane Studies in Zoology 


were sacrificed at the end of the experiment 
(21 days) and the incidence of gastroliths 
determined. The experiment was repeated 
three times and the results are presented in 
Fig. 5 in which Group I represents 49 ani- 


80 
70 
60 
50 
40 
30 


20 


PER CENT WITH GASTROLITHS 


Figure 5. Percentages of crayfish produc- 
ing gastroliths after appendages were re- 
moved. Crayfish with one cheliped removed 
(vertical line bar), two chelipeds missing 
(crosshatched bar), two chelipeds and two 
pairs of walking legs missing (horizontal 
ane bar), and no appendages missing (open 
ae) 


mals; Group II, 54; Group III, 57; and 
Group IV, 54. 

Group I showed a 39 per cent gastrolith 
incidence, Group II a 45 per cent gastrolith 
incidence, the control group showed a 39 
per cent gastrolith formation, and Group 
Ill, having two chelipeds and two pairs of 
walking legs missing, showed the highest 
gastrolith incidence, 61 per cent. The num- 
ber of actual molts that occurred were: 
Group I, one; Group II, one; Group III, 10; 
and the control, two. 

The above results were treated by a com- 
parison of frequencies using the Chi-square 
test. Group ILI, having two chelipeds and 
two pairs of walking legs missing, showed 
a statistically significant difference when 
compared to the control group (p. 0.01). 
Group II, having two chelipeds missing, did 
not show a statistically significant difference 


Vol. 11 


when compared to the control group (p. 
0.30). 

In a survey of 753 animals from random 
collections of groups between molting peaks 
(Table 3) the following observations were 
made: 18.98 per cent of the population was 


TABLE 3. 

A survey of 753 animals from collections 
made when the incidence of gastroliths 
was less than 60 per cent, showing the 
percentage with missing appendages or 

those in the process of regenerating 
limbs 


Number Percentage 


One Cheliped 

Missing 
Two Chelipeds 

Missing: 18 
Two Chelipeds and 

One Pair of 

Walking Legs 

Missing 1 
Regeneration of 

One Cheliped 16 
Regeneration of 

Two Chelipeds 2 


Total Number of 
Animals Missing: 
or Regenerating 
Limbs 


100 13.27 


2.39 


0.93 
2.12 
0.27 


143 18.98 


either regenerating or would be regenerat- 
ing limbs, 13.27 per cent had one cheliped 
missing, 2.39 per cent had two chelipeds 
missing, 0.93 per cent had two chelipeds and 
one pair of walking legs missing, 2.12 per 
cent were regenerating one cheliped, and 
0.27 per cent were regenerating two chelt- 
peds. 


Influence of Endocrine Factors 


Molt-accelerating Factor. The following 
experiments were conducted to determine 
if a molt-accelerating factor exists either in 
the eyestalk or in the supraesophageal gang- 
lia and circumesophageal connectives of F. 
clypeata. The experimental animals were 
selected in a postmolt condition and one 
eyestalk was removed from each animal 24 
hours prior to use in the experiment. Finger- 
man and Lowe (1957) have found that the 
chromatophore responses of one-eyed indi- 
viduals were greater than responses of intact 
specimens, presumably because the presence 
of both eyestalks made the crayfish more 
capable of antagonizing injected chromato- 
phorotropins. The eyestalk is a proven 
source of molt-inhibiting hormone in the 


No. 3 


fiddler crab Uca pugilator (Abramowitz and 
Abramowitz 1939, 1940), in the crayfishes 
Procambarus clarkit (R. Smith, 1940), Or- 
conectes virilis (Kyer, 1942), and Orconectes 
immunis (Scudamore, 1942). Preliminary 
experiments indicated that the eyestalk of 
F, clypeata was a source of a molt-inhibit- 
ing hormone and animals with one eyestalk 
did not molt any sooner than intact animals. 
The removal of one eyestalk would presum- 
ably reduce the titer of molt-inhibiting hor- 
mone in the blood and the animal would 
then be less capable of antagonizing injected 
molt-accelerating factor. 


Eyestalks and supraesophageal ganglia 
with the circumesophageal connectives at- 
tached were removed from only those donor 
animals having gastroliths. The assumptions 
were made that those animals having gastro- 
liths were in the premolt condition and if 
an accelerating factor was present it would 
be present in greatest titer during this period 
of the molt cycle. 


In the first experiment 40 eyestalks were 
triturated, suspended in 0.6 ml of van Har- 
reveld’s solution (van Harreveld, 1936) buf- 
fered to pH 4.8 and 0.02 ml (containing 
1.35 eyestalks) of this extract was injected 
into the third abdominal segment of each 
of 20 animals. The pH of 4.8 was chosen 
to determine if an acid solution would acti- 
vate the molt-accelerating factor. All extracts 
of eyestalks used in the experiments were 
centrifuged after trituration to remove the 
bits of exoskeleton and retinal pigments. 
Twenty supraesophageal ganglia with the 
circumesophageal connectives attached were 
likewise triturated, suspended in 0.5 ml of 
buffered van Harreveld’s solution (pH 4.8) 
and 0.02 ml (containing 0.8 supraesopha- 
geal ganglia with the circumesophageal con- 
nectives attached) of this extract was in- 
jected into each of 20 animals. The control 
was composed of 20 animals injected with 
0.02 ml each of buffered van Harreveld’s 
solution (pH 4.8). 


The animals were placed five to a pan 
(previously described) under identical light 
and temperature conditions and were not 
fed during the course of the experiment. 
The animals were injected again five and 
nine days after the original injection. On 
the 15th day the animals were sacrificed to 
determine if gastroliths were present. The 
presence of gastroliths in greater quantity 


Mobberly: Molting Cycle in Crayfish 87 


than in the control would indicate that the 
injections had initiated a premolt condition. 
The experiment was repeated twice. 

The second experiment was conducted in 
the same manner as described above with 
the exception that van Harreveld’s solution 
was not buffered and was used at its normal 
pH of 7.9, this being approximately the pH 
of the crayfish’s blood. 

The third experimental procedure was es- 
sentially the same as described for the above 
experiments with the exception that the van 
Harreveld’s solution was buffered to pH 9.6. 
This pH was chosen to determine if a basic 
solution would activate the molt-accelerating 
factor. 


The results are presented in Fig. 6. At a 
pH of 4.8 the injection of the extract of the 
supraesophageal ganglia with the circum- 


oe 
ra 


wn 
3e 
z 
= 5°F 
(e) 
= 
2 40F 
1o) — 
x 
& 30F 
= ma 
i 
& A10) 
2) 
a 
a 10f 
{eae eae N= 
pH 4.8 pH 7.9 pH 9.6 
Figure 6. Percentages of crayfish that 


produced gastroliths after injections of 
eyestalk extract (horizontal line bar) and 
supraesophageal ganglia with circumesoph- 
ageal connectives attached extract (cross- 
hatched bar). The control group (open 
bar) was injected with buffered van Harre- 
veld’s solution. 


esophageal connectives attached produced a 
gastrolith incidence of 63 per cent, the eye- 
stalk extract 45 per cent, and the control 37 
per cent. In the experiment using a pH of 
7.9, the extract of the supraesophageal gang- 
lia with the circumesophageal connectives 
attached had a 41 per cent gastrolith inci- 
dence, the eyestalk extract 39 per cent, and 
the control 31 per cent. At a pH of 9.6 the 
supraesophageal ganglia with the circum- 
esophageal connectives attached extract pro- 
duced a gastrolith incidence of 4 per cent, 
the eyestalk extract 2 per cent, and the con- 
trol 5 per cent. 


88 Tulane Studies in Zoology 


The above results were treated statistically 
by comparison of frequencies using the Chi- 
square test. Injection of the buffered (pH 
4.8) extract of the supraesophageal ganglia 
with the circumesophageal connectives at- 
tached showed a statistically significant dif- 
ference when compared with the control 
(p. 0.01). There was no statistically sig- 
nificant difference when the results from 
injection of eyestalk extract (pH 4.8) were 
compared with the control. The results from 
injection of extracts of eyestalks and supra- 
esophageal ganglia with the circumesopha- 
geal connectives attached did not show a 
significant difference compared with the 
controls at pH values of 7.9 and 9.6 (p. 
0.30 and p. 0.20 respectively ). 


Molt-inhibiting Factor. A molt-inhibiting 
hormone has been found in the eyestalk 
of several crustaceans (Abramowitz and 
Abramowitz, 1939, 1940; R. Smith, 1940; 
Kyer, 1942; and Scudamore, 1942) and 
some authors (Stephens, 1951; Carlisle, 
1954) have also obtained evidence for an 
inhibiting factor in the supraesophageal 
ganglia with the circumesophageal connec- 
tives attached. The following experiments 
were conducted to determine if a molt- 
inhibiting factor exists in the eyestalk or 
supraesophageal ganglia with the circum- 
esophageal connectives attached of F. cly- 
peda. 


Eyestalks and supraesophageal ganglia 
with the circumesophageal connectives at- 
tached were removed from only those donor 
animals not having gastroliths. The assump- 
tions were made that those animals not hav- 
ing gastroliths were either in the postmolt 
or intermolt condition and if a molt-inhibit- 
ing hormone was present it would be present 
in greatest titer during these periods of the 
molt cycle. 


Eyestalk extracts were prepared as follows: 
80 eyestalks were removed from the cray- 
fish, placed in a mortar and triturated. The 
triturated eyestalks were suspended in 0.6 ml 
of the appropriate buffered van Harreveld’s 
solution and centrifuged to remove the bits 
of exoskeleton and retinal pigments. Three 
buffers were prepared; 0.1 molar sodium 
phosphate and 0.05 molar citric acid at pH 
values of 3.5 and 7.6 and 0.1 molar sodium 
hydroxide and 0.1 molar boric acid to pro- 
duce a pH of 9.4. These three pH values 
were selected to determine: (1) if an acid 


Vol. 11 


solution would activate the molt-inhibiting 
hormone, (2) if this hormone could be de- 
tected at approximately the pH of the cray- 
fish's blood, and (3) if a basic solution 
would activate the molt-inhibiting hormone. 

Postmolt specimens of F. clypeata were 
selected because these animals did not con- 
tain gastroliths. The success of the experi- 
ment would be based on the ability of the 
injected molt-inhibiting hormone to prevent 
gastrolith production when the eyestalks are 
removed. The eyestalks were removed as 
described above. The animals were placed 
20 each into two steel tanks (previously de- 
scribed), kept side by side in an air-condi- 
tioned laboratory. Water, aerated by means 
of an air compressor, was changed every 
other day. The animals were not fed during 
the course of the experiment. 


The extract at pH 3.5 was taken up in a 
1 ml syringe and each of 20 animals was 
injected in the ventral abdominal region 
with 0.02 ml (containing 2.70 eyestalks). 
The control animals were injected with buf- 
fered van Harreveld’s solution (pH 3.5). 
On the sixth and tenth day the animals were 
injected again. The experiment was repeated 
twice. Extracts at pH 7.6 and pH 9.4 were 
prepared and injected in the same manner 
as described above. 


The procedure for preparing extracts of 
the supraesophageal ganglia with the cir- 
cumesophageal connectives attached was the 
same as for eyestalks with three exceptions: 
(1) 40 supraesophageal ganglia with cir- 
cumesophageal connectives attached were 
used instead of the 80 eyestalks, (2) the 
extract was not centrifuged, and (3) each 
0.02 ml contained 1.6 supraesophageal gang- 
lia and circumesophageal connectives. 


The data presented in Fig. 7 are the per- 
centages of animals that contained gastro- 
liths after 12 days other than those animals 
that died in the first 48 hours after injection. 
Animals that died after 48 hours and the live 
animals remaining at the termination of the 
experiment were dissected to determine the 
incidence of gastroliths. The presence of 
gastroliths in smaller quantity than in the 
controls would indicate that the injections 
had prevented a premolt condition. In the 
experiment with eyestalk extracts the re- 
sults were: pH 3.5, 79 per cent, control 92 
per cent; pH 7.6;.57 percent; control 98 per 
cent; and pH 9.4, 70 per cent, control 100 


No. 3 Mobberly: Molting Cycle in Crayfish 89 
” 90 
A= 
i 
5 
© 70 
| oa 
VY) 
© 

50 
IE 
is 
= 
= 30 
Zi 
Ww 
UO 
TIO 
Qa 

pH 3:5 pH 7.6 pH 9.4 pH 3.5 PH 7.6 PH 9.4 
EYESTALK BRAIN 

Figure 7. Inhibition of gastrolith formation in Faxonella as a response to injected ex- 


tracts of the eyestalk (black bar) and supraesophageal ganglia with the circumesophageal 
connectives attached (horizontal line bar). The control group (open bar) was injected 


with buffered van Harreveld’s solution. 


per cent. In the experiment with the supra- 
esophageal ganglia and circumesophageal 
connectives extract the results were: pH 3.5, 
69 per cent, control 100 per cent; pH 7.6, 
53 per cent, control 97 per cent; and at pH 
9.4, 74 per cent, control 98 per cent. 


The above results were treated by a com- 
parison of frequencies using the Chi-square 
test. In the eyestalk and the supraesophageal 
ganglia with circumesophageal connectives, 
the inhibiting factor of these extracts at pH 
values of 3.5, 7.6, and 9.4 showed a statis- 
tically significant difference in preventing 
gastrolith production when compared to the 
control groups (eyestalk extract, pH 3.5, p. 
0.02; remaining extracts p. 0.01). 


IV. DISCUSSION 


The collection data (Fig. 2), based on 
per cent of animals with gastroliths, indi- 
cated three major molting periods; Novem- 
ber-December-January, April-May, and July- 
August. E. Smith (1953) worked with the 
growth rate of F. clypeata from approxi- 
mately the same area. She found that ma- 
turing females as a group increased sig- 
nificantly in size from February to early 


June, mid-July to September, and December 
to January. The male growth pattern was 
similar to that of the females. Smith's data 
are substantiated by the gastrolith data in 
showing that F. clypeata undergoes three 
molts a year. 

Rainfall can influence the molting cycle 
to the extent that animals may be forced 
to burrow if rainfall is scanty. The animals 
do not molt in their burrows but wait until 
they again have ample surface water as was 
especially noticeable during the winter of 
1960-61 when the winter molting peak had 
shifted approximately 36 days from _ the 
period of the previous winter. The amount 
of rainfall during the months of October 
and November, 1960, was so small that no 
surface water collected in ditches, but when 
the water level again rose after the Decem- 
ber rains, the animals began to molt at once 
after coming out of their burrows (Fig. 1). 

R. Smith (1940) found that the intermolt 
period in the crayfish Procambarus clarku 
was shorter (average 8.1 days) in eyestalk- 
less animals than in intact ones (28.9 days). 
Scudamore (1942) determined that the pre- 
molt period in the crayfish Orconectes 1m- 
munis was 16.26 days for eyestalkless ani- 


90 Tulane Studies in Zoology 


mals. Intact animals molted only twice a 
year (in the spring and summer). The re- 
sults of R. Smith and Scudamore showed 
that the intermolt period of eyestalkless ani- 
mals was shorter than the natural intermolt 
period. In the crayfish F. clypeata the pre- 
molt period was 7.9 days after both eyestalks 
were removed (Fig. 3). The collection data 
(Fig. 2) indicated that the intermolt period 
was 60-70 days between the November- 
December-January molting peak and the 
April-May molting peak. The intermolt 
period was 30-40 days between the April- 
May peak and the July-August peak. Be- 
tween the July-August peak and the No- 
vember-December-January peak the inter- 
molt period was 60-70 days. In every case 
the intermolt period was much longer in 
intact animals than in eyestalkless ones. 


The photoperiod experiment (Fig. 4) 
showed that exposure of F, clypeata to con- 
stant light slowed the formation of gastro- 
liths. In reduced photoperiods of 6 and 12 
hours the animals had a larger per cent of 
gastroliths when compared to the animals in 
constant light. As mentioned above, the nor- 
mal intermolt period during warm temper- 
atures appeared to be 30-40 days. When a 
40 day intermolt period was marked off on 
Fig. 4, the gastrolith percentage peaks oc- 
curred approximately at this interval in 
groups II, II, and IV. Group I did not fol- 
low this pattern but seemed to have a longer 
intermolt period. This observation was in 
contrast to the results obtained by Stephens 
(1955) with a northern group of crayfish. 
A light period of 20 hours per day during 
the winter months resulted in an increased 
molting frequency in O. virilis. Stephens 
postulated that the spring molt in these ant- 
mals is the result of the transition from 
darkness or very short daily exposures of 
light to day-lengths of 12 hours or more 
when the animals emerge from their bur- 
rows in the spring. On the other hand, Bliss 
(1956) observed that growth in the crab 
Gecarcinus lateralis was inhibited in constant 
light. Premolt limb regeneration and pre- 
molt uptake and retention of water also 
ceased. Molting did not occur in a constant 
illumination of 100 ft-c. Suko (1958), 
working with histological changes of the 
developing ovaries, found that in the cray- 
fish Procambarus clarki the ovaries are in- 
fluenced by darkness. Secretion of a sub- 


Vol-at 


stance found in the sinus gland and central 
nervous system that controls ovarian de- 
velopment may have a periodicity. When 
the quantity of the secretory substance in- 
creases to a certain level in the sinus gland, 
light may be effective in inhibiting the 
function of the sinus gland. 

Constant light may inhibit the production 
of the molt-acceleration hormone and in re- 
duced light periods of 6 and 12 hours this 
inhibition would be absent. Once a crayfish 
begins premolt and then is placed in con- 
stant light, the animal will continue pre- 
molt but at a slower rate than crayfish at 
light periods of 6 and 12 hours per day. 
When the major molting peaks are plotted 
against day-length (U. S. Weather Bureau 
data) for the New Orleans area (Fig. 8) 
molting occurs at the period of short day- 
lengths and just before and after the longest 
day-lengths. The April-May molting peak 
occurs at a day-length of 13 hours and the 
July-August molting peak at the same length, 
however at a day-length of 14 hours molting 
peaks are not found. 


In the experiments on temperature, only 
its role in determining the length of the 
molting process was considered, not the role 
of temperature as an activator of molting. 
The results (Table 2) suggested that once 
premolt had begun, temperatures of 8-10° C 
inhibited the actual molting as a result of 
decreasing the metabolic rate. Mean num- 
ber of days in room temperature required 
for molting ranged from 7.1 to 8.6 days for 
the seven groups. The molting process was 
inhibited for the time period the crayfish 
were subjected to low temperatures. No 
regression in the molting process occurred 
while the crayfish were chilled. As soon as 
the animals were placed in room temper- 
atures again, the premolt process progressed 
until the time the animals were again ex- 
posed to an unfavorable temperature. 

Passano (1960b) introduced his discus- 
sion on temperature effects on crustacean 
molting with the statement that unlike most 
environmental variables such as light, tem- 
perature can influence both molting itself 
and molt-control processes. A number of 
investigators have shown that low temper- 
atures inhibit or lower the incidence of 
molting and high temperatures increase the 
incidence (Hess, 1941; Kyer, 1942; Hiatt, 
1948; and Passano, 1960a). 


Mobberly: Molting Cycle in Crayfish 91 


ND, JF 7k 


OpeN: oD Jo oF UNM Ag eM ed AS SO M A M 


1959 MONTHS 196] 


Figure 8. Annual variation in photoperiod at New Orleans, Louisiana, 50 miles south- 
west of the collection site, expressed as monthly mean number of hours from sunrise to 
sunset. The solid bars at the base of the figure indicate the period during which 60 per 


cent or more of the crayfish contained gastroliths. 


Diapause in insects is dependent on ex- 
ternal factors (photoperiod and temperature) 
and is followed by molting or hatching. 
Diapause occurs in any stage of the life 
cycle except the adult. In the silk worm, 
Bombyx mori, Muroga (1951, cited by Lees, 
1955) stated that 40 days chilling is re- 
quired before it would molt at room tem- 
perature. Lees (1955) suggested that from 
recent experience with the agrotid moth, 
Diataraxta oleracea, and the red spider mite, 
Metertetrancyhus ulmi, temperature should 
be regarded as a signal stimulus in the same 
sense as photoperiod. Andrewartha (1943, 
1952, cited by Lees, 1955) has shown that 
if eggs of the grasshopper Awstrotceles cru- 
ctata are chilled at 10° C for 60 days they 
hatch normally at an incubation of 25° C 
but if chilled at 6 or 13.5 ° C for the same 
length of time, fewer eggs hatch when in- 
cubated at 25° C. However, normal de- 
velopment occurred when the period was 
extended beyond the 60 days needed for the 
10° C temperature. Williams (1956), work- 
ing with diapause in the cecropia silk worm, 
suggested that low temperature served as a 


catalyst to the brain in restoring the en- 
docrine function. 

In the spiny lobster, Panulirus argus, 
Travis (1955a) postulated that molting fre- 
quency was a consequence of metabolism. 
Broekhuysen (1955) found an incubation 
period of one month was necessary for eggs 
of the crown crab, Hymenosoma orbiculare, 
at 16.5° C but 38-48 days were required at 
12-15° C. A temperature of 10° C blocked 
premolt in the crab Sesarma and when such 
blockage was removed, premolt proceeded 
normally (Jyssum and Passano, 1957). Vern- 
berg (1959) showed in the fiddler crabs 
Uca pugnax and Uca rapax that temperature 
affected metabolism. Passano (1960a) hy- 
pothesized that a metabolic event was 
blocked by temperature in the molting of 
the fiddler crab, Uca pugnax, 

F. clypeata molted throughout the year 
and temperatures were never low enough 
(8-10° C) to block molting completely for 
any extended period of time. Inasmuch as 
the U. S. Weather Bureau does not maintain 
a temperature station at Pearl River, the 
temperatures recorded (Fig. 9) are from 
Slidell, Louisiana, eight miles southwest of 


92 Tulane Studies in Zoology 


100 


90 


80 


DEGREES FAHRENHEIT 


ON wal), ames © Hecke Vana 
1959 


Min Au Mines} 


MONTHS 


Vol. 11 


A/S. (OV viN Dasa 


1961 


M A M 


Figure 9. Annual variation in temperature at Slidell, Louisiana, eight miles southwest of 
collection site, expressed in terms of mean monthly temperatures (circles, high and dots, 
low). The solid bars at the base of the figure indicate the period during which 60 per 


cent or more of the crayfish contained gastroliths. 


the collection area. Major molting peaks oc- 
curred when the temperatures were the 
lowest for the area and when the temper- 
atures were the highest. The molting periods 
in both warm and cool temperatures would 
preclude the conclusion that a chill period 
such as diapausing insects need is associated 
with molting in the crayfish F. clypeata. 
The November-December- January molting 
peak was longer in duration than the peaks 
occurring in the warmer months presumably 
because the cold weather (8-10° C) acted 
as a suppressor. Although a thin film of ice 
formed over the water surface in January, 
1960, and 1961, animals were still present 
in the water. In insects a cold shock may be 
necessary before the animal can complete 
diapause, and thus development, but in the 
crayfish temperature may act merely to de- 
crease the metabolic rate and not neces- 
sarily as an activator of hormonal secretions. 


The results (Fig. 5) with F. clypeata 
showed that drastic limb loss was associated 


with an increase in gastrolith formation. 
A regeneration stimulus was suggested by 
Darby (1938) as the reason for the molt- 
initiating effect of bilateral eyestalk removal. 
However, later workers have shown that 
molting was actually initiated by hormones 
(Brown and Cunningham, 1939; Scuda- 
more, 1942; and Passano, 1953). R. Smith 
(1940), working with the crayfish P. clarki, 
found that an eyestalk injury was the only 
type injury that would produce a molt- 
initiating effect during the early intermolt 
period. In a sample of five animals, Bliss 
(1956) produced molting in G. lateralis by 
removing six to eight limbs, but only one 
out of nine animals molted when she re- 
moved one or two limbs. Passano (1960b), 
however, did not find a molt-inducing stimu- 
lus in Carcinus on loss of six appendages. 


Brown and Cunningham (1939) have 
shown in the crayfish Orconectes immunis, 
that eyestalk tissue, under nervous control, 
liberates a hormonal substance into the blood 


No. 3 


which inhibits molting. In the crayfish 
Cambarus propinquus, Scudamore (1948) 
postulated that one reason egg carrying fe- 
males did not molt was because the presence 
of the eggs on the pleopods sent an impulse 
over nerve-reflex pathways that prolonged 
the molt-inhibiting action of the sinus 
glands. In F. clypeata appendage loss may 
result in nervous system stimulation which 
inhibits the sinus gland from releasing the 
molt-inhibiting hormone, consequently gas- 
troliths may then be formed. 


Explanation of activation of the neuro- 
secretory system by loss of appendages may 
be similar to the thesis used by Fingerman 
and Fitzpatrick (1956) in which they ex- 
plained why the fiddler crab Uca pugilator 
exhibited a sexual difference in coloration. 
In this situation removal of the male’s large 
chela reduced the blood volume to such an 
extent that the titer of darkening hormone 
was concentrated and thus produced darken- 
ing, however in the female, limb loss did 
not have as drastic an effect. Appendage 
loss in F. clypeata may have reduced the 
blood volume to such an extent that the 
titer of the molt-accelerating hormone was 
concentrated and thus caused  gastrolith 
formation. 


In F. clypeata it seems probable that a 
nervous mechanism is involved in the in- 
creased gastrolith formation following drastic 
loss of limbs. The hormonal mechanism de- 
scribed above requires a serious reduction 
in blood volume and the loss of two cheli- 
peds and two pairs of walking legs would 
not reduce the blood volume enough. In Uca 
the large claw was approximately one-third 
of the body volume. If the production of 
gastroliths was due to a nervous impulse. 
then it would seem likely that the loss of 
two chelipeds did not produce a sufficient 
nerve stimulus to initiate gastrolith forma- 
tion, whereas the loss of two chelipeds and 
two pairs of walking legs did produce a 
sufficient stimulus. 


The loss of appendages may be one of the 
factors influencing animals to molt between 
molting peaks. In collection samples from 
periods between molting peaks, 18.98 per 
cent of the population contained animals 
either missing appendages completely or re- 
generating them (Table 3). Although in- 
tact specimens with gastroliths were also 
found at the same time, limb regeneration 


Mobberly: Molting Cycle in Crayfish 93 


may explain why some animals were molting 
at times other than during the peak periods 
when the majority of the crayfish molted. 


The results from the experiments on the 
molt-accelerating factors are presented in 
Fig. 6. In the experiment using van Harre- 
veld’s solution buffered to pH 4.8, the re- 
sults indicated that a gastrolith producing 
factor existed in the extracts of the supra- 
esophageal ganglia with the circumesopha- 
geal connectives attached. Scudamore (1947) 
suggested the possibility of nervous or secre- 
tory factors in or near the central nervous 
system having a molt-accelerating action. 
When he injected extracts of central nervous 
tissue into eyestalkless O. immunis and O. 
virilis, stimulation of gastrolith formation 
and an increase in the rate of oxygen con- 
sumption were noted. Carlisle (1953) stated 
that evidence points to the existence of an 
eyestalk molt-accelerating hormone in the 
shrimp Palaemon serratus. In 1953, Car- 
lisle and Dohrn published a paper on the 
shrimp Lysmata seticaudata in which they 
intected intramuscularly an acidified extract 
of eyestalk tissue and found an accelerated 
rate of molting. 


The results of the injection of extracts of 
eyestalks and of the supraesophageal ganglia 
and circumesophageal connectives show that 
a molt-inhibiting hormone exists in these 
structures (Fig. 7). 


The inhibiting hormone may actually be 
a series of hormones instead of one hormone. 
Kyer (1942) suggested that in the crayfish 
O. virilis the inhibiting hormone inhibited 
an enzyme system involved in the removal 
from the exoskeleton of calcium and_ its 
deposition as gastroliths. Scudamore (1947) 
has shown that implants of sinus glands into 
evestalkless O. tmmunis and O. virilis in- 
hibited increase in water content and sinus 
gland extracts decreased the rate of oxygen 
consumption after eyestalk ablation. Travis 
(1951a, 1951b*) has shown that during pre- 
molt the shrimp Panulirus argus took in cal- 
cium from its environment, reduced calcium 
excretion to maintain a normal blood cal- 
cium balance, and that the eyestalk played a 
role in the regulation of phosphate metabo- 
lism. Blood proteins of P. argus increased 
prior to molt, declined following molt and 
reached a subnormal value by the third day. 
Below normal values remained throughout 
most of the 14 day postmolt observation 


94 Tulane Studies in Zoology 


(Travis, 1955b). Durand (1956) stated 
that there are four cytologically distinct 
types of neurosecretory cells in the eyestalk 
and brain of Orconectes virilis. The Type 2 
neurosecretory cells are the only neuro- 
secretory cells that undergo histologically 
demonstrable changes in secretory activity in 
relation to the molting cycle. 

Further experiments are necessary before 
one can make a positive statement that the 
inhibiting factors originating in the eyestalk 
and supraesophageal ganglia and the circum- 
esophageal connectives are the same or dif- 
fer physiologically. Inhibition of gastrolith 
formation is noted at pH values of 3.5, 7.6, 
and 9.4, thus showing that the molt-inhibit- 
ing hormone can be active over a wide 
range of pH values. This may indicate that 
a number of hormones are involved. The 
possibility also exists that these factors are 
the same substance but stored in the sinus 
gland and produced in the supraesophageal 
ganglia and the circumesophageal connec- 
tives. Another explanation could be that 
there are two different sources for the same 
hormone, similar to the situation in which 
the mammalian adrenal cortex produces hor- 
mones similar in structure and action to sex 
hormones, both male and female. 


V. SUMMARY 


1. The molting cycle of the crayfish F. 
clypeata is defined. While animals can be 
found with gastroliths throughout the year, 
three major peaks of molting, November- 
December-January, April-May, and July- 
August. occur. 

2. F. clvbeata molts, on the average, 7.9 

davs after bilateral evestalk ablation. 
3. Exposure of F. clypeata to constant 
light reduces gastrolith production, whereas 
reduced photoperiods of 6 and 12 hours 
increase gastrolith production. 

4. In eyestalkless crayfish subiected to a 
temperature of 8-10° C, the molting process 
is blocked but the process resumes on ex- 
posure to temperatures of 20-28° C. 

5. Limb loss of at least two chelipeds and 
two pairs of walking legs causes an increase 
in eastrolith production. 

6. An acidified extract (pH 4.8) of the 
supraesophageal ganglia with the circum- 
esophageal connectives attached accelerates 
gastrolith formation when injected intra- 
muscularly into F. clypeata. 

7. A molt-inhibiting factor occurs in the 


Voli 


eyestalks and the supraesophageal ganglia 
with the circumesophageal connectives at- 
tached. 


VI. ACKNOWLEDGEMENTS 


The author expresses his appreciation to 
Dr. Milton Fingerman, Committee Chair- 
man, for inspiration and advice. Sincere 
appreciation is also extended to the other 
members of the committee, Drs. D. Eugene 
Copeland, Norman C. Negus, and Stuart S. 
Bamforth. Gratitude is also expressed to 
Dr. Charles F. Lytle for his helpful sugges- 
tions concerning the statistical analysis of 
the data and to Dr. R. Nagabhushanam for 
his assistance on several field trips. 


VII. REFERENCES CITED 


ABRAMOWITZ, R. Z., and A. A. ABRAMOWITZ 
1939. Moulting and viability after re- 
moval of the eyestalks in Uca pugilator. 
Biol. Bull., 77: 326-327. 


z .....-.. 1940. Moulting, growth, 
and survival after eyestalk removal in 
Uca pugilator. Biol. Bull., 78: 179-188. 


BLACK, J .B. 1958. Ontogeny of the first 
and second pleopods of the male crawfish 
Orconectes clyneatus (Hay). Tulane Stud. 
Zool., 6: 190-208. 


Buiss, D. E. 1954. Light inhibition of re- 
generation and growth in the crab, Gec- 
arcinus lateralis. Anat. Rec., 120: 742-7438. 


2 1956. Neurosecretion and 
the control of. growth in a decapvod crus- 
tacean. Jn: “Bertil Hanstroém, Zoological 
Papers in Honour of His Sixty-fifth 
Birthday, November 20th, 1956.” (K. G. 
Wingstrand, ed.), pp. 56-75. Zoological 
Institute, Lund. 

BROEKHUYSEN, G. J. 1955. The breeding 
and growth of Hymenosoma orbiculare. 
Desm. Ann. S. African Museum, 41: 318- 
343. 

Brown, F. A. JR., and O. CUNNINGHAM 
1939. Influence of the sinus gland of 
crustaceans on normal viability and eedy- 
sis. Biol. Bull., 77: 104-114. 


CARLISLE, D. B. 1958. Moulting hormones 
in Leander (Crustacea, Decapoda). 
Mar. Biol. Assoc. U. K., 32: 289-296. 


1954. On the hormonal 
inhibition of moulting in decapod crusta- 
cea. J. Mar. Biol. Assoc. U. K., 33: 61-63. 


1957. On the hormonal 
inhibition of moultine in decapod Crusta- 
cea. II. The terminal aneedysis in crabs. 
J. Mar. Biol. Assoc. U. K., 86: 291-307. 


CARLISLE, D. B. and P. F. R. DoHRN 1953. 
Studies on Lysmata seticaudata Risso. 
Il. Experimental evidence for a growth 
and moult-accelerating factor obtainable 
from eyestalks. Pubbl. Staz. Zool. Napoli, 
24: 69-83. 


No. 3 


Darsy, H. H. 1938. Moulting in the crus- 
tacean, Crangon armillatus. Anat. Rec., 
72(Suppl.): 78. 

DrAcH, P. 1939. Mue et cycle d’intermue 
chez les crustacés décapodes. Ann. Inst. 
Oceanogr. Paris, 9: 103-391. 

DuRAND, J. B. 1956. Neurosecretory cell 
types and their secretory activity in the 
crayfish. Biol. Bull., 111: 62-76. 

_ 1960. Limb regeneration 
and endocrine activity in the crayfish. 
Biol. Bull., 118: 250-261. 

ECHALIER, G. 1959. L’organ Y et le déter- 
minisme de la croissance et da la mue 
chez Carcinus maenas (L), crustacé déca- 
pode. Ann. Sci. Nat. Zool., Ser. 12, 1: 
1-57. 

FINGERMAN, M., and C. FITZPATRICK 1956. 
An endocrine basis for the sexual differ- 
ence in melanin dispersion in Uca pugi- 
lator. Biol. Bull., 110: 138-148. 

FINGERMAN, M. and M. E. Lowek 1957. Hor- 
mones controlling the chromatophores of 
the dwarf crawfish, Cambarellus shu- 
feldti: their secretion, stability, and sep- 
aration by filter paper electrophoresis. 
Tulane Stud. Zool., 5: 151-171. 

GABE, M. 1956. Histologie comparée de la 
glande de mue (organe Y) des Crustaces 


Malocostracés. Ann. Sci. Nat. Zool., 18: 
145-152. 
Hess, W. N. 1941. Factors influencing 


moulting in the crustacean, Crangon ar- 
millatus. Biol. Bull., 81: 215-220. 

Hiatt, R. W. 1948. The biology of the lined 
shore crab, Pachygrapsus crassipes Ran- 
dalla Pactfie Sev., 2: 185-211. 

JYSSUM, S., and L. M. PASSANO 1957. Endo- 
erine regulation of preliminary limb re- 
generation and molting in the crab Sesar- 
ma. Anat. Rec., 128: 571-572. 

Kyer, D. L. 1942. The influence of the 
sinus glands on gastrolith formation in 
the crayfish. Biol. Bull., 82: 68-78. 

Lees, A. D. 1955. The Physiology of Dia- 
pause in Arthropods, Cambridge: Cam- 
bridge Univ. Press, 150 pp. 

PASSANO, L. M. 1953. Neurosecretory con- 
trol of molting in crabs by the X-organ 
sinus gland complex. Physiol. Comp. et 
Oecol., 3: 155-189. 

ee 1960a. Low temperature 


blockage of molting in Uca pugnax. Biol. 
Bull., 118: 129-1386. 


ee eT 1960b. Molting and its 
control. IN: The Physiology of Crusta- 
cea, Vol. I: Metabolism and Growth (T. 
H. Waterman, ed.), pp. 478-536. New 
York: Academic Press. 


ScupAMorRE, H. H. 1942. Hormonal regula- 
tion of molting and some related phe- 
nomena in the crayfish, Cambarus immu- 
nis. Anat. Rec., 84: 514-515. 


1947. The influence of 
the sinus glands upon molting and asso- 


Mobberly: Molting Cycle in Crayfish 95 


ciated changes in the crayfish. Physiol. 
Zool., 20: 187-208. 

ee OAS actors lntliwenc= 
ing molting and the sexual cycles in the 
crayfish. Biol. Bull., 95: 229-237, 

SMITH, R. I. 1940. Studies on the effects 
of eyestalk removal upon young crayfish 
(Cambarus clarkii Girard). Biol. Bull., 
79); 145-152. 

SMITH, E. W. 1953. The life history of the 
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Gl Dii2eonioe 


108: 235-241. 


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ee 19 tb. Physiological 

changes which occur in the blood and 

urine of Panulirus argus Latreille during 

the molting cycle. Anat. Rec., 111: 573. 


ie Be an oc 1955a. The molting cycle 
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ent ROW eeeR Se 1955b. The molting cycle 
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485-503. 

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diapausing pupa of the cecropia silkworm. 
Biol. Bull., 110: 201-218. 


ABSTRACT 


The molting cycle of the crayfish 
Faxonella clypeata was defined. Field 
collections were made at least twice 


96 


a month, except during periods of 
drought, from October 19, 1959, through 
May 7, 1961. While crayfish contain 
gastroliths throughout the year in Lou- 
isiana, three major molting peaks oc- 
cur in November - December - January, 
April - May, and July - August. 

The three major molting peaks occur 
during the warmest and coolest parts 
of the year. The November-December- 
January peak is longer in duration 
than the peaks occurring in warmer 
months due to the fact that cold (8- 
10° C) acts as a suppressor. In favor- 
able temperatures the molting process 
continues until the temperature falls 
below a value that allows the animal to 
continue the premolt process. 


Constant light reduced gastrolith pro- 
duction, whereas crayfish exposed to 
photoperiods of 6 and 12 hours per day 
had a higher incidence of gastroliths 
than the controls. 


Rainfall can influence the molting 
cycle to the extent that animals may be 
forced to burrow if rainfall is scant. 
Animals do not molt in their burrows 


Tulane Studies in Zoology 


but wait until they again have ample 
surface water. 

Loss of appendages may be one of 
the factors stimulating animals to molt 
during the period between molting 
peaks. When crayfish in the laboratory 
lose two chelipeds and two pairs of 
walking legs, the animals increase gas- 
trolith production in comparison with 
animals having only one cheliped, two 
chelipeds, or no appendages missing. 

A molt-accelerating factor appears 
to be present in the supraesophageal 
ganglia and circumesophageal connec- 
tives. An acidified extract (pH 4.8) of 
the supraesophageal ganglia with the 
circumesophageal connectives attached 
is active in accelerating gastrolith for- 
mation when injected intramuscularly 
into F’. clypeata. 


A molt-inhibiting hormone was found 
in the eyestalks and the supraesopha- 
geal ganglia plus the circumesophageal 
connectives. The experimental results 
suggest that this inhibiting hormone 
may in reality be composed of a num- 
ber of hormones. 


Vol. 11 


TULANE STUDIES IN ZOOLOGY 


Volume 11, No. 4 


March 24, 1964 


CONTENTS 


MURICIDAE (GASTROPODA ) FROM THE NORTHEAST COAST OF SOUTH 
AMERICA, WITH DESCRIPTIONS OF FOUR NEW SPECIES 


Var vey? Reeves Unllise reweeemrameere eeeere o_ BN A 99 


EDITORIAL COMMITTEE: 


R. TUCKER ABBOTT, Pilsbry Chair of Malacology, Academy of Natural Sciences of 
Philadelphia, Philadelphia, Pennsylvania 

HARALD A. REHDER, Curator, Division of Mollusks, United States National Museum, 
Washington, D. C. 

EmiLy H. VoKEs, Department of Geology, Tulane University, New Orleans, Lou- 
isiana 


CHIRONOMIDAE (DIPTERA) OF LOUISIANA I. SYSTEMATICS AND IM- 
._ MATURE STAGES OF SOME LENTIC CHIRONOMIDS OF WEST-CENTRAL 
LOUISIANA 


James E. Sublette________ aE TS) es re os RE Oy SO ons oo Ce 109 


CHIRONOMIDAE (DIPTERA) OF LOUISIANA. II. THE LIMNOLOGY OF 
THE UPPER PART OF CANE RIVER LAKE, NATCHITOCHES PARISH, LOU- 
ISIANA, WITH PARTICULAR REFERENCE TO THE EMERGENCE OF CHI- 
RONOMIDAE 


Burton RK: Buckley and) janes BxSuplettenass = eareeie ee ee 15a 


EDITORIAL COMMITTEE: 


M. W. BOESEL, Professor of Zoology, Miami University, Oxford, Ohio 
J. S. DENDy, Professor of Zoology, Auburn University, Auburn, Alabama 
HENRY TOWNES, Research Associate, University of Michigan, Ann Arbor, Michigan 


7 
ie 
- 7 La . 
. 3 . A 
eye 
"Cap ” + A 
Ee sey m= a - 
‘ i= - ia a Re 
oe i aes ; a 
‘ a 
i a s * om > 
oP Dee | ‘aon! att BAe gt eee ete oN = Aa j° 
: ; ; aoe ae le . eras _ 2 
He oo 
Re en ae 10g 
- — 


MURICIDAE (GASTROPODA) FROM THE NORTHEAST COAST OF SOUTH 
AMERICA, WITH DESCRIPTIONS OF FOUR NEW SPECIES 
HARVEY R. BULLIS, JR., 

Branch of Exploratory Fishing, 

Bureau of Commercial Fisheries, 

Pascagoula, Mississippi 


The U. S. Bureau of Commercial Fish- 
eries has conducted two exploratory trawling 
cruises of the M/V OREGON along the 
northeast coast of South America from Trint- 
dad to the Amazon River. This work was 
carried out during the fall of 1957 and the 
late summer of 1958: on these cruises 295 
shrimp trawling stations were completed at 
depths ranging from 6 to 400 fathoms. 

Prior to this work virtually nothing was 
known of the biological nature of the area 
traversed, particularly in depths beyond the 
25-fathom contour. Some 50,000 specimens 


collected on the cruises have greatly increased 


the study material available from this region. 
The mollusk population was found to be 
very rich, and the volume of material from 
the two trips almost equals the total material 
collected during the 76 other OREGON ex- 
ploratory cruises in the tropical and sub- 
tropical western north Atlantic. 

This paper covers the macromollusca be- 
longing to the Muricidae. The genus Murex 
was represented at 56 localities by 11 species 
taken in depths ranging from 10 to 275 
fathoms (living material down to 200 fath- 
oms). Of the eleven, four are new, four are 
range-extension records of considerable mag- 


nitude, and three have been previously re- 
ported from the area. A single species of 
Trophon was found at one locality at the 
upper edge of the continental slope. One 
species of Typhis was found at two localities 
in the 20 to 30 fathom range. 

The general pattern of depth distribution 
for the specimens of Murex found during 
these cruises is given in Table 1. Locations 
of OREGON stations corresponding to the 
numbers given under each species record aré 
listed in Table 2. 

Holotypes and paratypes of new species 
described herein, as well as material pertain: 
ing to the other species discussed are depos- 
ited in the United States National Museum. 
Paratypes have been placed in the following 
collections, Museum of Comparative Zoolo- 
gy, Academy of Natural Sciences of Phila- 
delphia, American Museum of Natural His- 
tory, Chicago Natural History Museum, Tu- 
lane University, and the Marine Laboratory 
Museum, University of Miami. 

I wish to acknowledge with thanks the 
courtesies extended to me by Dr. Harald A. 
Rehder, Curator of Mollusks, United States 
National Museum, and by Dr. William J. 
Clench, Curator of Mollusks, Museum of 


TABLE 1. 
Depth distribution records for the species of Murex collected off the northeast coast of 
South America. Depths with records of live specimens are denoted as 0; depths 
with records of dead (empty) shells only are denoted as +. 


11-20 
31-40 
41-50 


Murex brevifrons 
Murex messorius 
Murex donmoorei 
Murex springeri 
Murex thompsoni 
Murex pomum 
Murex cellulosus nuceus 
Murex consuela 
Murex tryoni 
Murex beaui 
Murex oregonia 


| —+-coococo 
| ocoo—-— | 


51-60 


depth in fathoms 


S me oS tt © mm C te © 

Se oe Fay = OA SW ES 

Sap Aree OG A aoa eh (one ONES SiN oe 

1 ! ' 

~ = Se a se wo Ss oe © ae 

S moa se q ia = S aw te = 

Om eC act yeah = 
at 

i 

Hg. ot) Os pst eee 

=e (Ot Oe AO Secs Ae 


100 


Tulane Studies n Zoology 


TABLE 2. 


Vol. 11 


Station list and localities of collection of Murex off north eastern South America, 


OREGON 
Station 
Number 


1981 
1983 
1984 
1985 
1988 
1989 
2002 
2015 
2022 
2023 
2038 
2049 
2050 
2051 
2061 
2063 
2068 
2080 
2084 
2230 
2232 
2236 
2254 
2255 
2267 
2268 
2269 
2271 
2272 
2274 
2275 
2276 
2284 
2285 
2286 
2289 
2290 
2291 
2292 
2293 
2294 
2295 
2296 
2303 
2307 
2309 
2321 
2322 
2324 
2327 
2328 
2329 
2331 
2333 
2334 
2335 
2337 
2344 


North 
Latitude 


10°03’ 
09°53’ 
09°45’ 
09°41’ 
09°24’ 
09°45’ 
07°52’ 
07°38’ 
07°15’ 
Onawot 
05°46’ 
04°02’ 
04°04’ 
04°05’ 
02°31’ 
02°35’ 
02°35’ 
02°04’ 
01°45’ 
08°33’ 
08°31’ 
08°09’ 
07°07’ 
07°09’ 
06°58’ 
06°53’ 
06°49’ 
06°34’ 
06°30’ 
06°54’ 
06°50’ 
06°42’ 
06°48’ 
07°27" 
07°26’ 
07°25’ 
O72 27 
07°27’ 
07° 28’ 
OT 27" 
07°25’ 
07°27’ 
06°29’ 
06°04’ 
05°57’ 
05°54’ 
06°52’ 
06°50’ 
06°46’ 
06°26’ 
06°33’ 
06°40’ 
06°55’ 
06°58’ 
06°56’ 
06°50’ 
06°50’ 
08°10’ 


1957 and 1958. 


West 


Longitude 


60°01’ 
59°59’ 
59° 45’ 
59°47’ 
59°41’ 
59°45’ 
DT ee. 
54°11’ 
53°25’ 
53°25’ 
53°00’ 
50°33’ 
50°32’ 
50°27’ 
48°48’ 
48°14’ 
47°48’ 
47°00’ 
46°46’ 
58°46’ 
58°37’ 
58°23’ 
57°08’ 
57°06’ 
56°02’ 
55°59’ 
55°57’ 
55°54’ 
55°52’ 
55°40’ 
55°39’ 
55°37’ 
55°12’ 
54°54’ 
54°49’ 
54°35’ 
54°27’ 
54°27’ 
54°21’ 
54°15’ 
54°08’ 
53°47’ 
52°30’ 
52°35’ 
52°20’ 
52°17’ 
53°18’ 
53°29’ 
54°24’ 
54°20’ 
54°23’ 
54°25’ 
55°04’ 
55°03’ 
54°55’ 
55°34’ 
55°23’ 
58°18’ 


General Area 


Off eastern Venezuela 
Off eastern Venezuela 
Off eastern Venezuela 
Off eastern Venezuela 
Off eastern Venezuela 
Off eastern Venezuela 
Off British Guiana 
Off Surinam 

Off Surinam 

Off French Guiana 
Off French Guiana 


Off Cabo Orange, Brazil 
Off Cabo Orange, Brazil 
Off Cabo Orange, Brazil 


Off Amazon River 
Off Amazon River 
Off Amazon River 
Off Amazon River 
Off Amazon River 
Off British Guiana 
Off British Guiana 
Off British Guiana 
Off British Guiana 
Off British Guiana 
Off Surinam 
Off Surinam 
Off Surinam 
Off Surinam 
Off Surinam 
Off Surinam 
Off Surinam 
Off Surinam 
Off Surinam 
Off Surinam 
Off Surinam 
Off Surinam 
Off Surinam 
Off Surinam 
Off Surinam 
Off Surinam 
Off Surinam 
Off French Guiana 
Off French Guiana 
Off French Guiana 
Off French Guiana 
Off French Guiana 
Off French Guiana 
Off French Guiana 
Off Surinam 
Off Surinam 
Off Surinam 
Off Surinam 
Off Surinam 
Off Surinam 
Off Surinam 
Off Surinam 
Off Surinam 
Off British Guiana 


Depth in 
Fathoms 


200 
125 
200 
150 


Date 


tty 
iy, 
11/ 
11/ 
11y, 
Ys 
117 
11/ 
117 
11/ 


3/57 
3/57 
3/57 
3/57 
4/57 
4/57 
6/57 
8/57 
9/57 
9/57 


11/11/57 
11/13/57 
11/13/57 
11/13/57 
11/15/57 
11/15/57 
11/15/57 
11/17/57 
11/18/57 


9/19/58 


No. 4 


Comparative Zoology, while referring to the 
collections in their care. Appreciation is also 
due to Dr. Rehder for critical review of the 
manuscript and to Emily H. Vokes for nu- 
merous helpful suggestions. 


MUREX (MUREX) DONMOOREI, 


sp. nov. 
Figures 1 and 2. 


Material: Station 2051, 1 dead; 2236, 1 
alive; 2254 (type locality, 45 miles north of 
St. Andrews Point, British Guiana, in 20 to 
22 fathoms), 1 alive; 2255, 1 dead; 2268, 
(alive: 2272, 2 alive: 2274, 2 alive: 2275, 
2 dead; 2284, 6 alive; 2309, 1 alive; 2321, 
2 alive, 2 dead; 2322, 2 dead; 2337, 4 alive. 

Holotype: height 50 mm, width without 
spines 22.5 mm, height of aperture 38 mm. 
Shell solid, heavily sculptured, with an 
elongate, spinose siphonal canal. Whorls 
7.3, including the protoconch of one and 
three-quarter whorls. Primary whorl slight- 
ly eroded, posteriorly flattened. Spire mod- 
erately extended. Suture deep, irregular, 
made undulous by the axial ridges. Aper- 
ture ovate and extending into the siphonal 
canal as a narrow slit. Palatal lip a thick 
varix, with six stout spines and intervening 
raised cords, one of which forms an imbri- 
cate scale on the face of the varix. VWarix 
incomplete, hollow, opened on the inner 
surface, behind which is a row of nine white 
rounded teeth. A single tooth persists in the 
upper end of the aperture, corresponding to 
the intruding varix of the penultimate 
whorl. Parietal lip slightly expanded, ad- 
hering above and erect below. Siphonal 
canal elongate, slightly more than half the 
length of the shell, and armed with three 
rows of spines. There are 4 spines in the 
first two rows and 3 in the third, all are 
curved slightly forward, and diminish in 
size anteriorly. Sculpturing on the body 
whorl consists of 17 distinct spiral cords 
that cross four or five axial intervarical 
ridges to form spirally elongate nodules 
across the ridges. On the posterior half of 
the canal are 13 spiral cords crossed with 
fine growth lines. First 3  post-nuclear 
whorls without varices and strongly can- 
cellate. The varices start on the fourth whorl 
and are radially aligned, the lower directly 
behind and adjoining the upper. Ground 
color light tan, varices brownish. Each spiral 
cord marked with a dark brown line. Tip of 


South American Muricids 


101 


canal mottled yellow, and surmounted by a 
dark, reddish-brown spot. Operculum thin, 
reddish-brown, with strong concentric ridges. 


Radula typical, formula te 1:0. U.S.N.M. 
635146. 


Discussion: This species was found from 
eastern British Guiana to eastern French 
Guiana. The southernmost record is a shell 
from off Cape Cassipore. OREGON rec- 
ords show a range of 17 to 50 fathoms, but 
the optimum depths appear to be between 
25 and 35 fathoms. 


In general shape and sculpturing M. don- 


mooret could be easily confused with M, 


cabritu Bernardi. The gross sculpturing 
agrees closely in form and structure. Many 
of the paratypes of M. donmoorei show 
similarly paired pallial crenulations. The 
dissimilarities are less obvious but are never- 
theless striking when series of both species 
are compared. The new species appears to 
be a much smaller form, although the pro- 
toconch is two to three times as large. The 
posterior flattening of the apical whorl is 
combined with a definite low-set supra- 
sutural cord, which differs from the tiny, 
unsculptured, bulbous protoconch in M. ca- 
britu. The anterior canal is proportionately 
shorter, equal to 50 to 55 percent of the 
maximum length. In this respect M. don- 
mooret more closely resembles M. elenensis 
Dall (Pacific coast of Central America). In 
M. cabritu the canal varies from 60 to 64 
percent of the length. The sharp brown 
threads that ride the spiral cords are a dis- 
tinctive color feature. Radular differences 
are equally distinct. The rachidian in M. 
cabr. tit is tricuspid, not typical of the genus, 
whereas there are five strong cusps in M. 
donmooret. 

Th's species is named for Donald R. 
Moore, University of Miami Marine Lab- 
oratory. 


MUREX (MUREX) TRYONI Hidalgo 

Records: Station 2230, 3 dead; 2236, 1 
dead. 

These records extend the observed range 
from the lower Antilles (Clench and Far- 
fante, 1945) to off French Guiana. The 
specimen from station 2291, measuring 50 
mm, ts the largest I have seen. 


Tulane Studies mm Zoolog) 


Figures 1-8. 1 and 2—Murex donmoorei, new species. Holotype. 3 and 4—Murex thomp- 
soni, new species. Holotype. 5 and 6—Murew oregonia, new species. Holotype. 7 and 8— 
Murex springeri, new species. Holotype. 


No. 4 


MUREX (MUREX) MESSORIUS 
Sowerby 


Records: Station 2230, 3 dead; 2236, 1 
dead; 2269, 1 dead; 2272, 1 alive; 2276, 1 
dead; 2327, 1 alive; 2328, 6 alive; 2329, 4 


alive; 2334, 2 alive. 

Recently Vokes (1963) has taken a firm 
position restricting M. recurvirostris (s.1.) 
to the Pacific coast, resurrecting M. mes- 
sorius Sowerby for the “recurvirostris” of 
the western Atlantic exclusive of Florida, 
and allying it with M. sallasi Rehder and 
Abbott (Yucatan) and M. rubidus Baker 
(Florida). One must regard this as a clart- 
fication of one of the most complex species 
groups in this region. The present material 
clearly falls within this group; however, it 
too displays rather consistent differences 
from the many described species and sub- 
specific forms. For the present it seems 
prudent to refer to the entire lot from off 
the Guianas as M. messorius and await the 
assemblage of material that will afford 
synoptic treatment. 

Specimens in this series reach a length 
of 65 mm, and the shell is noticeably more 
solid than in either M. sallasz or M. rubidus. 
The aperture is proportionately larger 
(about 10 percent longer and wider of the 
total length of the shell) than in M. rubidus. 
The coloration is a mottled brown-tan over 
cream. 

All of the living material collected was 
taken from catches that contained large 
amounts of mud and sand, evidence of hard 
digging by the trawl. Most of the above 
records came from off the Surinam River 
along the 30-fathom curve. 


MUREX (MUREX) CONSUELAE 
Verrill 

Record: Station 2063, 1 dead. 

Recently Clench (1959) recorded this 
species (as M. pulcher A. Adams) from 
Salvador (Bahia), Brazil, a range extension 
of over 2000 miles from Barbados. The 
present record from the offings of the 
Amazon River is intermediate. 


MUREX (MUREX) THOMPSONI, 
sp. nov. 
Figures 3 and 4. 
Material: Station 2061, 12 alive; 2230, 1 
dead; 2232, 2 alive; 2269, 1 dead; 2321 
(type locality, 75 miles NNE of Pte. Mana, 


South American Muricids 


103 


French Guiana, in 34 fathoms), 4 alive, 1 
dead; 2322, 7 alive; 2329, 1 dead; 2331, 23 
alive; 2333, 4 alive. 

Holotype: Height 35.8, width 18.9, height 
of aperture 22.9 mm. Shell solid and small. 
Whorls 7.5 including the protoconch. Spire 
acute, moderately extended. Suture ap- 
pressed, irregular. Aperture small, oval, por- 
celaneous white, with a well-developed tooth 
within the posterior margin. Aperture lead- 
ing into a narrow slit on the siphonal canal. 
Outer lip thick, erect, and crenulate. Behind 
and well-separated from the crenulations are 
eight elongate denticulations. Parietal lip 
adherent above, erect below, with four par- 
allel denticulations on the columella. Si- 
phonal canal moderately extended, recurved, 
and reflected to the right. There are one 
and one-half bulbous, shiny, unsculptured 
nuclear whorls. Below these are two and 
one-half strongly cancellate whorls, followed 
by the first varical ridge. Each remaining 
whorl bears three heavy corrugated varices, 
which are axially aligned and slightly behind 
the superior varix. Sculpturing consists of 
alternate strong and light spiral cords that 
traverse three intervarical ridges to form 
prominent nodules. These persist over the 
varices as raised rings which can best be 
described as tracheoid. The lower half of 
the outer forward margin of each varix sup- 
ports a low frill. On the last two varices is 
a short, open spine at the shoulder and an- 
other at the base of the canal. On the for- 
ward face of the varices there are numerous, 
fine v-shaped imbrications. Fine growth 
lines, present over the entire shell, are em- 
phasized by weak ridges of light yellow 
periostracum. The ground color is light 
cream. The stronger spirals each have a light, 
orange-brown thread. Operculum unguicu- 
late, thick reddish-brown, and heavily sculp- 
tured with a sub-apical nucleus. U.S.N.M. 
635147. 

Discussion: The type is somewhat more 
exotic than the rest of the series at hand but 
was selected for its size, condition, and ap- 
parent maturity. The only larger specimen 
(44 mm) is more typical but it is badly 
eroded and heavily encrusted with bryozoans, 
barnacles, and worm tubes. Generally, the 
body is more slender and elongate and with- 
out spines. The species characteristically 
shows a tendency to raise a varical web or 
frill as in M. cazlleti Petit, but to a lesser 


104 


degree. This is present in all specimens 
down to 22 mm and absent in a series of 
23 juveniles ranging from 9.5 to 19.5 mm. 
Ground coloration varies from whitish to 
mottled brown. One specimen is marked 
with two brown bands. The darker lines are 
present on all living specimens, but they ap- 
pear to fade quickly on dead shells. 


Murex thompsoni may be confused with 
small specimens of another new species, 
M. springeri. A comparison of their respec- 
tive characters is given under that species. 
In the western Atlantic fauna M. thompsoni 
can best be compared with M. cailleti (and 
its subspecies kuvgleri Clench and Farfante) , 
from which it differs in having a smaller 
aperture, a proportionately heavier varix, 
and a shorter, stouter siphonal canal. It is 
also a smaller species and occupies a much 
shallower depth range. The present series 
was taken between the Orinoco River and 
Maraca Island, a range of some 800 miles, 
in depths of 22 to 60 fathoms. The large 
series of juveniles came from a depth of 
30 fathoms. This species is named after 
John R. Thompson, colleague at the Bureau 
of Commercial Fisheries Exploratory Fish- 
ing Base, Pascagoula, Mississippi. 


MUREX (SIRATUS) BEAUI 
Fischer and Bernardi 
Records: 


Station 1981, 4 alive: 1983, 6 
alive; 1984, 2 dead; 1985, 7 alive: 1988, 1 
alive; 1989, 2 dead; 2080, 1 alive; 2290, 9 
alive; 2291, 2 alive; 2292, 1 alive; 2293, 7 


alive; 2294, 1 dead; 2296, 4 alive. 


These captures extend the known range 
of M. beaui some 1,000 miles, from Guade- 
loupe Island to northeastern Brazil. The 
larger specimens (above 100 mm) display 
an almost complete absence of webbing and 
are very similar to those caught on grey mud 
bottom in the north-central Gulf of Mexico. 
Smaller specimens have webbed varices. 


Popular belief is that extreme webbing 
is directly related to bottom type; numerous 
observations of this species do not confirm 
this. More likely the higher degree of vari- 
cal webbing is atypical. Truly exotic speci- 
mens can be found in most of the localities 
over its range if large series are obtained. 


Tulane Studies n Zoology 


Vol. 11 


MUREX (SIRATUS) SPRINGERI, 
sp. nov. 
Figures 7 and 8. 


Material: Station 2002, 1 dead; 2015, 2 
dead: 2061, 24 alive; 2230, 5 dead; 2232, 1 
alive; 2271, 1 alive; 2286, 1 dead; 2289 
(type locality, 95 miles NNE of Surinam 
River entrance in 75 to 80 fathoms), 23 
living and dead; 2290, 1 dead; 2322, 23 
alive: 2328, 1 alive; 2329, 1 dead; 2333, 7 
alive; 2344, 1 dead. 

Holotype: Height 70, width 32.7, height 
of aperture 45 mm. Shell of medium size 
and solid. Whorls 9.3 including protoconch. 
Nucleus white and smooth; whorls 2, the 
first a flattened planorboid coil giving the 
protoconch a truncate appearance. Spire 
moderately extended. Suture distinct, the 
lower whorl applied as a thin fold, below 
which there is a definite canaliculate de- 
pression. Aperture milky white, ovoid, with 
a slight anal canal bordered on the body wall 
by a small rounded tooth. Aperture leads 
into a narrow slit on the siphonal canal. 
Outer lip extending well beyond the last 
varix, erect, faintly denticulate within, and 
moderately crenulated. The crenulations on 
the outer lip are colored brownish on the 
inner surface. Parietal lip adherent over 
most of its length, erect, and lightly den- 
ticulate just ahead of where it turns sharply 
into the siphonal slit. Axial sculpture of 
three strong varices on each whorl. Each 
varix supports a single strong spine at the 
shoulder and two shorter spines on the canal. 
Shoulder spines imbricate and hollowed, but 
with a laminated “filling” to the varix edge. 
This “filling” has the appearance of having 
been “squeezed out” of the spinal slit. On 
the spine and along the forward outer edge 
of the varix there is a substantial low frill 
which is strongest on the basal portion. 
Numerous intervarical ridges, diffused and 
very slight, appear between the last two 
varices; from three to six on earlier sections. 
These and the varices are crossed by raised 
spiral cords, every second to sixth of which 
is heavier and more darkly colored. This ir- 
regular cording produces numerous small, 
irregularly shaped nodules which are strong- 
er on the spire than on the body whorl. 
Spirals cross the varices as raised rings or 
extend onto the frill. Anterior canal mod- 
erately extended, rather heavy, recurved, and 
reflected to the right. Spiral sculpturing 


No. 4 


crowded over and between the two spines 
on the canal but well separated below. 
Ground color cream, with a brownish band 
below the shoulder, most prominent on the 
varices. The heavier spirals are various 
shades of yellow-brown. The forward face 
of the siphonal canal is glossy white, but is 
heavily etched with growth lines. The speci- 
men was collected dead. U.S.N.M. 635148. 

Discussion: This species was found quite 
commonly from eastern British Guiana to 
Brazil in depths of 18 to 120 fathoms. Most 
of the deeper records of M. springeri are of 
dead specimens, which tend to be the largest 
individuals taken. 

At less than 30 mm M. springert may be 
confused with another new species, M. 
thompsoni, due to the latter's occasional 
tendency to carry a small shoulder spine, and 
rarely a spine or two on the anterior canal. 
A reliable way to separate these two is by 
nuclear characters. The first whorl of the 
nucleus is bulbous in M. thompsoni and pos- 
teriorly flattened in M. springert. Neither 
can be confused with M. beaui which at 
this shell size is very light and fragile, more 
elaborately frilled, and has a nucleus of one 
and one-half to three rounded but evenly 
tapering whorls. 

Radular characters also appear to separate 
these three species. The lateral teeth of all 
species vary so slightly as to be of little or 
no diagnostic use. The differences in the 
characteristics of the rachidian are more of 
degree than of basic structure. Typically, the 
muricid rachidian possesses five cusps; the 
central is usually the longest and heaviest, 
the outer cusps are somewhat smaller, and 
the intermediary ones are very small. The 
basal plate varies relative to the cusps both 
in shape and size; however the problem of 
preparing uniform slide mounts makes evalu- 
ation difficult. The general specific agree- 
ment in cusp formation encouraged me to 
use this character here. 

At a larger size, M. springeri resembles 
closely several specimens of M. beam that 
I have seen, but in addition to the radular 
and nuclear characters it differs in being 
much heavier and having a shorter siphonal 
canal. Both species are quite variable in 
shape. As in M. beaut, which frequently 
lacks any sign of spines in the larger forms, 
several specimens of M. springeri lack the 
shoulder spine. In the large series at hand, 
all show the spines on the canal. The degree 


South American Muricids 


105 


of frilling is also variable, but none of the 
present material even approaches the mag- 
nificent frilling that is seen occasionally in 
M, beaut, This species is named for Stewart 
Springer of the Bureau of Commercial 
Fisheries. 


MUREX (PHYLLONOTUS) POMUM 
Gmelin 


Records: Station 2050, 2 alive; 2061, 4 
alive; 2230, 2 dead; 2267, 2 alive; 2269, 1 
dead; 2275, 1 alive; 2284, 11 alive; 2289, 
1 dead; 2307, 1 alive; 2322, 5 dead, 1 alive; 
2325, 1 alive; 2337, 9 alive. 

Although these captures are well within 
the known geographic range of M. pomum, 
the present records of living specimens range 
from depths of 25 to 60 fathoms, and for 
dead shells from 22 to 80 fathoms, well be- 
yond the bathymetric range formerly con- 
sidered typical. This probably indicates sa- 
linity tolerances since the enormous river 
drainage along this entire coast must cer- 
tainly lower salinities appreciably out to the 
vicinity of the 20 fathom curve. 


Although the coloring of a few of these 
specimens resembled that of forms from the 
shores of United States, many were much 
lighter, or even whitish, with a light purplish 
sheen within the aperture. 


Assignment of these specimens to M. 
pomum has been done with some doubt fol- 
lowing the discussions of Abbott (1958) 
and Clench (loc. cit.) regarding M. mar- 
garitensis (s.s.) Abbott. The present ma- 
terial shows the same ambivalence to either 
name, except for the deep pink coloration 
of the aperture which is quite striking. How- 
ever, the brown patch at the posterior st- 
phonal notch is strong. Since my observa- 
tions of the radulae indicate no discernible 
differences I have conservatively assigned 
all of the above specimens to M. pomum. 


MUREX (CHICOREUS) BREVIFRONS 


Lamark 


Records: Station 2038, 3 alive; 2049, 5 
alive: 2050, 2 alive; 2272, 10 alive; 2276, 
1 dead; 2303, 2 alive; 2327, 4 alive. 

These extend the range of M. brevifrons 
from the Guianas to the offings of the Ama- 
zon River. Depths at the collecting sites 
ranged from 15 to 40 fathoms. Most of the 
specimens were very elaborately spined and 


106 


not at all dissimilar to Murex 
and Farfante. 


argo Clench 


MUREX (POIRIERIA) OREGONIA, 


sp. nov. 
Figures 5 and 6. 


Material: Station 1981, 1 alive; 2022, 2 
alive; 2023 (type locality, 95 miles north of 
Pte. Mana, French Guiana, in 135 fathoms), 
2 alive; 2084, 1 dead; 2285, 2 alive; 2286, 
5 dead; 2291, 1 alive; 2292, 1 alive; 2293, 
7 alive; 2294, 5 alive; 2295, 3 alive; 2296, 
150 dead and living. 


Holotype: Height 90.2, width 37.8 (not 
including spines); height of aperture 54.9 
mm. Shell heavy, elongate, and strongly 
sculptured. Apex eroded (as are all mature 
specimens in the series), 8 whorls remain- 
ing. Aperture elongate, leading into an ex- 
tended siphonal canal. Outer lip flared, 
sharp, somewhat crenulate on the edge and 
with 13 teeth on the inner margin; the lip 
supports a strong, upturned, imbricate and 
hollowed spine at the shoulder. Parietal wall 
with a raised, erect, shield-like callus. Spire 
turreted, with a deep, irregular suture. Spiral 
sculpturing of eight strong cords between 
the base and shoulder. Axial sculpture of 
nine varices supporting eight upturned im- 
bricate spines which correspond to the cords, 
plus the shoulder spine, and a pair of long 
spines on the base of the siphonal canal. The 
penultimate whorl has eight varices with the 
sub-shoulder spines absent, but with nodules 
at the crossing of the cords and varices, with 
three cords left exposed. On the rest of the 
spire only two cords are exposed. The varices 
cross the shoulder with a step- -like over- 
lapping of the succeeding section. Growth 
lines in the form of undulating wrinkles are 
most evident on the shoulder. The eight 
spines below the shoulder on the last varix 
are imbricately doubled and not directly con- 
nected to the margin of the lip. The second 
varix is similar but a few of the forward 
spines directly receive involutions of the 
former lip. This is more so on the third 
varix and entirely the case on the fourth. 
Five previous anterior canals are present. 
Operculum dark reddish-brown, unguiculate, 
broadly oval with a terminal apex, and 
strongly sculptured with growth lines. The 
radula is large; the rachidian has five cusps 
of about equal length. The middle and outer 


Tulane Studies n Zoology 


Voladd. 


two cusps are slightly stouter. The laterals 
are triangulate and hooked. Radular formula 


is 0:1:°:1:0. US.NM. 635149. 


Discussion: This is a striking species that 
forms a closely related group with M. pazi 
Crosse and M. nuttingi Dall, and bears some 
affinities to M. atlantis Clench and Farfante, 
and M. carnicolor Clench and Farfante: the 
latter to a lesser degree. It differs from all 
of the above in having paired siphonal 
spines, and in the length/whorl relationship, 
being from two and one-quarter to four 
times greater in size by whorl count. Murex 
pazi and M. nuttingi are more similar in 
shell characteristics than has been previously 
indicated, with the former showing a great 
range of sculpturing which may eventually 
provide a basis for a geographical subspecific 
distinction. Bahamian and Cuban M. pazi, 
which represent the typical Antillean species, 
have the intermediary sculpturing between 
the shoulder and siphonal canal spines great- 
ly reduced. This is exemplified by the figure 
given by Clench and Farfante (1945, p. 44). 
Along southern Florida and extending per- 
imetrically around the Gulf of Mexico shelf 
in the 100 fathom range, M. paz: has heavy 
intermediary sculpturing. M. nuttimgi might 
be confused with this group but it can be 
differentiated by the denticulate inner sur- 
face of the pallial lip and the more elongate 
spines. 

The large series of paratypes was collected 
over a depth range of 105 to 275 fathoms 
from positions due east of Galeota Point, 
Trinidad, east of the Amazon River, and at 
several points between, indicating a con- 
tinuous range from the equator to 10° North 
Latitude. 


The series from station 2296 (off the 
Maroni River) included 126 living and 24 
dead examples of this species. The speci- 
mens range from 40 to 75 mm in height, 
and provide an excellent series for review- 
ing some of the juvenile characters. A few 
of the smallest specimens possess a complete 
planorboid protoconch of about two whorls 
that are posteriorly flattened and truncate 
and bear faint axial riblets on the second 
whorl. The original protoconch was almost 
certainly a fragile, deciduous structure of 
which no trace remains; the protoconch pres- 
ent is a shelly dome, secreted within and 


No. 4 


closing off the embryological apical chamber. 
The coil starts in the form of a “tuck” below 
the following coil, creating a minute apical 
pit. 

Under 60 mm there are no indications of 
parietal thickening. The callus is thin and 
adhering. All spines except the shoulder 
spines are greatly reduced, and the body ap- 
pears more attenuate anteriorly. The shells 
are thin and fragile, marked with four to 
eight spirals, and have eight to nine varices 
on the body whorl. They resemble in a gen- 
eral way, M. atlanticus except the siphonal 
spines differ and they do not have a bent 
canal. The five smallest specimens from sta- 
tion 2296 all have paired siphonal spines. 
A 44 mm specimen from station 2291 has 
only a single row, as do six of the entire 
series at hand. Above 60 mm the spiral 
cording remains constant at seven or eight 
lines although some are irregular and faint. 


MUREX (FAVARTIA) CELLULOSUS 
NUCEUS Morch 


Record: Station 2050, 1 dead. 


This record extends the southward range 
of M. cellulosus nuceus from St. Thomas, 
Virgin Islands (Clench and Farfante, 1945), 
to northern Brazil. 


TYPHIS ALATUS Sowerby 1850 


Records: Station 2324, 1 alive: 2331, 2 
dead. 

This species has been reported from the 
Miocene of the Dominican Republic (type 
locality), the Chipola formation of Florida, 
the Gatun formation of Toro Cay, Panama, 
and from Bowden, Jamaica. Woodring 
(1928) points out that the entire subgenus 
Talityphis has been considered extinct in 
the Caribbean region. 


South American Muricids 


107 


A comparison of the present material with 
the Bowden species and the description and 
figures of the subspecies T. a. obesws Gabb, 
substantiated Woodring’s suspicions as to the 
validity of obesus. The entire group of live 
and fossil material does not show consistent 
tangible differences and the present material 
must be relegated to T. alatus. 


TROPHON ACTINOPHORUS Dall 


Record: Station 2068, 7 alive. 

These specimens extend the southward 
distributional record for this species from 
south of Barbados to Brazil. 


REFERENCES CITED 


ABBOTT, R. T. 1958 The marine mollusks of 
Grand Cayman Island, British West In- 
dies. Acad. Nat. Sci. Philad. Mono. 11: 
61-62. 

CLENCH, W. J. 1959 The genus Murex in 
the Western Atlantic. Johnsonia, 3(39): 
333-334. 

CLENCH, W. J. and J. PEREZ FARFANTE 1945 
The genus Murex in the Western Atlan- 
tic. Johnsonia, 1(17): 1-58. 

VoKES, E. H. 1963 Cenozoic Muricidae of 
the Western Atlantic Region. Pt. 1— 
Murex sensu stricto. Tulane Stud. Geol., 
1L(3)) 8 OB =a 

WoopRING, W. P. 1928 Miocene mollusks 
from Bowden, Jamaica. Part II, Gastro- 
pods and discussion of results. Carn. Inst. 
Wash. Publ. 385: 294-295. 


ABSTRACT 


Eleven species of Murex were col- 
lected off the northeast coast of South 
America by the M/V OREGON. The 
material came from 56 lecalities be- 
tween Trinidad and the Amazon River 
in depths ranging from 10 to 275 fath- 
oms. Four of the species are consider- 
ed new; J. donmoorei, M. thompsoni, 
M. springeri, and M. oregonia. Four 
species represent first records for the 
area. Records for live specimens are 
also given for Trophon actinophous 
Doll and Typhis alatus Sowerby. 


CHIRONOMIDAE (DIPTERA) OF LOUISIANA 
I. SYSTEMATICS AND IMMATURE STAGES OF SOME LENTIC 
CHIRONOMIDS OF WEST-CENTRAL LOUISIANA 


JAMES E. SUBLETTE, 
Eastern New Mexico University, 
Portales, New Mexico 


NOMENCLATURE 


The chironomid fauna of the Southern 
United States was poorly known until the 
publication of Henry K. Townes’ monu- 
mental work on the Nearctic Chironomini 
(=Tendipedini) in 1945. Since that time 
several published works have added to the 
knowledge of a regional chironomid fauna. 
These are reviewed and the species synono- 
mized as they apply to the present study in 
the systematic treatment which follows. 
Since Townes reviewed the synonomy of 
each species of Chironomini, I shall not 
duplicate here his lists but rather cite only 
those relevant contributions since 1945. For 
those subfamilies and tribes not included in 
Townes’ work, as complete a synonomy as 
is known to me is given. Citations of the list 
in my 1955 paper are not given in the 
synonomies which follow as they were dup- 
licated in my 1957 paper, nor are the species 
listed by Townes (in Johannsen and Townes, 
1952) given since this publication is an 
abridgment of his 1945 paper. 

Nomenclature of the Chironomidae is in 
an extremely confused state. Notable points 
of controversy are the Meigen 1800 versus 
1803 names and the application of Tany- 
tarsus by Townes in a very different sense 
from customary usage of approximately the 
previous half-century. 


The Meigen names controversy as well as 
that of Tanytarsus are now before the Inter- 
national Commission on Zoological Nomen- 
clature. In the interim, I am following usage 
that appears to be consistent with the opin- 
ion of a majority of dipterologists, as evi- 
denced by publication and personal cor- 
respondence. 


Two recent publications, Brundin (1956) 
and Fittkau (1962), have greatly clarified 
the status and position of many taxa of the 
Orthocladiinae and Tanypodinae, respective- 
ly. Unfortunately, an application of these 
works to the Nearctic fauna would necessi- 
tate a re-examination of most of the types 
of North American chironomids. To have a 
solid systematic treatment I am following 


mostly the taxa of Freeman (1955-1961) 
that are based largely on adults. I have at- 
tempted to indicate position of appropriate 
species in the Brundin-Fittkau nomenclature. 

The following genera and subgenera as 
used in this paper are compared with those 
given in Johannsen and Townes (1952), 
the most inclusive modern work on adult 


Chironomidae of North America. 


Present Usage 


Subfamily Tanypodinae 


Tanypus 
Procladius 

(Psilotany pus) 
Procladins (Procladinus) 
Coelotany pus 


Pentaneura (Pentaneura) 


Ablabesmyia 
Subfamily 
Orthocladiinae 
Cricotopus 
Nanocladius 


Psectrocladius 


Smittia 
Subfamily 
Chironominae 
Tribe Tanytarsini 
Tanytarsus 
(Tanytarsus) 
Tanytarsus 
(Cladotanytarsus) 
Micropsectra 
Tribe Chironomini 
Psendochironomus 
Lauterborniella 
Parclauterborniella 
Polypedilum 
(Poly pedilum) 


Stenochironomus 
Chironomus 
(Endochironomus) 
Chironomus 
(Xenochironomus ) 
Chironomus 
(Cryptochironomus) 


Chironomus 
(Chironomus) 


Chironomus 
(Dicrotendipes) 

Gly ptotendipes 
(Phytotendipes) 


Johannsen and 
Townes (1952) 
Pelopiinae 

Pelopia 

Procladins 


Coelotany pus 
Pentaneura Group C-E 
Pentaneura Group A 


Hydrobaeninae 
Cricotopus 
Hydrobaenus 
(Eukiefferiella) 
Hydrobsenus 
(Psectrocladius) 
Hydrobaenus (Smittia) 


Tendipedinae 
Calopsectrini 


Calopsectra 


Tendipedini 
Pseudochironomus 
Lauterborniella 
Apedilum 
Polypedilum 
(Tripodura) 
Poly pedilum 
(Polypedilum ) 
Stenochironomus 
Tanytarsus 
(Endochironomus) 
Xenochironomus 


Cryptochironomus 
Harnischia 


Tendipes (Tendipes) 
Tendipes (Einfeldia) 
Tendipes 
(Limnochironomus ) 
Glyptotendipes 
(Phytotendipes) 


110 


Scope 

Most of the material included in this re- 
port was collected while the writer and two 
of his former graduate students, Burton R. 
Buckley and Robert F. Tyler, were at North- 
western State College, Natchitoches, Louisi- 
ana. Most specimens were collected from 
Cane River Lake, Chaplain’s Lake, and the 
holding ponds at the United States Fish 
Hatchery at Natchitoches, Louisiana. Col- 
lecting methods included tent and funnel 
traps (Sublette and Dendy, 1958 (1959) ) 
rearing of larvae and pupae, and light traps. 
From the latter, specimens included were of 
species known not to occur exclusively in 
lotic water. 

A small amount of the material presented 
here was collected while I was engaged in 
research projects supported by the National 
Institutes of Health (RG 4594 and 6829) 
and the Atomic Energy Commission (AT- 
(40-1 )-2596). Most of the results of these 
researches will appear elsewhere. 

Disposition of material is given in paren- 
thesis after the collection data of each spe- 
cies. The following abbreviations are used: 

U.S.N.M.—United States National Muse- 
um Collection, Washington, D. C. 

C.N.C.—Canadian National Collections, 
Ottawa. 

A.N.S.P.—Academy of Natural Sciences, 
Philadelphia, Pennsylvania. 

I.N.H.S.—Illinois National History Sur- 
vey, Urbana, Illinois. 

Duplicate material unless otherwise listed 
is in my personal collection. 

The localities Cane River Lake, Chap- 
lain’s Lake, and the United States Fish 
Hatchery ponds in Natchitoches Parish, 
Louisiana are abbreviated C.R.L., Ch.L. and 
US.F.H., respectively. 

Grateful acknowledgment is made to my 
wife, Mary Smith Sublette, for assistance in 
preparation of study material and the manu- 
script. 


SUBFAMILY TANYPODINAE 
TANYPUS STELLATUS Coguillett 


Tanypus stellatus Coquillett, 1902: 
of adult. 


89, description 


Protenthes stellatus (Coquillett); Malloch, 1915a; 
383, description of pupa and adult. 
Tanypus stellatus Coquillett; Johannsen, 1937: 20, 


description of larva and pupa. 
Tanypus stellatus Coquillett; Morrissey, 1950; 90, 
distribution; description of pupa; phenology. 


Tulane Studies n Zoology 


Vol. 11 


Pelopia stellata (Coquilletc) ; 
sen and Townes), 1952: 


Johannsen (in Johann- 
10, adult, in key. 


Pelopia stellata (Coquillett); Neff, 1955: 5, descrip- 
tion of larva, pupa and adult. 
Pelopia stellata (Coquillett) ; Tebo, 1955: 96, ecology. 


Pelopia stellata (Coquillett); Paine and Gaufin, 
1956: 296, ecology. 

Pelopia stellata (Coquillett); Sublette, 1957: 381, 
ecology; phenology. 

Pelopia stellata (Coquillett) ; Roback, 1957c: 47, 48, 
description of larva and pupa. 

Tanypus stellata (Coquillett); Beck and Beck, 1959: 


91, adult. 


Pelopia stellata (Coquillett); Davis, 1960: 71 and 
following pages, ecology. 

Pelopia stellata (Coquillett); Judd, 1960: 206, phe- 
nology. 

Pelopis stellata (Coquillett); Judd, 1961: 95, phe- 
nology. 

Males: Wing length 2.34-2.43, mean 2.37 
mm (3); leg ratio 0.70-0.81, mean 0.76 
(3); antennal ratio 2.12-2.40, mean 2.23 
(3). 

Material examined: Four males, 7-[X-56; 
1 female, 11-IX-56; 1 male, 6-VII-57; 1 
female, 10-[X-57; 1 male, 17-573) 1 


male, 14-X-57; C.R.L. One male, 
(UES dese 


TANYPUS new species 1 


Description of this new species is given 
by Sublette (in press in Proc. U.S. Natl. 
Mus.). 

Material examined: Specimens from 
Natchitoches and environs were included in 
the type series. In addition to these I have 
examined 2 males, 6-[X-58; U.S.F.H. 


20-TX-58; 


TANYPUS new species 2 
Protenthes punctipennis (Meigen) Malloch, 1915: 

389 (in part), description of pupa and adult, 

misidentification of punctipennis Meigen. 
Tanypus punctipennis Meigen; Morrissey, 1950: 90, 

phenology; misidentification of punctipennis Meigen. 

Description of this new species is in press 
in Proc. U.S. Natl. Mus. 

Larva: Described from exuviae of reared 
adults. Head pale yellowish, with tips of 
mandibles, lingula support, and posterior 
border of the head blackish; head length 
0.54 mm. Lingula (Fig. 1) yellowish; super- 
lingulae (Fig. 1) colorless. Labium (Fig. 
2) dark brown. Antenna, Figure 3; mandi- 
ble, Figure 4. Maxillary palpus 2.5 times as 
long as wide. Body with numerous hairs. 
Preanal papillae about 5 times as long as 
wide, slightly curved and colorless; each 
bears 12 long, colorless bristles; a heavy 
seta on anterior face. Anal prolegs with 
about 12 long yellowish claws which are 


No. 4 


only slightly curved and which lie parallel 
to one another. 


Pupa: Described from exuviae of reared 
adults. Pale yellowish except for blackish 
respiratory organs. Total length 6.98 mm. 
Length of pupal respiratory organs 0.68 
mm. Pupal respiratory organ as shown by 
Malloch (1915a, Pl. XXVI, figure 13) ex- 
cept that the long hairs shown by him are 
not visible on my specimens. Abdominal 
tergites covered with a dense uniform 
shagreen. Laterally on each tergite are two 
sets of bristles, the alveoli of which are 
surrounded by a brownish spot; the anterior 
pair is anterolateral in position; the posterior 
set is more medial, in a posterolateral posi- 
tion. Lateral margins of all segments ciliate 
with a fringe of long colorless bristles which 
become finer and denser on posterior seg- 
ments. Anal lobe as figured by Malloch 
(1915a, Pl. XXVI, figure 4). 

The type series was, in part, taken from 
Cane River Lake and will not be listed here. 


Additional material examined: One male, 
7-IX-56; 1 female, 11-III-57; 1 male, 23- 
MS 7ael male, 15-VIl-57: 4 males, 2° fe- 
males, 6-VIII-57; C.R.L. One female, 28- 
X-54; 3 males, 3-II-57; 2 males, 4-JI-57; 1 
male, 9-II-57; Ch.L. Three males, 1 female, 
£5159: US.F.H: 


The larva keys to stellatus in Johannsen 
(1937a, page 19), but appears to differ in 
the form of the paralabial plates (cf. Jo- 
hannsen 1937a, figure 45, Pl. IV). 


The pupa keys in Johannsen (op. cit.) to 
punctipennis Meigen but differs in that the 
eighth segment has only the fringe of finer 
bristles. 


PROCLADIUS (PROCLADIUS ) 
species | 

Procladius culiciformis, American authors, nec Linné 
(Part?). 

Procladius culiciformis (Linné); Darby, 1962: 37, 
BiSeee So mrs Ono SEE Oe zee Olen OOF hO> 
113, 114, 121-128; description of larva, pupa, and 
adult; ecology; misidentification of culiciformis 
(Linné). 


new 


Description of this new species is in press 
in Proc. U.S. Natl. Mus. 

Males: Wing length 2.61, 2.70 mm (2); 
leg ratio 0.74-0.78, mean 0.75 (3); an- 
tennal‘ratio 2.05, 2.36 (2). 

Additional material examined: One fe- 
male, 7-II-56; Ch.L. One male, 21-IV-58; 


Chironomids of West-Central Louisiana 


111 


C.R.L. One male, 9-IV-57; 1 male, 16-IV- 
57; at light, Natchitoches, La. 


PROCLADIUS (PSILOTANYPUS) 
BELLUS (Loew ) 


Tanypus bellus Loew, 1866; 4, description of adult. 

Procladinus bellus (Loew); 1905: 128, 
redescription of adult (after Loew). 

[Procladius] bellus [(Loew) ]; Johannsen, 1908: 270, 
subfamily position; genus indeterminate. 

Protenthes bellus (Loew); Malloch, 1915a: 388, de- 
scription of larva, pupa, and adult. 

Procladius bellus (Loew); Johannsen, 1937a: 23, re- 
description of larva and pupa, after Malloch. 
Procladius bellus (Loew); Judd, 1949: 8, phenology. 
Procladius bellus (Loew); Morrissey, 1950: 90, phe- 

nology; ecology. 
Procladius bellus (Loew); Johannsen (In Johannsen 
and Townes), 1952: 10, 11, adult, in key. 


Johannsen, 


Procladius bellus (Loew); Judd, 1953: 813, phe- 
nology. 

Procladius bellus (Loew); Sublette, 1957: 381, 
ecology; phenology. 

Procladius bellus (Loew); Judd, 1957: 400, phe- 


nology. 
Procladius bellus (Loew); Roback, 1957c: 48, larva 

and pupa, in key. 
Procladius belius (Loew): 

distribution. 
Procladius bellus (Loew) ; 

lowing pages, ecology. 
Procladius bellus (Loew); Judd, 1961: 96, phenology. 

Males: Wing length 1.53-2.04, mean 1.72 
mm (4); leg ratio 0.67-0.73, mean 0.74 
(4); antennal ratio 1.52-1.84, mean 1.69 
(ee 

Material examined: Five males, 2-III-57: 
1 male, 5 females, 11-III-57; 2 males, 1 fe- 
male, 12-III-57; 4 males, 16-III-57; 1 male, 
28-III-57; 2 males, 5-[V-57; 1 male, 8-IV- 
57; 1 male, 9-IV-57; 1 female, 12-IV-57; 1 
male, 6-V-57; 1 male, 1 female, 14-V-57; 
1 male, 15-V-57; 1 male, 16-V-57; 3 males, 
1 female, 21-V-57; 1 male, 3-VI-57; at light, 
Natchitoches, La. One female, 11-VI-57; 1 
male, 19-VI-57; 1 male, 1 female, 25-VI-57; 
1 male, 18-VII-57; 6 males, 4 females, 13- 
VIII-57; 1 male, 10-[X-57; 2 males, 3-X- 
572) deamale, axon I imales I4xe57- eh 
male, 19/21-III-58; 1 male, 5-V-58, 1 male, 
12-V-58; C.R.L. One female, 20-VIII-58; 
WES 


Beck and Beck, 1959: 91, 


Davis, 1960: 71, and fol- 


COELOTANYPUS TRICOLOR (Loew ) 


Tanypus tricolor Loew, 1861: 309, description of 
adult. 

Procladius tricolor (Loew); Johannsen, 
redescription of female, after Loew. 

[Coelotanypus] tricolor [ (Loew) ]; Johannsen, 1908: 
270, subfamily position; genus indeterminate. 

Coelotanypus tricolor (Loew); Malloch, 1915a: 396, 


adult. 


1905: 130, 


Piz 


Coelotanypus tricolor (Loew); Johannsen, 1926: 
generic position. 

Coelotanypus tricolor (Loew); Johannsen, 1934: 348, 
generic position. 

Coelotanypus tricolor (Loew); Johannsen, 1937a: 25, 
generic position. 

Coelotanypus tricolor (Loew); Johannsen (in Johann- 
sen and Townes), 1952: 11, adult in key. 

Coclotanypus tricolor (Loew); Sublette, 1957: 382, 
ecology; phenology. 

Coelotanypus tricolor (Loew); Roback, 1957a: 1-2, 
description of larva. 

Coelotanypus tricolor (Loew); Beck and Beck, 1959: 
91, distribution of adult. 


27.5)5 


Female: Wing length 3.24 mm; leg ratio 


0.66. 


Material examined: One female, 11-[X- 
HO-ar light GRE 


COELOTANYPUS SCAPULARIS 
( Loew ) 


Tanypus scapularis Loew, 1866: 2, description of 
adult. 

Procladius scapularis (Loew); Johannsen, 1905: 134, 
description of adult. 

Procladius scapularis [(Loew)]; Johannsen, 
270, generic position. 

Procladius scapularis (Loew); Malloch, 1915a: 393, 


description of adults. 


1908: 


Coelotanypus scapularis (Loew); Johannsen, 1934: 
348, generic position. 

Coelotanypus scapularis (Loew); Johannsen, 1937a: 
25, generic position. 

Procladius scapularis (Loew); Adams, 1940: 127, 


distribution. 

Coelotanypus scapularis (Loew); Morrissey, 1950: 89, 
distribution; phenology. 

Coelotanypus scapularis (Loew); Johannsen (in 
Johannsen and Townes), 1952: 12, adult, in key. 

Coelotanypus scapularis (Loew); Neff, 1955: 7, 
adult; ecology. 

Coelotanypus scapularis (Loew); Sublette, 1957: 382, 
ecology; phenology. 

Coelotanypus scapularis (Loew) ; 
1959: 91, distribution of adult. 


Male: Wing length 2.49 mm; leg ratio 
0.70; antennal ratio 3.05. 


Material examined: One male, 11-IX-56; 
at light, C.R.L. 


Beck and Beck, 


COELOTANYPUS CONCINNUS 


(Coquillett ) 


Tanypus concinnus Coquillett, 1895: 
tion of adult female. 

Procladius concinnus (Coquillett); Johannsen, 1905: 
129, redescription of female. 

Procladius concinnus (Coquillett); Johannsen, 1908: 
270, generic position. 

Procladius concinnus (Coquillett); Malloch, 1915a: 
394, description of larva, pupa and adult. 


308, descrip- 


Coelotanypus  concinnus  (Coquillett); Johannsen, 
1934: 348, generic position. 
Coelotanvbus  concinnus  (Coquillett); Johannsen, 


Tulane Studies n Zoology 


Vol. 11 


1937a: 25, description of larva and pupa. 

Coelotanypus  concinnus  (Coquillett); Morrissey, 
1950: 89, distribution and phenology. 

Coelotanypus concinnus (Coquillett); Johannsen (in 
Johannsen and Townes), 1952: 12, adult, in key. 

Coclotanypus concinnus (Coquillett); Roback, 1953: 
108, ecology. 

Coelotanypus concinnus (Coquillett); Wurtz and 
Roback, 1955: 199, distribution; ecology. 

Coelotanypus  concinnus (Coquillett); Paine and 
Gaufin, 1956: 296, ecology. 

Coelotany pus concinnus (Coquillett) ; Roback, 1957a: 
47, ecology. 

Coclotanypus concinnus (Coquillett); Sublette, 1957: 
382, ecology; phenology. 

Coelotanypus concinnus (Coquillett); Beck and Beck, 


1959: 91, distribution of adults. 

Coelotanypus concinnus (Coquillett); Dendy and 
Sublette, 1959: 510, adults. 

Coelotanypus concinnus (Coquillett); Davis, 1960: 


71, ecology. 


Male: Wing length 3.27 mm (1); leg 
ratio 0.76 (1); antennal ratio 2.88 (1). 

Females: Wing length 3.06-3.74, mean 
3.50 mm (3); leg ratio 0.63-0.70, mean 
O67 3): 

Material examined: One female, 18-V-54; 
2 females, 29-IV-57; 1 male, 1 female, 14- 
V-57; 1 female, 15-V-57; 1 female, 21-V-57; 
at light, Natchitoches, La. 


PENTANEURA (PENTANEURA) 
PLANENSIS Johannsen 

Pentaneura nigropunctata (Staeger); Hauber, 1945: 
§02, description of adult (genitalia); phenology; 
misidentification of nigropunctate (Staeger). 

Pentaneura planensis Johannsen, 1946: 282, 
description of adult. 

Pentaneura planensis Johannsen; Morrissey, 1950: 88, 
description of pupa and adult; phenology; syn- 
onomy. 

This species places in the genus Larsia 
Fittkau. For reasons listed earlier, I am 
using the more inclusive taxon, the genus 
Pentaneura, in a broad sense. 

Males: Wing length 1.78-1.80, mean 1.79 
mm (4); leg ratio 0.77-0.82, mean 0.79 
(4); antennal ratio 1.30-1.60, mean 1.45 
(4); 

Material examined: One male, 13-VIII-57; 
1 male, 9-VII-58; C.R.L. Three males, 12- 
IX-58; 4 males, 27-[X-58; U.S.F.H. 


PENTANEURA (PENTANEURA) 
PILOSELLA (Loew) 


Tanypus pilosellus Loew, 1866: 5, description of adult. 

Tanypus pilosellus Loew; Malloch, 1915a: 372, de- 
scription of ?larva and pupa; adult. The descrip- 
tion of the pupa does not agree with my material. 

Tanypus pilosellus Loew; Walley, 1928: 583, adult, 
in key. 

?Pentancura pilosella (Loew); Johannsen, 1937a: 13, 
redescription of larva and pupa, after Malloch. 


284, 


No. 4 


Pentaneura pilosella (Loew) ; 
description of adult. 

Penteneura pilosellus (Loew); Johannsen, 1946: 282- 
283, adult, in key; description of adult. 

Pentaneura pilosella (Loew); Beck and Beck, 1959: 
91, distribution of adults. 


This species was placed by Fittkau (1962 ) 
in his genus Labrundinia, and with this 
placement I concur. For reasons given earlier 
I am not employing Fittkau’s generic units 
but rather a broader, more inclusive concept 
of the genus Pentaneura. From Fittkau’s ex- 
haustive description of Labrundinia longt- 
palpis (Goetghebuer) I suspect that it may 
be synonymous with pélosella (Loew). I 
hesitate to synonomize it, however, without 
examining European material. Dr. Fittkau 
has examined my material and agrees that 
only a critical comparison of material from 
Europe and North America can resolve the 
specific identities. 

Males: Wing length 1.26-1.36, mean 1.32 
mm (3); leg ratio 0.60-0.80, mean 0.68 
(3): antennal ratio 1.21-1.30, mean 1:24 
): 

Material examined: One male, 23-VII- 
57; 1 male, 13-VIII-57; 3 males, 20-VIII-57; 
1 male, 2 females, 27-VIII-57; 1 male, 17- 
Peay teimale, 7-X-5/7; 1 male, 19-V-58; 
1 male, 10-VI-58; 1 male, 19-VI-58; C.R.L. 
One male, 12-VIII-58; 1 male, 20-VIII-58; 
30 males, 2 females, 28-VIII-58; 5 males, 
6-IX-58; 35 males, 12-IX-58; 9 males, 20- 
IX-58; 16 males, 27-IX-58; 5 males, 4-X-58; 
6 males, 15-X-59; U.S.F.H. 


Hauber, 1945: 502, 


ABLABESMYIA AEQUIFASCIATA 


(Dendy and Sublette), new 
combination. 


Pentaneura (Ablabesmyia) aequifasciata Dendy and 
Sublette, 1959: 507, description of adult. 

Pentaneura (Ablabesmyia) aequifascista Dendy and 
Sublette; Darby, 1962: 37, 40, 41, 58, 73, 76, 101, 
116-121, description of larva, pupa, and adult; 
ecology. 


Males: Wing length 2.84, 2.84 mm (2); 
foreleg ratio 0.78, 0.80 (2); antennal ratio 
2.16, 237 (2); fore tibial band 0:54 (1); 
basitarsal band 0.40 (1). 

Material examined: One male, 28-III-57; 
1 male, 5-IV-57; light trap, Natchitoches, La. 


ABLABESMYIA PELEENSIS (Walley ) 


peleensis Walley, 1926: 64, description of 
1928: 585, 590, adult, in key; notes on 


Tanypus 
adult; 
adults. 

Tanypus peleensis Walley; Adams, 1940: 126, distri- 
bution. 


Chironomids of West-Central Louisiana 


WS 


Pentaneurs peleensis (Walley); Hauber, 1945: 496, 
499-500, description of pupa; adult, in key. 

Pentaneurs peleensis (Walley); Johannsen, 1946: 270, 
274, redescription of adult; distribution. 

Pentaneura peleensis (Walley); Roback, 1957c: 41, 
description of larva and pupa. 

Pentaneura (Ablabesmyia) peleensis (Walley); Dendy 
and Sublette, 1959: 508, adult, in key. 

Pentaneura (Ablabesmyia) peleensis (Walley); Ro- 
back, 1959: 122, added description of adult. 


I have compared my material with para- 
types from the Canadian National Collection 
kindly loaned to me by Dr. J. R. Vockeroth. 

Males: Wing length 2.48-2.59, mean 2.52 
mm (3); leg ratio 0.77-0.84, mean 0.81 
mm (3); antennal ratio 2.27-2.50, mean 
2.35 (3); fore tibial band 0.47; basitarsal 
band 0.37. 

Material examined: Two males, 18-V-54, 
Natchitoches, La. One male, 14-X-57; 1 
male, 21-IV-58; C.R.L. One male, 12-IX-58; 
1 male, 1 female, 27-[X-58; 1 male, 6-XI-58; 
WES:E ae 


ABLABESMYIA ILLINOENSIS 
(Malloch ) 


Tanybus illinoensis Malloch, 1915a: 376, description 
of pupa and adult. 

Tanypus illinoensis Malloch; Walley, 1925: 272, 
adult, in key; 1928: 585, 589, adult, in key; dis- 
tribution. 

Pentaneura illinoensis (Malloch); Johannsen, 1937a: 
12, redescription of pupa. 

Pentaneura illinoensis (Malloch); Miller, 1941: 19, 
and following pages; ecology. 

Pentaneura illinoensis (Malloch); Hauber, 1945: 496, 
adult, in key. 

Pentaneura illinoensis 
270, 273, adult. 

Pentaneura  illinoensis (Malloch); Johannsen (in 
Johannsen and Townes), 1952: 6, adult, in key. 

Pentaneura illinoensis (Malloch); Paine and Gaufin, 
1956: 295, ecology. 


(Malloch); Johannsen, 1946: 


Pentaneura illinoensis (Malloch); Judd, 1957: 399, 
phenology. 
Pentaneura illinoensis (Malloch); Beck and Beck, 


1959: 90, distribution of adult. 

Pentaneura (Ablabesmyia) illinoensis (Malloch) ; 
Dendy and Sublette, 1959: 508, adult, in key. 
Pentaneura (Ablabesmyia) illinoensis (Malloch); Ro- 

back, 1959: 121, redescription of adult. 

Males: Wing length 2.48-3.02, mean 2.75 
mm (4); foreleg ratio 0.76-0.83, mean 0.79 
(4); antennal ratio 2.20-2.52, mean 2.31 
(3); fore tibial band 0.54; basitarsal band 
0.38. 

Material examined: One female, 12-III- 
57; 2 males, 16-III-57; 1 female, 5-IV-57; 
1. male, 8-IV-57; 1 male, 6-V-57; 1 male, 
1 female, 21-V-57; 1 female, 16-VII-57; light 
trap, Natchitoches, La. One male, 2 females, 
15-X-58; 1 male, 6-XI-58; U.S.F.H. 


114 


ABLABESMYIA RHAMPHE 
new species 
Pentaneura basalis Walley?; Sublette, 1957: 38, larval 
ecology; misidentification of basalis Walley. 
?Tanypus sp. A, Malloch, 1915a: 397, larva. 

Holotype male: U.S.N.M., No. 66454, 
collected at light, Natchitoches, Louisiana, 
16-VI-57, by James E. Sublette. 

Postocular bristles partially in two rows; 
palpal proportions 20:20:20:42 (paratype) ; 
antennal ratio 2.00. Head, including anten- 
nal pedicels, and thorax dark brown over- 
lain with a conspicuous greenish pruines- 
cence; prescutellum shining; halteres pale. 
Wing length 1.89 mm; venarum ratio 0.78; 
prothorax with about 12 fine lateral bristles; 
supra-alar bristles 2; prealar bristles, 1 row 
staggered, becoming 2 rows and dividing 
around presuctellar area to join dorsolateral 
bristles where the row then appears doubled 
back to scutellum. Scutellar bristles about 
40, 12 of which are large and erect, forming 
a straight transverse row, the remainder 
more or less strewn anteriorly. Dorsolateral 
bristles in one row. Anterolateral bristles 12. 

Fore tarsus not bearded; legs banded; 
white bands of legs wider than narrow 
brown bands which they separate; middle 
brown band of fore tibia about equidistant 
between basal and apical bands (0.53); fore 
femur with a broad basal brown band and 
a narrow apical one; mid and hind femora 
with only the apical one; remainder of seg- 
ments on all legs banded as follows: tibia 
with three bands; basitarsus with one band 
slightly before center (0.40) and one at 
apex; Ta, to 4, each with an apical band 
only; Ta; entirely dark. Tibial spur length 
and ratio of spur length to diameter of apex 
of tibia: foreleg, 0.03 mm, 8:8; middle leg, 
0.04 mm, 9:8; hind leg, 0.04 mm, 10:9. 


Leg proportions: 


Leg 

Eetietat as) 2s 4. 5 tatio 

Foreleg AP 50) 40) 28 19) 12" 7 0.80 

Middleleg 45 438 38 20 15 9 8 0.88* 

Hind leg 43° 55> 49° 27 19 11 6 0:89 
* (paratype) 


Wings patterned very much like aspera 
Roback (cf. Roback, 1959, pl. XIV, figure 
1) except that the spots are heavier and 
broader and with those under Rs 13, sub- 
basal in M, at end of Cu; and in cell Cu, 
confluent at edges. 

Tergite I of abdomen largely pale, ter- 
gites II-V with an anterior brown transverse 


Tulane Studies 1m Zoology 


Vol. 11 


fascia, which occupies one-half of segment 
Il becoming wider posteriorly; tergites VI- 
VII almost entirely dark; genitalia dark. 
The basistyle of the genitalia has a con- 
spicuous outward bulge which led me 
(1957) to identify this species as basalis?. 
At that time I observed that the species 
differed from basalis in some particulars, 
hence the query. The genitalia most closely 
resemble johannsent Roback but differ sig- 
nificantly in lacking serrations along the ac- 
cessary blade (Fig. 5, aedeagus in normal 
position; Fig. 6, everted position). 
Allotype: In U.S. National Museum; col- 
lected in a tent trap, C.R.L., Natchitoches 
Parish, Louisiana, 10-VII-57; B. R. Buckley. 
Colored as male. Antennal flagellum pale 
except terminal segment which is dark and 
bears a terminal bristle; proportions of seg- 
ments: 13:6:6:6:7:7:6:6:6:8:19 (paratype). 
Ratio of last segment to remainder, 0.26. 
Postocular bristles in 2 rows. Palpal propor- 
tions 13:15:13:30. Clypeus densely haired 
with about 45-50 bristles. 
Prothorax with 14 fine lateral bristles. 
Wing length 1.58 mm; supra-alar bristles 2; 
prealar bristles 25; dorsomedial and dorso- 


lateral bristles as in male; anterolateral 
bristles 25. 
Leg proportions: 
& proportio Leg 


F Ti Tay oo 9 Sa ere 
Foreleg 35942) 355200 15) 0 aeOrss 
Middlelee 45 40 36 20 18 9 5 0.90 
Hind leg 40) B00) = = = = = (ister 
* (paratype) 


Wings heavily haired; wing spots more 
diffuse and coalesced than in male. 

Abdomen fasciate, the bands becoming 
progressively broader posteriorly. Genitalia 
with very small lamellae (Fig. 7); 3 sperma- 
thecae, large and oval; blackish-brown ex- 
cept at junction of duct (Fig. 8). 

The species keys in Roback (1959, p. 
120) to couplet 6 where it can be distin- 
guished from janta Roback by lacking a 
brush and from johannseni Roback by lack- 
ing serrations. 

Paratypes: Wing length 1.62-2.12, mean 
1.85 mm (9); leg ratio 0.76-0.85, mean 0.81 
(4); antennal ratio 1.96-2.35, mean 2.16 
(75 

Paratypes examined. One male, 3-1V-57; 
1 male, 16-VI-57; 1 male, 25-VI-57; 2 males, 
10-VII-57; 1 male, 11-VII-57; 1 male, 18- 
VII-57; 2 males, 2 females, 23-VII-57; 1 


No. 4 


male, 25-VII-57; 1 male, 1 female, 26-VII- 
57; 1 female, 27-VII-57; 2 females, 29- 
VII-57; 2 males, 1 female, 30-VII-57; 1 
male, 6-VIII-57; 1 male, 1 female, 13-VIII- 
oye Pomale, 17-[X-57; 1 male; 14-x-57: 1 
male, 12-V-58; 1 male, 4-VI-58; C.R.L. One 
male, 4 females, 28-VI-57; light trap, Natchi- 
toches, La. Five males, 14-V1-62; University 
of Oklahoma Biological Station, Willis, 
Oklahoma. In the collections of U.S.N.M., 
Cornell University, INAS, ANSP. 
C.N.C., and Florida State Board of Health. 


Larva: Described from exuvia of reared 
specimen. Head length 0.72 mm; width, 
0.44 mm. Mandible (Fig. 9) length 0.12 
mm; mandible:head length 0.17. Antenna 
(Fig. 10) length 0.38 mm, antenna:head 
length 0.52. Head entirely yellow except for 
lingula (Fig. 11) and posterior margin of 
occiput. Maxillary palpus, Figure 12. 


Posterior prolegs each with two dark 
curved hooked claws: with one brown 
hooked claw; and with 12 slightly curved 
yellow claws. Preanal papilla about 0.41 as 
wide as long; each with 7 bristles. 

The larva keys in Roback (1957c) to 
couplet 11 of the key to Pentaneura (page 
29). It may be distinguished from peleenszs 
Walley by having a lower antennal ratio 
(about 4.5:1) and from monzilis (Linné) 
Johannsen (americana Fittkau) by the 
basal two maxillary palpal segments being 
subequal. 


Pupa: Described from exuvia of reared 
female and from male pupa with visible 
genitalia. Pupa length 4.05 mm; exuviae 
length 3.33 mm. Entirely pale yellow except 
for dark respiratory organs (Fig. 13) which 
are 0.36 mm long. Seventh segment with 4 
lateral flattened filaments which have the 
following positions in terms of distance from 
the anterior margin of the segment: first, 
0.41; second, 0.56; third, 0.75; fourth, 0.96. 
Eighth segment with 5 lateral flattened fila- 
ments which have the following positions in 
terms of distance from the anterior margin: 
first, 0.28; second, 0.49; third, 0.70; fourth, 
0.85; fifth, 1.00. Swim fin, Figure 14. 

The pupa keys in Roback (op. cit.) to 
couplet 11 then no longer fits the key well; 
it appears to be distinctive in the structure 
of the respiratory organ (Fig. 13). 


Chironomids of West-Central Louisiana 


115 


SUBFAMILY ORTHOCLADIINAE 
CRICOTOPUS BICINCTUS (Meigen) 


Chironomus bicinctus Meigen, 1818: 41, description 
of adult. 

Cricotopus bicinctus (Meigen) ; Johannsen, 1905: 256, 
redescription of adult after van der Wulp, 1874: 
132% 

Trichocladius Kieffer; Potthast, 
243, description of immature stages. 

Cricolopus bicinctus (Meigen); Malloch, 1915a: 505 
redescription of adult; distribution. 

Cricotopus bicinctus (Meigen); Edwards, 1929: 321 
redescription of adult; synonomy. 

Cricotopus bicinctus (Meigen); Goetghebuer, 
29, 34, description of adult; distribution. 

Cricotopus bicinctus (Meigen); Tokunaga, 1936: 16 
redescription of adult. 

Cricotopus bicinctus (Meigen); Johannsen, 
54, redescription of immature stages. 

Cricotopus bicinctus (Meigen); Miller, 
and following pages, ecology. 

Trichocladius bicinctus (Meigen); Brundin, 1949: 
461, 497, 498, 506, 728; ecology; distribution. 

Cricotopus bicinctus (Meigen); — Johannsen (in 
Johannsen and Townes), 1952: 18, adult, in key. 

Trichocladius bicinctus (Meigen); Thieneman, 1954: 
Uti WS 252), 266; 2E75 25 32P5 342: 35), 360, 
367, 369, 370, 456, 459, 461, 468, 477, 510256 01S 
511, 525, 592; biology. 


eterimanus LOSS 


> 
1932: 
1937a: 


1941: 19, 


Cricotopus bicinctus (Meigen); Paine and Gaufin, 
1956: 295, ecology. 

Cricotopus bicinctus (Meigen); Judd, 1957: 400, 
phenology. 

Cricotopus bicinctus (Meigen); Mundie, 1957: 164, 
ecology. 


Cricotopus bicinctus (Meigen); Roback, 1957c: 7A 
redescription of larva; distribution. 
Cricotopus bicinctus (Meigen); Gaufin, 

ecology. 
Cricotopus bicinctus (Meigen); Beck and Beck, 1959: 
91, distribution of adults. 


1958: 205, 


Cricotopus bicinctus (Meigen); Surber, 1959: 111, 
ecology. 

Cricotopus bicinctus (Meigen); Judd, 1960: 207, 
phenology. 


Males: Wing length 1.58-1.94, mean 1.71 
mm (3); leg ratio 0.53-0.63, mean 0.57 
(6) = antennal rato 4. e520) (2), 

Material examined: One male, 3-III-57: 3 
females, 6-V-57: at light, Natchitoches, La. 
One male, 21-IV-58; 1 male, 5-V-58: C.RLL. 


CRICOTOPUS REMUS new species 
Cricotopus tricinctus (Meigen); Sublette, 1957: 384, 

ecology; phenology; misidentification of fricinctus 

(Meigen). Occurrence of fricincfus in the 

Nearctic region is doubtful. 

Holotype male: U.S.N.M. No. 66455; 
collected in a funnel trap set in 0.5 meters 
of water, C.R.L., Natchitoches Parish, Lou- 
isiana, 21-IV-58, B. R. Buckley. 

Antennal pedicel dark brown, antennal 
flagellum pale brown; postocular bristles in 
a single staggered row; proportions of palpal 


116 Tulane Studies in Zoology Volt 


Figures 1-4. Tanypus n. sp. 2. 1. lingula and supralingula of larva; 2. labium; 3. anten- 
na; 4, mandible. Figures 5-14. Ablabesmyia ramphe new species. 5. male genitalia, 
aedaegus in normal position; 6. aedaegus in everted position; 7. female genitalia; 8. 
spermatheca; 9. mandible of larva; 10. antenna; 11. lingula; 12. maxillary palpus; 13. 


pupal respiratory organs; 14. swim fin. 


No. 4 


segments, 5:7:12:20; antennal ratio 1.27. 

Prothorax entirely yellow; broad, almost 
parallel-sided, with 3 fine lateral bristles. 
Mesothorax ground color yellow separated 
by the dark brown vittae; prescutellum yel- 
low; scutellum brown; postnotum dark 
brown; sternopleuron  blackish-brown on 
venter; halteres pale. Wing length 1.69 mm; 
prealar bristles 4, 2 large ones posteriorly, 
2 small ones anteriorly; dorsomedial and 
dorsolateral bristles minute, suberect; scu- 
tellar bristles 2, large, erect. 


Forelegs with femur dark on distal one- 
third; tibia on basal one-third, and distal 
one-tenth; tarsi entirely dark. Middle legs 
with femur dark on distal one-third; tibia, 
basal one-third; basitarsus largely pale, sec- 
ond tarsal joint dark on distal one-third. 
Hind legs with femur dark on distal one- 
third; tibia, basal one-sixth; tarsal joints 1 
and 2 pale; 3 dark only on distal one-fourth, 
4 and 5 entirely dark. Single long slender 
spur on fore tibia 1.23 times as long as 
apical diameter of tibia. Spurs of middle 
tibia very short, of equal length. Hind leg 
with inner spur 2.25 times length of outer; 
tibial comb with 18 bristles; row tapered 
toward center. 


Leg proportions: 
Bue Nay. 5: sa os 


o 


Leg 
; ratio 


Foreleg 77100 50 30 20 18 10 0.50 
Middleleg 76 80 35 20 15 10 10 0.44 
Hind leg 80 92 47 25 20 If 11 0-51 


Wings: C slightly produced; R415 ter- 
minates proximal to M; f-Cu distal to r-m; 
An reaches middle of Cus. 

Abdomen illustrated in Figure 15. 

Genitalia illustrated in Figure 16. The 
broad spatulate dististyle is distinctive among 
American Cricotopus. 

Allotype: In U.S. National Museum. Col- 
lected C.R.L., 4-VI-58, B. R. Buckley. 

Antennal proportions, 11:7:6:6:17. Ratio 
of terminal segment to remainder, 0.71. Pal- 
pal proportions, 8:12:15:26. Wing length 
135mm. 


Leg proportions: 
Fae Ti sans eae 


Leg 
= ratio 


4 
Foreleg 55 70 380 -—- - — — 0.43 
Middleleg 60 55 29 12 10 7 9 0.58 
Hind leg 55 6b 32 1b 14> 7 + 9 0:50 


Abdomen similar to male except for sex- 
ual differences. Color pattern less intense 


Chironomids of West-Central Louisiana 


117 


than male so that dark bands are incomplete 
on all segments leaving a pale fascia anterior 
and posterior to each main dark fascia; en- 
tire abdomen appearing thus vittate. Sperma- 
theca (Fig. 17); genitalia (Fig. 18). 

Paratypes: Wing length 1.46-1.58, mean 
1.54 mm (3); foreleg ratio 0.48-0.53, mean 
0.51 (3); middle leg ratio 0.39-0.41, mean 
0.40 (3); hind leg ratio 0.50-0.52, mean 
0.51 (3); antennal ratio 1.20-1.33, mean 
EZE=()e 

Paratypes examined: One female, 2-II-57; 
1 female, 7-II-57; 2 females, 12-IV-57: 1 
female, 16-IV-57; 2 males, 2 females, 29- 
IV-57; 4 females, 6-V-57; 1 male, 2 females, 
21-V-57; 1 male, 3-VI-57; 4 males, 6 fe- 
males, 14-IV-58; 12 males, 27 females, 21- 
IV-58; 1 male, 5 females, 5-V-58; 1 male, 
3 females, 30-IV-58; at light, Natchitoches, 
La. In the collections of U.S.N.M., C.N.C., 
A.NS.P., I.N.H.S., Cornell University and 
the Florida State Board of Health. 


This species keys in Johannsen (in Jo- 
hannsen and Townes, 1952) to ¢trifasciatus 
(Panzer) or tricinctus (Meigen) depending 
upon the degree to which the femora are 
infuscate. Neither of these Palearctic species 
has been adequately described and separation 
of the species has been entirely on color 
characteristics (cf. Edwards, 1929, page 
319). Edwards (op. cit.) describes both spe- 
cies as having abdominal segments 1, 4 and 
7 mostly yellow; only the posterior margin 
of remus is yellow. The species tricinctus 
is described as having tergites, 2, 3, and 5 
entirely black, while ¢rifasciatus has 2, 3, 
and 5 narrowly yellow at base. In remus 
new species 2 and 5 are narrowly yellow 
but 3 is entirely dark. 


Larva: Described from exuvia of para- 
type male. Head, yellowish-brown; only oc- 
ciput, labial plate, tips of mandibles and pre- 
mandibles darker brown; head length 0.53 
mm. Labial plate (Fig. 19) and mandible 
(Fig. 20) very similar to that figured by 
Johannsen (1937) for Cricotopus trifaciatus 
(Panzer). Antenna (Fig. 21) 0.08 mm 
long. Mandible 0.16 mm long. In addition 
to the bristles shown posterior to the labial 
plate, the venter of the head bears two pairs 
of closely spaced bristles; one pair lateral to 
the last tooth of the labial plate and about 
one-third the width of the plate from the 
last tooth; the other pair almost at the mid- 
dle of the head near the lateral margin. In 


118 


each set the two bristle alveoli are contig- 
uous. Venter of labrum and epipharyngeal 
area of typical pattern (cf. Fig. 37 for ter- 
minology which follows): near the anterior 
margin are two strong bristles; seta III, 
flanked on either side by shorter bristles; 
posterior to seta III and slightly medial are 
two strong bifurcate bristles, seta II; lateral 
to these on each side is a dense clump of 
10 to 12 moderately strong bristles, the chae- 
tae. The pecten epipharyngis is formed from 
three large, triangular, blunt teeth; below 
these on each side are three long slender 
teeth, the chaetulae basales. Premandible 
(torma) bifurcate, yellowish-brown at tips. 

The body bears characteristic hair pencils 
(Johannsen, 1937a, Pl. XV, figure 184). 
The preanal papillae are about as long as 
broad, each is blackish-brown on posterior 
face; on the anterior surface is a fine, pale 
bristle; each papilla bears six long yellowish- 
brown terminal bristles. Posterior prolegs 
with 14-16 slender, gently curved hooks. 

Pupa: Described from exuvia of paratype 
male. Exuvia length 4.13 mm; pale yellow- 
ish, slightly darker on lateral margin of last 
three segments. Respiratory organs (Fig. 
22) gradually tapering, 0.24 mm long, with- 
out spinules, slightly darker than remainder 
of integument. Anterior to respiratory or- 
gans are three bristles, two anterior large 
ones and a much smaller posterior one. 
Dorsum of thorax strongly papillose. Ab- 
dominal chaetotaxy very similar to other spe- 
cies of Cricotopus. Tergites I to VI with 
a more or less continuous field of coarse 
shagreen; intertergal membrane between III 
and IV, IV and V, and V and VI, with fine 
shagreen in multiple rows. Tergites VII and 
VIII not shagreened. Anal lobes with three 
short, heavy, almost straight bristles. Pos- 
terolateral margins of sternites TV-VII with 
a patch of spinules; sternites 1V-VIHI with 
sparse fine shagreen which becomes pro- 
gressively denser towards posterior segments 
so that sternites VI and VUHI are almost 
completely shagreened. 

The larva keys in Roback (1957c, pages 
68, 69) to couplet 8, where it cannot be dis- 
tinguished from C. sylvestris, trifasctatus, or 
tricinctus because of inadequate descriptions. 

The pupa keys to couplet 11 (loc. cit., page 
70) where, as in the larval stage, it cannot 
be distinguished from C. ¢rifasciatus, syl- 
vestris, or tricinctus due to inadequacies of 
descriptions of those species. 


Tulane Studies in Zoology 


Vol. 11 


CRICOTOPUS LEBETIS new species 


Cricotopus tricinctus (Meigen), American authors, 


in part. 

Holotype male: U.S.N.M., No. 66456. 
Collected at U. S. Fish Hatchery, Natchi- 
toches, Louisiana, 29-X-58, Robert F. Tyler. 

Head, thoracic vittae, sternopleuron, scu- 
tellum, postnotum and abdominal tergites 
II, Ill, V, VI and VIII blackish-brown; 
pronotum, narrow mesothoracic ground col- 
or and abdominal tergites 1, IV and VII 
yellowish. Antennae and mouth parts black- 
ish. Halteres pale. Coxa of foreleg pale, 
that of middle and hind leg black; trochant- 
ers of all legs yellowish; all femora dark 
brown becoming still darker apically; fore 
tibia dark brown basally and apically, paler 
brown in the middle; middle and hind tibiae 
darkened apically and basally, infuscate 
white in the middle; fore tarsi entirely dark; 
middle tarsi paler brown; hind tarsi paler 
brown. Fore tarsi with hairs no more than 
2 times tarsal diameter. Empodium as long 
as Claws; pulvilli minute (clearly visible only 
at 430 magnification). Fore tibia with a 
single spur; middle tibia with two short sub- 
equal spurs; hind tibia with the usual comb 
and with two spurs, of which the outer is 
about one-half the length of the inner. Ratio 
of spur length to diameter of tibia: foreleg, 
8:8, middle leg, 3:8; hind leg (inner spur), 
8:9. 

Prothorax broad, almost parallel-sided to 
the apex where it projects slightly; halves 
distinctly notched; lateral margin with about 
8 fine bristles. Wing length 1.62 mm; no 
supra-alar bristles; 2 prealar bristles; dorso- 
medial and dorsolateral bristles minute, in 
a single row, depressed; scutellar bristles 8, 
moderate sized, erect. Chaetotaxy of ab- 
dominal tergite II shown in Figure 25. 


Leg proportions: 


Leg 

F Ti Ta, 5» 93 4 Renato 

Foreleg $2 43 22-11 "9 ‘o-aeone 
Middlelee 388 38 17 10 7 5 5 0.45 
Hindleg 37 45 25 12 10 6 5 0.56 


Wing structure appears to be the same as 
the allotype. 

Genitalia, Figure 23. The strong, almost 
right angled basal lobe on the basistyle as 
well as the shape of the dististyle seem dis- 
tinctive; dististyle in dorsal view, Figure 24. 

Allotype: In the U. S. National Museum 
Collection. Collected at light, Natchitoches, 
Louisiana, 6-V-57. 


No. 4 


Colored as the holotype male except that 
the dark areas are more intense; thoracic vit- 
tae fused completely so that mesothorax is 
almost solid black; pronotum also infuscate. 
Abdomen colored as male. Eyes reniform, 
more abbreviate dorsally. Legs dark, vittate 
appearance not discernible. Palpal propor- 
tions, 7:16:16:22; wing length 2.48 mm; 
venarum ratio 1.17; 4 prealar bristles; dorso- 
medial bristles apparently in a single row; 
dorsolateral bristles in a rather broad stag- 
gered row; scutellum with about 10 bristles 
in a slightly staggered row. Antennal pro- 
portions 12:8:8:8:22; ratio of terminal seg- 
ment to remainder, 0.41. 


Leg proportions: 


Leg 

Be Pas 25) eee a saadio 

Foreleg 40 47 25 15 11 8 5 0.58 
Middleleg 38 40 19 11 8 5 5 0.47 
Hind leg 35 43 24 12 10 6 5 0.56 


Wing membrane greyish-brown by trans- 
mitted light, veins dark brown. C produced 
beyond R4,5, terminating proximal to M 
and distal to Cuy. Ro.3 terminates at about 
the middle of the ends of R; and R4,;5. The 
anal vein terminates beyond f-Cu. 

Genital lamella, Figure 26. 

Paratype males: Wing length 1.31-1.44, 
mean 1.40 mm (3); leg ratio 0.51-0.52, 
mean 0.52 (3); antennal ratio 0.86-1.09, 
mean 0.97 (3). 

Paratypes: One male, 6-IX-58; 3 males, 
12-IX-58; 1 male, 4-X-58; 7 males, 29-X- 
58; 5 males,,6-XI-58; 1 male, 10-XI-58: 2 
males. 271-56: 3) males, 12-X11-58 3 
males, 10-I-59; U.S.F.H. In the collections 
of CN.C., A.N:S.P., I.N.H.S., and Cornell 
University. 

This species keys in Johannsen (in Jo- 
hannsen and Townes), 1952, to tricinctus 
Meigen. Separation of Jebetis from Pale- 
arctic material on the basis of color pat- 
terns is difficult. The genitalia as figured 
by Tokunaga (1936) is distinctively dif- 
ferent. 


PSECTROCLADIUS VERNALIS 
Malloch 


Psectrocladius vernalis Malloch, 1915a: 520, de- 
scription of adult. 
Psectrocladinus vernalis Malloch; Dendy and Sublette, 


1959: 513, added description of adult. 

Males: Wing length 1.71-2.25, mean 1.87 
mm (5); leg ratio 0.67-0.70, mean 0.69 
(5S); antennal ratio 1.00-1.26, mean 1.13 


Chironomids of West-Central Louisiana 


119 


Material examined: One male, 4-X-58; 
1 male, 10-X-58; 2 males, 15-X-58; 7 males, 
29-X-58; 41 males, 6-XI-58; 1 male, 26- 
XI-58; 2 males, 27-XI-58; 109 males, 12- 
XII-58; 2 males, 10-I-59; 1 male, 15-I-59; 
8 males, 25-I-59; 6 males, 5-II-59; 2 males, 
9-II-59; 9 males, 15-II-59; US.F.H. One 
male, 3-II-57; 1 male, 5-II-57; 1 male, 6- 
I-57; C._R.L. on Typha. One male, 3-III-58; 
1 mile south of Farmersville, La., borrow 
ditch. 


Larva: Described from exuvia of reared 
male. Head length, 0.50 mm; head capsule 
pale yellow except for tips of mandibles and 
labial plate which is dark brown and occiput 
which is dark yellow. Labial plate, Figure 
27; mandible, Figure 28; antenna, Figure 
29; ventral surface of labrum somewhat ob- 
scured; epipharyngeal apparatus, Figure 30; 
Sm apparently absent; seta I palmate deeply 
and coarsely incised; seta II, simple, slightly 
anterior and lateral to seta I; seta III, small 
and simple, medial to seta II; about 8 chae- 
tae; torma (premandible), Figure 31; max- 
illa with about 8 long medial blades; palpus 
about as long as wide; with about 8-10 low 
pointed scales medial to the palpus. Pos- 
terior end of exuvia lost. 


The larva does not fit well into Roback’s 
key (1957c, page 86), the most compre- 
hensive treatment of Nearctic species. The 
labial plate resembles that of the Palearctic 
species stratiotis Kieffer (Goetghebuer, 
1914, vide Roback, op. cit.) but differs in 
having the medial teeth distinctly less pro- 
jecting. 

Pupa: Described from exuvia of reared 
male. Length 4.5 mm. Entirely pale yellow; 
respiratory organs darker (Fig. 32). Cep- 
halic tubercles low conical, each with a 
terminal seta. Tergites II-VI with a pos- 
terior transverse band of coarse spines and 
spinulae; tergites IV-VI with a central disc 
of coarse spines, consisting of 7 spines on 
IV, 13 spines of V, and 13 spines of VI. 
Tergite IV is shown in Figure 33. Segments 
VI and VII have 4 lateral flattened bristles, 
2 anterior and 2 posterior; segment VII has 
5 flattened bristles, 2 anterior and 3 pos- 
terior. Swim fin with about 28 fringe 
bristles and, in addition, 3 heavy, posteriorly 
directed spines. No intersegmental spinulae. 

The pupa keys to elatws Roback in Ro- 
back’s key (op. cit., page 87). It may be 
distinguished by the structure of the respir- 


120 Tulane Studies in Zoology Vol. 11 


Figures 15-22. Cricotopus remus new species. 15. color pattern of male abdomen, lat- 
eral view; 16. male genitalia; 17. spermatheca of female; 18. genitalia of female; 19. 
labial plate of larva; 20. mandible; 21. antenna; 22. respiratory organ of pupa. Fig- 
ures 23-26. Cricotopus lebetis new species. 23. male genitalia; 24. dististyle of male 
genitalia, dorsal view; 25. second tergite of male abdomen, lateral view; 26. female 
genitalia, microtrichae omitted. 


No. 4 


atory organ (compare Fig. 32 and Roback’s 
figure 244). 


NANOCLADIUS ALTERNANTHERAE 
Dendy and Sublette 


Nanocladius alternantherae Dendy and Sublette, 1959: 
510, description of larva, pupa and adult; ecology, 
phenology. 

To the original description should be 
added: eyes hairy; postocular bristles two, 
dorsolateral to the eyes; dorsomedial bristles 
two, rather short and heavy, on a very slight 
mesonotal tubercle; middle tibia with two 
spines, the inner slightly shorter than the 
outer (in the original description the outer 
mesotibial spur apparently was obscured or 
missing); small pulvilli present; scutellum 
with four bristles. 


As Freeman (1961) pointed out, his 
earlier (1954) synonomy of Nanocladius 
was too broad. I consider his synonomy of 
Microcricotopus with Nanocladius to be val- 
id but the other genera which he synono- 
mized not so. The valid genera then appear 
to be (after Brundin, 1956): Nanocladius 
Kieffer (synonym Microcricotopus Thiene- 
mann and Harnisch); Evkiefferiella Thiene- 
mann (synonym Akzefferiella Thienemann); 
Parakieffertella (Thienemann) Brundin; 
Krenosmittia Thienemann (synonym Camp- 
tokiefferiella Goetghebuer ). 

The two other Nearctic species sordens 
Johannsen and brevinervis Malloch, pro- 
visionally placed in Nanocladius by Dendy 
and Sublette (1959), do not appear to be- 
long here in the restricted genus. The posi- 
tion of brevinmervis most probably is in 
Euktefferiella, while that of sordens is un- 
certain. Johannsen’s very brief original de- 
scription did not mention hairy eyes so pos- 
sibly this is also a species of Ewkiefferiella 
in its restricted sense. A reexamination of 
the type will be needed before a positive 
placement can be given. 

Material examined: Two females, 2-II-57; 
2 females, 6-II-57; 1 male, 6-V-57; 1 male, 
25-VI-57; 2 females, 2-VII-57; 1 female, 
)-VII-57; 1 male, 11-VII-57: 1 male, 27- 
VII-57; 1 male, 30-VII-57; 5 males, 8 fe- 
males, 24-IX-57; 1 male, 30-[X-57; 1 male, 
1-X-57; 1 male, 29-X-57; 1 female, 11- 
XI-57; 2 males, 25-XI-57; 1 male, 1 female, 
2-X1I-57; 1 female, 23-XII-57; 1 male, 28- 
1-58; C.R.L. One female, 25-VII-56; Ch.L. 


Chironomids of West-Central Loutsiana 


SUBFAMILY CHIRONOMINAE 
Tribe Chironomini 
PSEUDOCHIRONOMUS AIX Townes 


Pseudochironomus aix Townes, 1945: 19, description 
of adult. 
Psendochironomus aix Townes; Beck and Beck, 1959: 
92, distribution; phenology. 
Immature stages were reared from speci- 
mens collected on alligator weed ( Alternan- 
thera philoxerotdes (Mart.) Standl.). 


Larva: Head pale except for brownish 
teeth of labial plate and mandible; gular 
area yellowish. Head capsule length 0.53 
mm. Labial plate (Fig. 34) very similar to 
that figured for Tanytarsus (sens. lat.) sp. 
J (Johannsen, 1937a) (?—Pseudochirono- 
mus pseudoviridis (Malloch), cf. Sublette, 
1957) and fulviventris (Johannsen) (cf. 
Hauber, 1947). Mandible (Fig. 35) also 
very similar to the two species mentioned. 
Maxilla, epipharyngeal apparatus, and an- 
tenna are shown in Figures 36, 37 and 38. 
Preanal tubercles more or less quadrate, 
about as high as wide. Each supports 9 long 
yellowish bristles. On the anterior face of 
each tubercle are 2 fine bristles while the 
membrane posterior and inferior to the 
tubercle supports one longer, heavier bristle. 
Anal legs each with about 14 strongly 
hooked, yellowish claws. 


Pupal exuvia: Length 4.44 mm. Thorax 
yellowish-brown, somewhat papillose on dor- 
sal surface; low spinules on either side of 
anterior margin of raphe. Respiratory organs 
not discernible. 


Tergal chaetotaxy of abdomen as follows: 
Segment I, devoid of spinules. Segment I], 
with a transverse band of shagreen towards 
the anterior margin; posteriorly there is a 
less definite triangular band of finer sha- 
green; along the midline the two bands are 
joined by fine shagreen. At the posterior 
margin is the usual heavy band of blackish 
upturned hooks, which is made up of 95 
hooks in an even straight row. Segment III, 
similar to II, with an anterior definite band 
joined to a fainter posterior area of sha- 
green. Intersegmental membrane with a few 
fine black spinulae. Segment IV, similar to 
III but with the tergal surface behind the 
anterior band almost uniformly and finely 
shagreened. Intersegmental membrane _be- 
tween IV and V with several rows of black- 
ish spinulae. Segment V, anterior band com- 
posed of a lateral area of heavy shagreen, 


122 


joined across middle by fine shagreen; pos- 
terior half of tergite finely shagreened ex- 
cept for bare lateral areas; margin of seg- 
ment with 3 large flattened bristles, 2 close 
together near anterior one-fourth and the 
third beyond the middle. Segment VI, as V 
except spinulae less dense and posterior area 
narrower, leaving wider marginal clear areas; 


Tulane Studies in Zoology 


Vol. 11 


lateral bristles as V. Segments VII and VIII, 
with an anterior band only (Tergite VIII, 
Fig. 39). The lateral bristles of VII as the 
preceding two segments. Segment IX, cen- 
tral part of disc with a band of coarse spinu- 
lae (Fig. 39); swim fin with a fringe of 37 
to 40 bristles. 

Males: Wing length 1.66-2.18, mean 1.91 


ai y 


! re 
Ke NEN ACE EY 1, 
SECU eros weeny! Cee ’ 


———— 
— ee 


33 


Figures 27-33. Psectrocladius vernalis Malloch. 27. labial plate of larva; 28. mandible; 
29. antenna; 30. epipharyngeal apparatus; 31. torma (premandible) ; 32. pupal respira- 
tory organ; 33. fourth abdominal tergite of pupa. Figures 34-39. Pseudochironomus aix 
Townes. 34. labial plate of larva; 35. mandible; 36. maxilla; 37. epipharyngeal apparat- 
us—abbreviations as follows: Pe = pecten epipharyngis Chb = chaetulae basales Tm 

- torma (= premandible) Ch = chaetae Sm = squama platia I = seta I II = seta 
II III = seta III Sp = spinulae DLS = dorsal labral setae; 38. antenna; 39. tergite 
VIII of pupa, and anal fin. 


No. 4 


mm (10); leg ratio 0.84-0.93, mean 0.89 
(10); antennal ratio 2.00-2.47, mean 2.22 
CSE 

Material examined: One female, 11-VII- 
572 1 male, 29-VIII-5/7; 1 male, 1 female, 
Zox-57- 1 male, 10°1X-57: 1 female, 14- 
X-57: 1 female, 21-X-57; 1 male, 7-IV-58; 
1 female, 30-IV-58; 4 males, 4 females, 
5-V-58:; 3 males, 6 females, 12-V-58; 3 
males, 2 females, 4-VI-58; 3 males, 3 fe- 
males, 6-VI-58; 3 males, 3 females, 19-VI- 
58; 3 females, 2-VII-58; on alligator weed, 
ERA. 


CHIRONOMUS (CHIRONOMUS) 
STIGMATERUS Say 


Chironomus stigmaterus Say, 1823: 15, description 
of adults. 
Tendipes (Tendipes) stigmaterus (Say); Townes, 


1945: 120, added description of adult; distribution 

and phenology. 

Chironomus (Chironomus) stigmaterus Say; Beck 
and Beck, 1959: 94, distribution and phenology of 
adults. 

Tendipes (Tendipes) 
1960: 211, adults. 
Males: Wing length 4.55, 4.95 mm (2); 

leg ratio 1.44 (1); antennal ratio 4.55, 5.00 

CZs): 

Material examined: One pupal exuvia, 
10-X-58; 1 male, 1 female, 27-XI-58; 1 
pupal exuvia, 12-XII-58; 1 pupal exuvia, 
25-I-59; 1 pupal exuvia, 15-II-59; U.S.F.H. 
One male, 14-V-57; at light, Natchitoches, 
La. 


stigmaterus Sublette, 


(Say) ; 


CHIRONOMUS (CHIRONOMUS ) 
ATTENUATUS Walker 


Chironomus attenuatus Walker, 1848: 20, descrip- 
tion of adult. 

Tendipes (Tendipes) decorus (Johannsen) ‘Townes, 
1945: 20, adults. 


Tendipes decorus [(Johannsen) ]; Gerry, 1951: 241- 
244, ecology; control. 
Tendipes (Tendipes) decorus 

1953: 129, ecology. 
Tendipes decorus (Johannsen); Jamnback, 1954: 1- 
36, ecology; control. 
Tendipes decorus (Johannsen); Jamnback and Col- 
lins, 1955: 1 ecology; control. 


(Johannsen) ; Roback, 


Tendipes (Tendipes) decorus (Johannsen); Neff, 
1955: 10, larva, pupa and adult; ecology. 
Tendipes (Tendipes) decorus (Johannsen); Tebo, 


1955: 96, ecology. 

Tendipes decorus (Johannsen); Jamnback and Col- 
lins, 1956: 1-5, ecology; control. 

Chironomus decorus Johannsen; Paine and Gaufin, 
1956: 296, ecology. 

Tendipes decorus (Johannsen) ; 
phenology. 


Tendipes (Tendipes) 


Judd, 1957: 401, 


decorus Roback, 


(Johannsen) 


Chironomids of West-Central Loutsiana 


123 


1957b: 17, distribution of adult; 113, de- 
scription of larva and pupa. 
Tendipes (Tendipes) decorus (Johannsen); Sublette, 


1957: 390, ecology; phenology. 


OM ics 


Chironomus decorus (Johannsen); Gaufin, 1958: 202, 
and following pages, ecology. 

Tendipes (Tendipes) decorus (Johannsen); Dendy 
and Sublette, 1959: 514, adults. 


Chironomus (Chironomus) decorus Johannsen; Beck 
and Beck, 1959: 94, distribution of adults; phe- 
nology. 

Tendipes decorus [(Johannsen) ]; Provost and Branch, 
1959: 49-62, ecology. 

Tendipes (Tendipes) attennatus 
1959: 135, synonomy; 

Tendipes (Tendipes) decorus (Johannsen); Davis, 
1960: 212, distribution of adults; phenology. 

Tendipes (Tendipes) attenuatus (Walker); Darby, 
1962: 161, ecology. 

Males: Wing length 2.93-4.05, mean 3.41 
mm (4); leg ratio 1.61-1.84, mean 1.70 
(4); antennal ratio 3.75-4.10, mean 3.92 
(ye 

Material examined: One male, 27-X-55; 
1 female, 8-II-56; 1 female, 9-II-56; 1 fe- 
male, 3-II-57; 2 females, 5-II-57; 1 female, 
17-IV-57, Ch.L. One male, 6-II-57; 1 male, 
DI Xe malesAS eo: ll males 19-2 1- 
IlI-58; 1 male, 12-V-58; C.R.L. One male, 
18-V-54; 1 male, 11-X-56; 3 males, 14-I-57; 
2 males, 14-II-57; 2 males, 15-II-57; 1 male, 
28-II-57: 3 males, 3 females, 2-III-57; 2 
males, 1 female, 3-III-57; 1 male, 10-III-57; 
1 male, 11-III-57; 5 males, 2 females, 12- 
III-57: 3 males, 13-III-57; 2 males, 28-III- 
57; 4 males, 5-IV-57; 3 males, 8-IV-57; 1 
male, 9-IV-57; 4 males, 12-IV-57; 2 males, 
16-IV-57; 1 male, 29-IV-57; 1 male, 6-V-57; 
4 males, 14-V-57; 2 males, 15-V-5/;, 1 male, 
21=V-57: 2 males, 16-VI-57: 12 «males; 26- 
VI-57; at light, Natchitoches, La. 


(Walker); Townes, 
notes on type. 


CHIRONOMUS (CHIRONOMUS ) 
FULVIPILUS Rempel 
Chironomus fulvipilus Rempel, 1939: 210, description 
of adult. 


Tendipes (Tendipes) fulvipilus (Rempel); Townes, 
1945: 119, adults. 

Tendipes (Tendipes) fulvipilus (Rempel); Dendy 
and Sublette, 1959: 514, adults. 

Chironomus (Chironomus) fulvipilus Rempel; Beck 
and Beck, 1959: 94, adults; distribution; phe- 
nology. 

Tendipes (Tendipes) fulvipilus (Rempel); Sublette, 


1960: 211, adults. 


Larva and pupa have been associated by 
rearing in other studies by the author and 
will be described elsewhere. 

Material examined: One male, 7-IX-56; 
Chale 


124 


CHIRONOMUS (CHIRONOMUS ) 
NATCHITOCHEAE new species 
(Einfeldia group) 

Holotype male: U.S.N.M., No. 66457. 
Collected at the U. S. Fish Hatchery, Natchi- 
toches, Louisiana, 20-VIII-58, R. F. Tyler. 

Head brownish; antennal pedicel blackish- 
brown; antennal flagellum dark brown; an- 
tennal ratio 3.00; frontal tubercles large and 
conspicuous, 0.07 mm long; palpi dark; 
ratio, 5:12:20:27. Mesothorax yellowish- 
brown; vittae, postnotum and sternopleuron 
blackish-brown; mesonotum with a slight 
central hump; scutellum infuscate; halteres 
dark; wing length 2.61 mm; dorsolateral 
bristles in a single row, sparse. Forelegs 
darkened on distal one-third of femur, and 
tarsal segments 2 to 5; middle and hind legs 
darkened on distal two-thirds of femur, 
basal one-third of tibia and tarsal segments 
3 to 5; combs of middle and hind legs with 
two short spurs of about equal length. Fore- 
tarsus without a beard. Wings without 
macrotrichiae but with conspicuous micro- 
trichiae. Abdomen entirely blackish-brown. 


Leg proportions: ee 

etrotay o¢? 31.4 -5 ratio 
Foreleg 57 40 88 44 34 28 11 2:20 
Middle leg 58 50 30 15 10 5 4 0.60 
Hind leg 60 65 48 23 17 10 6 0.74 

Genitalia very similar to chelonia Townes 
but differs in having a more clavate disti- 
style. Figure 40 shows the normal view; 
Figure 41 shows anal point and superior 
appendage in lateral view. 

Allotype: In the Collection of the U. S. 
National Museum. Collected at the U. S. 
Fish Hatchery, 4-X-58, R. F. Tyler. 

Coloration and other features similar to 
male except for sexual differences. The 
mesonotal hump is more conspicuous than 
in the male. Wing length 3.15 mm. 


Leg proportions: 


Leg: 

eT Tas; oo 9 a>. & ratio 

Foreleg 60 45 98 38 33 27 12 2.18 
Middleleg 65 55 32 16 12 6 5 0.58 
Hind leg 68 70 50 23 18 11 8 0.71 


Lamellae of genitalia scarcely produced 
ventrally, almost quadrate in outline, (Fig. 
42). 

This species keys to brunneipennis (Jo- 
hannsen) in the key given by Townes (1945, 
page 111). It may be distinguished from 
that species by having the anal point of the 


Tulane Studies m Zoology 


Vol. 11 


genitalia broadened rather than narrow and 
by differences in the superior appendage. 

Paratype males: Wing length 2.39-2.70, 
mean 2.54 mm (4); leg ratio 1.85, 2.00 (2); 
antennal ratio 2.82-3.10, mean 2.93 (4). 

Paratypes: Two females, 3-V-57; C.R.L. 
One male, 12-VIII-58; 7 males, 20-VIII-58; 
5 males, 6-IX-58; 9 males, 12-IX-58; 21 
males, 20-I[X-58; 1 male, 1 pupal exuvia, 
27-[X-58; 3 males, 2 females, 2 pupal exu- 
viae, 4-X-58; 2 males, 1 pupal exuvia, 15- 
X-58;: U.S.F.H. In the collections of the 
USN.M., CN.C, ANSP) TNESSGer 
nell University, and Florida State Board of 
Health. 

Larva: Described from exuvia associated 
with reared adult. Head length 0.59 mm; 
capsule ventrally darkened; tips of man- 
dibles, torma, labial plate, and narrow oc- 
cipital border, black. Labial plate as in 
Figure 43; mandible, Figure 44; antenna, 
Figure 45. Epipharyngeal area similar to 
other Chironomini: Pe, somewhat pulvilli- 
form, Figure 46; Chb, 7 in number, finely 
serrate; torma, black tipped, distally with the 
usual bifurcation; Ch, 6 in number; Sm, 
finely pectinate; teeth not clearly discernible 
but in excess of 25 on each side; seta I, 
palmate, with very fine teeth; seta II, long, 
curving and unbranched; seta III, minute, 
medial to Il and almost contiguous with it; 
Sp, small and inconspicuous. 

Preanal papillae very short, each with 5 
to 6 rather short, terminal yellowish bristles. 
Anal prolegs with about 18 long, curved, 
yellow claws. 

Pupa: Described from exuviae. Length 
8.00 mm; almost entirely blackish. Cephalic 
tubercle long and pointed with a conspicu- 
ous preapical bristle. Respiratory organs 
white with numerous branches, apparently 
arising from 3 main branches. Tergites II- 
VI with an anterior and posterior transverse 
band of spinulae, each tergite with a central, 
laterally more restricted patch of spinulae 
(Fig. 47). Segment II with the usual pos- 
terior row of upturned hooks; about 81 
hooks in the row. Intersegmentalia between 
III-IV, with a few spinulae; between IV-V 
and V-VI, heavily spinose. Posterolateral 
comb of segment VII shown in Figure 48; 
caudal fin with about 116 fringe bristles in 
an irregular double row. 

The larva cannot clearly be distinguished 
from the Palearctic species Chironomus 
(Chironomus) insolita Kieffer (Einfeldia 


so 


No. 4 Chironomids of West-Central Louisiana 125 
basis of having a posterolateral comb of 
only 3 teeth. It might be confused with 
certain species of Chironomus (Crypto- 
chironomus) but these are usually smaller 


group) on the basis of the description avail- 


able to me. 
The pupa is distinguished from all other 


Nearctic Chironomus (Chironomus) on the 


ZA 
| 
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A \ \) \ 
¥ q i] 
“\\ \ 
2 Wes ~ 
r \S { 
\ > 
/ 4 = % 
‘ ae 
if f x \\ — 
Me) : 
) 98 / NON 
f © \ } & oh. 
\ 40 ° ) \ \V = Ae 
{ K Lh 
\ 2 Sa) \ 
es Veal \ ~ 
0 | i) \ 
h y 
? o \ 
2 f 41 \ 
4 48 ) \ 
0 | a 
0 | 
0 
SER 


iS SRY Zig 
Z NW 
s . } 43 
Sx 
Se ; 
| FT { = Fe 
Zi) |) || y AS 
SKA, Wy} \N — 8 
ZY Ar 
o yy | y | \ L - 
A fff} | ff \ k ip ie are 
Yf}}%,\) 0 A RG | 
Oi |; < 42 G ye 
pho = | 
\ SS UJ | 
re. 1 | 
a Pn | 
: i Hen Ol 
. oa \ ay 
i > \ a 
4 Q = tog 
— x 
Uy 
fay) | fae) ) 
Ife | ladies a f ) 
46 : ; \ 5 
s =) 
\ = * S / 
3 
| \ ~ \ = 
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} 45 ce . > . Z ————— 
| \ — ee { 
—, SSA SS 


Figures 40-48. Chironomus (Chironomus) natchitocheae new species. 40. genitalia of 
male; 41. anal point and superior appendage in lateral view; 42. female genitalia; 43. la- 
bial plate of larva; 44. mandible; 45. antenna; 46. pecten epipharyngis; 47. tergite chae- 


totaxy of pupa; 48. posterolateral margin of Segment VIII. 


126 


in size and the spines of the posterolateral 
comb are finer and paler. 


CHIRONOMUS (DICROTENDIPES ) 
MODESTUS Say 


Chironomus modestus Say, 1823: 13, 
adult. 

Tendipes (Limnochironomus) modestus (Say); 
Townes, 1945: 106, adults; generic position. 

Tendipes (Limnochironomus) modestus (Say); 
Hauber and Morrissey, 1945: 288, description of 
larva, pupa and adult; phenology. 

Tendipes (Limnochironomus) modestus (Say); 
Neff, 1955: 9, larva, pupa and adult; ecology. 


description of 


Tendipes (Limnochironomus) modestus (Say); 
Tebo, 1955: 96, ecology. 

Tendipes  (Limnochironomus) modestus (Say); 
Roback, 1957c: 111, larva and pupa. 

Chironomus (Limnochironomus)  modestus (Say); 
Beck and Beck; 1959: 94, distribution and_ phe- 


nology of adults. 
Tendipes (Dicrotendipes) 

and Sublette, 1959: 

generic position. 


modestus 


Dendy 
adult; 


(Say) 
514, description of 


Tendipes (Dicrotendipes) modestus (Say); Sublette, 
1960: 218, adult. 
Tendipes (Dicrotendipes) modestus (Say); Darby, 


1962: 38, 51, 54, 101, 158, adult. 


Males: Wing length 2.11-2.70, mean 2.40 
mm (4); leg ratio 1.66-1.86, mean 1.78 
(3); antennal ratio 2.60-2.87, mean 2.74 
(4). 

Material examined: Four males, 29-III- 
54; 1 male, 12-IV-57; 1 male, 16-IV-57; 1 
male, 6-V-57; 1 male, 16-VII-57; Natchi- 
toches, La. One male, 5-II-57; Ch.L. 


CHIRONOMUS (DICROTENDIPES ) 
NERVOSUS Staeger 


Chironomus nervosus Staeger, 1839: 567, description 
of adult. 

Tendipes (Limnochironomus) lucifer (Johannsen) ; 
Hauber and Morrissey, 1945: 288, description of 
larva, pupa and adult; phenology. 

Tendipes (Limnochironomus)  nervosus  (Staeger) ; 
Townes, 1945: 102, 103, 108, taxonomy. 

Limnochironomus nervosus [ (Staeger) ], Wohlschlag, 
USO 954515 ecology. 


Limnochironomus nervosus [(Staeger)], Mundie, 
1955: 578, ecology. 
Limnochironomus  nervosus  [(Staeger)], Palmen, 


1955: 20, ecology. 

Tendipes (Limnochironomus) (Staeger) ; 
Anderson and Hooper, 1956: 262, ecology. 

Limnochironomus nervosus (Staeger); Mundie, 1957: 
165, ecology. 

Tendipes (Limnochironomus) 
Sublette, 1957: 390, ecology. 

Tendipes (Limnochironomus)  nervosus  (Staeger) ; 
Roback, 1957c: 110-111, larva, pupa, in key. 

Chironomus (Limnochironomus) nervosus Staeger; 
Beck and Beck; 1959: 94, distribution and phen- 
nology of adults. 


nNerVOSUS 


nervosus (Staeger) ; 


Tulane Studies m Zoology 


Vola 


Tendipes (Dicrotendipes) nervosus (Staeger); Dendy 
and Sublette, 1959: 514, generic position. 

Tendipes nervosus (Staeger); Judd, 1960: 207, phe- 
nology. 

Tendipes (Limnochironomus)  nervosus  (Staeger) ; 
Davis, 1960: 71 and following pages, ecology. 
Tendipes (Dicrotendipes) nervosus (Staeger); Sub- 
lette, 1960: 220, adult. 
Tendipes (Limnochironomus) 
Buscemi, 1961: 294, ecology. 
Tendipes nervosus (Staeger); Judd, 1961: 96, phe- 


nervosus (Staeger) ; 


nology. 
Tendibes (Limnochironomus) nervosus  (Staeger) ; 
Darby, 1962: 158, ecology. 


Males: Wing length 1.70-1.79, mean 1.76 
mm (3); leg ratio 1.87-2.17, mean e200 
(3); antennal ratio 2.27-2.67, mean 2.46 
CS). 

Material examined: Seven males, 7-I[X-56; 
2 males, 11-[X-56; 2 males, 6-II-57; 2 males, 
2-VII-57; 1 male, 2 females, 16-VII-57; 2 
males, 18-VII-57; 2 males, 20-VIII-57; 2 
males, 22-VII-57; 1 male, 23-VII-57; 1 fe- 
male, 27-VII-57; 1 male, 1 female, 30-VII- 
57: 1 male, 1-VIII-57; 4 males, G-Villesg. 
2 males, 13-VIII-57; 1 male, 20-VIII-57; 
2 males, 27-VIII-57; 1 female, 29-VIII-57; 
4 males, 30-[X-57; 1 male, 7-X-57; 4 males, 
14-X-57; 7 males, 21-X-57; 2 males, 5-V- 
58; 1 male, 12-V-58; 1 male, 10-VI-58; 
C.R.L. Two males, 29-III-54; 1 male, 18- 
V-54; 1 male, 28-II-57; 1 male, 2-III-57; 
1 male, 12-III-57; 2 males, 13-III-57; 1 male, 
26-III-57; 2 males, 28-III-57; 2 males, 5- 
IV-57; 1 male, 8-IV-57; 1 male, 12-IV-57; 
1 male, 14-V-57; 2 males, 21-V-57; 2 males, 
16-VI-57; 2 males, 28-VI-57; 2 males, 16- 
VII-57; 2 males, 22-VII-57; at light, Natchi- 
toches, La. 


CHIRONOMUS (DICROTENDIPES ) 
INCURVUS new species 


Holotype male: U.S.N.M., No. 66458; 
U. S. Fish Hatchery, Natchitoches, Louisiana 
27-IX-58, R. F. Tyler. 

Head and thorax pale stramineous; ab- 
domen pale; antennal flagellum and front 
tarsus beyond middle of basitarsus darkened. 
Pronotum scarcely narrowed medially, only 
slightly inferior to anterior edge of meso- 
notum; mesoscutum with a slight hump. 
Frontal tubercles exceedingly small, scarcely 
visible at 100 magnification. Palpal propor- 
tions 7:10:15:22. Antennal ratio 1.88. 


Wing length 1.80 mm; no supra-alar 
bristles; prealar bristles 3; dorsomedial bris- 
tles decumbent, apparently in one row; 
dorsolateral bristles in one row, erect; scu- 


No. 4 


tellar bristles 8, large, in a straight trans- 
verse row, anteriorly are 2 smaller bristles, 
one on either side of the midline. 


Leg proportions: 


Leg 

tie Ray eo se ae oon ratio 

Foreleg 48 35 66 30 25 22 10 1.89 
Middleleg 42 35 21 11 7 5 5 0.60 
Hind leg AS 52 32 18 ds) 98) 5 O:6L 


Fore tarsus without a beard. 


Wing veins scarcely darkened, membrane 
clear; venarum ratio 1.12. 


Genitalia very similar to nervosus Staeger 
but with an inturned superior appendage 
which is distinctive (Fig. 49). 


Allotype female: In the U. S. National 
Museum Collection; collected at the U. S. 
Fish Hatchery, 6-XI-58, R. F. Tyler. 


Head somewhat darkened, thoracic vittae, 
postnotum and mesosternum brown; fore- 
legs beyond middle of femora, middle and 
hind tarsi, antennal flagellum, palpi and 
genital lamellae infuscate. Thoracic chaeto- 
taxy as holotype male. Palpal proportions, 
5:14:15:25. Antennal segment proportions, 
1510-10210" 15: 

Wing length 2.48 mm; venarum ratio 
Veale 


Leg proportions: 


; Leg 

HeLa 5 hoe} 4 ge ratio 

Foreleg 55> 42 75 31 26 238 11 1.79 
Middleleg 50 47 25 12 8 5 5 0.58 
Hind leg DpOoals to) 1G. 8) 50°57 


Abdomen pale. Genital lamellae quadrate 
(Fig. 50); spermatheca ovoid, 0.11 x 0.08 
mm, seminal duct emerging eccentric; ducts 
and spermatheca colorless (Fig. 51). 


Paratype males: Wing length 1.62-1.80, 
mean 1.73 mm (3); leg ratio 1.90-1.94, 
mean 1.92 (3); antennal ratio 2.00-2.21, 
mean 2.11 (3). 


Paratypes: Three males, 20-VIII-58; 8 
males, 12-IX-58; 6 males, 20-[X-58; 5 males, 
27-IX-58: 9 males, 15-X-58; 9 males, 29- 
X-58; 14 males, 1 female, 6-XI-58; 1 male, 
15-11-59; US.F.H. One male, 24-XI-59; 
Chivary Dam Spillway, Natchitoches Parish, 
La. In the collection of A.N.S.P., Florida 
State Board of Health, C.N.C., I.N.HLS., and 
Cornell University. 

This species may be distinguished from 
all other Nearctic species by the male geni- 
talia, which are similar to mervosus but dif- 


Chironomids of West-Central Loutstana 


7 


fer in having an inturned superior append- 
age. 

Larva: Described from exuvia of reared 
male. Head length 0.50 mm. Head capsule 
yellowish except for tips of mandibles and 
labial plate. Mandible (Fig. 52) not at all 
like mervosus but rather more like modestus 
(cf. Hauber and Morrissey, 1945, figure 10). 
Mandible length 0.16 mm. Labial plate, 
(Fig. 53) also similar to modestus. In the 
single larva available, the details of the 
epipharyngeal area were obscured. The pre- 
mandible (torma) is yellowish and appears 
to be bifurcate. The antennae are also ob- 
scured in that they do not lie in a flat 
plane. Segment 3 appears shorter than 2 
and 4 which are approximately subequal in 
length. The posterior part of the larval 
exuviae was lost. 

Pupa: Described from exuvia of reared 
male and other exuviae compared with it. 
Exuvia dark yellowish-brown; length 4.5 
mm. Respiratory organs a tuft of many 
white filaments. Cephalic tubercles short 
(0.04 mm) and conical, each with a fine 
preapical bristle. Abdominal tergites II to 
VI with fine shagreen, that of IJ occurring 
in a longitudinal band on either side of the 
midline, that of II to VI occupying most of 
the tergite. Segment II with the usual row 
of hooks along the posterior margin; the row 
consists of about 80 fine, yellowish hooks. 
Sternites I and II with a posterior band of 
colorless needle-like elongate spines; sternite 
III with a similar band of less elongate spines 
at about the anterior one-third; lateral mar- 
gins of segments VI to VIII each with 4; 
flattened bristles, 2 in the anterior half and 
2 in the posterior. Abdominal segment 
VIII with a conspicuous double spur ( Figs. 
54, 55 for variation). Swim fin with about 
65 flattened filaments. 

The larva keys in Roback (1957c, pages 
109, 110) to fwmidus Johannsen. It differs 
from fumidus by having the most laterad 
tooth of the labial plate appearing as a 
lateral shelf on the fifth tooth, and by hav- 
ing all mandibular teeth darkened. 

The pupa keys to couplet 16 (page 111, 
op. cit.). It may be distinguished from 
modestus Say and neomodestus Malloch by 
a higher number of filaments in the swim 
fin (about 65) and by the distinctively dif- 
ferent caudolateral spur of segment 8. 


128 


ry 
> 


eee 


49 


51 


\ 


) 


Figures 49-55. Chironomus (Dicrotendipes) incurvus new species. 


Tulane Studies n Zoology 


Vol. 11 


52 


ZB 

by Page 

| / YA 
‘a: 


49. male genitalia; 


50. female genitalia; 51. spermatheca and duct; 52. larval mandible; 53. labial plate; 
54. posterolateral spur of Segment VIII of pupa; 55. variation of posterolateral spur. 


CHIRONOMUS (XENOCHIRONOMUS) 
XENOLABIS Kieffer 

Chironomus xenolabis Kieffer (in Thienemann and 
Kieffer, 1916): 526, description of adult. 

Xenochironomus xenolabis (Kieffer); Townes, 1945: 
92, description of adult. 

Xenochironomus xenolabis (Kieffer); Beck and Beck, 
1959: 93, distribution and phenology of adult. 


Males: Wing length 2.25-2.47, mean 2.38 
mm (3); leg ratio 1.70 (1); antennal ratio 
2.40-2.70, mean 2.46 (3). 

Material examined: Three males, 2 fe- 
males, 17-IX-57; 1 male, 24-IX-57; C.R.L. 
One male, 16-VII-57; at light, Natchitoches, 


La. 


CHIRONOMUS (ENDOCHIRONOMUS) 
NIGRICANS Johannsen 


Chironomus nigricans Johannsen, 1905: 219, descrip- 
tion of larva, pupa and adult. 
Tanytarsus (Endochironomus) nigricans (Johann- 


sen); Townes, 1945: 64, description of adult. 


Tanytarsus nigricans (Johannsen); Judd, 1949: 9, 
phenology. 
Tanytarsus (Endochironomus) nigricans  (Johann- 


sen); Berg, 1950: 97-98, ecology. 
Tanytarsus nigricans (Johannsen); Judd, 1953: 813, 
phenology. 


Tanytarsus (Endochironomus) nigricans  (Johann- 
sen); Roback, 1953: 129, ecology. 
Tanytarsus (Endochironomus) nigricans (Johann- 


sen); Anderson and Hooper, 1956: 282, ecology. 


Endochironomus nigricans (Johannsen); Paine and 
Gaufin, 1956: 296, ecology. 
Tanytarsus (Endochironomus) nigricans (Johann- 


sen); Roback, 1957c: 120, larva and pupa. 
Tanytarsus nigricans (Johannsen); Judd, 1957: 401, 
phenology. 
Endochironomus (Endochironomus) nigricans (Jo- 
hannsen); Beck and Beck, 1959: 93, distribution 
and phenology of adult; generic position. 


Tendipes (Endochironomus) nigricans (Johannsen) ; 
Dendy and Sublette, 1959: 514, adult; generic 
position. 

Tanytarsus nigricans (Johannsen); Judd, 1960: 207, 
phenology. 

Tanytarsus Endochironomus) nigricans (Johann- 


sen); Davis, 1960: 71 and following pages, ecology. 

Tendipes (Endochironomus) nigricans (Johannsen) ; 
Sublette, 1960: 216, adult. 

Tanytarsus nigricans (Johannsen); Judd, 1961: 96, 
phenology. 

Tendipes (Endochironomus) 
Darby, 1962: 157, ecology. 


Males: Wing length 2.70-3.15, mean 2.93 
mm (3); leg ratio 1.30, 1.32 (2); antennal 
ratio 2.67-3.05, mean 2.88 (3). 

Material examined: Three males, 1 fe- 
male, 17-IV-57; 4 males, 13-VIII-57; 1 


nigricans (Johannsen) ; 


No. 4 


male, 27-VIII-57; 1 male, 4-XI-57; 1 male, 
10-IX-57; 1 male, 7-X-57; 5 males, 21-IV- 
58: 3 males, 4-VI-58; 1 male, 6-VI-58; 1 
male, 10-VI-58; C.R.L. Two males, 20-IX- 
jeu males. 27-[X-58; USF: Two males, 
28 IIJ-57; 1 male, 9-IV-57; 5 males, 29-IV- 
5/2 2 males, 6-V-57; 5 males, 15-V-57; 2 
males, 21-V-57; 1 male, 28-VI-57; at light, 
Natchitoches, La. 


CHIRONOMUS (CRYPTOCHIRONO- 
MUS) PONDEROSUS new species 


Holotype male: U.S.N.M. No. 66459. 
Collected in a funnel trap, Cane River Lake, 
Natchitoches Parish, Loutsiana, 19-V-58, 
B. R. Buckley. 

Antennal pedicel yellowish-brown; anten- 
nal flagellum dark; antennal ratio 3.25; 
frontal tubercles absent; palpi slightly dark- 
ened, proportions, 7:25:20:32. 

Thorax yellowish with vittae, postnotum, 
and sternopleuron golden brown; wing 
length 2.70 mm; prealar bristles 7; dorso- 
medial bristles in one row, erect; scutellar 
bristles about 14 in straight transverse row. 

Legs yellowish with fore tibia, entire fore 
tarsus, and last 4 segments of middle tarsus 
darkened; hind tarsi missing. Fore tarsal 
beard absent. 


Leg proportions: 


Chironomids of West-Central Louisiana 


Leg 

Heist as i 94 5 Eatio 

Fore leg 65 53 95 48 388 21 12 1.79 
Middle leg 65 57 35 16 11 6 5 0.61 


Hind leg i 1. > SS 


Abdomen yellowish-green. Genitalia very 
similar to fwlvuws (Johannsen) Townes, but 
differs in having a spatulate anal point and 
by having the basistyle and dististyle less 
massive, with the latter slightly curved 
GBigs.556, 57). 

Allotype female: In U.S.N.M. Lake Ober- 
lin, Bryan Co., Oklahoma, 21-VII-62, reared 
from larva collected in silty-mud at 0.80 
meters. 

Antennal pedicel and antennal flagellum 
pale except terminal segment which is dark; 
palpi infuscate; proportions 6:20:16:30. 

Thorax ground color stramineous; vittae, 
sternopleuron, and postnotum  ocherous. 
Mesothorax with a conspicuous hump. Wing 
length 2.25 mm; | supra-alar bristle; dorso- 
medial bristles long, erect, in one row; dorso- 
lateral bristles long, erect, in one staggered 
row; scutellar bristles 15, in a long, pos- 
terior, straight, transverse row; 8 slightly 


129 


shorter in anterior straight row, antero- 
lateral bristles 4. Halteres pale. 

Fore femora and tibiae dark; remainder 
of forelegs missing; middle and hind legs 
pale except for terminal 2 tarsal segments 
which are dark. 


Leg proportions: 


Leg 
By iia 5 4 5 ratio 

Fore leg Ny AR SS SS Se 
Middleleg 52 46 29 13 9 5 £5 0.68 
Hind leg 5b pps 40, se 5 8) 6 ONS 


Abdomen pale greenish. Genitalia, Fig- 
ure 53: 

Paratype 
mean 2.36 mm (3); leg ratio 1.76-1.80, 
mean 1.78 (3); antennal ratio 3.22-3.50, 
mean 3.24 (3). 

Paratypes: One male, 1 pupal exuvia, 
13-VIII-57; 1 male, 2 pupal exuviae; 28- 
Will-57;, | pupal exuviay 7-X-57/> CR.L- 
One male, 14-VI-62; 1 male, 18-VI-62; 4 
males, 29-VI-62; 1 male, 20-VII-62; 3 pupal 
exuviae, 25-VII-62; at light, University of 
Oklahoma Biological Station, Willis, Okla. 
In the collections of C.N.C., A.N.S.P. and 
J.N-HLS. 

This species is one of the fwlvus group; 
the adult male is distinguishable by the 
genitalia with their long anal point, which 
is slightly spatulate apically. 


males: 


Larva: Described from exuvia of reared 
allotype; associated by distinctive pupal 
armature. 


Head dark yellow; tips of labium and 
mandible strongly contrasting black; head 
capsule 5.95 mm long. Labium (Fig. 59) 
very similar to digitatus (cf. Curry, 1958). 
Paralabial plates as other members of this 
group. Mandible (Fig. 60) also very simi- 
lar to that of digitatus but with the ac- 
cessory tooth longer and more attenuate. 
Antenna (Fig. 61) almost identical with 
that of digitatus but with a basal segment 
shorter relative to that of dzgitatus; ratio, 
50:30:23:2:1. Premandible (torma) ob- 
scured on the single specimen available. The 
epipharyngeal area does not differ signifi- 
cantly from that illustrated by Curry (1958, 
figure 1) for digitatus. The jointed labral 
bristles are apparently only two segmented. 
Maxilla, Figure 62. 

Anal prolegs each with about 15 curved 
yellowish spines; preanal papillae each with 
8 bristles. 


Pupa: Exuviae pale yellowish; the ce- 


130 


phalic tubercles (Figs. 63, 64) darker yel- 
low. Exuviae length 6.5 mm. Respiratory 
organs consist of numerous white filaments. 
Abdominal tergite I with a conspicuous 
anterior lateral spinose tubercle on each side; 
tergite devoid of shagreen; tergite II with 
fine shagreen which is coarser near the lat- 
eral and ventral margins; posterior margin 
of II with the usual row of hooks number- 
ing about 80, the row interrupted near the 
midline; tergites HI to VII with a par- 
tially doubled row of tubercle spines, the 
row being slightly interrupted near the mid- 
line; posteriorly the spines on each segment 
decrease in number and progressively be- 
come more acutely tipped; segments VI to 


Figures 56-64. Chironomus (Cryptochironomus) ponderosus 


Tulane Studies nr Zoology 


Vole id. 


VII with 4 lateral flattened filaments which 
are about evenly spaced; segment VHI with 
5 Jateral filaments, 2 in the anterior half and 
3 in the posterior half. Swim fin with a 
fringe of 72 filaments. Genital sacs with 
an acutely tipped terminal constriction; ter- 
gite IX with the usual bifurcate process. 

The larva keys in Curry (1958: 431-433) 
to couplet 4. It may be distinguished from 
sorex (Townes) and digitatus Malloch by 
different features of the mandible and labial 
plate (cf. Figures 59, 60). 

The pupa can be distinguished from all 
other species of this group by the distinc- 
tively different cephalic tubercles (cf. Fig- 
ures 63, 64). 


new species. 56. male 


genitalia; 57. details of inferior and superior appendage; 58. female genitalia; 59. lar- 
val labium; 60. mandible; 61. antenna; 62. maxilla; 63. cephalic tubercles of pupa, dor- 
sal view; 64. cephalic tubercles of pupa, lateral view. 


No. 4 


CHIRONOMUS (CRYPTOCHIRONO- 
MUS) FULVUS Johannsen (Townes) 


Chironomus fulvus Johannsen, 1405: 224, descrip- 
tion of pupa and adult female. Townes (1945) 
reports the pupa to have probably been misiden- 
tified. 

Chironomus sp. c Malloch, 
pupa (fide, Townes, 1945). 

Cryptochironomus  fulvus (Johannsen); Townes, 
1945: 98, adult male and female; synonomy. 

Cryptochironomus fulvus (Johannsen); in part; Sub- 
lette, 1957: 389, ecology. 

Tendipes ‘Cryptochironomus) fulvus (Johannsen) ; 
Dendy and Sublette, 1959: 516, generic position; 
adult male. 

?Cryptochironomus fulvus (Johannsen); Beck and 
Beck, 1959: 93, adult distribution; phenology. 
Tendipes (Cryptochironomus) fulvus (Johannsen) ; 
Sublette, 1960: 223, adult male and female (part). 
Nec!Chironomus (Cryptochironomus)  fulvus  Jo- 

hannsen, 1937b: 39 larva and pupa. 

Nec!Cryptochironomus fulvus (Johannsen); Curry, 
1958: 435, larva and pupa; ecology. 

Nec! Tendipes (Cryptochironomus) fulvus (Johann- 
sen); Darby, 1962: 162, description of larva and 
pupa; ecology. 


1915a: 529, larva and 


Larva: This species was briefly described 
by Malloch (1915a: 529) as Chironomus spe- 
cies c. During the present study several 
larvae were reared, the adults of which agreed 
with the description given by Townes, 1945, 
and the specimens from the Hauber collec- 
tion identified by Townes. 

Head 0.38-0.40, mean 0.39 mm long (5) 
by 0.26-0.29, mean 0.27 mm. wide (2). 
Head capsule yellow except for tips of man- 
dibles and labial plate. Labial plate (Fig. 
65) very similar to that of other species of 
the fwlvus group, most closely resembling 
that of C. digitatus (Malloch) (cf. Curry, 
1958, figure 19). The antenna (Fig. 66) 
is similar to that figured by Malloch (1915, 
Pl. XXX, figure 2) but differs in the de- 
tails of the last two segments. The mandibles 
(Fig. 67) are also similar to that shown by 
Curry (op. cit.) for digitatus. The pre- 
mandibles (torma) are shown in Figure 68. 
The epipharyngeal area is indistinguishable 
from that of digitatus (cf. Curry, 1958, 
figure 1). Preanal papillae short, each with 
6 to 7 pale terminal bristles. Anal prolegs 
with about 20 yellowish claws. 

Pupa: Briefly described by Malloch 
(1915a) as Chironomus sp. c and keyed by 
Roback (1957c) as Cryptochironomus sp. 3. 

The cephalic tubercles (dorsal view, Figs. 
69 and 70; lateral view, Fig. 71) are dis- 
tinctively different. There is some variation 
in this structure among different popula- 
tions: it is construed as varietal. Exuviae 


Chironomids of West-Central Loutsiana 


dark yellowish-brown; length 5.18-8.18, 
mean 6.23 mm (5). Tergite I devoid of 
spines or shagreen, but with two large ven- 
tral tubercles beset with denticles, a feature 
apparently characteristic of the fwlvus group. 
Tergite II with the usual row of posterior 
marginal spines interrupted for about the 
middle one-fourth of the row length; the 
row composed of about 48 yellowish curved 
spines. Tergites III to VII each with a pos- 
terior row of denticles, the rows progessively 
decreasing in size posteriorly. Immediately 
in front of the posterior rows of segments 
III to V is a row of 4 fine bristles on each 
side of the midline; on segments VI and 
VII there are only 2 bristles instead of 4. 
Tergite VII has only a single bristle on 
each side of the midline at the posterior 
margin. Between the genital sacs is the 
usual bifurcate process typical of this group. 
Swim fin with the bristle fringe in a single 
layer anteriorly, the posterior part doubled 
dorso-ventrally. Number of fringe bristles 
50-67, mean 60 (3). 

Adult: Chironomus fulvus Johannsen was 
described from a female, the characteristics 
of which are insufficient for specific recog- 
nition, as they are for most members of this 
family. The male was first described by 
Malloch (1915a) but Townes (1945) re- 
ported Malloch’s series to have been mixed. 
The description of the male by Townes (op. 
cit.) is then the first authoritative one. 
Townes synonomizes with fawlvws Johannsen 
the following: fwlvws Johannsen, Malloch 
(in part); parvilamellatus Malloch; and mal- 
lochi Kieffer (=abbreviatus Malloch, nec 
Kieffer). As it now stands Townes’ inter- 
pretation of fwlvus must be accepted as 
definitive with his illustration of the male 
genitalia serving in lieu of type specimen. 
I have examined adult males in the Hauber 
Collection determined by Townes and be- 
lieve the material at hand is conspecific 
with it. However, the adults which I listed 
for California (1960) also agree with the 
Hauber specimens in every feature exam- 
ined. Darby (1962), on the basis of adults 
determined by me, described a larva and 
pupa which is identical with that illustrated 
by Curry (1958) for falvws. Thus at least 
two distinct larval and pupal types appear 
to exist under the name of fw/vus. Further, 
I have reared specimens of three additional 
pupal types (to be described elsewhere), 
the adults of which are indistinguishable 


from fwlvws in Townes’ key. Thus falvus 
probably is a complex of closely related spe- 
cies which are best defined by pupal char- 
acteristics. Distinctive adult characteristics 
remain to be demonstrated. 

The pupa questionably associated with the 
holotype female is undoubtedly a member 
of the Harnischia group of Cryptochirono- 
mus as was pointed out by Townes (op. cit.) 
and Darby (op. cit.). 

On the basis of Townes’ synonomy of 
Malloch’s Chironomus species c with fulvus 
I am taking it to represent the pupa of 
fuluus. With this reasoning Curry’s and 
Darby's falvus will need to be described as 
a new species. 

The adult males before me agree well 
with the color description given by Townes 
(1945). The frontal tubercles are small and 
inconspicuous but clearly visible on a well 
mounted specimen. Postocular bristles in 
two rows; immediately behind the eyes is a 
straight row of large bristles; posterior to 
that is a smaller second row much more 
closely spaced. The hump at the end of the 
mesonotal median vitta is distinct. Dorso- 
medial and dorsolateral bristles long and 


Bae L J 67 


V1 s 
| re. 
} s 
J is / ~~ 
, F / 
Ue i 2 ~~ 
je | / 
Mee | f, i 
(rene | 
\ bs WY \ /| os } 
es ¥ Me ae a ena 2 
f r \\ , on / "4 | 
} 4 p ‘< c Vier bY k Li 
\ { 5 \ » 47 
\; .< 


Figures 65-75. Chironomus (Cryptochironomus) fulvus (Johannsen) Townes. 
bial plate of larva; 66. antenna; 67. mandible; 6& 


Tulane Studies n Zoolog) 


Voleaial 


erect, the latter in a partial double row. 
Prealar bristles 5; supra-alar bristles absent; 
posterior scutellar bristles large and erect, 
about 10, in a straight transverse row; an- 
terior scutellar bristles about 6, smaller, in 
a strewn pattern. 

Antennal and leg ratios, as well as size 
(wing length) are highly variable among dif- 
ferent populations. In all instances, however, 
values overlap. Wing length 1.94-3.15, mean 
2.39 mm (29); leg ratio 1.60-2.02, mean 
1.78 (27); antennal ratio 2.75-3.35, mean 
3.04 (20). 

The genitalia are quite variable as to ratio 
of width to length of dististyle: ratio 2.00- 
3.30, mean: 2.60 (34). Two principal types 
are apparent, a light bodied one as shown 
in Figure 72 and a heavier one shown in 
Figure 73. Intermediates are shown in Fig- 
ures 74 and 75. 

Material examined: One male, 8-VI-57; 
1 male, 2-VII-57; 1 pupal exuvia, 23-VII-57; 
1 male, 30-VII-57; 2 males, 6-VIII-57; 3 
males, 13-VIII-57; 1 male, 10-[X-57; 1 male, 
25-II-58: 1 male, 7-IV-58: 1 male, 5=V-58; 
1 pupal exuvia, 10-VI-58; C.R.L. One pupal 
exuvia, 9-VI-51; 1 male, 29-VI-62; Uni- 


( é 
ul 15 
13 
65. la- 
68. premandible (torma); 69-70. ce- 


phalic tubercles, dorsal views; 71. cephalic tubercles, lateral view; 72. male genitalia; 


73-75. variations of male genitalia. 


No. 4 


versity of Oklahoma Biological Station. Two 
males, 29-IV-57; 2 males, 21-V-57; 2 males, 
14-V-57; 1 male, 15-V-57; at light, Natchi- 
foches, La. One male, 5-X-59: 1 male, 10- 
X-59:; Sabine River at La.-Tex. line, west of 
Many, La. One male, 16-[I-60; Bayou Pierre, 
5 miles north of Natchitoches, La. One male, 
25-VI-62; Forguson’s Pond, west of Willis, 
Okla. One male, 16-VI-60; small stream 7 
miles east of Liberty, Tex. One male, 20- 
VI-41; 2 males, 14-VII-41: 1 male, 23- 
VII-41; 1 male, 27-VII-41; 1 male, 2-V-42: 
1 male, 22-V-42; Davenport, Iowa. One 
male, 18-VII-40, Lake Okobojii, Iowa, from 
Hauber Collection. Two males, 11-V-60: 
Sabine Bayou east of Clarence, La. One 
pupal exuvia, 24-IV-60; Trinity River, 13 
miles west of Livingstone, Tex. One male, 
21-VII-62; Lake Oberlin, Bryan Co., Okla. 
One male, 16-III-60; Old River, Natchi- 
toches Parish, La. 


CHIRONOMUS (CRYPTOCHIRONO- 
MUS) NIGROVITTATUS Malloch 


Chironomus nigrovittatus Malloch, 1915a: 456, adult. 

Harnischia (Harnischis) nigrovittata (Malloch) ; 
Townes, 1945: 163, adult; generic position. 

Harnischia (Harnischia)  nigrovittata (Malloch) ; 
Sublette, 1957: 393, description of pupa; phe- 
nology. 

Harnischia (Harnischiz)  nigrovittata (Malloch) ; 
Beck and Beck, 1959: 95, phenology and distribu- 
tion of adult. 

Tendipes (Cryptochironomus) nigrovittatus  (Mal- 
loch); Sublette, 1960: 224, description of adults; 
generic position. 


6-1.70, mean 1.59 
1.62, mean 1.53 
222 meat. 200 


= . ’ 


Males: Wing length 1.3 
mm (3); leg ratio 1.40- 
(3); antennal ratio 2.03- 
(4). 

Material examined: Two males, 18-VI- 
57; 1 female, 27-VIII-57; 1 male, 30-IX-57; 
6. males, 11-XI-57. 


CHIRONOMUS (CRYPTOCHIRONO- 
MUS) CARINATUS (Townes) new 
combination 


Harnischia (Harnischia) carinata Townes, 1945: 
158, adult. 

Harnischia (Harnischia) cerinata Townes; Sublette, 
1957: 393, adult. 

Harnischia (Harnischia) carinata Townes; Beck and 


Beck, 1959: 95, adult distribution; phenology. 

Larva and pupa unknown. 

Males: Wing length 1.36-1.80, mean 1.57 
mm (4); leg ratio 2.00-2.07, mean 2.02 
(3); antennal ratio 2.06-2.10, mean 2.05 


(Coe 


Chironomids of West-Central Loutsiana 


133 


Material examined: One male, 16-VII- 
57; 2 males, 27-VIII-57; 2 males, 10-IX-57: 
2 maless50-LX-5i7/5 Iimales7xe5i7-) male: 
14-X-57; CR.L. One male, 3-VI-57: at 
light, Natchitoches, La. One male, 15-X-58; 
WS EE: 


CHIRONOMUS (CRYPTOCHIRONO- 
MUS) CHAETOALA (Sublette) new 


combination 
Tendipes (Cryptochironomus)  chaetoala Sublette, 
1960: 220, description of adult. 
Tendipes (Cryptochironomus) chaetoala Sublette; 


Darby 1962: 35, 38) 50; 51; 59s 66, 735 82, 83, 
101, 160, 161, description of immature stages; 
ecology. 

Males: Wing length 1.79-1.87, mean 1.85 
mm (4); leg ratio 1.79-1.96, mean 1.88 
(4); antennal ratio 2.14-2.50, mean 2.24 
(4). 

Material examined: One male, 29-III-54: 
1 male, 11-[X-56; at light, Natchitoches, La. 
One male, 20-VIII-58; 5 males, 28-VIII-58; 
4 Smales, (G-DXe58= 5s males\ 1251x584 
males, 20-[X-58; 9 males, 27-[X-58; 1 male, 
29-IX-58: 6 males, 4-X-58: 5 males, 15-X- 
Sie WAS eh 


CHIRONOMUS (CRYPTOCHIRONO- 
MUS) DIRECTUS (Dendy and Sublette) 
new combination 
Tendipes (Cryptochironomus) directus Dendy and 

Sublette, 1959: 514, description of adult. 

Material examined: In addition to the 
original type series from this locality, I have 
examined the following: Three males, 11- 
DPX256; P imale, 27-Vill-57= 1 males 17-ixe 
S72 males, 245bx-57>" f male, 40257; 
f malesn/-<-ois 5. males lA x-=5 7G Rel 
Two males, 6-IX-58; 3 males, 12-IX-58; 2 
males, 20-IX-58; 7 males, 27-I[X-58; 1 male, 
15-X-58; 1 male, 29-X-58; 1 male, 6-XI-58; 
CES EE 


CHIRONOMUS (CRYPTOCHIRONO- 
MUS) EMORSUS (Townes) new 
combination 


Harnischia (Harnischis) emorsa Townes, 1945: 161, 
description of adults. 

Chironomus fulvus Johannsen, 1905: 224, descrip- 
tion of pupa, misdetermined. 

Chironomus (Limnochironomus) sp. Johannsen, 1938: 
(1937b) 44, description of pupa. 

Harnischia (Harnischia) emorsa Townes; 
1957c: 101, pupa, in key. 

Harnischia (Harnischia) 
Beck, 1959: 95, adult. 


Beck and 


Roback, 


emorsa Townes; 


134 


Males: Wing length 1.46, 1.61 mm (2); 
leg ratio 2.14, 2.25 (2); antennal ratio 1.89, 
224» (2:)8 

An adult with an adherent pupal exuvia 
and a pupa with visible male genitalia col- 
lected during the study agree with the de- 
scriptions listed in Townes’ synonomy given 
in parenthesis above, thus confirming his 
association. 

Material examined: One male, 1 pupa, 
6-VIII-57: 1 male, 10-VIII-57; CR.L.. One 
pupal exuvia, 20-X-55; Ch.L. 


CHIRONOMUS (CRYPTOCHIRONO- 
MUS) GALEATOR (Townes) new 
combination 
Harnischia (Harnischia) galeator Townes, 1945: 170, 

description of adult male. 

Harnischia (Harnischia) galeator Townes; Beck and 
Beck, 1959: 95, adult distribution and phenology. 
Male: Wing length 1.62-1.75, mean 1.69 

mm (4); leg ratio 1.70-2.34, mean 1.95 

(4); antennal ratio 2.11-2.33, mean 2.22 

(4). 

Material examined: One male, 17-XI-55; 
Ch.L. Seventeen males, 27-VIII-57; 2 males, 
3-IX-57; 2 males, 10-[X-57; 1 male, 30- 
[X-57:, 4 males, 7-X-57; 1 male, 29-X-57: 
C.R.L. Two males, 2-III-57: 1 male, 29- 
IV-57; 1 male, 14-V-57; 1 male, 10-VII-57; 
at light, Natchitoches, La. 


CHIRONOMUS (CRYPTOCHIRONO- 
MUS) MONOCHROMUS van der Wulp 


Chironomus unicolor van der Wulp, 1858: 5, descrip- 
tion of adult. 

Chironomus monochromus van der Wulp, 1874: 129 
(new name for unicolor van der Wulp nec Walker, 
1848). 

Harnischia  (Harnischia) 
Wulp); Townes, 1945: 
tion. 

Harnischia (Harnischia) 
Wulp); Sublette, 1957: 
mature stages; ecology. 

Harnischia (Harnischia) monochromus 
Wulp); Beck and Beck, 1959: 95, 
and phenology of adult. 


monochromus (van der 
160, adult; generic posi- 


(van der 
391, description of im- 


monochromus 


(van der 
distribution 


Tendipes (Cryptochironomus) monochromus (van 
der Wulp); Dendy and Sublette, 1959: 516, 
generic position. 

Tendipes (Cryptochironomus) monochromus (van 
der Wulp); Sublette, 1960: 223, adults. 

Tendipes (Cryptochironomus) monochromus (van 


der Wulp); Darby, 1962: 38: 50, 53, 164, ecology; 
adult, in key. 


Males: Wing length 1.49-1.94, mean 1.74 
mm (4); leg ratio 1.50-1.80, mean 1.66 
(4); antennal ratio 1.80-2.10, mean 1.97 
(Sn 

Material examined: One male, 19-21-III- 


Tulane Studies in Zoology 


Vol. 11 


58: 1 male, 7-IV-58; 9 males, 21-IV-58; 2 
males, 30-IV-58; 3 males, 5-V-58; 2 males, 
12-V-58; 3 males, 6-VI-58; C.R.L. One male, 
3-IlI-57: 1 male, 11-III-57: 1 male, 122i 
57: 1 male, 16-III-57; 1 male, 28-IIJ-57; 4 
males, 8-IV-57; 2 males, 9-[V-57; 3 males, 
12-IV-57; 5 males, 16-IV-57; 3 males, 29- 
IV-57; 1 male, 6-V-57; 2 males, 14-V-57; 
at light, Natchitoches, La. One male, 20- 
[X-58; 1 male, 15-X-58; U.S.F.H. 


CHIRONOMUS (CRYPTOCHIRONO- 
MUS) EDWARSDI (Kruseman ) 


Chironomus (Chironomus) virescens Meigen; Ed- 


wards, 1929: 391, misidentification of virescens 
Meigen. 

Tendipes (Parachironomus) edwardsi Kruseman, 1933: 
194, new name for virescens Meigen of Edwards; 
adult male. 

Harnischia —(Harnischia) 
Townes, 1945: 167, adult. 

Harnischia (Harnischia) edwardsi (Kruseman), Beck 
and Beck, 1959: 95, adult distribution and_phe- 
nology. 

Tendipes (Cryptochironomus 
Sublette, 1960: 224, adult. 


edwardsi (Kruseman) ; 


edwardsi Kruseman; 

Larva: Described from exuviae. 

Head length 0.34 mm; mandible length, 
0.10 mm; head darkened on posterior gular 
region, tips of labial plate and mandibles; 
the latter slender, acutely tipped with basal 
teeth rather conspicuous, Figure 76. Labial 
plate, Figure 77; antenna, Figure 78; pre- 
mandible (torma), Figure 79. The epi- 
pharyngeal area differs somewhat from 
that illustrated for Pseduochironomus aix 
(Figure 37). Setae I and II, which are in 
line with one another, are large, simple and 
attentuate; seta II is slightly the larger; seta 
III is very small and inconspicuous and is 
located anterior and medial to II; the lateral 
spinulae are apparently absent, as are the 
pecten eppiharyngis, the chaetae, and the 
squama platia; there are about 5 chaetulae 
basales on each side. There is a_ single 
jointed dorsal labral bristle. 

Pupa: Total length 2.92 mm; exuviae 
length 3.38 mm, exuviae pale yellowish- 
brown. Respiratory organ with about 20 
fine branches. Cephalic tubercles prominent, 
acutely tipped (Figure 80). Tergite II with 
the usual posterior row of recurved hooks, 
the row interrupted along the midline and 
with about 12 hooks in each half. Tergites 
III to V with a partially doubled row of 
tubercle-like spines; tergite VI with a small 
mace-like tubercle (Figure 81) on midline, 
which is beset with the low tubercle-like 


No. 4 


spines of the preceding segments. Tergites 
VII and VIII devoid of spines. Posterolateral 
margin of segment VIII with a short atten- 
uate spine, Figure 82. Swim fin with a 
fringe of 48 bristles. 

Males: Wing length 1.39-1.70, mean 1.50 
mm (4); leg ratio 1.80-2.00, mean 1.91 
(4); antennal ratio 1.88-2.02, mean 1.97 
>) 

Material examined: One male, 21-IV-57; 
3 males, 23-VIII-57; 1 male, 13-VIII-57; 6 
males, 20-VIII-57; 3 males, 27-VIII-57; 1 
male, 3-IX-57; 4 males, 10-IX-57; 2 males, 
24-IX-57; 4 males, 30-[X-57; 10 males, 7- 
X-57; 21 males, 14-X-57; 4 males, 21-X-57; 
1 male, 29-X-57; 7 males, 4-XI-57; 2 males, 
11-XI-57; 1 male, 5-V-58; 1 male, 4-VI-58; 
C.R.L. Two males, 21-X-59; 1 male, 11- 
XI-59; Many, La. Two males, 20-X-55; 
Chk. 


GLYPITOTENDIPES (PHYTOTENDIPES) 
MERIDIONALIS Dendy and Sublette 


Glyptotendipes (Phytotendipes) paripes (Edwards) ; 
Sublette, 1957: 391, description of larva and 
pupa; ecology; misidentification of paripes (Ed- 
wards). 

Glyptotendipes (Phytotendipes) meridionalis Dendy 
and Sublette, 1959: 517, description of adult. 

Glyptotendipes (Phytotendipes) meridionalis Dendy 
and Sublette; Beck, 1961: 126, distribtution; phe- 
nology. 


The larva of meridionalis may be distin- 
guished from that of /obiferus, the only other 
Nearctic species of this subgenus known in 
the larval stage, by the accessory tooth of 
the mandible which is simple in meridtonalis 
and notched in /obiferus, and by the antenna 
the third segment of which is 0.60 as long 
as the second in meridionalis and 0.75 as 
long as lobiferus. 

The pupa of meridtonalis differs from 
that of lobiferus as follows (based on ma- 
terial before me) : 


exuviae length 

mean length times mean width of 
maces on abdominal 2nd segment 

3rd segment 

4th segment 

5th segment 

6th segment 

Color of spines of abdominal maces 

Mean number of spines on maces 
2nd segment 

3rd segment 

4th segment 

5th segment 

6th segment 

Number of caudo-lateral spines 
on segment 8 


Chironomids of West-Central Louisiana 


£35 


Material examined: One male, 12-IX-58; 
U.S.F.H. One male, 9-XI-56; North Shore 
of Red River, Grande Ecore, Natchitoches, 
La. Five males, 28-IJ-57; 1 male, 3-III-57; 
female, 9 12-IN-5:7593) males, 13-573 3 
males, 29-IV-57; 1 male, 3-V-57; 2 males, 
G-V-5/: 2 males, 14-V-57; 1 male, 15-V-57: 
3 males, 21-V-57; 1 male, 3-VI-57; 3 males, 
25-VI-57; 1 male, 12-IV-58; at light, Natchi- 
toches, La. One male, 7-[X-56; 4 males, 
11-IX-56; 2 males, 22-X-56; 1. male, 6-II- 
57; 1 female, 8-VI-57; 1 male, 18-VI-57; 
i amale; 19-Vil-57;, 1 male, 30-VIIL-57° 1 
male, 7-[X-57; 1 female, 17-IX-57; 1 male, 
1-X-57; 4 males, 7-X-57; 3 males, 1 female, 
14-X-57; 1 male, 4-XI-57; 1 male, 11-XI- 
57; 1 male, 26-II-58; 1 female, 1-III-58; 
1 male, 9-III-58; 2 males, 19-21-III-58; 2 
females, 7-IV-58; 1 male, 12-V-58; 7 males, 
2 females, 4-VI-58; 6 males, 6-VI-58; 1 
male, 10-VI-58; 2 males, 10-VII-58; C.R.L. 


GLYPTOTENDIPES (PHYTOTENDIPES) 
LOBIFERUS (Say) 


Chironomus lobiferus Say, 1823: 12, description of 
adult. 

Glyptotendipes (Phytotendipes)  lobiferus (Say) ; 
Townes, 1945: 142, description of adult; generic 
position. 


Glyptotendipes lobiferus (Say); Judd, 1949: 9, 
phenology. 
Glyptotendipes (Phytotendipes) lobiferus (Say); 


Berg, 1950: 92-94, description of larva and pupa; 
ecology. 
Glyptotendipes lobiferus [(Say)]; Gerry, 1951: 141- 
144, ecology. 
Glyptotendipes lobiferus 
phenology. 
Glyptotendipes lobiferus [(Say)]; Gerry, 1954: 148, 
ecology; control. 
Glyptotendipes lobiferus 
1956: 296, ecology. 
Glyptotendipes (Phytotendipes) lobiferus (Say); Ro- 
back, 1957c: 123, larva and pupa, in key. 


(Say); Judd, 1953: 813, 


Paine and Gaufin, 


(Say) ; 


Glyptotendipes lobiferus (Say); Judd, 1957: 401, 
phenology. 
meridionalis lobiferus 


8.00 mm (7-8.5) 10.00 mm (9-11) 
0.10 x 0.07 mm 
0.15 x 0.07 mm 
0.17 x 0.08 mm 
0.22 x 0.08 mm 
0.32 x 0.12 mm 


0.16 x 0.10 mm 
0.22 x 0.12 mm 
0.24 x 0.12 mm 
0.381 x 0.14 mm 
0.53 x 0.22 mm 


brown yellowish 
ff 8 

7 8 

a 8 

6 9 

11 19 

0-2 4-9 


Tulane Studies rn Zoology 


Vela 


Figures 76-82. Chironomus (Cryptochironomus) edwardsi (Kruseman). 76. larval man- 
dible; 77. labial plate; 78. antenna; 79. premandible (torma) ; 80. cephalic tubercles of 
pupa; 81. chaetotaxy of tergite VI; 82. spine of posterolateral margin of Segment VIII. 


Glyptotendipes (Phytotendipes) lobiferus (Say); 
Beck and Beck, 1959: 95, distribution and_phe- 
nology of adult. 

Glyptotendipes (Phytotendipes) lobiferus (Say): 
Dendy and Sublette, 1959: 518, adult, in table. 


Glyptotendipes lobiferus (Say); Judd, 1960: 207, 
phenology. 

Glyptotendipes (Phytotendipes)  lobiferus (Say); 
Sublette, 1960: 225, adult. 

Glyptotendipes lobiferus (Say); Judd, 1961: 96, 
phenology. 

Glyptotendipes (Phytotendipes) lobiferus (Say) ; 


Darby, 1962: 38, 39, 47, 59, 68, 74, 89, 101, 114, 
170-172; description of larva, pupa and adult; 
ecology. 

Males: Wing length 3.38-4.05, mean 3.78 
mm (5); leg ratio 1.40-1.66, mean 1.47 
(5); antennal ratio 4.12-5.10, mean 4.68 
(5); body length 7.40-8.00, mean 7.51 mm 
(6): Tao: Pag! 21-131 mean 1:26 (5). 

Material examined: One male, 23-VII-57; 
1 female, 18-VII-57; 1 male, 6-VIII-57; 2 
males, 17-IX-57; 1 female, 24-IX-57; 1 
female, 3-X-57; 1 male, 14-X-57; 1 female, 
1-IJI-58; 1 male, 1 female, 3-III-58; 1 male, 
3-IV-58: 1 male, 7-IV-58; 1 male, 14-IV-58; 
1 male, 21-IV-58; 2 males, 1 female, 12-V- 
58; 1 female, 19-V-58; 1 male, 19-VI-58; 
C.R.L. One male, 6-X-56; 1 male, 2-III-57; 
1 male, 3-III-57; 1 male, 29-IV-57; 2 males, 
14-V-57; 1 male, 15-V-57; 2 females, 28- 
VI-57; at light, Natchitoches, La. One male, 
12-IX-58; 1 male, 27-[X-58; 1 male, 15-X- 
5S; Were 


PARALAUTERBORNIELLA ELACHISTA 


( Townes ) 


Apedilum elachistus Townes, 1945: 33, description 
of adult. 

Apedilum elachistus Townes; Gerry, 1954: 146, ecol- 
ogy; control. 

Apedilum elachistus Townes; Beck and Beck, 1959: 
92, distribution and phenology of adults. 

Paralauterborniella elachistus (Townes); Dendy and 
Sublette, 1959: 513, generic position. 

nec!Paralauterborniella elachistus (Townes); Darby, 
1962: 46, and following, description of larva, 
pupa and adult; ecology. I consider Darby’s species 
to be a variety of Paralauterborniella subcincta 
subcincte. 


Males: Wing length 1.22-1.56, mean 1.39 
mm (4); leg ratio 1.29, 1.31 (2); antennal 
ratio 1.00-1.05, mean 1.03 (4). 

Material examined: One male, 28-VII-57; 
1 female, 13-VIII-57; C.R.L. One male, 20- 
VIII-58; 3 males, 28-VIII-58; 1 male, 12- 
IX-58; 1 male, 29-X-58; 1 male, 6-XI-58; 
1 male, 27-XI-58; 1 male, 27-X1I-59; U.S.F.H. 


LAUTERBORNIELLA VARIPENNIS 
( Coquillett ) 


Chironomus varipennis Coquillett, 1902: 94, descrip- 
tion of adult. 


Lauterborniella  varipennis (Coquillett); Townes, 
1945: 21, description of adult. 
Lauterborniella varipennis (Coquillett); |Hauber, 


1947: 459, description of larva and pupa; _phe- 
nology; ecology. 

Lauterborniella varipennis (Coquillett); Beck and 
Beck, 1959: 92, distribution and phenology of 
adult. 


No. 4 


Males: Wing length 1.45-1.69, mean 1.59 
mm (4); antennal ratio 1.35-1.66, mean 
SC). 

Material examined. Two females, 3-VII- 
57; 1 male, 30-VII-57; 2 males, 1 female, 
6-VIII-57; 1 female, 20-VIII-57; C.R.L. One 
female, 28-VIII-58; 1 male, 6-IX-58; 3 
males, 12-IX-58; 1 male, 20-[X-58; 1 male, 
27-IX-58; U.S.F.H. 


STENOCHIRONOMUS MACATEEI 
( Malloch ) 


Chironomus macateei Malloch, 1915b: 45, description 
of adult. 

Stenochironomus macateei (Malloch); Townes, 1945: 
89, adults. 

Stenochironomus maecateei (Malloch) ; 
Gaufin, 1956: 296, ecology. 
Males: Wing length 1.67-1.85, mean 1.77 

mm (3); leg ratio 1.13, 1.21 (2); antennal 

ratio 1.70-2.00, mean 1.83 (3). 


Material examined: Four males, 1 female, 
30-VII-57; light trap, Natchitoches, La. 


Paine and 


PEDIONOMUS gen. nov. 


Type species: Pedionomus beckae new 
species. 

Male antenna composed of 13 segments; 
female with 5 (2 basal segments fused im- 
movably together, thus considered as 1). 
Fork of the cubitus distinctly distal to r-m 
crossvein. Ry, parallel with Ros for most 
of length then slightly divergent at tip; dis- 
tinctly separated. Wing membrane devoid 
of macrotricha but with conspicuous micro- 
tricha at 100 magnification. Pronotum nar- 
rowed dorsally, inferior to anterior pro- 
jection of mesonotum. Dorsocentral and 
dorsolateral bristles conspicuous, in partial 
biserial rows. The dorsocentral bristles be- 
gin at the anterior margin of the mesoscu- 
tum and extend posteriorly to the conspicu- 
ous hump of the mesonotum. Anterior tibia 
with a projecting scale which bears, in ad- 
dition to a terminal small spine, 2 conspicu- 
ous setae (Fig. 83). Middle and hind tibiae 
with combs that are contiguous or over- 
lapping, not fused (middle tibial combs, 
Figure 84). The middle tibia has one long 
spine on the inner comb and none on the 
outer, while the hind tibia has a still longer 
spine on the outer but none on the inner 
comb; spines on both combs with slightly 
recurved tips. Pulvilli present, deeply pecti- 
nate but not bilobed as in Polypedilum. 


Chironomids of West-Central Louisiana 


13y/ 


Eighth abdominal segment not basally con- 
stricted as is Polypedilum. 

Male genitalia, Figure 85, similar to 
Stictochironomus and Polypedilum but lack- 
ing inner row of setae at apex of dististyle; 
basal part of superior appendage more elon- 
gate than in these genera. 

Etymology. Dwelling on the plains. 


PEDIONOMUS BECKAE new species 


Holotype male: U.S.N.M. No. 66460. 
Reared from a larva collected on floating 
wood, Cane River Lake, Natchitoches Parish, 
Louisiana, 1-X-57, J.ES. 

Head and antennal pedicel yellowish- 
brown, concolorous with ground color of 
thorax; antennal flagellum, narrow vittae 
and sternopleuron darker brown. Postocular 
bristles in a single row, reaching a point level 
with dorsal extension of eyes. Frontal tu- 
bercles absent; palpi normal, proportions 
7:15:15:25, clypeus with 20 bristles. An- 
tennal ratio 2.31. 


Pedicel of haltere pale; knob black; wing 
length 2.04 mm; venarum ratio 1.12; supra- 
alar bristles absent; prealar bristles 6; dorso- 
median bristles partially in 2 rows, long and 
erect; dorsolateral bristles partially in 2 
long, erect rows; scutellum with a posterior 
row of 18 heavy bristles in straight, trans- 
verse row; anteriorly with 8 fine bristles in 
a slightly staggered row. 


Leg proportions: 


Leg 

ER Dans oie Seat 

Foreleg 55 42 70 45 35 27 12 1.66 
Middle leg 60 50 33 20 15 8 5 0.66 
Hind leg 6357 Al 27-20. 1256 0.83 


Wing hair fringe long and dark; squama 
with about 10 hairs. Wing membrane de- 
void of macrotricha but with conspicuous 
microtricha visible at 100 magnification. 
R, parallel with Ro, to near tip where the 
two diverge slightly; tips distinctly separated. 
Ry,5 terminates over M; fork of Cu distal 
to r-m crossvein. 


Abdominal incisures pale; each tergite oc- 
cupied by a broad dark brown fascia; ab- 
domen thus with conspicuous vittate pattern. 

Genitalia (Fig. 85) superficially resemble 
that of several species of Polypedilum, 
Stictochironomus, and Tribelos but differs 
in lacking an inner apical row of bristles on 
dististyle, by having abbreviate dististyles, 
and by the distinctively shaped superior ap- 


138 
pendage. The almost parallel-sided eighth 
abdominal segment is in strong contrast 


with the triangular shaped one of Poly- 
pedilum. 

Allotype female: U.S.N.M. Reared from 
larva collected from floating wood, Cane 
River Lake, Natchitoches Parish, Loutsiana, 
1-X-57. 

Similar to the male except for sexual dif- 
ferences and a generally darker coloration. 
Head and thorax dark cinnamon brown, 
the thoracic vittae slightly darker; abdomen 
fasciate, the dark bands broader than in the 
male. Clypeus with 23 bristles; palpi pro- 
portions 5:13:13:24. 

Wing length 2.18 mm; venarum ratio 
1.19; prealar bristles 5 (paratype female); 
scutellar bristles 15 (paratype female). 


Leg proportions: 


Leg 

Betis ha ako ae ee LALO 

Foreleg 65 47 716) 42635. 21 12 62 
Middleleg 68 58 32 18 19 9 5 0.55 
Hind leg: 68 64 45 25 20 12 5 0.70 


Genitalia, Figures 86 and 87. Sperma- 
thecae very pale but discernible; spherical, 
each 0.08 mm in diameter. 


Larva: Described from exuviae. Head 
length 0.49 mm; mandible length 0.13 mm. 
Head yellowish except for tips of mandibles, 
labial plate and narrow occipital margin. 


Labial plate, Figure 88; antenna 5 seg- 
mented, Figure 89; mandible, Figure 90. 

Epipharyngeal apparatus similar to that 
illustrated for Psewdochironomus aix Townes 
(Figure 37). The pecten epipharyngis is 
composed of 3 palmate, contiguous plates; 
chaetulae basales about 7 in number, tips 
of at least 2 members finely pectinate; 
Squama platia with about 21 fine teeth form- 
ing a comb on each side; seta I palmate, 
shorter than in P. azx, distally fringed; seta 
II long, curved and undissected; seta III as 
in P, ax; seta [V present as 2 small peg-like 
structures on each side; chaetae about 6 on 
each side, finely pectinate; spinulae low and 
inconspicuous, about 4 on each side. Torma 
(premandible) yellowish, not distally dark- 
ened: with 3 terminal teeth which are pro- 
gressively reduced in size basally. 

Preanal papillae each with 8 long, yellow- 
ish bristles; on the anterior face of each 
papilla is the usual small bristle; below each 
papilla is a long yellow bristle. Each anal 
proleg with about 15 yellow claws. 


Tulane Studies n Zoology 


Vol. 11 


Pupal exuviae: Total length 4.97 mm. 
Thorax, Ist two abdominal segments, and 
caudal lobe infuscate, remainder of exuvia 
pale except for dark spines and spinulae and 
lateral margin markings. 


Cephalic tubercles scarcely discernible on 
the frontal plate; each low, rounded tubercle 
with a fine bristle. Respiratory organs long 
and finely branched, each with apparently 3 
main basal branches. 


Abdominal tergites as follows: I, devoid 
of shagreen. I, along the lateral margins 
are 2 fine, pale bristles each side. In the 
anterior one-third of the segment is a broad 
band of black spines like those illustrated 
on segment IV (Figure 91); at the posterior 
margin is the usual row of recurved black 
hooks numbering 37. In the center of the 
tergite on either side of the midline is an 
oval patch of fine shagreen. 


III to V, pattern of bristles, spines, spin- 
ulae and shagreen essentially identical (cf. 
Figure 91 for segment IV). Intersegmental 
membrane between segments IV and V with 
a band of black spines (Figure 91). Sha- 
green of segment III less contiguous along 
anterior margin. 


VI, anterior band of black spines absent. 
In the center of the tergite on either side 
of the midline is an ovoid patch of fine 
shagreen; near the posterior margin on either 
side of the midline is a patch of 10 to 12 
fine spinulae. 


VII, completely devoid of spines and 
shagreen. 


VII, posterolateral margin shown in Fig- 
ure 92. Swim fin with about 82 uniformly 
arranged lateral filaments. 


Paratype males: Wing length 1.78- 
mean 2.04 mm (9); leg ratio 1.51- 
mean 1.70 (5); antennal ratio 2.04- 
mean 2.34 (5). 


Paratypes: One male, 1 female, 3-VII-57; 
1 male, 13-VII-57; 2 males, 18-VII-57; 1 
female, 22-VII-57; 1 male, 13-VIII-57; 1 
female, 20-VIII-57; 1 female, 10-[X-57; 1 
male, 17-[X-57; 3 males, 3 females, 24-IX- 
57; 1 male, 1-X-57; 2 males, 14-X-57; 
C.R.L. One male, 21-V-57; 1 male, 3-VI-57; 
5 males, 30-VII-57; at light, Natchitoches, 
La. One male, 30-X-59; Old River, Cypress, 
La. One male, 1 female, 24-X-57; below 
Chivary Dam near Clarence, La. One male, 
23-XI-59; Bayou Pierre, 5 miles north of 


b= 00 
Neh 
WN 0 NO 
NNN 


Figures 83-92. Pedionomus beckae new species. 


Natchitoches, La. One male, 23-VI-57; 1 
male, 30-VI-57; Polk Co., Lakeland, Fla. 
One male, 20-IX-60; 2 males, 16-VI-61; 
Jackson Co., Fla. One male, 13-VII-56; 1 
male, 29-IX-60; Gadsen Co., Fla., Chatta- 
hoochee. One male, 16-X-57; Glades Co., 
Moore Haven, Fla. One male, 17-IX-60; 2 
males, 28-IX-60; 1 male, 15-X-60; We- 
wahitchka, Fla., Gulf Co. One male, 6-IX- 
57; Seminole Co., Geneva, Fla. One male, 
3-VIII-57; Miami, Dade Co. One male, 
8-XI-60; Indian River, Vero Beach, Fla. 
One male, 23-IX-60; 1 male, 4-X-60; Bro- 
ward Co., Andytown, Fla. One male, 6-IX- 


Chironomids of West-Central Louisiana 


139 


\ 


ew 


é . 7 


resi 
for 


Sad. Ba ee 


a 0OFY | 
84. middle tibial 


alheecers 
83. foretibial apex; 
combs; 85. male genitalia; 86. female genitalia; 87. variation of female genitalia; 88. 
labial plate of larva; 89. antenna of larva; 90. mandible; 91 chaetotaxy of tergite IV 
of pupa; 92. posterolateral margin of Segment VIII. 


57; Winter Park, Orange Co., Fla. One male, 
16-[X-57; Nokomis, Sarasota, Fla. One male, 
6-VII-57; Lake Worth, Palm Beach Co., Fla. 
One male, 9-VUI-57; Jacksonville Branch, 
Duval Co., Fla. Seven males, 8-V-62; Bayou 
George, Bay Co., Fla. Two males, 16-VI-60; 
Concho River Lake, San Angelo, Tex. One 
male, 14-VI-60; Lubbock, Tex. In the col- 
lections of USN.M. CNC, A.N:SP., 
I.N.H.S., Cornell University and Florida 
State Board of Health. 

Pedionomus beckae, new genus and spe- 
cies, differs from Polypedilum by having 
the ninth tergite of the male subtruncate 


140 


rather than triangular in outline; the disti- 
style lacks long bristles; and the pulvilli are 
not bifid. It differs from Stictochironomus 
by having a conspicuous spine at the apex of 
the fore tibia and in details of the male 
genitalia. 

The larva keys in Roback (1957c, pp. 96- 
98) to Polypedilum. It closely resembles the 
fallax group in having a labial plate with 
teeth of about equal length. It may be dis- 
tinguished by the paralabial plates which are 
about 4 times as broad as long. 

The pupa keys in Roback (loc. cit.) to 
Tanytarsus (part). This species does not fit 
well in his key to species of Tanytarsus. It 
can be distinguished from the species he 
described by the low rounded cephalic tu- 
bercle bearing a fine bristle. 

I take pleasure in naming this species in 
honor of Mrs. Elisabeth Beck, Florida State 
Board of Health, who supplied the paratype 
material from Florida. 


POLYPEDILUM (POLYPEDILUM) 
TRIGONUM Townes 


Polypedilum (Polypedilum) 
49, description of adult. 

Polypedilum (Polypedilum) trigonus Townes; Hauber, 
1947: 462, adult. 

Polypedilum (Polypedilum) trigonus Townes; Dendy 
and Sublette, 1959: 513, adult. 


trigonus Yownes, 1945: 


Polypedilum (Polypedilum) trigonus Townes; Beck 
and Beck, 1959: 92, distribution; phenology of 
adult. 


Males: Wing length 1.71-1.98, mean 1.83 
mm (3); leg ratio 1.73 (1); antennal ratio 
205-225, Mean ZoLom( oy): 

Material examined: One male, 3-IX-57; 
1 male, 17-IX-57; 1 male, 24-IX-57; 1 male, 
30-IV-58; C.R.L. 


POLYPEDILUM (POLYPEDILUM) 
ILLINOENSE (Malloch) 


Chironomus illinoensis Malloch, 1915a: 471, descrip- 
tion of adult. 


Polypedilum (Polypedilum) illinoense (Malloch) ; 
Townes, 1945: 57, adult. 

Polypedilum  (Polypedilum) illinoense (Malloch) ; 
Hauber, 1947: 462, description of larva and pupa; 
ecology. 

Polypedilum  (Polypedilum) illinoense (Malloch) ; 
Berg, 1950: 91-92, description of larva and pupa; 
ecology. 

Polybedilum  (Polypedilum) illinoense (Malloch) ; 


Roback, 1953: 124, ecology. 
Polypedilum illinoense (Malloch); Wurtz and 
back, 1955: 200, distribution; ecology. 


Ro- 


Polypedilum  (Polypedilum)  illinoense (Malloch) ; 
Tebo, 1955: 97, ecology. 
Polypedilum  (Polypedilum) illinoense (Malloch) ; 


Paine and Gaufin, 1956: 296, ecology. 


Tulane Studies in Zoology 


Vol. 11 
Polypedilum  (Polypedilum)  illinoense (Malloch) ; 
Sublette, 1957: 387, ecology; phenology. 
Polypedilum (Polypedilum) illinoense (Malloch) ; 
Roback, 1957c: 117, description of larva and pupa. 
Polybedilum (Polypedilum)  illinoense (Malloch) ; 
Dendy and Sublette, 1959: 513, adult. 
Polypedilum (Polypedilum)  illinoense (Malloch); 


Beck and Beck; 1959: 93, distribution and phe- 

nology of adult. 
Polypedilum  (Polypedilum) 

Sublette, 1960: 207, adult. 

Males: Wing length 1.67-2.34, mean 1.98 
mm (3); leg ratio 1.57-1.80, mean 1.68 
(3); antennal ratio 1.73-2.00, mean 1.91 
(oy) 

Material examined: One male, 5-II-57; 
1 male, 28-III-57; 1 male, 9-IV-57; 2 males, 
6-V-57; 1 male, 14-V-57; 1 male, 21-V-57; 
2 males, 3-VI-57; 1 male, 11-VII-57; 1 male, 
13-VIII-57; 1 male, 1-XI-57; 1 male, 5-V- 
Doe ele: 


POLYPEDILUM (POLYPEDILUM) 
DIGITIFER Townes 

Polypedilum (Tripodura) digitifer Townes, 1945: 45, 
description of adult. 

Polypedilum (Tripodura)  digitifer Townes; Sub- 
lette, 1957: 386, description of larva and pupa; 
ecology; phenology. 

Polypedilum (Tripodura) digitifer Vownes; Beck 
and Beck, 1959: 92, distribution and phenology 
of adult. 

Polypedilum (Polypedilum) digitifer Townes; Dendy 
and Sublette, 1959: 513, adults. 

Polypedilum (Polypedilum) digitifer Townes; 
lette, 1960: 206, adults. 

Polypedilum (Polypedilum) digitifer Townes; Darby, 
1962: 38, 48, 49, 146, 150, ecology. 

Males: Wing length 1.56-1.80, mean 1.69 
mm (4); leg ratio 1.83-2.08, mean 1.94 
(4); antennal ratio 1.54-2.00, mean 1.81 
(42D) 

Material examined: Four males, 20-X-55; 
3 females, 21-X-55: 1 male, 27-X-55: Gh. 
One male, 7-[X-56; 1 male, 5 females, 22- 
X-56; 1 male, 2 females, 10-XII-56; 2 males, 
26-II-57; 1 male, 3-IIJ-57; 1 male, 28-II- 
57; 3 males, 5-[X-57; 2 males, 8-IV-57; 1 
male, 12-[V-57; 1 male, 14-IV-57; 4 males, 
16-IV-57; 1 male, 29-IV-57; 1 female, 30- 
IV-57; 1 male, 6-V-57; 1 male, 14-V-57; 
1 male, 21-V-57; 1 intersex, 30-V-57; 1 
male, 11 females, 3 intersexes, 18-VI-57; 6 
males, 26 females, 5 intersexes, 25-VI-57; 
1 male, 28-VI-57; 2 females, 2-VII-57; 1 
male, 10-VII-57; 1 female, 1 intersex, 13- 
VII-57; 5 males, 7 females, 15-VII-57; 2 
females, 16-VII-57; 3 females, 18-VII-57; 3 
males, 23-VII-57; 1 male, 6-VIII-57; 2 males, 
20-VIII-57; 2 males, 2 females, 27-VIII-57; 


(Malloch) ; 


illinoense 


Sub- 


No. 4 


3 females, 28-VIII-57; 1 female, 10-[X-57; 
1 male, 7-IX-57; 1 female, 30-IX-57; 1 
intersex, 3-X-57; 1 male, 5-X-57; 4 males, 
21-X-57; 3 males, 1 female, 30-XI-57; 1 
male, 12-V-58; 1 male, 10-VI-58; C.R.L. 
Two males, 21-III-54; 1 male, 3-JII-57; 1 
male, 10-III-57; 4 males, 12-III-57; 2 males, 
13-I11-57; 3 males, 16-III-57; 2 males, 28- 
I-57; 3 males, 5-IV-57; 1 male, 21-V-57; 
1 male, 1 intersex, 3-VI-57; 2 males, 16- 
VII-57; at light, Natchitoches, La. 


Tribe Tanytarsini 


TANYTARSUS (CLADOTANYTARSUS) 
VIRDIVENTRIS Malloch 


Tenytarsus virdiventris Malloch, 1915a: 491, descrip- 
tion of adult. I have examined Malloch’s type 
series. 

Tanytarsus Walker; Hauber, 1944: 456, 
description of pupa and adult, misidentification of 
mancus Walker. I have examined Hauber’s mate- 
rial. 

Calopsectra viridiventris (Malloch); Johannsen (in 
Johannsen and Townes), 1952: 26, adult, in key. 

Tanytarsus (Cladotanytarsus) viridiventris Malloch; 
Dendy and Sublette, 1959: 513, adult. 

Tanytarsus (Cladotanytarsus) viridiventris Malloch; 
Warbyael 962.5385) 55 S65 19 707A Oe OBR 1 Olle 
110, 111, 172-179, description of larva, pupa and 
adult; ecology. 


Males: Wing length 1.40-1.80, mean 1.61 
mm (3); leg ratio 1.73-2.00, mean 1.88 
(3); amtennal ratio 1.21-1.39, mean 1.30 
(3). 

Material examined: Five males, 25-VI-57; 
1 male, 4-XI-57; 1 male, 19-VI-58; C.R.L. 
One male, 12-VIII-58; 1 male, 20-VIII-58; 
1 male, 28-VIII-58; 1 male, 4-X-58; U.S.F.H. 


Mancus 


TANYTARSUS (TANYTARSUS) 
CONFUSUS Malloch 


Tanytarsus confusus Malloch, 1915a: 490, descrip- 
tion of adult. I have examined Malloch’s type 
series. 

Tanytarsus (Calopsectra) sp. B Hauber, 1944: 454. 
I have examined Hauber’s material and it appears 
to belong here. 

Calopsectra confusa (Malloch); Johannsen (in Jo- 
hannsen and Townes), 1952: 26, adult, in key. 

Calopsectra confusa (Malloch); Roback, 1956: 113- 
116, description of larva, pupa and adult; 1957c: 
131-132, larva and pupa, in key. 

Calopsectra neoflavellus (Malloch); Sublette, 1957: 
385, description of larva and pupa; ecology; phe- 
nology: misidentification of mneoflavellus Malloch. 

Tanytarsus (Tanytarsus) confusus Malloch; Dendy 
and Sublette, 1959: 513, adult. 


Males: Wing length 1.80-2.12, mean 1.90 
mm (4); leg ratio 2.67-2.73 (2); antennal 
ratio 1.33-1.61, mean 1.42 (5). 

' Material examined: One male, 20-VIII- 


Chironomids of West-Central Louisiana 


141 


57; 1 male, 30-IX-57; 1 male, 7-X-57; 
C.R.L. Four males, 7-VIII-39; Lake Oko- 
boji, Iowa, U. A. Hauber. 


TANYTARSUS (TANYTARSUS) 
DENDYI new species 
Tanytarsus ejuncidus Walker (?); Hauber, 1944: 

455, description of pupa and adult, misidentifica- 

tion of ejuncidus. 

Holotype male: U.S.N.M. No. 66461; 
collected in a tent trap, Cane River Lake, 
Natchitoches, La., 11-XI-57; B. R. Buckley. 

Antennal pedicel, thoracic vittae, postno- 
tum and _ sternopleuron — blackish-brown; 
ground color of head, thorax and entire ab- 
domen yellowish-green; antennal flagellum 
and legs infuscate; halteres pale; postoculars 
in a single staggered row; frontal tubercles 
present, 0.03 mm long; palpal proportions: 
10:15:15:25; antennal ratio 1.44. 

Wing length 2.07 mm; venarum ratio 
1.11. No supra-alar bristles; dorsomedial 
and dorsolateral bristles in a single row; 
scutellum with about 6 erect bristles in a 
straight transverse row. 

Combs of middle and hind tibiae well 
separated, both inner and outer on each with 
a spur. Pulvilli not visible at 100 magnifi- 
cation. 


Leg proportions: 


Leg 

ie as fo el gk pera tO 

Foreleg AG 26) 65) 32002725 108250 
Middlelee 52 44 26 15 10 8 5 0.59 
Hind leg Smo 4222 bee OL70 
Wing membrane well haired on distal 


half; f-Cu distal to r-m. 

Genitalia (Figure 93) very similar to 
xanthus new species, but differing in having 
appendage la less than half the length of the 
apex, and by having the group of bristles on 
the ninth tergite. Figure 94 shows a variant 
collected at the type locality. 

Paratype males: Wing length 1.62-2.25; 
mean 1.87 mm (8); leg ratio 2.40-3.00, 
mean 2.60 (7); antennal ratio 1.21-1.76, 
mean 1.39 (7). 

Paratypes: One male, 3-II-58; 1 male, 
5-V-58; C.R.L. One male, 23-II-60; below 
Chivary Dam, Natchitoches Parish, La. One 
male, 16-III-57; at light, Natchitoches, La. 
Two males, 5-VI-56; 1 male, 6-VI-56; 4 
males, 8-VI-56; 1 male, 15-XI-56; 1 male, 
15-I11-57; 1 male, 28-10-57; Auburn, Ala.. 
J.S. Dendy. One male, 5-V-41; 1 male, 12- 
II-43; 2 males, 28-IV-43; 2 males, 25-V-43; 


142 


1 male, 5-VI-43; Davenport, Iowa. One 
male, 15-IV-42; 1 male, 12-II-43; Credit 
Island, Iowa. One male, 1-IV-42; 1 male, 
10-IV-42; Duck Creek Park, Iowa. One 
male, 16-VI-60; Concho River Lake, San 
Angelo, Tex. In the collections of C.N.C., 
A.N.S.P., I.N.H.S., and Cornell University. 

This species keys in Johannsen (in Jo- 
hannsen and Townes, 1952, page 25) to neo- 
flavellus Malloch; it may be distinguished 
by the distinctively different superior ap- 
pendage of the male genitalia which in 
neoflavellus has a strong lateral tubercle. 
Also, the punctae of the anal point in dendyi 
n. sp. are in a single row; in neoflavellus 
they become multiple basally. 

This species is named for Dr. J. S. Dendy, 
Auburn University, Auburn, Alabama, who 
contributed the paratype series from Ala- 
bama. 


TANYTARSUS (TANYTARSUS) 
XANTHUS new species 
Tenytarsus (Tanytarsus) neoflavellus Malloch; Dendy 
and Sublette, 1959; 613, adult. Misidentification 
of neoflavellus Malloch. This identification had 
been based on an examination of a specimen in 
the JNHS Collection, determined by Malloch. 

A recent examination of the lectotype by the 

author has revealed Malloch’s type series to be 

of one species and the named specimen (slide no. 

3075) to be another species, described here as new. 

Holotype male: U.S.N.M. No. 66462. Col- 
lected from the hatchery ponds at the U. S. 
Fish Hatchery, Natchitoches, Louisiana, 12- 
VIII-58; funnel trap, R. F. Tyler. 

Head, thorax and abdomen pale yellow, 
the thorax slightly darker because of the 
wing musculature. 

Eyes somewhat reniform, the dorsal ex- 
tension rather short and broad. Frontal tu- 
bercles present, length, 0.041 mm. Palpal 
proportions, 10:21:22:40. Antennal ratio, 
1.53. Postocular bristles long and erect, in 
a single row of about 16 bristles. 

Pronotum considerably below the rounded, 
projecting apex of the mesonotum; prono- 
tum halves slightly notched. Dorsolateral 
and dorsomedial bristles long and erect, in 
a single row, the latter slightly staggered at 
mesonotal apex. A single heavy prealar bris- 
tle present. Scutellum with 6 bristles. 

Wing membrane well haired almost to 
base. R45 ends proximal to M and distal 
to Cu;. Wing length, 1.69 mm. 

Fore tarsus not bearded. Pulvilli absent; 
empodium finely dissected, almost as long 


Tulane Studies in Zoology 


Volania 


as claws which are almost straight being 
curved slightly only near tip. 


Leg proportions: 

FT Tay a eee 
Foreleg 70 41105 47 40 32 18 
Middle leg 77 62 38 21 16 10 7 0.61 
Hind leg 85 73 56 35 32 19 

Both combs of middle and hind leg 
spurred; spurs of middle leg of unequal 
length; ratio, 15:10; longer spur curved near 
tip. Spurs of hind tibia almost equal in 
length; outer spur more strongly curved near 
tip than middle tibial spur. 

Genitalia, Figures 98 and 99, with anal 
point sparsely and coarsely punctate; append- 
age la almost as long as apex of superior 
appendage; ninth tergite with bristles. 

Female not associated. 

Paratype males: Wing length 
mean 2.07 mm (4); leg ratio 
mean 2.35 (4); antennal ratio 
meam 1:57" (>))F 

Paratypes: Louisiana: one male, 12-VIII- 
58; 1 male, 6-EX-58; 1 male, 12-IX-58: 2 
males, 20-IX-58; 1 male, 27-[X-58; 2 males, 
4-X-58; 2 males, 15-X-58; 1 male, 29-X-58; 
5 males, 6-XI-58; 6 males, 4 pupal exuviae, 
27-XI-58; 1 pupal exuvia, 12-XII-58; 1 
pupal exuvia, 10-I-59; 1 male, 15-I-59; 2 
males, 25-I-59; 8 males, 5-II-59; 2 males, 
7-II-59; 1 male, 9-II-59; 10 males, 15-II-59; 
U.S.F.H. One male, 8-IV-57; 2 males, 9-IV- 
57; 1 male, 28-VI-57; at light, Natchitoches, 
La. Illinois: one male, Peoria, Ill., 22-X-14, 
slide 3075 (In IN.HS. sub meoflavellus 
Malloch; determined J. R. Malloch). 

Larva: Described from exuvia of reared 
male. 

Head capsule pale except for the yellow- 
ish mandibular tips, labial plate and narrow 
occipital margin. Head length 0.40 mm. 
Mandible (Fig. 95) 0.12 mm long. An- 
tenna (Fig. 96) 0.20 mm long; basal tu- 
bercle without a spur. Epipharyngeal area 
similar to other Chironominae; labial plate, 
Figure 97; pecten epipharyngis composed 
of 3 digitate blades; chaetulae basales finely 
pectinate distally, apparently 5 on each side; 
torma (premandible) gently curved distally, 
yellowish at tip, somewhat obscured on the 
mount before me so that bifurcation not 
visible; 5 long curved filiform chaetae on 
each side; squama platia with about 24 ex- 
ceedingly fine teeth: seta I rather closely 
spaced together, coarsely pectinate; seta II 


No. 4 


anter:or to I and directly in line with it, 
long curved filiform seta reaching posteriorly 
to the squama platia; seta HI minute, an- 
terior to II and in line with it; spinulae 
somewhat obscured; dorsal labral bristles 
not evident. 


Preanal papillae short, each bearing 8 con- 
spicuous long blackened bristles. Posterior 
prolegs with 14 hooked yellowish claws. 


The larva may be distinguished from re- 
lated Nearctic species by the paralabial plates 
lying close together at the midline; the an- 
tennal tubercle lacking a spine; and the 
petiole of the Lauterborn organs being about 
as long as the last 3 antennal segments. 


Pupa: Described from exuviae of reared 
males and from exuviae found in funnel 
traps in which adult males were taken. 


Exuviae length 5.18 mm; cephalic tuber- 
cles, respiratory organs and abdominal chae- 
totaxy as illustrated by Roback (1957c) for 
neoflavellus; comb of segment VIII similar 
to that figured by Roback (op. cit.) but 
more spinose (5-10 large marginal spines; 
about 20-25 smaller disc spines); swim fin 
with a fringe of 39-40 flattened bristles on 
either side; disc finely shagreened. 


Paratypes in the collections of C.N.C., 
A.NS.P., IIN.H.S. and Cornell University. 


TANYTARSUS (TANYTARSUS) 
NEOFLAVELLUS Malloch 


Tanytarsus neoflavellus Malloch, 1915: 
tion of adult. 

Tanytarsus neoflavellus Malloch; 
distribution and phenology. 
Calopsectrs neoflavella (Mallcch) ; 
hannsen and Townes), 1952: 26, adult, in key. 
Tanytarsus neoflavellus Malloch; Hauber, 1944: 455. 
Nec! Tanytarsus (Tanytarsus) neoflavellus Malloch; 


489, descrip- 
Boesel, 1940: 19, 


Johannsen (in Jo- 


Dendy and Sublette, 1959: 513, adult, misiden- 

tification. 

Nec! Calopsectra neoflavellus (Malloch) Sublette, 

1957: 385, misidentification. 

Males: Wing length 1.67-2.25, mean 1.94 
mm (5); leg ratio 2.56-3.33, mean 2.93 
(3): antennal ratio 1.20-1.44, mean 1.30 
(oe 


Material examined: One male, 11-VI-57; 
4 males, 18-VI-57; 3 males, 25-VI-57; 1 
male, 15-VII-57; 3 males, 23-VII-57; 1 male, 
24-IX-57; 1 male, 7-X-57; 1 male, 11-XI-57; 
imate. 5-V-58: 1.male. 10-ViI-5S: G.R.L. 
Five males, 16-III-57; 1 male, 8-IV-57; 
1 male, 29-IV-57; at light, Natchitoches, La. 
One male, 6-XI-58; US.F.H. One male, 


Chironomids of West-Central Loutstana 


143, 


6-VI-56; 2 males, 8-VI-56; 1 male, 28-III-57; 
1 male, 8-IV-57; Auburn, Ala. One male, 
8-VIII-39; 5 males, 7/8-VIII-39; 2 males, 
6-VIII-49; Lake Okoboji, Iowa. One male, 
2-IX-62; Stratford, Conn. 

This species resembles xanthus new spe- 
cies as well as dendyi new species. It differs. 
significantly in genitalia characteristics. The 
tubercle on the superior appendage and the 
details of the anal point are distinctive. 


TANYTARSUS (TANYTARSUS) 
RECENS new species 

Holotype male: U.S N.M. No. 66463. Col- 
lected from Cane River Lake, Natchitoches, 
Louisiana, 12-V-58, B. R. Buckley. 

Head, thorax and abdomen pale stramine- 
ous; frontal tubercles small, conical, length 
0.02 mm; antennal ratio 1.09. Palpal pro- 
portions 7:17:19:30: 

Wing length 1.62 mm; one prealar bristle; 
halteres pale. 

Combs of middle and hind legs separated; 
without spurs. Tip of legs obscured or miss- 
ing so pulvilli not observed. 


Leg proportions: 


Leg 
B Die lage | coe Wea dp ey Eaule. 
Foreleg 75 48108 - - — — 2.25 


Middle leg 80 
Hind leg’ 95 85 


Wing membrane well haired. 


The genitalia, Figure 100, are very similar 
to guadratus new species but with the fol- 
lowing differences: appendage la evenly at- 
tenuate, not swollen near center; anal point 
with the basal ornamentation closed below 
(basally); tip of anal point more rounded; 
appendage 2 (inferior appendage) strongly 
capitate with 4 to 5 posteriorly directed bris- 
tles; appendage 2a with long terminal lamel- 
lae; ninth tergite with only 2 rather in- 
conspicuous bristles. 

This species is very similar to varela (Ro- 
back) (cf. Roback, 1957c, page 128, fig. 
455) but if his specimen is accurately fig- 
urged this is a distinct new species. 


TANYTARSUS (TANYTARSUS) 
BUCKLEYI new species 
Holotype male: U.S.N.M. No. 66464. Col- 
lected from Cane River Lake, Natchitoches 
Parish, Louisiana, 17-IX-57, B. R. Buckley. 
Frontal tubercles small, cylindrical; length. 


144 


0.03 mm; palpal proportions 8:15:19:28. 
Antennal ratio 0.96. 


Ground color of head and thorax yellow- 
ish; antennal pedicels, pronotum, mesonotal 
vittae, postnotum and sternopleuron cinna- 
mon brown; halteres pale. Wing length 1.36 
mm. Prealar bristles 1; dorsolateral and 
dorsomedial bristles in one row, large and 
erect; scutellar bristles 6, large and erect. 


Except for the coxae the legs are pale; 
forelegs slightly darker; tarsal beard absent. 
Tibial combs of middle and hind legs well 
separated; those of the hind leg each with 
a spur; those of middle leg with a spur only 
on the inner comb. Pulvilli not visible at 
100 magnification. 


e€ tions: 
Leg proportions ee 
Dit Pai ou ke Secale 


Foreleg 60 34 67 38 30 20 10 1.97 
Middlelege 60 48 26 15 10 6 5 0.54 
Hind leg 68 63 40 25 22 14 10 0.63 


Wing membrane sparsely haired only on 
distal one-sixth; Ry,; terminates proximal 
to M and distal to Cu. 


Abdomen greenish-yellow. Genitalia, Fig- 
ure 101. 


Paratype males: Wing length 1.19-1.80, 
mean 1.47 mm (5); leg ratio 1.87-2.10, 
mean 1.96 (3); antennal ratio 0.90-1.21, 
mean 1.07 (4). 


Paratypes: One male, 13-VII-57; 1 male, 
20-VIII-57; 2 males, 3-IX-57; 2 males, 30- 
IX-57; 1 male, 14-X-57; 2 males, 11-XI-57: 
Lvmale, 9=XUI-57: 1 dale, “WG=xXIL57: 4 
males, 23-XII-57; C.R.L. In the collections 
OCG. 2A Nise: 

This species keys to pusio (Meigen) in 
Johannsen’s key (in Johannsen and Townes, 
1952, page 26). I have examined Johann- 
sen’s material at Cornell University and the 
species which he identified as pusso Meigen 
is totally different from buckleyi new spe- 
cies. Further, his material does not agree 
with the scanty description of pwsio in Euro- 
pean literature. Pwszo probably does not 
occur in the Nearctic region. Tanytarsus 
(Tanytarsus) buckleyi new species is very 
similar to glabrescens Edwards, a Palaearctic 
species, but appears to differ in details of 
genitalic structure (cf. Edwards, 1929, fig- 
ure 15f; Brundin, 1947, figure 112). 


This species is named in honor of Burton 
R. Buckley, Natchitoches, Louisiana, who 


Tulane Studies n Zoology 


Vol. 11 


collected the specimen designated here as 
holotype. 


TANYTARSUS (TANYTARSUS) 
QUADRATUS new species 


Holotype male: U.S.N.M. No. 66466. Col- 
lected from Cane River Lake, Natchitoches 
Parish, Louisiana, 2-II-58, funnel trap, B. R. 
Buckley. 

Frontal tubercles small, cylindrical, length 
0.25 mm; antennal ratio 1.16. Palpal pro- 
portions 5:13:15:23. 

Ground color yellowish; antennal pedicels, 
pronotum, mesonotal vittae, sternopleuron 
and postnotum dark cinnamon brown. Scu- 
tellum infuscate. Mesothorax with a dis- 
tinct hump; halteres pale. Wing length 1.72 
mm. Prealar bristles 2; dorsolateral and 
dorsomedial bristles in one row;  scutellar 
bristles 8, large and erect; in a straight, 
transverse row. 

Legs pale; posterior 4 tibial combs small, 
well separated, without spurs; forelegs not 
bearded. Pulvilli not visible at 100 mag- 
nification. 


Leg proportions: tae 
F Ti Tay “5 39 eeeeeee 


Foreleg 85 63 86 50 40 27 138 1.36 
72 45 25 20 7 T 0.63 


Middle leg 90 
Hind leg 95 95 62) 40) 25 ombe0265 


Wing membrane well haired almost to 
wing base. 

Abdomen with yellowish ground color, 
each tergite almost completely covered by a 
brown band, thus giving abdomen vittate 
appearance. 

Genitalia, Figure 106, very similar to dzs- 
stmulis Johannsen and recens new species. 

Paratype males: Wing length 1.28-1.99, 
mean 1.59 mm (5); leg ratio Laieiea 
(2); antennal ratio 1.05-1.26, mean 1.12 
(Geple 

Paratypes: One male 27-VIII-57; 1 male, 
3-J1-58; 1 male, 22-II-58; 1 male, 2-III-58; 
3 males, 12-V-58; C.R.L. In the collections 
Of GIN. G., ACN Sub: 

This species runs in Johannsen’s key (in 
Johannsen and Townes, 1952, pages 25, 26) 
to dissimilis Johannsen. It may be differ- 
entiated from that species by having a more 
setose ninth tergite, a shorter more quadrate 
anal point, and by differently shaped ap- 
pendages la and 2a. It differs from recens 
new species in several features as noted 
under that species. 


No. 4 Chironomids of West-Central Louisiana 1 


<i 
i ¢ 
t A 
7s “A \ 
f 6 J | 
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i \ | \ 
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f 
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oer een 
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X @ L 96 a 
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93 
7 
SS SSS Fie Lf 
aS = 9 ; 
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Le p ( 
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a i | 
A Le 
0 7S ) | fos y a 
aS 8 (Ne naam) 1Q\ 
4, ~\\ 
/ Hes | i 
/ ¢ / Wasi Ss) iD Al 2 \ 
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A f ‘\ _¥ PS a \ 
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Figures 93-94. Tanytarsus (Tanytarsus) dendyi new species. 93. male genitalia; 94. 
variation of male genitalia. Figures 95-99. Tanytarsus (Tanytarsus) xanthus new spe- 
cies. 95. larval mandible; 96. antenna; 97. labial plate; 98. male genitalia; 99. details 
of appendage-2a. Figure 100. Tanytarsus (Tanytarsus) recens new species. 100. male 


genitalia. Figure 101. Tanytarsus (Tanytarsus) buckleyi new species. 101. male gen- 
italia. 


aN 


WN 


146 


TANYTARSUS (TANYTARSUS) 
ALLICIS new species 


Holotype male: U.S.N.M. No. 66465; U. S. 
Fish Hatchery, Natchitoches, Louisiana, 29- 
X-58, R. F. Tyler. 

Antennal pedicel, thoracic vittae, postno- 
tum and mesosternum dark brown; ground 
color of head, thorax and abdomen stramine- 
ous; legs, antennal flagellum and _halteres 
scarcely darkened. Antennal ratio 1.00. 
Frontal tubercles small, conical; length 0.02 
mm. Palpal proportions 6:15:19:30. 

Wing length 1.49 mm. Dorsomedial and 
dorsolateral bristles in one row, erect, scu- 
tellum with 5 long erect bristles. 

Fore tarsus not bearded; middle tibial 
combs with a single spur; hind tibiae with a 
spur on each comb. Pulvilli not visible at 
100 magnification. 


Leg proportions: 


Leg 

Per Ua, os: sata etatio 

Foreleg 66 385 58 42 32 25 12 1.66 
Middleleg 70 56 33 17 11 8 7 0.59 
Hind leg 75 75 50 28 25 18 10 0.67 


Wing membrane with macrotrichia on dis- 
tal third only; longitudinal veins with macro- 
trichia almost the entire length. 

Genitalia, Figure 102, mounting variant, 
Figure 103, very similar to buckleyi new spe- 
cies but differing in the narrower anal point, 
and the strikingly different appendage 2a; 
appendage la is also different. 

Paratype males: Wing length 1.26-1.58, 
mean 140 mm (3); leg ratio 2.09 (1); 
antennal ratio 0.91-1.06, mean 0.97 (3) 

Paratypes: Two males, 27-I[X-58; 1 male, 
6-XI-58; U.S.F.H. One male, 16-X-57; San 
Jose Creek, Many, La. One male, 13-III-57; 
at light, Natchitoches, La. In the collections 
OL CONC 7 AEN: S Ee. 

This species is very similar to the pale- 
arctic species, Tanytarsus (Tanytarsus) re- 
curvatus Brundin (cf. Brundin, 1947, page 
75, figure 113). It differs only in slight 
features of appendages la and 2a. Among 
the Nearctic fauna it most closely resembles 
buckleyi new species but is differentiated as 
was described under that species. 

This species was reared from larvae col- 
lected in aquatic vegetation. 

Larva: Exuviae not recovered. 

Pupa: Described from exuvia of reared 
male. Exuvia approximately 2.48 mm long, 
yellowish-brown with blackish-brown lateral 
longitudinal markings. Respiratory organ a 


Tulane Studies n Zoology 


Vokwit 


simple tubular filament, approximately 0.32 
mm long. Abdominal chaetotaxy shown dia- 
gramatically in Figure 104. Posterolateral 
comb of segment VIII as in Figure 105. 
Swim fin with about 18 to 20 filaments. 

The pupa keys in Roback (1957c, page 
132) to couplet 7. It may be distinguished 
from the species designated by Roback as 
Calopsectra (1.e., Tanytarsus) neoflavella? 
(Malloch) by the shorter respiratory organs 
(0.32 mm versus 0.53 mm); by a different 
pattern of chaetotaxy (cf. Figure 104 and 
Roback’s figure 516) and by different 
caudo-lateral combs (cf. Figure 105 and 
Roback’s figure 486). 


TANYTARSUS (TANYTARSUS) 
LIMNETICUS new species 


Holotype male: U.S.N.M. No. 66467. Col- 
lected from the U. S. Fish Hatchery, Natchi- 
toches, Louisiana, 20-VIII-58, tent trap, R. F. 
Tyler. 

Postocular bristles in one row, beginning 
medial to dorsal extension of eyes; dorsal 
extension of eyes rather broad, extension 
having 4 to 5 facets in a transverse row; 
minute but distinct frontal tubercles, length 
0.02 mm (paratype); palpal proportions 
5:10:10:22; antennal ratio 1.44. Head, 
thorax and abdomen pale; vittae, sterno- 
pleuron and postnotum pale ocherous. Wing 
length 1.71 mm; venarum ratio 1.06. Pre- 
alar bristles 3; dorsomedian bristles long 
and erect; dorsolateral bristles long, erect, in 
one row; about 14 scutellar bristles in a 
transverse row. 

Legs with tibial combs separate, each 
comb with a spur; forelegs darkened beyond 


tibia. Small pulvilli visible at 100 mag- 
nification. 

Leg proportions: Tee 

F Ti Ta, «9° 4 nes 

Foreleg 55) 28° 70) 32° 4 =) SeSee50 

Middleleg 50 40 26 16 13 8 6 0.65 

Hind leg 57 62-385 22°17 10a yeoieT 


Wings with Ry,; terminating distal to 
M; f-Cu slightly distal to r-m; wing mem- 
brane well haired; C and R with long con- 
spicuous pale hairs. 

Genitalia, Figures 108, 109, 110. 

Paratype males: Wing length 1.67-2.03, 
mean 1.83 mm. (3); leg ratio) 225-20) 
mean 2.37 (3); antennal ratio 1.40-1.50, 
mean 1.45 (3). 


Chironomids of West-Central Loutstana 


147 


Figures 102-105. Tanytarsus (Tanytarsus) allicis new species. 102. male genitalia; 103. 
variation of male genitalia; 104. pupal chaetotaxy; 105. posterolateral comb of Segment 
VIII. Figures 106-107. Tanytarsus (Tanytarsus) quadratus new species. 106. male gen- 
italia; 107. variation of ninth tergite. Figures 108-110. Tanytarsus (Tanytarsus) lim- 
neticus new species. 108. male genitalia; 109. superior appendage; 110. appendage 2a. 


Paratypes: Five males, 12-VIII-58; 
males, 20-VIII-58; 1 male, 28-VIII-58; 
males, 6-IX-58; 7 males, 12-I[X-58; 3 males, 
20-1X-58: 2 males, 15-X-58: U:SFH. In 
the collections of A.N.S.P., LN.H.S., C.N.C., 
and Cornell University. 


3 
Z, 


This species may be distinguished from 
the remainder of the Nearctic fauna by its 
distinctive male genitalia. It keys to neo- 


flavellus Malloch in Johannsen (Johannsen 
and Townes, 1952, pages 25, 26) but the 
punctate anal point of that species is totally 
different. 
SUMMARY 

1. Fifty-seven species of chironomid mid- 
ges are reported from lentic situations in 
Louisiana. 

2. Of the species described, one is a new 


148 


genus and fourteen are species new to 
science. 


3. The immature stages of twelve species 
are described. 


4. Five new taxonomic combinations are 
given. 


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ABSTRACT 
SUBLETTE, JAMES E. (Eastern New 
Mexico U., Portales). Chironomidae 


(Diptera) of Louisiana I. Systematics 
and immature stages of some lentic 
chironomids of West-Central Louisiana. 
Tulane Stud. Zool. 

Fifty-seven species of chironomid 
midges are reported, including the fol- 
lowing fourteen new species and one 
new genus: Ablabesmyia rhamphe, Cri- 
cotopus remus, lebetis, Chironomus 
(Chironomus) natchitocheae, (Dicro- 
tendipes) incurvus, (Cryptochironomus) 
ponderosus, Pedionomus beckae, Tany- 
tarsus dendyi, xanthus, recens, quad- 
ratus, buckleyi, allicis, limneticus. The 
immature stages of twelve species are 
described. Five new combinations are 
given: Ablabesmyia aequifasciata (Pen- 
taneura (Ablabesmyia) aequifasciata 
Dendy and Sublette) , Chironomus (Cryp- 
tochironomus) chaetoala (Tendipes 
(Cryptochironomus)  chaetoala Sub- 
lette), Chironomus (Cryptochironomus) 
directus (Tendipes (Cryptochironomus) 
directus Dendy and Sublette), Chirono- 
mus (Cryptochironomus) emorsus (Har- 
nischia (Harnischia) emorsa Townes), 
Chironomus (Cryptochironomus) galea- 
tor (Harnischia (Harnischia) galeator 
Townes). 


CHIRONOMIDAE (DIPTERA) OF LOUISIANA 
De hoe LIMNOLOGY OF THE UPPER PARE OF GANE RIVER LAIKE, 
NATCHITOCHES PARISH, LOUISIANA, WITH PARTICULAR 
REFERENCE TO THE EMERGENCE OF CHIRONOMIDAE! 


BURTON R. BUCKLEY 
Northwestern State College, 
Natchitoches, Louisiana 
and 
JAMES E. SUBLETTE 
Eastern New Mexico University, 
Portales, New Mexico 


Much of what is known about aquatic 
insect populations has been learned from 
bottom faunal studies using conventional 
bottom samplers. Quantitative and quali- 
tative data have been secured in this man- 
ner but investigators have experienced dif- 
ficulty in making positive identifications of 
the immature forms taken. Consequently, 
many limnologists have not attempted to 
make identifications below the family level, 
or at most to genus. This is particularly true 
for the Family Chironomidae which com- 
prises one of the major components of the 
benthos. The problem of generic and spe- 
cific determination is considerably reduced 
when tent and funnel traps are used since 
the adults which are taken in these traps can 
be more positively identified. 


Recent reviews of the literature concern- 
ing tent and funnel traps for capturing 
emerging adults of aquatic insects have been 
made by Wohlschlag (1950), Jonasson 
(1954), and Guyer and Hutson (1955). In 
addition, Sublette and Dendy (1959), and 
Buscemi (1961) have described modifica- 
tions of trapping devices. 


In view of the difficulty encountered in 
making positive identification of larval 
midges, and the scarcity of studies on lakes 
in southern United States, this research was 
done to secure quantitative and qualitative 
data on the benthic organisms, particularly 
the Chironomidae, through conventional col- 
lections made from larval populations and 
from samples of adults taken by tent and 
funnel traps. 


1 This paper represents the greater por- 
tion of a thesis submitted by the senior 
author to the faculty of the Graduate School 
of Northwestern State College, Natchi- 
toches, Louisiara, in partial fulfillment of 
the requirements for the degree of Master 
of Science. 


Appreciation is expressed to Mary Smith 
Sublette for the preparation of genitalia 
mounts from pinned specimens and to Judith 
Jones Buckley for assistance in making the 
graphs. 


HISTORY AND PHYSIOGRAPHY OF 
CANE RIVER LAKE 


At the time of settlement by European 
immigrants of the lower Red River Valley, 
the river and surrounding flood plain were 
in a unique stage of development. For rea- 
sons only partially understood, tremendous 
numbers of logs, much debris, and silt 
choked the stream. This resulted in the 
formation of what was locally referred to as 
the “Great Raft.” The damming action of 
the silt-laden log jams or “raft” caused the 
river to spill over into the back water 
swamps which are so characteristic of agrad- 
ing mature river systems. This produced a 
series of “braided” channels and connecting 
shallow lakes (cf. Hutchinson, 1957, page 
115). According to John Sibley, 1808, “An 
account of the Red River and country ad- 
jacent’”, American Register, Volume IV, fide 
Guardia (1927), there were four channels 
in the vicinity of the town of Natchitoches. 
These streams were, from east to west: 
Rigolets du Pon Dieu (now the main chan- 
nel of Red River). Atoho (now Little 
River), Cane River (now Cane River Lake, 
an impoundment) and False River (now 
Old River). At his writing, Cane River was 
the boat channel. False River was navigable 
but had low banks. The Rigolets du Bon 
Dieu was the smallest of the four, but sig- 
nificantly (as it was later to steal the main 
channel) it was the swiftest. At the time 
of Sibley’s report the Rigolets was just be- 
coming navigable (Guardia, 1927). Over a 
period of a few years (the time ts contro- 


versial, but probably about 1830-1840) the 
Rigolets du Bon Dieu channel degraded as 
a result of successive high water periods and 
captured the major part of the water flow 
below Grand Ecore (four miles east of 
Natchitoches). This left Cane River as a 
flood water divergence channel navigable by 
shallow draft boats only during time of high 
water. During summer months the stream 
dried up to small stagnant pools. In the 
spring of 1916 (fide, The Golden Jubilee 
Issue, Natchitoches Times, 1953) two earth- 
en dams were constructed on the channel, 
one at the upper end northeast of the city 
of Natchitoches and one located about two 
and one-half miles northwest of Derry, 
Louisiana, thus forming Cane River Lake. 
The origin of the lake is evidenced by its 
narrowness and great length, the meandering 
path it takes through the parish, its regular 
shoreline and bottom contours, and the many 
remnants of sandbars. In addition, precipi- 
tous slopes forming the shore of much of 
the lake and the existence of terraces some 
distance away indicate its stream origin. 


Throughout its length the lake is bordered 
by rich farm land. Consequently, the chief 
supply of water is run-off from culti- 
vated fields. The main axis of the lake lies 
northwest-southeast. 

The following morphometric features 
were listed by Geagan and Allen (1961): 
maximum length, 34.5 miles (55.5 km); 
mean width, 250 feet (76.2 meters); surface 
area, 1,044 acres (423.5 hectares); mean 
depth, 11.5 feet (3.4 meters); maximum 
depth near the spillway, 25.0 feet (7.6 
meters ). 


DESCRIPTION OF THE STUDY SITE 


Samples taken at irregular intervals and 
sites during a preliminary survey of Cane 
River Lake from February 5, 1957, to July 
12, 1957, showed the bottom contours and 
shoreline to be fairly uniform in the upper 
part of the lake. This survey was made while 
the junior author was conducting a research 
program subsidized by a National Institutes 
of Health research grant, RG-4594. Water 
samples taken at several sites in the upper 
end of the lake did not differ markedly 
from each other in physico-chemical char- 
acteristics. 

Beginning in June, 1957, a program of 
seasonal study of the lake was begun at a 


Tulane Studies mm Zoology 


Vol. 11 


site approximately one and one-half miles 
above the “new bridge” (bridge at Church 
Street crossing) located in downtown Natch- 
itoches, and approximately two and one- 
fourth miles below the upper dam. In this 
report only the results obtained during the 
seasonal study are presented. Since the water 
quality and bottom features were found to 
be very similar in the several areas in the 
upper part of the lake only one transect was 
used for the seasonal study. 


A three-quarter mile stretch of the lake 
on which the transect site was located lies 
in a northeast-southwesterly direction. The 
shoreline is quite regular, the eastern side 
being less precipitous than the western. The 
soil on the eastern side is Yahola Sandy 
Loam and on the western side is Yahola Clay. 
A few scattered willows, Salix nigra Marsh, 
are located along the shore above and below 
the sample site. The lake has a well-devel- 
oped zone of emergent vegetation composed 
principally of alligator weed, Alternanthera 
philoxeroides (Mart.) Standl., and cutgrass, 
Zizantopsis miliacea (Michx.) Doll and Asch. 


The bottom contours of the seasonal 
sampling site were very regular, with each 
lying closely parallel to the shoreline. The 
one meter contour was located about four 
meters out from the shore and the two meter 
contour was about six meters beyond the 
first. The width of the lake at this point 
was forty-one meters and the maximum 
depth 2.8 meters. 


MATERIALS AND METHODS 


Collection of adults. Samples of emerg- 
ing adults were obtained at approximately 
weekly intervals from June, 1957, to July, 
1958, using the traps and techniques de- 
scribed by Sublette and Dendy (1959). Two 
funnel traps were set on the bottom within 
each contour zone. Those set within the 
zero to one meter zone were located in a 
small cove formed by the outgrowth of alli- 
gator weeds. A conical tent trap was sus- 
pended over a bed of alligator weeds about 
fifty feet from the funnel traps. Except for 
three attempts to evaluate diel periodicity, 
the traps were lifted at the end of a twenty- 
four hour period. At the time of trap setting 
the temperature of each half meter depth of 
water was taken. 


The flasks containing the insects were re- 
moved to the laboratory where adult chi- 


No. 4 


ronomids which had been “wet down” and 
the exuviae were separated from the other 
forms and preserved in 70 per cent ethyl 
alcohol. A slide was later prepared of each 
adult and exuvia. Dry specimens were 
pinned on minuten nadeln. Insects other 
than the Chironomidae were preserved in 
70 per cent alcohol and stored. 

Collection of larvae. Bottom samples were 
taken with a six inch Ekman-Birge dredge at 
approximately monthly intervals. From three 
to five dredgings were taken at random 
within each meter contour. All sampling 
was done during daylight hours. Samples 
were washed in the field using a screen with 
25 meshes per inch, then preserved in 10 per 
cent formalin. In the laboratory the or- 
ganisms were removed from the debris, pre- 
served in 70 per cent alcohol, and subse- 
quently separated, counted, and measured 
volumetrically using Anderson and Hooper's 
(1956) modification of the technique given 
by Ball (1948). 

Physical and. chemical data. Monthly water 
samples were taken using an APHA sewage 
sampler. A Whitney electrical resistance 
thermometer was used to measure water 
temperature. The pH of the water was de- 
termined during a part of the study by a 
laboratory, line-operated Beckman pH meter. 
A Taylor block comparator was used in the 
field for the pH determination during the 
remaining time. A six inch Secchi’s disc 
was used to estimate light penetration. The 
turbidity of the water was assayed using a 
Bausch and Lomb Spectronic 20 colorimeter 
following the procedure outlined by the 
Hach Chemical Company, Ames, Iowa. 

The Alsterberg modification of the Wink- 
ler Method of dissolved oxygen determina- 
tion was used following the procedure out- 
lined in the 10th edition of Standard Meth- 
ods for the Examination of Water, Sewage, 
and Industrial Wastes (1955). Phenolph- 
thalein and methy! alkalinities were deter- 
mined following the procedure given by 
Welch (1948). Methyl organge alkalinity 
was obtained by using M-Alka Ver indi- 
cator supplied by the Hach Chemical Com- 
pany, Ames, Iowa. 


PHYSICAL FEATURES 


Bottom sediments, The 0-1.0 meter zone 
was characterized by sandy silt with much 
detritus. Plant fragments, especially bits of 
alligator weed and cutgrass, were concen- 


Limnology of Cane River Lake 15 


ws) 


trated in this zone. Near the end of Decem- 
ber, 1957, and throughout much of January, 
1958, leaves from sycamore, Platanus occi- 
dentalis L., located nearby became so abun- 
dant upon the bottom that the dredge fre- 
quently was prevented from closing properly 
and had to be reset. 


A very distinct shell zone existed in the 
1.1-2.0 meter zone. Sublette and Sublette 
(1958) and Moore (1950, 1952) did not 
find such a zone in other Louisiana im- 
poundments. Less detritus was present in the 
second zone and a considerable amount of 
sand and sandy silt was evident. 

The sediment of the 2.1-2.8 meters zone 
was a soft, brown, flocculent mud with some 
plant fragments. 


Thermal characteristics, In general, the 
lake showed a strong tendency toward ther- 
mal stratification from April to September. 
However, due to the shallow nature of the 
lake, the exposure to wind, and the direction 
of the lake’s main axis, thermally stratified 
waters with an epilimnion, thermocline, and 
hypolimnion which fit the criteria used by 
limnologists for other North American lakes 
were not observed. 

On the occasions when a thermocline oc- 
curred (July 2, July 16, July 30, September 
3, October 14, November 2, December 4, 
December 16, and April 7) it was present 
from the surface to a point near the middle 
or extended from the surface to the bottom 
(May 17). Isothermal water was frequently 
observed in January and February. Similar 
conditions were observed by Moore (1952) 
on Lake Chicot and Sublette and Sublette 
(1958) on Chaplain’s Lake. 

Although only incipient stratification was 
observed, oxygen depletion in bottom waters 
persisted throughout the summer months. 

Transparency. The maximum Secchi’s disc 
reading was 1.0 meter with a turbidity of 15 
ppm, recorded on July 13, and the minimum 
of 0.3 meter with a turbidity of 58 ppm 
was obtained December 4. These findings 
agree well with the result obtained by Gea- 
gan and Allen (1961) on lower sections of 
the lake where a three year average gave 


1.99 feet (0.66 meter ). 


The lake manifested a _ well-developed 
phytoplankton but allochthonous sediments 
were the apparent cause of the much reduced 
transparency. 


154 


CHEMICAL FEATURES 


Dissolved oxygen. Although a complete 
oxygen depletion was recorded only on three 
occasions (July 13, August 17, and May 17), 
in direct accordance with the thermal gradi- 
ents recorded on those dates, there was a 
general tendency for bottom waters to be 
depleted of oxygen during the warm seasons. 
The lowest concentration of oxygen at the 
surface was 6.2 ppm on August 17, and the 
highest value of 16.0 ppm was observed on 
February 22. Geagan and Allen (1961) did 
not find values as low at the transects where 
they sampled in the lower (southern) part 
of the lake. 


That a thorough mixing of the water oc- 
curred in January is reflected by the homo- 
geneity of water temperatures from surface 
to bottom and by only a slight difference in 
surface and bottom oxygen values for the 
month. 


Alkalinity. Normal carbonate was lacking 
in both surface and bottom waters the entire 
year. Bicarbonates were the sole source of 
alkalinity. During most of the study only 
slight variations were observed between 
amounts recorded from surface and bottom 
waters. With two exceptions (September 
30, January 4) larger amounts were recorded 
from bottom water. The highest value ob- 
tained from the surface was 172.0 ppm 
(August 17) with a low of 20.0 ppm (May 
17). On May 17, when Cane River Lake 
showed strong thermal gradients, a value of 
234.0 ppm was recorded for bottom water. 
This is in agreement with Moore’s (1950) 
observations on Lake Providence that higher 
alkalinity occurred in the lower stratum dur- 
ing periods of thermal stratification. 


Tulane Studies n Zoolog 


Vol. 11 


Hydrogen 1on concentration. Surface hy- 
drogen ion concentration varied from pH 
6.7 to 8.5 and that of the bottom from pH 
6.7 to 7.7. The bottom pH dropped slightly 
during May, June, July and August which 
seemed to be in accordance with other 
changes that occurred. 


RESULTS AND DISCUSSION 


Composition and seasonal changes in the 
bottom fauna. Conventionally, three major 
life-zones on the floor of thermally stratified 
lakes have been recognized: the littoral, sub- 
littoral, and profundal. As these terms have 
been employed, the littoral extends from the 
water's edge to the lakeward limits of rooted 
aquatic vegetation; the profundal occupies 
all of the lake floor bounded by the hypolim- 
nion; and the sublittoral lies in an inter- 
mediate position between the two and repre- 
sents a zone of transition. 

While these have proved useful in pre- 
vious studies, they do not adequately present 
the limits of the life-zones found in Cane 
River Lake. Instead, zonation of the lake 
floor which approximated meter contours 
proved more useful and accurate. The lit- 
toral zone as evidenced by bottom sedi- 
ments and associated animal assemblages, 
encompassed the area between the shoreline 
and the 1.0 meter contour, although the 
rooted aquatic plants extended only to about 
0.5 meter depth. The 1.1-2.0 meters zone 
included the sublittoral (—ecotone, cf. Sub- 
lette, 1957) or zone of transition from coarse 
littoral to finer profundal sediments. The 
profundal zone, with its finely divided, some- 
what flocculent yellowish-brown sediments, 
was located beyond the 2.0 meters contour 
and made up most of the lake floor. 


TABLE 1. 
Mean annual standing crop of bottom fauna, Cane River Lake, Louisiana. 

0-1.0 M 1.1-2.0 M 2.1-3.0 M 
ORGANISMS No./M? Vol./M? No./M? Vol./M? No./M? Vol./M? 
Oligochaeta 38295 3.9 8922 6.7 754 3.4 
Chironomidae 2201 1.9 841 1.95 184 2.8 
Chaoborus = — 25 0.09 945 eis 
Ceratopogonidae 6 0.01 8 0.01 4 0.01 
Ephemeroptera 5 0.04 3 0.34 ~ = 
Zygoptera 29 0.45 = - = = 
Anisoptera = — 10 0.33 - = 
Corixidae 14 0.01 - -- - = 
Gastropoda 538 - 705 — 22, _ 
Pelecypoda 54 — — _ - - 
Hirudinea 26 0.17 - = = = 
Turbellaria 14 0.01 = _ = = 
Mean Total 
Organisms 6192 6.49 5514 9.42 1909 7.96 


No. 4 


Qualitatively and quantitatively, a diversi- 
fied fauna composed principally of oli- 
gochaetes, fewer chironomids, and still small- 
er numbers of snails and small clams existed 
in the 0-1.0 meter zone (Table 1). A 
graphic representation of the total organisms 
(exclusive of Mollusca) for each zone is 
presented in Figure 1. 

The fauna of the 2.1-3.0 meters zone was 
made up of relatively large numbers of 
Chaoborus (Culicidae: Diptera), restricted 
almost entirely to this zone, and oligochaetes, 
with a very small number of chironomids. 
A few gastropods occasionally were taken 
there. These two zones lost their distinctive- 
ness in the transition area and merged to- 
gether. Oligochaetes, chironomids, and gas- 
tropods, in that order, were the major com- 
ponents. 


nosu® 


| 


0-10 METER 


11-20 METERS 


wosm 21-30 METERS 


acme sree es ea ee ae 
pet = = = == 
: = eoaplieeenteil 
f = ee ne if } t ; ~ | 
Figure 1. Seasonal abundance of all organ- 


isms averaged together. (mean number, sol- 
id line; mean volume, dashed line). 


Limnology of Cane River Lake 


1555) 


The relationship between numbers and 
volumes of the aquatic earthworms is shown 
in Table 1. An inverse relationship between 
numbers and volumes of chironomids was 
observed in the littoral and profundal zones 
with the largest number but smallest volume 
recorded in the Q-1.0 meter zone and the 
smallest number and largest volume in the 
2.1-3.0 meters zone. 

The predaceous ceratapogonids were dis- 
tributed almost uniformly. Immature forms 


nose? 0-10 METER 
12,000 


+ . | SI 


o4 >_> == SS 1? 


vou-cc/m? 
a 


nome 11-20 METERS 


Sa <== T v Tr a 


no su® 21-30 METERS 
7 


se Sa =< ae { 
= aS 


° = a == - - 
= Y + + 7 = Yr Y r 7 


~ «@ 
Te 

2-x14, —— 
:) = 

| 1 
—— 
en 

a 


2 z ‘ ‘ 
id a els t a —— eS 
Figure 2. Seasonal occurrence of Oligo- 


chaeta (mean number, solid line; mean vol- 
ume, dashed line). 


O-|0 mETen 


<a 


most 11-80 METERS 


4 


vevit 
17ewiit 
bee 


Figure 3. Seasonal abundance of larval 
Chironomidae, all species averaged together 
(mean number, solid line; mean volume, 
dashed line). 


MEAN WO/te 0-10 METERS 
Looe, 


—————————— ~ = a 
\1-2.0 METERS 
sss 


vo.- cosm® 


enn 


2.1-3.0 METERS 


0004 4 = | 
alle a t+—_ 

ol. . = : : — - = 
VOL-CC/mE 

* 

cr 

.) \ + 

hea 

ay | | 

as 

4 

4 

Oy Y co 1 r r Y Y + ¥ i 

ine fo “4 i ee eo 
Figure 4. Depth distribution and season- 


al abundance of Chaoborus punctipennis 
(Say), (mean number, solid line; mean vol- 
ume, dashed line). 


Tulane Studies n Zoology 


Vol. 11 


of the three gastropods, Campeloma lewisi 
(Walker), Physa integra (Haldeman), and 
Helisoma trivolvis (Say), were collected fre- 
quently in the deeper water as well as the 
shallow zone. The gastropods were identi- 
fied by a comparison with specimens identi- 
fied by Dr. R. Tucker Abbott, The Academy 
of Natural Sciences of Philadelphia. 

Seasonal variations in the benthic assem- 
blages are shown in Fig. 1. Figures 2, 3, and 
4 show seasonal trends for each of the three 
major components: Oligochaeta, Chirono- 
midae, and Chaoborus. 

Population increases occurred in late 
winter and again in the spring. Frequently 
we observed that the volumes increased fol- 
lowing a peak in the numbers. This was 
apparently due to the maturation of the 
organisms. 

The distribution of larval Chironomidae 
(Fig. 3) during the cold and warm seasons 
reflected the physical and chemical condi- 
tions of the lake during those periods. The 
virtual absence of chironomids in the deep 
water during June, July, August, and Sep- 
tember is a manifestation of the tendency 
toward thermal stratification of the lake and 
the associated poor supply of oxygen. At all 
other seasons the lack of prolonged thermal 
and accompanying chemical stratification al- 
lowed at least some species to distribute 
themselves within the profundal zone. 

Chaoborus punctipennis (Say) was the 
most characteristic animal of the profundal 
bottom during the daylight hours. It was 
not collected from the 0-1.0 meter zone at 
any time during the study and was recorded 
from the 1.1-2.0 meters only on seven oc- 
casions (Fig. 4) with the largest number 
taken August 17, 1957. On that date none 
were collected from the deeper water. The 
increase in numbers within the transition 
zone, presumably resulting from an upward 
migration to avoid stagnation effects, did 
not account for all of the profundal zone 
inhabiting Chaoborus that previously oc- 
curred there. After the August decline the 
number gradually increased until a peak was 
reached about the first of December. 


THE EMERGENCE OF CHIRONOMIDAE 


Forty-six species of Chironomidae were 
collected during the study. These have been 
treated taxonomically in Part I of this 
series. Table 2, which is a checklist of the 
chironomids found in Cane River Lake, sum- 


No. 4 Limnology of Cane River Lake 157 


TABLE 2. 
A checklist of the chironomids of Cane River Lake together with the depth 
distribution based on emergence of adults (x = presence) 


0-1.0 1.1-2.0 2.1-3.0 
Species Vegetation Meter Meters Meters 


TANYPODINAE 


Tanypus stellatus Coquillett 
Tanypus n. sp. 1 
Tanypus n. sp. 2 
Procladius n. sp. 1 Xe 
Procladius (Psilotanypus) 

bellus (Loew) oR x x 
Pentaneura (Pentaneura) 

planensis Johannsen x 
Pentaneura (Pentaneura) 

pilosella (Loew) x 
Ablabesmyia peleensis (Walley) x 
Ablabesmyia rhamphe Sublette 3 % 


ORTHOCLADIINAE 
Cricotopus bicinctus (Meigen) x x 
Cricotopus remus Sublette x x 
Nanocladius alternantherae 
Dendy and Sublette x axe x 


CHIRONOMINAE 
CHIRONOMINI 


Pseudochironomus aix Townes exe axe 
Chironomus (Chironomus ) 

natchitocheae Sublette xX x 
Chironomus (Chironomus) 

attenuatus Walker ee x x 
Chironomus (Chironomus) 

fulvipilus Rempel Xe 
Chironomus (Dicrotendipes) 

modestus Say x 
Chironomus (Dicrotendipes ) 

nervosus Staeger x xe De 
Chironomus (Xenochironomus ) 

xenolabis Kieffer os 
Chironomus (Endochironomus) 

nigricans Johannsen x Xs 
Chironomus (Cryptochironomus ) 

monochromus v.d. Wulp 3 x 
Chironomus (Cryptochironomus ) 

nigrovittatus Malloch x ae 
Chironomus (Cryptochironomus ) 

carinatus (Townes) x x 
Chironomus (Cryptochironomus ) 

directus (Dendy and Sublette) x x 
Chironomus (Cryptochironomus ) 

emorsus (Townes) 3 x x 
Chironomus (Cryptochironomus ) 

galeator (Townes) x x x 
Chironomus (Cryptochironomus ) 

edwardsi (Kruseman) x x x 
Chironomus (Cryptochironomus ) 

ponderosus Sublette ss x 
Chironomus (Cryptochironomus ) 

fulvus Johannsen x x x 
Glyptotendipes (Phytotendipes) 

lobiferus (Say) x x or 
Glyptotendipes (Phytotendipes) 

meridionalis Dendy and Sublette x x3 
Paralauterborniella elachista 

(Townes) x 
Lauterborniella varipennis 

(Coquillett) x x 


x 


mmm 


158 


TABLE 2. 


Tulane Studies in Zoology 


Vol. 11 


(Continued) 


A checklist of the chironomids of Cane River Lake together with tke depth 


distribution based on emergence of adults (« = Lresence) 


(1250 


0-1.0 2.1-3.0 
Species Vegetation Meter Meters Meters 
Stenochironomus macateei 
(Malloch) x 
Pedionomus beckae Sublette x x 
Polypedilum (Polypedilum) 
trigonum Townes me 
Polypedilum (Polypedilum) 
illinoense (Malloch) x x 
Polypedilum (Polypedilum ) 
digitifer Townes x xe x x 
TANYTARSINI 
Tanytarsus (Cladotanytarsus ) 
viridiventris Malloch x 
Tanytarsus (Tanytarsus) 
confusus Malloch x 
Tanytarsus (Tanytarsus) 
xanthus Sublette x x 
Tanytarsus (Tanytarsus) 
neoflavellus Malloch x x >¢ 
Tanytarsus (Tanytarsus ) 
dendyi Sublette x XK x 
Tanytarsus (Tanytarsus) 
buckleyi Sublette x 
Tanytarsus (Tanytarsus) 
quadratus Sublette x x x 
Tanytarsus (Tanytarsus ) 
recens Sublette x 


marizes depth distribution, while phenology 
by months is presented in Table 3. Emer- 
gence of adult chironomids in relationship 
to water temperature is given in Table 4. 


Seasonal variation. The very mild winters 
of this latitude were reflected in the repro- 
ductive behavior of the Chironomidae, with 
some adults emerging, and presumably mat- 
ing, in every month of the year. However, 
the emergence from the lake did not occur 
uniformly. Two distinct periods were dis- 
cernible in the total emergence for the year. 
One was from early April to late November 
(Fig. 5), coinciding with a marked rise in 
water temperature which occurred beginning 
about the first of March. The rate of emer- 
gence gradually increased throughout the 
early spring and summer, becoming highest 
in late September and early October. As 
water temperatures declined during the fall 
there was a parallel drop in numbers of 
midges emerging. The second period of 
emergence, or rather the lack of it, was from 
December to April during which time the 
low water temperatures were associated with 
a Sparse emergence that was irregular in dis- 
tribution. The two periods were particularly 


noticeable for those midges trapped over 
beds of aquatic vegetation (alligator weed). 


Adults taken in tent traps over beds of 
mat vegetation perhaps represent two larval 
populations; namely, those from the under- 
lying littoral floor and from the vegetation 
itself. These may be distinguished in the fol- 
lowing manner: larvae inhabiting the vege- 
tation beds would be represented by adults 
in tent traps above the vegetation beds only, 
whereas adults transforming from larvae on 
the littoral bottom would be taken not only 
in traps set over bare bottoms but also by 
traps set above vegetation. 

However, the funnel traps over bare bot- 
toms would take a preponderance of adults 
of the littoral bottom-inhabiting larvae, since 
those living on the bottom under the vege- 
tation would tend to migrate laterally (as 
pupae) to avoid the dark, shadow-producing 
mass of vegetation above (Scott and Opdyke, 
1941). If adults of a species occur in ap- 
proximately equal numbers in traps from 
both the littoral floor and from vegetation, 
the species is assumed to have no substratum 
preference. 

The emergence of chironomids from the 


No. 4 Limnology of Cane River Lake 159 


TABLE 3. 7. 
Phenology of emergence of adult chironomids by months 


Species ee MTV Ve VV Tiny Wee eT 


TANYPODINAE 


Tanypus stellatus Coquillett Xie eX 
Tanypus n. sp. 1 x 
Tanypus n. sp. 2 x Ks 
Procltadius (Procladius) n. sp. 1 x 
Procladius (Psilotanypus) 

bellus (Loew ) SUE XC me KOU MONO AG? EXe eX XA LK 
Pentaneura (Pentaneura) 

planensis Johannsen 2s 2 
Pentaneura (Pentaneura) 

pilosella (Loew) Ky te EXC KG) AK 
Alabesmyia peleensis (Walley) | Oe Ki Xs) OX 
Ablabesmyia rhamphe Sublette Be BSF BS G8 Oya 


ORTHOCLADIINAE 
Cricotopus bicinctus (Meigen) re Sk 
Cricotopus remus Sublette Xda Xe X 
Nanocladius alternantherae 
Dendy and Sublette x ee CU CME Cm es SS Oicee 1 5.ce, Trp Game 


CHIRONOMINAE 
CHIRONOMINI 


Pseudochironomus aix Townes KK KT GTX eK 
Chironomus (Chironomus) 

natchitocheae Sublette eK 
Chironomus (Chironomus ) 

attenuatus Walker KG FXG SCR pKa Ke XOX x 
Chironomus (Chironomus ) 

fulvipilus Rempel x 
Chironomus (Dicrotendipes ) 

modestus Say xe 
Chironomus (Dicrotendipes) 

nervosus Staeger eR De Se GS DC DG ge 
Chironomus (Xenochironomus ) 

venolabis Kieffer xx 
Chironomus (Endochironomus ) 

nigricans Jchannsen 3 Xi EKG Xe XO Xeon Ko RN, 
Chironomus (Cryptochironomus ) 

monochromus v.d. Wulp ie KEP KCK FRX 
Chironomus (Cryptochironomus ) 

nigrovittatus Malloch X0 eX eX ee 
Chironomus (Cryptochironomus ) 

carinatus (Townes) Kae Xe EKG x 
Chironomus (Cryptochironomus ) 

directus (Dendy and Sublette) Noe OXI Xa EK) XK 
Chironomus (Cryptochironomus ) 

emorsus (Townes) SX oe GG Xo Xe eX 
Chironomus (Cryptochironomus ) 

galeator (Townes) x Dee? Gale? Cee < 
Chironomus (Cryptochironomus ) 

edwardsi (Kruseman) eee AEDS Oe) een Gas Oe GE NO 
Chironomus (Cryptochironomus ) 

ponderosus Sublette x x x 
Chironomus (Cryptochironomus ) 

fulvus Johannsen 5 Gale i <a>. GI Ce Eb GaP Wap. >.< 
Glyptotendipes (Phytotendipes) 

lobiferus (Say) Sons SKN Sct x eK 
Glyptotendines (Phytotendipes) 

meridionalis Dendy and Sublette a a eee CPD Cre CES CE 'C ee aeo 
Paralauterborniella elachista 

(Townes) OK 
Lauterborniella 

varipennis (Coquillett) xa eX 
Stenochironomus 

macateei (Malloch) x 


160 Tulane Studies n Zoology Vola 
TABLE 3. (Continued) 
- Phenology of emergence of adult chironomids by months 
Species 1 It IV V. VIViIl Vx Sean 
Pedionomus beckae Sublette x X. x eee 
Polypedilum (Polypedilum) 
trigonum Townes x x 
Polypedilum (Polypedilum) 
illinoense (Malloch) Xo ek se > Gyee.< 


Polypedilum (Polypedilum) 
digitifer Townes 


TANYTARSINI 


Tanytarsus (Cladotanytarsus) 
viridiventris Malloch 

Tanytarsus (Tanytarsus ) 
confusus Malloch 

Tanytarsus (Tanytarsus) 


vanthus Sublette XE 
Tanytarsus (Tanytarsus) 
neoflavellus Malloch 2:8 


Tanytarsus (Tanytarsus) 
dendyi Sublette 

Tanytarsus (Tanytarsus ) 
buckleyi Sublette 

Tanytarsus (Tanytarsus) 
quadratus Sublette 

Tanytarsus (Tanytarsus) 
recens Sublette 


x x ».¢ 
x x x 
x x x x x x x 
x x x x xX x x x 
x x x 
x x x x x x 
x x x x 
x 


littoral benthos (0-1.0 meter) occurred 
through the warmer months of the year with 
only slight peaks of abundance observed 
(Fig. 5). The mean annual emergence of 
adults per square meter from that area was 
about half the mean number which emerged 
per square meter from the vegetation (Table 
1 and Figs) 


VEGETATION 


0-10 METER 


1957 1958 


Figure 5. Emergence of adult Chironomi- 
dae, all species averaged together. 


The number of adults emerging from the 
littoral zone was much greater than from 
the profundal (2.1-3.0 meters) and tran- 
sition zones (1.1-2.0 meters) combined. The 
emergence of profundal chironomids oc- 
curred from July through November, 1957, 
and in March and April of 1958. The num- 
ber of individuals arising from that area was 
small in comparison to the total emergence. 
In general, emergence decreased with an in- 
crease in depth. 

Table 5 shows the mean annual emer- 
gence per square meter for all insects taken 
in traps. The caddisflies and mayflies were 
recorded primarily from the vegetation in 
April and May, 1958. A few emerged dur- 
ing late August and September, 1957. Chao- 
borus and ceratopogonids made up a very 
small part of the total organisms trapped 
during the year. 

Species emerging from aquatic vegetation. 
Thirty-four species of Chironomidae were 
collected from tent traps set over vegetation. 
While most of these were also recorded in 
funnel traps set in shallow water, certain 
species showed a preference for vegetation 
habitats. Five species constituted about two- 
thirds of the total individuals from that 
location. In order of their abundance they 
were: Chironomus (Cryptochironomus) di- 
rectus Dendy and Sublette, Nanocladims 


No. 4 Limnology of Cane River Lake 161 


Species 8 10 12 14 16 18 20 22 24 26 28 30 32 34 


TANY PODINAE 


Tanypus stellatus Coquillet 


Tanypus n. sp. 1 


Tanypus n. sp. 2 


Procladius (Procladius) n. sp. 1 

Procladius (Psilotanypus) bellus (Loew) 
Pentaneura (Pentaneura) planensis Johannsen 
Pentaneura (Pentaneura) pilosella (Loew) 
Ablabesmyia peleensis (Walley) 


Ablabesmyia ramphe Sublette 


ORTHOC LADIINAE 


Cricotopus bicinctus (Meigen) 
Cricotopus remus Sublette 


Nanocladius alternantherae Dendy& Sublette 


CHIRONOMINAE 


Chironomini 


bot 
Ge Soa 
Ps acl 
SEES le 
eae 
Pseudochironomus aix Townes 
| 
pa) 
me 
[aaa 


Chironomus (Chironomus) natchitoceae Sublette 
Chironomus (Chironomus) attenuatus Walker 

Chironomus (Chironomus) fulvipilus (Rempel) 
Chironomus (Dicrotendipes) nervosus Staeger 
Chironomus (Dicrotendipes) modestus (Say) 

Chironomus (Xenochironomus) xenolabis Kieffer 
Chironomus (Endochironomus) nigricans Johannsen 
Chironomus (Cryptochironomus) monochromus v.d. Wulp 


Chironomus (Cryptochironomus) nigrovittatus Malloch 


Chironomus (Cryptochironomus) carinatus (Townes) 
Chironomus (Cryptochironomus) directus (Dendy& Sublette) 


Chironomus (Cryptochironomus) emorsus (Townes) 


Chironomus (Cryptochironomus) galeator (Townes) 


Chironomus (Cryptochironomus) edwardsi (Kruseman) 


Chironomus (Cryptochironomus) ponderosus Sublette 


Pe 
REAR: 
EN 
eee 
aE 
Chironomus (Cryptochironomus) fulvus Johannsen 
Glyptotendipes (Phytotendipes) lobiferus (Say) 
Glyptotenipes (Phytotendipes) meridionalis Dendy& Sublette 
Paralauterborniella elachista (Townes) Pec] 
Lauterborniella varipennis (Coquillet) [<2 einer | 
Stenochironomus macateei (Malloch) a 
Pedionomus beckae Sublette — 
Polupedilum (Polypedilum) trigonum Townes [a | 
Polypedilum (Polypedilum) illinoense (Malloch) ; 
Polypedilum (Polypedilum) digitifer Townes intersex Ef 2 ee 
fa 
eT ae 7 ee 
ES CE 
BS A 
eae Err ea 


normal 


Tanytarsini 
Tanytarsus (Cladotanytarsus) viridiventris Malloch 
Tanytarsus (Tanytarsus) confusus Malloch 
Tanytarsus (Tanytarsus) xanthus Sublette 
Tanytarsus (Tanytarsus) neoflavellus Malloch 
Tanytarsus (Tanytarsus) dendyi Sublette 
Tanytarsus (Tanytarsus) buckleyi Sublette 


Tanytarsus (Tanytarsus) quadratus Sublette 


Tanytarsus (Tanytarsus) recens Sublette | ca 


TABLE 4. EMERGENCE OF ADULT CHIRONOMIDS IN RELATIONSHIP TO. WATER 
TEMPERATURE ( DEGREES CENTIGRADE) 


162 Tulane Studies n Zoology Vol. 11 
TABLE 5. 
; Mean annual emergence of aquatic insects, No./M? 

Mean 

Organism Vegetation 0-1.0 M 1.1-2.0 M 2.1-3.0 M Total 
Chironomidae 217.9 107.2 21.5 9.6 486.1 
Trichoptera 19.4 3.0 0.56 - 22.96 
Ephemeroptera Lat} PEP 0.2 - 13.84 
Chaoborus 0.66 0.2 0.19 1.9 2.96 
Ceratopogonidae 0.64 0.59 — —- TAs: 


alternantherae Dendy and Sublette, Psedo- 
chironomus ax Townes, Chironomus (Dz1- 
crotendipes) nervosus (Staeger) and Crz- 
cotopus remus Sublette. With one excep- 
tion, Chironomus (Cryptochironomus) dt- 
rectus was taken entirely from the aquatic 
vegetation (Fig. 6) and comprised one-third 


VEGETATION 


0-10 METER 


1-20 METERS 


ou —_— — - 


21-30 METERS 


VEGETATION 


0-10 METER 


«0 — = ie r 

: ee ee eee a ze 
40 21-30 METERS 

VEGETATION 

20) 

100: h } 

ea | \ | \ 

“| mia 

die 2< Yee - 

a, 0-10 METER pS 

o ea 

9) 1-20 METERS : 

Al 

40, 21-30 METERS Z 

ol — —— = = ——— ae ~s 

J J A s [o} N 7} F 7 K 5) 
1957 1958 

Figure 6. Emergence of adult males of 


Chironomus (Cryptochironomus) edwardsi 
(above); Chironomus (Cryptochironomus) 
galeator (middle); and Chironomus (Cryp- 
tochironomus) directus (below). 


of the total number of chironomids emerg- 
ing from vegetation. Nanocladius alternan- 
therae and Pseudochironomus aix were im- 
portant littoral forms which occurred pri- 
marily in the tent traps, but were occasionally 
collected in the funnel traps set on the bot- 


tom in the shallow water (Figs. 7 and 8). 
The occurrence of large number of Pseudo- 
chironomus ax in the spring suggests that 
the species has one generation each year. 


VEGETATION 


morme 
#0 


> 


0-1 OMETER 
—— tee 


11-20 METERS 


21-30 METERS 


J J A 8 0 N ) J F ‘ A 

1957 1958 

Figure 7. Emergence of adults of Nano- 

cladius alternantherae (males, above abscis- 
sa; females, below). 


a 
Api VEGETATION 


#0) A 


0-10 METER 


1+2.0 METERS 


> 21-30 METERS 


$ 0 N J F Ae 

1957 1958 
Figure 8. Emergence of adults of Pseudo- 
chironomus aix (males, above abscissa; fe- 
males, below). 


J 


Cricotopus remus, like Chironomus (Crypto- 
chironomus) directus, was limited to the 
vegetation but was taken in a funnel trap 
on one occasion. Chironomus (Dircroten- 
dipes) nervosus was collected in about equal 
numbers from the tent and funnel traps. 

Of the remaining twenty-nine species, 
only one or two specimens of Chirono- 


No. 4 
mus (Xenochironomus) xenolabis (Kieffer), 
Stenochironomus macateet (Malloch) and 


Chironomus (Chironomus) fulvipilus (Rem- 
pel) were collected during the study. The 
other species occurred more frequently but 
were not numerous (Tables 2, 3, and 4) 


Species emerging from the littoral bei 
thos. The predominant species taken by the 
shallow water funnel traps was Chironomus 
(Cryptochtronomus) edwardst (Kruseman) 
(Fig. 6) followed closely by Polypedilum 
(Tripodura) digitifer Townes (Fig. 9). The 


ova VEGETATION 
“0 


0-1 O METER 


SSeS 


aa 11-20 METERS 


2.1-3.0 METERS 


VEGETATION 


0-10 METER 


11-20 METERS 


21-30 METERS 


eo 8 o 8 o 8 o 8 


Fee La ae a eee Ja] aL eT ey) pee 
1957 1958 


Figure 9. Emergence of adults of Polype- 
dilum (Polypedilum) digitifer. Top figure, 
males (above abscissa) and females (be- 
low) ; bottom figure, intersexes produced by 
mermethid nematode infections. 


relatively small number of Chironomus ed- 
wards: taken from vegetation, when com- 
pared to the large number which emerged 
from the littoral bottom, suggests that the 
species is restricted to the bottom and that 
the individuals which were taken in the 
tent traps emerged from the bottom and 
passed up through the vegetation. 


Other species which were frequently 
trapped were Glyptotendipes (Phytotendipes) 
meridionalis Dendy and Sublette, Chirono- 
mus (Cryptochtronomus) galeator (Townes), 
Nanocladius alternantherae Dendy and Sub- 
lette, and Tanytarsus neoflavellus Malloch. 


Thirty-six species were collected as they 
emerged from the littoral zone. Other than 
the species mentioned above, most occurred 
in small numbers (Tables 2, 3 and 4). 


Limnology of Cane River Lake 163 


Species emerging from the sublittoral and 
profundal. The predominant species trapped 
was Procladius bellus (Loew) (Fig. 10) 
and, in keeping with its predaceous feeding 
habits, it was found widely distributed. Dur- 


mo.snt VEGETATION 
20) 


0-0 METER 


11-20 METERS 


2|-3.0 METERS 


STL [nay Rak Tis uD Ge SL 


1957 1958 
Figure 10. Emergence of adults of Procla- 
dius bellus (males above abscissa; females, 
below). 


ey oleae 


ing the year most of the individuals were 
taken in water 1 to 2 meters in depth. A 
few were trapped in the littoral zone during 
June and July, 1957, and in the spring of 
1958. 

Only one other species, Tanypus stellatus 
Coquillett, occurred more than once in the 
profundal zone and it emerged in small 
numbers. Chironomus (Cryptochironomus) 
galeator (Townes) occurred once in a fun- 
nel trap set in the profundal zone (Fig. 6) 
In general, emergence from depths greater 
than 1.0 meter was small and sporadic and 
represented the lower fringes of typical lit- 
toral inhabiting populations. A total of nine- 
teen species was taken in small numbers 
below 1.0 meter depth (Table 2). 

Miller (1941) reported Cricotopus br- 
cinctus (Meigen) as a typical species found 
living below the thermocline in Costello 
Lake, Ontario. Only five individuals were 
trapped from: Cane River Lake and those 
were taken from the littoral. 

Sexual differences. Miller (1941)  ob- 
served slight differences in the time of 
emergence and distribution of the sexes. 
Males showed a tendency to reach their peak 
of emergence a short time before females, 
and females were found to be more abundant 
in deeper water. These differences, he postu- 
lated, were in some way correlated with 
temperature. 

Only slight sexual differences were noted 


164 


for three of the more common species found 
in Cane River Lake, Nanocladius alternan- 
therae Dendy and Sublette, Polypedilum 
digitifer Townes, and Procladius bellus 
(Loew) (Figs. 7, 9, and 10). The ratio of 
total males to females of Psewdochironomus 
aix Townes for the year was about 1:1 
(Fig. 8). However, a difference in the 
ratio of approximately 3:1 was recorded 
during the fall and a 1:3 ratio occurred in 
the spring. Glyptotendipes meridionalts 
Dendy and Sublette also showed an unusual 
ratio of males to females (Fig. 11). A ratio 
of three males to one female occurred 
throughout the year. 


Ya VEGETATION 


Tp lea] o-Pa 1 oaal Teel eT LS, LST Vic cL 
1957 1958 


Figure 11. Emergence of adults of Glyp- 
totendipes (Phytotendipes)  meridionalis 
(males, above abscissa; females, below). 


Intersexuality in a species of Chironomt- 
dae. Intersexes of Polypedilum digitifer 
Townes were trapped on several occasions, 
as has been reported for other chironomids 
(Wiilker, 1961, gives a review of mermi- 
thid parasitism in the Chironomidae). The 
individuals collected showed various de- 
grees of intersexuality and typically bore 
the characteristic female antennae, and a 
short, thickened abdomen with much re- 
duced male genitalia. Occasionally, a speci- 
men was taken with an antenna approach- 
ing the typical plumose condition of the 
male. 


Diel periodicity. The results of the three 
attempts to determine daily rhythms of emer- 
gence are presented in Figure 12. The great- 
est emergence of Chironomidae, July 9, 
1957, and June 10, 1958, took place between 
5 and 9 P.M. A decline in emergence oc- 
curred during the next four hours. On June 
19, 1958, the traps were lifted at two-hour 


Tulane Studies in Zoology 


Vol. 11 


9-VII-57 
NO./M@ 
120 


100 


80 


10-ViI-58 


19-ViI-58 


s ae 
re} — ~ ~v — =a 
» bt o = 


1PMy 
1AM 


Figure 12. Diel cycle of emergence, all spe- 
cies together, at three selected dates. 


intervals from 5 to 1 A.M. in order that 
more precise periods of emergence might 
be determined. No adults emerged between 
5 and 7 P.M. and equal numbers emerged 
between 7 and 9 P.M. and 9 and 11 P.M. 
Emergences declined between 11 P.M. and 
1 A.M. No emerging adults were trapped 
during hours of high light intensity. 


SUMMARY 


1. This study of the benthic faunal as- 
semblages and the emergence of Chironomi- 
dae from Cane River Lake was made from 
June, 1957, to July, 1958. It is the fourth 
bottom faunal study which has been made 
on lakes in Louisiana and the first study in 
Southern United States in which the emer- 
gence of Chironomidae was investigated. 


No. 4 


2. The purpose of the study was to se- 
cure quantitative and qualitative data on the 
benthic organisms, particularly the Chirono- 
midae, through collections made from larval 
populations and from adults taken by tent 
and funnel trappings. Collections were made 
using such devices to facilitate positive 
identifications. 


3. Cane River Lake is an impoundment 
of what was once a channel of Red River. 
The lake has a length of 34.5 miles, a mean 
width of 250 feet and an area of 1,044 
acres. Run-off is the principal supply of 
water. 


4. The lake has a well developed zone 
of emergent vegetation. It exhibited a strong 
tendency toward thermal stratification from 
April to September but did not stratify 
stably. Complete oxygen depletion in bot- 
tom water occurred on several occasions in 
accordance with the strong thermal gradi- 
ents and partial stratification. Bicarbonates 
were the sole source of alkalinity. 


5. Qualitatively and quantitatively, a di- 
versified fauna occurred in the littoral zone. 
Fewer groups occurred in the profundal. 
The principal components of the benthos 
were Oligochaeta, Chironomidae, and Chao- 
borus in that order. 


6. The distribution of Chironomidae dur- 
ing the cold and warm seasons reflected the 
physical and chemical conditions of the lake 
during those periods. Some chironomids 
were found distributed in the profundal zone 
during the cold season but were virtually 
absent during the warm season. 


7. Forty-six species of Chironomidae 
were collected during the study, including 
nine Tanypodinae, three Orthocladiinae, 
twenty-six Chironomini, and eight Tany- 
tarsini. 


8. Some adults emerged during every 
month of the year but two distinct peaks of 
emergence occurred. One peak was observed 
during the spring and early summer and the 
other in the fall. 


9. Emergence from the littoral zone was 
much greater than emergence from the sub- 
littoral and profundal zones combined. A 
greater number of adults emerged from the 
aquatic vegetation than from the littoral 
benthos. Emergence from the littoral bot- 
tom occurred from May to November with 
only slight peaks of numbers observed. 


Limnology of Cane River Lake 


165 


10. In general, the period of greatest 
emergence of chironomids coincided with a 
low population of larvae. With an increase 
in depth the number of larvae recorded and 
the number of adults emerging decreased. 


11. Of the thirty-four species of chirono- 
mids taken from the vegetation, five species, 
Chironomus (Cryptochtronomus) directus, 
Nanocladius alternantherae, Pseudochirono- 
mus ax, Chironomus (Dicrotendipes) ner- 
vosus, and Cricotopus remus constituted 
about two-thirds of the total number of in- 
dividuals. 


12. Thirty-six species of chironomids 
were trapped from the littoral benthos. The 
predominant species encountered were Chi- 
ronomus (Cryptochironomus) edwardst and 
Polypedilum dtgitifer. 


13. Three species were trapped from the 
profundal with Procladius bellus the pre- 
dominant species. 


14. Only slight differences in the emer- 
gence of males and females were observed 
for the three most common species, Nano- 
cladius alternantherae, Polypedilum digitifer, 
and Procladius bellus. Differences in the 
ratio of males to females of Psewdochtrono- 
mus aix and Glyptotentdipes meridionalts 
were recorded. 


15. Intersexes of Polypedilum digitifer 
showing various degrees of intersexuality 
were trapped on several occasions. 


16. Limited diel sampling showed that 
the greatest emergence of chironomids oc- 
curred between 5 and 11 P.M. 


REFERENCES CITED 


ANDERSON, RICHARD O. and FRANK F. Ho0o- 
PER 1956 Seasonal abundance and pro- 
duction of littoral bottom fauna in a 
southern Michigan lake. Trans. Amer. 

__Microscop. Soc. 75(8) : 259-270. 

BALL, Ropert C. 1948 Relationship between 
available fish food, feeding habits of fish 
and total fish production in a Michigan 
lake. Agric. Exp. Sta. Tech. Bull. 206: 
59. 


BusceMI, PHILIP 1961 Ecology of the bot- 
tom fauna of Parvin Lake, Colorado. 
Trans. Amer. Microscop. Soc. 80(3) : 266- 
307. 

GEAGAN, DONALD W. and THoMAsS D. ALLEN 
1961 An ecological survey of factors af- 
fecting fish production in Louisiana wa- 
ters. Publ. La. Wildlife Fish. Comm. (un- 
numbered), 100 pp. 


166 . 


GUARDIA, JOHN EDWARD 1927. Successive 
human adjustments to raft conditions in 
Lower Red River Valley. M.S. Thesis, 
University of Chicago. 

GUYER, GORDON and RAy HuTSON 1955 A 
comparison of sampling techniques uti- 
lized in an ecological study of aquatic in- 
sects. Jour. Heon. Entomol. 5(9): 321- 
426. 

HUTCHINSON, G. EVELYN 1957 A Treatise 
on Limnology. Vol. I. John Wiley and 
Sons, Inc., N. Y. 1015 pp. 


JONASSON, PETUR M. 1954 An improved 
funnel trap for capturing emerging aquat- 
ic insects, with some preliminary results. 
Oikos 5(2): 179-188. 

MILLER, RICHARD B. 1941 A contribution 
to the ecology of the Chironomidae of Cos- 
tello Lake, Algonquin Park, Ontario. 
Publ. Ontario Fish. Res. Lab. No. 60 
Univ. Toronto Stud., Biol. Ser. No. 49: 
1-63. 

Moore, WALTER G. 1950 Limnological stud- 
ies of Louisiana lakes. 1. Lake Provi- 
dence. Heology. 31: 86-99. 

__..... 1952 Limnological stud- 
ies of Louisiana lakes. 2. Lake Chicot. 
Proc. La. Acad. Sci. 15; 37-49: 

Scott, WILL and DAvip F. OppykKE 1941 
The emergence of insects from Winona 
Lake. Invest. Indiana Lakes and Streams 
2 (1) 7 3-14: 

SUBLETTE, JAMES E. 1957 The ecology of 
the macroscopic bottom fauna in Lake 
Texoma (Denison Reservoir) Oklahoma 
and Texas. Amer. Midl. Nat. 57(2): 371- 
402. 


for the construction of simplified tent and 
funnel traps. S.W. Nat. 3: 220-223. 

ee _........ and Mary SMITH SuvupB- 
LETTE 1958 The physico-chemical features 
and bottom fauna of Chaplain’s Lake, 


Tulane Studies n Zoology 


Vol. 11 


Natchitoches Parish, Louisiana. Proc. La. 
Acad. Sci. 20: 85-94. 

WELCH, P. S. 1948 Limnological Methods. 
The Blakiston Co., Philadelphia. 

WOHLSCHLAG, DONALD E. 1950 Vegetation 
and invertebrate life in a marl lake. Jn- 
vest. Indiana Lakes and Streams 3(9): 
BVA Bye 

WULKER, WOLFGANG 1961 Untersuchungen 
uber die Intersexualitat der Chironomiden 
(Dipt.) nach Paramermis-Infektion. Arch. 
Hydrobiol./Suppl. 25: 127-181. 


ABSTRACT 


BUCKLEY, BURTON R. (Northwest- 
ern State College, Natchitoches, Louisi- 
ana) and JAMES E. SUBLETTE (Eastern 
New Mexico U., Portales). II. The lim- 
nology of the upper part of Cane River 
Lake, Natchitoches Parish, Louisiana, 
with particular reference to the emer- 
gence of Chironomidae. 


Cane River Lake is a shallow im- 
poundment of what was once a channel 
of Red River. It has a well developed 
zone of emergent vegetation. While it 
does not develop stable thermal strati- 
fication, it does show occasional oxygen 
depletion from bottom water during 
summer months. The principal compo- 
nents of the benthos are Oligochaeta, 
Chironomidae, and Chaoborus, in that 
order. Forty-six species of chironomids 
were collected with emergence cf some 
species occurring during every month 
of the year. Greatest emergence peri- 
ods coincided with lowest larval popu- 
lation levels. Number of species by site 
of emergence was thirty-four from veg- 
etation, thirty-six from littoral benthos, 
and three from the profundal benthos. 
Intersexuality of Polypedilum digitifer 
Townes is reported. Greatest period of 
diel emergence was between five and 
eleven P.M. 


TULANE STUDIES IN ZOOLOGY 


Volume 11, Number 5 May 25, 1964 
CONTENTS 
Per NC ON EDC NUR Ge ee meme SOS, en A Dl oe) On 2 169 
LES INTRODUCTION AND EISTORICALIRBVIEW. = 169 
TdT TM ITSRTEI SO BAS ac I a ee 170 
IV. DESCRIPTION AND DISCUSSION OF SPECIES... eaten ae ie | ae 170 
iaratlypaspidocastridae Pochesl9 2s. iis 2 rien ee Mv 170 
eset EAT TES Gee LN lg ee ar a Se ee ee) 170 
iBevaayl 71 Re) Fove eS oven sa Veet ot ne YO ee ea ee nO ee Oe ee 174 
Bamiuly Bivesiculidae OE A re AO, RD Re hae a OE 178 
panatlvaarampinstOmatidae ss. ie 88 su eee ee Se a 178 
Bamallygeromocep halicld@c.i “Ok ee eee ee) ee A 179 
Fn ype lao © POG Ae | <coe sees OE ere oa Bee 179 
anallveaplosplanchnidae: 2 Asee ese = Sls ee a ee 179 
Family Megaperidae___________ See ee eee ne eerie ee ee ee Ses 183 
mt pt aL CC cl i lac a Oe SN 184 
Ppreamilyglis Ore AC ti la Ciz.2 ot tae eae eee Ts ee es eS ee 184 
arn O pec Oc] (aes co. ko ee ot ean Mame ETE ee See 194 
fica @ pistiiolebeticla Ce ts aaa eee See SS 198 
eral yg Gx Ol Tel Cm Sas ae a 198 
Rp AV AMOO COMiCae = val cin SSS eee ee 199 
aan otonGlii dae 6 eet Meena t ee Ne 200 
ecitanlliyg @ay PLO P OMIM IAC Mane eee we eee ag 209 
eerste Cte TO plays dacs santero De 209 
Heconlliype NCATE O CO] [3 aC meme wemmenrenen et ke ee a Ee 209 
ea ap Fed GOTT AS a wee ee 211 
Family, Accacoeltidae____- = = 5 A Cn een ea er ee Ii DR ERIS, ETSY 216 
acy ab linie iiellidac awaeee meee 8 216 
Ramally Sclerodistomitdac eee cs Win SO ea 216 
Emm lyaProsogconotrematidacmeem st. sos 5 Te a ee 216 
War GEOGRAPHICAT, DISTRIB UMTO Nise tothe kes ee 216 
SRE OST OPECTEICHI YS <0 eemmmemmemmnm nes.) ATU | Coe hme Mh A 218 
“HUTS is CRS TeR RDO ig wt 500) DS aM ST a SET aL 220 
Phe ISiOE NEGATIVE RISHES memeber mero 5 Si « be Ee A 224 
DPxePeDERATORE-CUlHl: tei a enee weer CE ee eee 224 
DOME NS in RAG eo wat ence ae Pee ee x Ee 6 oe Be te: a eon mal eee 226 
Dee INDEX OF TRENATODE\SPECIES mre a. 08 ea ee 227 


EDITORIAL COMMITTEE: 


MARY HANSON PRITCHARD, Research Associate, Department of Zoology and Physi- 
ology, University of Nebraska, Lincoln, Nebraska 


FRANKLIN SOGANDARES-BERNAL, Associate Professor of Zoology, Department of 
Zoology, Tulane University, New Orleans, Louisiana 


Horace W. STUNKARD, Research Associate, The American Museum of Natural His- 
tory, New York, New York 


DIGENETIC AND ASPIDOGASTRID TREMATODES FROM 
MARINE FISHES OF CURACAO AND JAMAICA+ 


F. M. NAHHAS 
and 
R. M. CABLE, 
Department of Biological Sciences, 
Purdue University, Lafayette, Indiana 


I. ACKNOWLEDGMENTS 


This study was supported by National Sci- 
ence Foundation Grant G-14691, and was fa- 
cilitated by the cooperative assistance of per- 
sonnel of the Caraibisch Marien-Biologisch 
Instituut, Curacao, and the Marine Biological 
Laboratory, Jamaica, W. I. Thanks are due 
especially to Dr. Ingvar Kristensen, Director 
of the laboratory in Curacao, and to Dr. Ivan 
Goodbody, Department of Zoology, Univer- 
sity College of the West Indies. We wish to 
express our appreciation also to Dr. J. E. 
Randall, University of Puerto Rico, for the 
identification of certain fishes and to Mrs. 
Mary Hanson Pritchard and Professor H. W. 
Manter of the University of Nebraska, Dr. 
Franklin Sogandares of Tulane University, 
and Dr. Allen McIntosh of the Beltsville 
Parasitology Laboratory for the loan of speci- 
mens and many helpful suggestions. 


II. INTRODUCTION AND HISTORICAL 
REVIEW 


This paper concerns adult digenetic trem- 
atodes parasitizing marine fishes of Curacao 
and Jamaica and is a continuation of a study 
begun by one of us (R.M.C.) in Puerto 
Rico in 1951 and reported in a series of 
publications (Cable, 1954a, 1954b, 1956a, 
1956b; LeZotte Jr., 1954; Siddiqi and Cable, 
1960). Results of that investigation indi- 
cated need for further study of larval and 
adult trematodes of shallow waters adjacent 
to Carribbean islands separated by deep 
water which could serve to isolate certain 
populations of intermediate and definitive 
host species. 

During 1961 the authors spent 7 months 
in the Caribbean region, 3 in Curacao and 
4 in Jamaica. The purpose of the trip was 
twofold: (1) the examination of as many 
shallow-water fishes as possible with em- 
phasis on duplicate or congeneric species 


from the two localities, and (2) the study 
of cercariae and as many of their life his- 
tories as time and facilities permitted. Four 
papers reporting life cycles and larval trem- 
atodes have appeared (Cable, 1962; Cable 
and Nahhas, 1962, 1963; Cable, 1963). 
The work on digenetic trematodes in the 
Western Atlantic-Gulf-Caribbean region was 
pioneered by Linton who reported about 75 
species from Woods Hole, Mass. (1898, 
1900, 1901, 1940) and several additional 
ones from Beaufort, North Carolina (1905), 
Bermuda (1907) and Tortugas, Florida 
(1910). Those areas were restudied by a 
number of investigators and especially by 
Manter, whose papers appearing from 1925 
to 1949 have made the region one of the 
better known so far as its trematode fauna 
is concerned. Other contributions have been 
made by Hanson (1950) who studied Bark- 
ker’s collection from Bermuda, and by Sparks 
(1957, 1958, 1960) who investigated the 
trematode fauna of the Bahama Islands and 
Grand Isle, Louisiana. Sogandares-Bernal 
(1959) compared the faunas of Bimini and 
the Panama Pacific and, in a series of papers 
in collaboration with others (Sogandares- 
Bernal and Hutton, 1959a, 1959b, 1959c; 
and Sogandares-Bernal and Sogandares, 
1961), studied the helminth parasites from 
Tampa and Boca Ciega Bays, Florida, and 
the Atlantic Coast of Panama. In Cuba, 
Pérez Vigueras (1940a, 1940b, 1955a, 1955b, 
1955c, 1956, 1958) reported a number of 
trematodes from several vertebrates includ- 
ing marine fishes. Major studies on larval 
forms have been made by Cable (1956b) 
who reported 51 marine cercariae from 
Puerto Rico, and by Holliman (1961) who 
described 28 species from the Apalachee Bay 
area of Florida. Less comprehensive studies 
will be cited in the discussion of species. 
During this study, we examined 1527 


7 Based on a thesis of the first author, submitted in partial fulfillment of the re 
quirements for the degree of Doctor of Philosophy, Purdue University, August, 1963. 


170 


fishes representing 185 species. Of these, 
698 individuals belonging to 124 species 
were collected in Curagao, and 829 repre- 
senting 127 species in Jamaica. Common to 
both localities were 68 species, of which 5 
harbored no endoparasitic trematodes. The 
same was true of 34 species in Curacao and 
21 in Jamaica. Usually, but not always, spe- 
cies negative for parasites were represented 
by individuals that were immature or few 
in number. 


Fishes from both localities yielded a total 
of 178 species of mature aspidogastrid and 
digenetic trematodes. Of these, 100 were 
found in Curacao and 140 in Jamaica, with 
62 common to both localities. Included were 
40 new species, of which 20 occurred in 
Curacao and 25 in Jamaica, with 5 common 
to both. One of the 5 was reported in con- 
nection with its life history (Cable and 
Nahhas, 1962); this paper includes descrip- 
tions of the remaining 39 new species in 
presenting and discussing data concerning 
the trematode fauna of Caribbean fishes. 
The system of LaRue (1957) is used for 
higher taxa, with a major revision concern- 
ing the position of the acanthocolpids in 
his system. 


Ill. MErTHODS 


Fishes were obtained from several sources 
including traps, nets, spear-fishing, use of 
rotenone, and commercial fishermen. When- 
ever possible, fish were kept alive in tanks 
and killed shortly before examination; others 
were examined as soon as possible, usually 
within two hours of death. When time per- 
mitted, the trematodes were studied alive, 
particularly to determine the extent of the 
bladder and other aspects of the excretory 
system. After removal from the host, they 
were washed in 0.7% saline and fixed with 
corrosive sublimate-acetic acid solution under 
light coverglass pressure. The worms were 
transferred to a dish, left in fixative for up 
to several hours, depending on the size of 
the worms, washed in several changes of 
water, dehydrated to 70%, left in 70% 
iodine-alcohol overnight to remove excess 
mercury, and stored in small vials of 80% 
alcohol. Most of the specimens were stained 
with Semichon’s carmine, cleared in methyl 
salicylate and mounted in damar. 


Most of the figures and their scales were 
drawn by microprojection but a few were 


Tulane Studies in Zoology 


Vok Ut 


free-hand. Differences in depth of focus 
may result in some dimensions not corre- 
sponding with text figures: the measure- 
ments given in text are exact. Measure- 
ments are in millimeters except where indi- 
cated as being in microns. Sucker ratios 
were calculated from the average of the 
length plus the width, and are expressed with 
the oral sucker taken as 1. 

Type specimens of all new species and 
representatives of a few previously described 
ones are deposited in the Helminthological 
Collection of the United States National Mu- 
seum. One asterisk indicates a new host rec- 
ord; two a new synonymy. The letters C and 
J indicate localities, Curagao and Jamaica, 
respectively. 


IV. DESCRIPTION AND DISCUSSION OF 
SPECIES 


FAMILY ASPIDOGASTRIDAE Poche, 
1925 

Cotylogaster basiri Siddiqi & Cable, 1960 

Host: *Calamus bajanado (J). 

Site; intestine. 


Lobatostoma ringens (Linton, 1905) 
Eckmann, 1932 

Synonyms: Aspidogaster ringens Linton, 
1905; Cotylogaster chaetodiptert MacCallum, 
1921. 

Host: *Micropogon furnieri (J). 

Site; intestine. 

The above species, like the freshwater 
aspidogastrids, probably are primarily para- 
sites of mollusks but are able to survive and 
even flourish in the intestine of vertebrates 
that feed on those hosts. Quantities of mol- 
lusk shells were found in the digestive tract 
of all of the fishes that harbored aspido- 
gastrids. 

Superorder Anepitheliocystidia La Rue, 1957 
Order Strigeatoidea La Rue, 1926 
Suborder Brachylaimata La Rue, 1957 
“Superfamily Bucephaloidea La Rue, 1926 
FAMILY BUCEPHALIDAE Poche, 1907 


Bucephalus varicus Manter, 1940 
Hosts: Caranx bartholomaei (J); C. ery- 
sos (C); C. hippos (C, J); C. latus (C, J); 
C. ruber (C, J); *Chloroscombrus chrysurus 
(J); *Seriola dumerili (J). 
Site: ceca and intestine. 


No. 4 


A great deal of variation was observed in 
the specimens from the several hosts and 
from individuals of the same host species. 
These variations may be explained by the 
condition of the worms at the time of fix- 
ation, their age, and possibly development 
in different host species. 


Bucephalus scorpaenae Manter, 1940 
Host: Scorpaena plumieri (C, J). 
Site: ceca and intestine. 


The Curacao material is in almost com- 
plete agreement with Manter’s description. 
The specimens from Jamaica have a some- 
what longer cirrus sac which sometimes ex- 
tends to the anterior margin of the anterior 
testis. 

The next species is figured and described 
but not named because only a single speci- 
men was found and it does not show such 
diagnostic features as the shape of the ten- 
tacles and extent of the excretory vesicle. 


Bucephalus sp. 
Figure 1 
Host: Centropomus undecimalis (J). 
Sze: Ceca. 


Deposited specimen: US.N.M. 60248. 


Description: Body elongate, tapering pos- 
teriorly, truncate anteriorly, 1.03 long, 0.20 
wide at level of ovary. Entrie cuticle spinose. 
Rhynchus sucker-like, 0.173 long, 0.20 wide, 
its tentacles retracted, probably 7 in number. 
Pharynx about equatorial in position, 0.053 
long, 0.060 wide; esophagus not evident; 
cecum subtriangular. Gonads in posterior 
half of body. Testes tandem, contiguous, to 
right of mid-line, 0.090-0.117 long, 0.083 
wide; cirrus sac 0.347 long, 0.058 wide, con- 
taining ovoid seminal vesicle and long pars 
prostatica surrounded by prostate cells. Geni- 
tal atrium wide, spherical. Vitellaria 12-14 
follicles on each side, anterior to pharynx. 
Ovary 0.060 long, 0.100 wide, anterior to 
testes, at about level of seminal vesicle; uter- 
ine seminal receptacle present; uterus ex- 
tending anterior to vitellaria, containing 
few normal eggs, 15-17 by 10-12 p. Excre- 
tory vesicle tubular, anterior extent not de- 
termined. 

The next species, although previously re- 
ported, is redescribed for reasons given 
below. 


Trematodes of Marine Fishes 


171 


Alcicornis carangis MacCallum, 1917 
Figure 2 
Host: Caranx ruber (C). 
Site: intestine. 
Deposited specimen: US.N.M. 60249. 


> 


Description based on 3 specimens. Body 
elongate, 1.58-2.1 long, 0.30-0.40 wide. En- 
tire cuticle spinose. Rhynchus wedge-shaped, 
0.566-0.633 long, 0.167-0.200 in greatest 
width; tentacles 7, varying in shape and 
length with degree of extension, 0.180-0.267 
long exclusive of filament, 0.030-0.045 wide 
at base; each with 2 lateral prongs and 
terminal filament, proximal prong more than 
twice as long as distal one; filament may be 
lost. Pharynx, seen in only one specimen, at 
level of anterior testis, 0.060 in diameter; 
esophagus not evident; cecum mostly an- 
terior to anterior testis. Testes tandem, con- 
tiguous, to right of midline, level variable, 
0.120-0.186 in diameter; cirrus sac 0.567- 
0.580 long, 0.100-0.133 wide, on left side 
of body, containing ovoid seminal vesicle, 
long pars prostatica and prostate cells. Ovary 
entire, anterior to testes, 0.133-0.146 long, 
0.080-0.100 wide; uterus voluminous, ex- 
tending from rhynchus to posterior end of 
body, sometimes overlapping posterior end 
of rhynchus. Genital atrium wide; genital 
pore at a distance from posterior end of 
body. Vitellaria in 2 lateral groups of 12-17 
follicles each, mostly in anterior half of 
body. Eggs 18-22 by 13-15 pw. Excretory 
vesicle not seen; excretory pore terminal. 

This species was known only from its in- 
correct description by MacCallum (1917) 
until it was found again by Pérez Vigueras 
(1955a). Our specimens agree with his re- 
description and show tentacles which have 
2 lateral prongs. MacCallum described the 
tentacles as being “branched like the antlers 
of a deer” and his figure shows a single 
prong on each tentacle. In our specimens, 
the number of prongs visible depends on 
the degree to which the tentacle is extended. 
Thus in the same individual, one tentacle 
may show both prongs and another only the 
distal one. 

Siddigi and Cable (1960) reported A. 
carangis from Caranx ruber in Puerto Rico. 
Comparing their specimens with ours indi- 
cates that their material represents a new 
species of Alctcornis, for which the name A. 
siddiqiu is proposed. The following descrip- 
tion was included by Siddiqi and Cable in 


172 


their original manuscript from which re- 
descriptions of known species were deleted 
before publication. 


Alcicornis Sid diqii n.sp. 
Figure 3 


Synonym: ** Alcicornis carangis of Sid- 
digi & Cable, 1960, nec MacCallum, 1917. 

Host: Caranx ruber (Puerto Rico). 

Site: stomach. 

Holotype: U.S.N.M. 39302 (deposited by 
Siddiqi & Cable). 


“Description based on 10 specimens: Body 
0.884-1.293 long, 0.165-0.198 wide; cylind- 
rical posteriorly, tapering anteriorly. Cuticle 
spinose. Rhynchus wedge-shaped, 0.118- 
0.147 by 0.067-0.099 exclusive of tentacles 
of which there are 7, each with 2 processes; 
ciltum absent. Pharynx spherical. 0.039- 
0.045 in diameter, somewhat posterior to 
midlevel and submedian; intestinal sac me- 
dian, small, equatorial. Testes 2, entire, 
0.082-0.097 by 0.075-0.090, tandem, con- 
tiguous, submedian to right, overlapped by 
pharynx and cirrus sac. Cirrus sac within 
posterior half of body, contaning sac-like 
seminal vesicle, long pars prostatica and 
prostate cells; genital pore ventral, a short 
distance from posterior end of body. Ovary 
entire, 0.070-0.096 by 0.066-0.075, sub- 
median, anterior to testes and intestine; 
seminal receptacle absent. Vitellaria scanty, 
in 2 short lateral bands of small follicles 
immediately anterior to ovarian level. Uterus 
voluminous, confined to posterior 2/3 of 
body, extending slightly anterior to vitellaria. 
Eggs numerous, 0.024-0.026 by 0.012-0.014. 
Excretory vesicle tubular, extending to level 
of vitellaria; excretory pore terminal, with- 
out evident sphincter.” 


Of the 4 species of Alcicornis that have 
been previously recognized, A. siddigii dif- 
fers from A. carangis in size of rhynchus, 
in the anterior extent of the uterus and evi- 
dently by lacking tentacular filaments; from 
A. baylisi Nagaty, 1937, in having a smaller 
rhyncus, shorter excretory vesicle and more 
anterior vitellaria; from A. longicornutus 
Manter, 1954, in having relatively much 
shorter tentacles with 2 prongs each; and 
from A. cirrudiscoides Velasquez, 1959, in 
having much smaller eggs. 


Tulane Studies tn Zoology 


Volant: 


Dollfustrema macracanthum Hanson, 1950 
Hosts: Gymnothorax moringa (C); *G. 
vicinus (C). 
Site; intestine. 


Deposited specimen: U.S.N.M. 60250. 


This species was described by Hanson as 
having one testis. Her material included 80 
specimens but her description is based on 
“eleven larger specimens, and particularly 
the holotype.” A reexamination of the type 
specimen reveals to us what looks like the 
faint outline of an anterior testis located at 
about the same level as the one seen distinct- 
ly in our material. 


Dollfustrema muraenae Sogandares- 


Bernal, 1959 


Hosts: *Gymnothorax funebris (J); *G. 
moringa (C, J). 

Site: intestine. 

Deposited specimen: U.S.N.M. 60251. 


The most distinctive features of this spe- 
cies are the position of the vitellaria and the 
3 rows of slender spines on the rhynchus. 
In our 37 specimens, topography of the 
gonads varies. The testes are usually sym- 
metrical but may be diagonal, often with 
the right testis anteriormost. The ovary is 
usually nearer the left testis and either an- 
terior to it or between the testes. The uterus 
occupies all available space between the 
rhynchus and the posterior end of the body. 
In contracted specimens, the rhynchus is 
drawn into the expanded anterior end of the 
body so that the vitellaria form a semicircle 
just posterior to the rhynchus and the cecum 
is drawn closer to that organ. Eggs in the 
Curacao specimens measure 24-26 by 15- 
18 », and in the Jamaican material 24-28 b 
16-21; Sogandares-Bernal (1959) gave a 
range of 24-33 by 19-27. 


Dollfustrema gymnothoracis n.sp. 
Figure 4 


Host: Gymnothorax vicinus (C). 

Site: upper intestine. 

Holotype: U.S.N.M. 60252. 

Description based on 18 specimens. Body 
elongated oval, more pointed posteriorly, 
1.03-1.66 long, 0.400-0.714 wide. Entire 
cuticle spinose. Rhynchus 0.098-0.120 long, 
0.120-0.165 wide, sucker-like when inverted; 
with 5 or 6 rows of spines, 5 » long, which, 
except for being closer together than those 
on adjacent cuticle, are not noticeably differ- 


No. 4 


ent from spines elsewhere. Pharynx at level 
of gonads, 0.060-0.075 in diameter; esopha- 
gus about same length as pharynx, cecum di- 
rected anteriorly. Testes symmetrical in 
younger specimens, diagonal in larger ones, 
0.135-0.200 long, 0.075-0.113 wide; cirrus 
sac 0.293-0.333 long, 0.093-0.146 wide, con- 
taining elongated seminal vesicle, long pars 
prostatica and prostate cells. Ovary inter- 
testicular, close to right testis, 0.100-0.113 
long, 0.068-0.105 wide; uterus extending an- 
teriorly in front of vitellaria, but never reach- 
ing rhynchus, and posteriorly almost to end 
of cirrus sac. Genital atrium spacious; geni- 
tal pore ventral, well removed from posterior 
end of body. Vitelline follicles form an in- 
verted U extending anteriorly from midlevel 
of testes in immature specimens, distinctly 
anterior to testes in older ones, but never 
intruding into anterior fifth of body length. 
Eggs thick-shelled, 33-42 by 22-27 pw. Ex- 
cretory vesicle tubular extending to about 
anterior end of cirrus sac. 

The arcuate arrangement of the vitelline 
follicles and the more posterior testes dis- 
tinguish this species from all others in the 
genus except D. echtnatum (Komiya and 
Tajimi, 1941). That species, known only 
from the metacercaria, differs from D. gym- 
nothoracis in having a conical rather than 
cup-shaped rhynchus. In other respects, D. 
gymnothoracis is similar to D. muranae and 
D. bipapillosum Manter and Pritchard, 1961. 
In those species, however, the vitellaria and 
uterus extend to the rhynchus which is larger 
than in D. gymnothoracis, and has rows of 
long spines well differentiated from the ad- 
jacent body spines. 

Nearly all of our specimens of D. gymno- 
thoracis were massively infected with coccus- 
like granules. 


Rhipidocotyle baculum (Linton, 1905 ) 
Eckmann, 1932 
Synonyms: Gasterostomum baculum Lin- 
ton, 1905; Gasterostomum sp. Linton, 1901; 
Nannoenterum baculum (Linton, 1905). 
Host: *Scomberomorus cavalla (C). 
Site: intestine and ceca. 


Our material is referred to Rhtpidocotyle 
baculum as described by Linton (1901, 
1905); his later paper (1940) evidently in- 
cludes more than one species as Nannoen- 
terum baculum. Rhipidocotyle baculum is 
very similar to R. adbaculum Manter, 1940, 
which Manter (1940c) distinguished on the 


Trematodes of Marine Fishes 


73 


basis of “size and shape of the cephalic disc 
which in R. adbaculum is larger and has a 
dorsal point. The body is larger and more 
elongate.” In body shape, our material is 
more like R. baculum. The shape of the 
cephalic disc is variable but in no case was 
a dorsal point evident nor did the cephalic 
disc extend laterally beyond the edges of the 
body. In some specimens, the uterus extends 
to the anterior level of the vitellaria. 


Siddigi and Cable (1960) reported Proso- 
rhynchus stunkardi from Scomberomorus sp. 
A reexamination of paratypes indicates that 
P. stunkardi is a synonym of either R. bacu- 
lum or R. adbaculum. Because the speci- 
mens were dead when removed from the 
host, the body and cephalic disc are not 
normal in shape. 


Bucephaloides longoviferus (Manter, 
1940) Hopkins, 1954 
Synonyms: **Bucephalopsis longoviferus 
Manter, 1940; Gasterostomum sp. Linton, 
1910 in part. 
Host: Sphyraéna barracuda (C, J). 
Site: ceca and intestine. 


Bucephaloides longicirrus (Nagaty, 
1937) Hopkins, 1954 
Synonyms: Bucephalopsis arcuatus of 
Manter, 1940, nec Linton, 1900; B. arcuvatus 
of Siddiqi & Cable, 1960, nec Linton, 1900. 
Host: Sphyraena barracuda (C, J). 
Site: ceca and intestine. 


Discussion: Manter 1963b draws attention 
to the fact that in the specimens from 
Sphyraena barracuda teferred by him 
(1940c) and by Siddigi and Cable (1960) 
to Bucephaloides arcuatus, the excretory 
vesicle does not extend beyond the pharynx 
whereas it terminates well anterior to that 
structure in the species as originally de- 
scribed by Linton (1900). We have con- 
firmed that difference between B. longicirrus 
and B. arcuwatus in both the Puerto Rican 
specimens and the present ones. 


Bucephalotdes arcuatus (Linton, 1900) 
Hopkins, 1954 

Synonyms: Gasterostomum arcuatum Lin- 
ton, 1900; Gasterostomum sp. Linton, 1900; 
Bucephalopsis arcuatus (Linton, 1900) Eck- 
man, 1932. 

Host: *Scomberomorus cavalla (C, J). 

Site: intestine and ceca. 


174 


Prosorhynchus atlanticum Manter, 1940 

Synonyms: Gasterostomum sp. Linton, 
1910 from M. bonaci; Gasterostomum sp. 
Linton, 1910 from M. venonosa, 

Hosts: Mycteroperca bonact (C, J); *M. 
falcata (C); M. venonosa (C). 

Site: ceca and intestine. 

Manter (1940c) distinguished this species 
from P. pacificum by the larger eggs with 
thicker and darker shells. Hanson (1950) 
was of the opinion that the 2 overlapped in 
that respect and synonymized them. P. pa- 
cificum has eggs 24-27 by 12-17 whereas 
they measure 29-36 by 18-24 in our speci- 
mens and 31-36 by 21-24 in those of Siddiqi 
and Cable (unpublished data). We regard 
that difference as being of specific magni- 
tude. 


Prosorhynchus aguayoi Pérez Vigueras, 
1955 

Host: Rypticus saponaceus (C, J). 

Site: intestine. 

Deposited specimen: US.N.M. 60253. 

Eggs from the Curacao material tend to 
be shorter and wider (38-43 by 24-27 yp) 
than those from Jamaica (40-46 by 21-26) 
but their measurements overlap. Pérez Vi- 
gueras gave an average egg size of 40 by 26. 


Prosorhynchus promicropsi Manter, 1940 
Host: Promicrops ttaiara (J). 
Site; intestine. 


Prosorhynchus ozaku Manter, 1934 
Host: *Mycteroperca bonaci (C). 
Site: intestine. 


Superfamily Fellodistomatoidea La Rue, 
1957 
FAMILY FELLODISTOMATIDAE 
Nicoll, 1913 


Antorchis urna (Linton, 1910) Linton, 
19 id 
Synonym: Mesorchis urna Linton, 1910. 
Hosts: Pomacanthus arcuatus (C, J); P. 
paru (C, J). 


Site: ceca and intestine. 


Antorchis holacanthi Siddiqi & Cable, 
1960 
Host: Holacanthus tricolor (J). 
Site: ceca. 


Mesolecitha linearis Linton, 1910 
Synonym:  **Proctoeces neomagnorus 


Siddiqi & Cable, 1960. 


Tulane Studies n Zoology 


Volui4 


Host: Acanthurus coeruleus (J). 
Site: intestine. 


Reexamination of the type of Proctoeces 
neomagnorus revealed the presence of cir- 
rus spines which in combination with a 
spherical rather than tubular seminal vesicle 
distinguish the genus Mesolecitha from Proc- 
toeces. P. neomagnorus thus agrees with 
Mesolecitha linearis as does our single speci- 
men and is reduced to synonymy with that 
species. 


Proctoeces maculatus (Looss, 1901) 
Odhner, 1911 

Synonyms: Distomum subtenue Linton, 
1907; Proctoeces subtenuis (Linton) Han- 
son, 1950; Proctoeces erythraeus Odhner, 
Oia 

Host: *Lactophrys tricornis (J). 

Site: intestine. 

Heretofore, this trematode has been re- 
ported from species of Calamus. Our single 
specimen from a trunkfish may represent an 
accidental infection. 


Proctoeces lintoni Siddiqi & Cable, 1960 
Hosts: *Calamus arctifrons (J); *C. ba- 
janado ch ) 3 


Site: intestine. 


Tergestia acuta Manter, 1947 
Hosts: Caranx bartholomaei (J); *C. 


crysos (J). 
Site: intestine. 


Tergestta laticollis (Rud., 1819) 
Stossich, 1899 
Synonyms: Distoma laticolle Rudolphi, 
1819; Pharyngora polonit Molin of Olsson, 
1869. 
Host: *Clepticus parrae (C, J). 
Site; intestine. 


Tergestia pectinata (Linton, 1905) 
Manter, 1940 

Synonyms: Distomum pectinatum Linton, 
1905; Theledra pectinata (Linton, 1905) 
Linton, 1910. 

Hosts: *Caranx bartholomaei (J); *C. 
hippos (J); C. latus (C, J); *Oligoplitis 
saurus (J); *Opisthonema oglinum (J); 
Selar crumenophthalmus (J). 

The next species closely resembles the 
genus Tergestia except that the oral sucker 
does not have lobes and the cervical region 
lacks the lateral folds characteristic of that 


No. 4 Trematodes of Marine Fishes 175 


2 


Figure 1. Bucephalus sp., dorsal view. Figure 2. Alcicornis carangis, dorsal view. 
Figure 3. Alcicornis siddiqii, holotype, ventral view (from Siddiqi and Cable, 1960). 
Figure 4. Dollfustrema gymnothoracis, holotype, ventral view. Figure 5. Gymnoterges- 
tia chaetodipteri, holotype, ventral view. Figure 6. Same, forebody enlarged. 


176 


genus. To receive the species, a new genus 
is proposed and characterized as follows; the 
generic name refers to the absence of orna- 
mentation of the forebody: 


Gymnotergestia n.g. 


Diagnosts: Fellodistomatidae. Distomes 
with elongated body; cuticle unarmed, an- 
nulated, especially in forebody. Oral sucker 
cup-shaped, without lobes. Prepharynx ab- 
sent; pharynx elongated, conical; ceca long. 
Testes 2, diagonal or tandem; cirrus sac 
well-developed, at level of acetabulum; ex- 
ternal seminal vesicle absent. Ovary pretes- 
ticular; true seminal receptacle absent; uterus 
voluminous, extending posterior to gonads; 
metraterm muscular. Vitelline follicles nu- 
merous, in lateral fields posterior to acetabu- 
um. Genital pore median, anterior to ace- 
tabulum. Excretory vesicle Y-shaped. Para- 
sites in intestine of marine fishes. 

Type and only species: 


Gymnotergestia chaetodipteri 
n.g., N.sp. 
Figures 5 and 6 
Host: Chaetodipterus faber (J). 
Site: lower intestine. 


Holotype: U.S.N.M. 60254. 


> 


Description based on 3 specimens. Body 
slender, 1.66-4.28 long, 0.380-0.567 wide at 
level of acetabulum. Cuticular rings of fore- 
body giving lateral margins a serrated ap- 
pearance. Oral sucker 0.140-0.187 long, 
0.200-0.280 wide; ventral sucker 0.320-0.440 
long, 0.247-0.340 wide, with longitudinal 
aperture; sucker ratio 1:1.67. Hindbody 
2-3 times length of forebody. Pharynx 
0.300-0.413 long, 0.123-0.173 in greatest 
width; esophagus short; ceca extending to 
posterior end of body. Testes 2, diagonal or 
tandem, 0.173-0.286 long, 0.140-0.173 wide, 
separated by uterine coils; posttesticular 
space one-half as long to equal length of 
forebody. Cirrus sac large, to left of ventral 
sucker, containing saccular seminal vesicle, 
large pars prostatica and long folded cirrus. 
Ovary entire, pretesticular, 0.126-0.186 long, 
0.106-0.133 wide, separated from anterior 
testis by uterine coils. Uterine seminal re- 
ceptacle present; uterus voluminous, occupy- 
ing most of hindbody; metraterm half as 
long as cirrus sac, muscular. Genital atrium 
small; genital pore median, a short distance 
anterior to ventral sucker. Eggs thick-shelled, 


Tulane Studies in Zoology 


Vol. 11 


30-36 by 20-25 p. Vitelline follicles in Jat- 
eral fields between acetabulum and posterior 
testis. Excretory vesicle bifurcating at ova- 
rian level, its arms extending to sides of 
pharynx to receive main collecting tubules. 


The larva of this species probably is simi- 
lar to Cercaria cartbbea XL which Cable 
(1956b) described from Puerto Rico. It 
has the elongated pharynx characteristic of 
Gymnotergestia chaetodipteri and lacks the 
oral lobes that are known to be developed 
in at least some cercariae of the genus Ter- 
gestta. 


Infundibulostomum anisotremi n.sp. 
Figure 7 
Host: Antsotremus virginicus (J). 
Site: intestine. 


Holotype: U.S.N.M. 60255. 


Description based on a single specimen. 
Body ovoid, more broadly rounded anteriorly 
than posteriorly, 0.767 long, 0.374 wide. 
Cuticle spinose, spines extending to pos- 
terior level of testis. Eye-spot pigment ab- 
sent. Oral sucker 0.078 long, 0.083 wide; 
ventral sucker about equatorial, 0.063 long, 
0.069 wide; sucker ratio 1:0.71. Prepharynx 
0.013 long; pharynx 0.040 long, 0.033 wide; 
esophagus slightly shorter than pharynx; 
ceca short, extending to near posterior level 
of vitellaria. Testis somewhat irregular, sub- 
median, 0.105 long, 0.159 wide; cirrus sac 
arcuate, to left of ventral sucker, containing 
large internal seminal vesicle and prostatic 
complex; external seminal vesicle immedi- 
ately posterior to ventral sucker. Ovary en- 
tire, to left of midline, at level of testis, 
0.111 by 0.075; seminal receptacle median 
to ovary; uterus filling posttesticular space, 
extending anteriorly to form convolutions 
on right side of body between testis and 
level of intestinal bifurcation, metraterm 
simple. Genital pore median, immediately 
anterior to acetabulum. Eggs numerous, 18- 
21 by 10-13 p». Vitellaria in 2 lateral clusters 
of 6 or 7 follicles each, at level of acetabu- 
lum. Excretory vesicle not observed. 

Infundtbulostomum anisotremi, the sec- 
ond species in its genus, differs from I. 
spinatum Siddiqi & Cable, 1960, in the 
shape of the body, extent of spination, shape 
and size of the oral sucker, and the sucker 
ratio. 


The systematic position of species assigned 
to the genus Bacciger is confused by the ab- 


No. 4 


sence of a distinct cirrus in B. harengulae 
which led Yamaguti (1958) to assign that 
genus to the family Cryptogonimidae. He 
stated that a cirrus sac is absent because 
Stossich did not show that structure in B. 
bacciger even though both Nicoll (1914) 
and Palombi (1934) clearly described a cir- 
rus sac in species in Bacciger. We have 
found a well-developed cirrus sac in a spe- 
cies from Opisthonema oglinum but it is 
absent in another from Sardinella macroph- 
thalmus. Evidently the two cannot be as- 
signed to the same genus and therefore we 
restrict to the genus Bacciger those species 
having a cirrus sac and erect a new genus as 
follows for those in which it is absent: 


Pseudobacciger n.g. 


Diagnosis: Fellodistomatidae. Body short; 
cuticle spinose; eye-spot pigment absent; 
distomate, ventral sucker in anterior half of 
body. Pharynx and esophagus present; ceca 
not extending to posterior end of body. 
Testes 2, symmetrical, postacetabular; cirrus 
sac abseat. Ovary intertesticular; seminal re- 
ceptacle present; uterus chiefly posttesticular. 
Vitellaria compact, lateral masses in acetabu- 
lar region. Eggs small. Excretory vesicle V- 
shaped. Parasites in intestine and ceca of 
marine fishes. Type species: Psewdobacciger 
harengulae (Yamaguti, 1938)  n.comb.; 
(Synonym: Bacciger harengulae). other spe- 
cies: Pseudobacciger mantert n.sp. 


Pseudobacciger manteri n.g., n.sp. 
Figure 8 

Synonym: ** Bacciger harengulae of Man- 
ter, 1947, nec Yamaguti, 1938. 

Host: Sardinella macrophthalmus (J). 

Site: ceca. 

Holotype: U.S.N.M. 60256. 

Description based on 20 specimens. Bod 
oval to pyriform, 0.373-0.747 long, 0.266- 
0.420 wide. Entire cuticle spinose. Oral 
sucker terminal, 0.045-0.068 in diameter; 
ventral sucker preequatorial, 0.057-0.090 in 
diameter; sucker ratio 1:1-1.3. Prepharynx 
not evident; pharynx 0.033-0.045 in diam- 
eter; esophagus 1-2 times length of pharynx; 
ceca extending a short distance posterior to 
testes. Testes 2, equatorial to somewhat pre- 
equatorial, symmetrical, 0.060-0.090 long, 
0.065-0.105 wide; cirrus sac absent; seminal 
vesicle indistinctly bipartite, extending 
slightly posterior to ventral sucker; pars 


Trematodes of Marine Fishes 


177 


prostatica indistinct. Ovary irregular to 
lobed, 0.060-0.090 in diameter, intertesticular, 
median, at about posterior level of testes; 
seminal receptacle small, overlapping ovary 
or not; uterus chiefly posttesticular; metra- 
term simple. Genital atrium small; genital 
pore median, near intestinal bifurcation. Vi- 
telline follicles in 2 largely extracecal groups 
of 7-10 each at about level of ventral sucker. 
Eggs 21-24 by 15-20 pw. Excretory vesicle V- 
shaped, with arms passing ventral to ceca, 
to terminate at esophageal level; excretory 
pore terminal. 


Manter (1947) reported this species as 
Bacciger harengulae Yamaguti, 1938, from 
3 specimens of which one was partly crushed. 
He noted its resemblance to Yamaguti’s 
material but indicated that it differed in 
having more rounded eggs and somewhat 
longer ceca; he was unable to see the ex- 
cretory vesicle. Measurements given by Ya- 
maguti overlap those of the present species 
which has consistently longer ceca and arms 
of the bladder extending farther anteriorly 
than in the Japanese species. 


The next species is named with hesitation 
because it may be the one that Price (1934) 
described as Steringotrema ovata from a 
single specimen. Both occur in the same 
host species and agree in all respects except 
the topography of the gonads. Price’s speci- 
men was examined but it was faded and the 
gonads were so indistinct that their identity 
and arrangement could not be determined 
with certainty. 


Bacciger opisthonemae n.sp. 
Figure 9 
Host: Opisthonema oglinum (J). 


Site: ceca and intestine. 
Holotype: U.S.N.M. 60257. 


Description based on 6 specimens. Body 
oval to somewhat pyriform, 0.386-0.493 
long, 0.240-0.300 wide. Entire cuticle with 
fine spines; eye-spot pigment absent. Suck- 
ers subequal, 0.042-0.067 in diameter, oral 
sucker slightly subterminal, ventral sucker 
about equatorial. Prepharynx not evident; 
pharynx 0.033-0.037 in diameter; esophagus 
very short; ceca extend to or slightly beyond 
midlevel of hindbody. Testes 0.040-0.060 
in diameter, smooth or somewhat irregular, 
symmetrical to slightly diagonal, near pos- 
terior margin of ventral sucker; cirrus sac 
pyriform, 0.090-0.112 long by 0.051-0.060 


178 


wide, median, anterior to or slightly over- 
lapping ventral sucker, containing saccate 
seminal vesicle, pars prostatica, conspicuous 
prostate cells and cirrus. Ovary 0.040-0.068 
in diameter, median to submedian, inter- 
testicular or slightly posterior to level of 
testes; uterus mostly postovarian; metraterm 
simple. Vitelline follicles in 2 groups of 
5-8 each, mostly extra-cecal, near level of 
acetabulum. Genital pore median or slightly 
submedian, near posterior edge of pharynx. 
Excretory vesicle V- or Y-shaped with short 
stem, arms extending to pharyngeal level. 
Removing from the genus Bacciger spe- 
cies lacking a cirrus sac leaves only B. bac- 
ciger (Rud., 1819) and B. nicolli Palombi, 
1934. Both differ from the present species 
in having shorter ceca and more extensive 
vitellaria with follicles extending well into 


the forebody. 


Suborder Azygiata La Rue, 1957 
Superfamily Azygioidea Skrjabin & 
Guschanskaja, 1956 
FAMILY BIVESICULIDAE Yamaguti, 
1939 


Bivestcula caribbensis Cable & Nahhas, 
1962 
Host: Myripristis jacobus (C, J). 
Site: pyloric ceca. 


Order Echinostomida La Rue, 1957 
Suborder Paramphistomata Szidat, 1936 
Superfamily Paramphistomatoidea Stiles & 
Goldberger, 1910 
FAMILY PARAMPHISTOMATIDAE 
Fischoeder, 1901 


The following species is identified as the 
one Pérez Vigueras (1940a) described as 
Macrorchitrema havanensis. In erecting the 
genus Macrorchitrema, he discussed various 
genera of amphistomes but did not men- 
tion Cleptodiscus with which Macrorchi- 
trema is Clearly synonymous. Our specimens 
and his were from the same host species 
and because his description is not complete 
or generally available, the species is re- 
described as follows: 


Cleptodiscus havanensis (Vigueras, 
1940) n.comb. 
Synonym: **Macrorchitrema havanensis 
Vigueras, 1940. 
Host: Holacanthus tricolor (J). 
Site: intestine. 


Deposited specimen: U.S.N.M. 60258. 


Tulane Studies in Zoology 


Vol. 11 


Description based on 3 specimens. Body 
broadly rounded posteriorly, tapering an- 
teriorly, 4.57-5.79 long, 1.35-1.64 in maxi- 
mum width. Cuticle thin, with a few minute 
spines near anterior end of body; eye-spot 
pigment present. Pharynx 0.233-0.266 long, 
0.166-0.200 wide, with 2 retrodorsal diver- 
ticula. Ventral sucker at posterior end of 
body, 1.20-1.35 long, 0.965-1.06 wide, with 
longitudinal aperture. Esophagus 0.868- 
1.013 long, 0.040-0.060 in maximum width 
at muscular bulb near intestinal bifurcation, 
surrounded by gland cells along entire 
length; ceca not extending posterior to ovary. 
Testes 2, irregular to lobed, 0.466-0.714 long, 
0.366-0.667 wide, diagonal, close together 
or separated by coils of uterus; anterior 
testis and cirrus sac near intestinal bifurca- 
tion; cirrus sac pyriform to spherical, 0.185- 
0.225 long, 0.135-0.180 wide, containing 
sinuous internal seminal vesicle, pars pro- 
statica, prostate cells, and relatively short 
cirrus; sac followed by very long, convoluted 
seminal vesicle. Ovary smooth, 0.220-0.266 
in diameter, near anterior edge of acetabu- 
lum, to right or left (in one specimen) of 
midline; seminal receptacle not evident; 
Mehlis’ gland posterior to ovary; uterus 
mostly dorsal, between acetabulum and an- 
terior testis. Genital pore midventral, at or 
slightly anterior to intestinal bifurcation. 
Eggs thick-shelled, more pointed at one end, 
60-86 by 40-51 p. Vitelline follicles in lat- 
eral fields, extending from anterior edge of 
acetabulum to about midlevel of anterior 
testis. Excretory system not observed. Lym- 
phatic channels two, extracecal, extending 
from near posterior end of body to sides of 
oral sucker. 

This species is distinguished from C. re- 
ticulatus Linton, 1910, by the much larger 
ventral sucker, its longitudinal aperture and 
the more anterior position of the testes and 
vitellaria. Hanson (1955) described a bi- 
partite cirrus sac in C. bulbosus, a species 
which is more like C. reticulatus than the 
present one. In our specimens, the external 
seminal vesicle may be compactly coiled or 
not and, because of the adjacent membranes 
of the lymph channels and parenchymal 
vesicles, appears to be embedded in a rela- 
tively much denser tissue that could be mis- 
taken for the posterior division of a bi- 
partite cirrus sac. 


No. 4 


Superfamily Notocotyloidea La Rue, 1957 


FAMILY PRONOCEPHALIDAE Looss, 
1902 
Glyphicephalus candidulus (Linton, 1910) 
Siddiqi and Cable, 1960 

Synonyms: Himasomum candidulum Lin- 
ton, 1910; Barisomum candidulum (\Lin- 

ton) Price, 1931; Plewrogonius candidulus 
(Linton) Manter, 1947. 

Hosts: Angelichthys ciliaris (J); Poma- 
canthus arcuatus (J). 

Site: intestine. 


Suborder Echinostomata Szidat, 1939 
Superfamily Haploporoidea Mehra, 1961 


Mehra (1961) proposed the superfamily 
Haploporoidea to include the Haploporidae, 
Waretrematidae, Haplosplanchnidae and 
Megaperidae. Although the first 2 families 
are questionably distinct, Cable (1962) gave 
evidence that they are closely related to the 
Haplosplanchnidae. As to the Megaperidae, 
its placement with those families in a com- 
mon superfamily is less certain because the 
development of the excretory system was 
not observed when Cable (1954b) reported 
the only cercaria known to be a megaperid 
larva. Thus it is not certain that the primary 
excretory pores are in the tail and that the 
bladder lacks an epithelium as would be ex- 
pected if the megaperids were closely re- 
lated to the other families that Mehra has 
placed in the Haploporoidea. Although 
Manter erected the family Megaperidae, he 
later (1963a) reduced it to the rank of a 
subfamily in the Lepocreadiidae; Cable 
(1954b) had previously suggested a close 
affinity with that family. However, it seems 
likely that further studies may support 
Mehra and for that reason, his view is ten- 
tatively accepted here by including the 
Megaperidae in the Haploporoidea. At this 
point it may be noted that the genera 
Enenterum, Cadenatella, and Jeancadenatia 
which generally are placed in the Lepocre- 
adiidae, have many features in common with 
those of families included in the Haplo- 
poroidea and ultimately may be transferred 
to that superfamily. In the absence of life 
history studies, however, those genera are 
tentatively retained in the Lepocreadiidae. 


FAMILY HAPLOPORIDAE Nicoll, 1914 


Hapladena varia Linton, 1910 
Hosts: Acanthurus hepatus (C, J); A. 
coeruleus (J). 


Trematodes of Marine Fishes 


179 


Site: intestine. 

The specimens from Curacao were mostly 
immature but a few were sufficiently de- 
veloped to make identification possible. 
Cable (1962) described the first haploporid 
cercaria and suggested on morphological and 
ecological evidence that it may be the cer- 
caria of H. varta. 


Hapladena ovalis (Linton, 1910) 
Manter, 1947 
Synonym: Deradena ovalis Linton, 1910. 
Hosts: *Sparisoma brachiale (J); S. fla- 
vescens (J); *Pseudoscarus guacamata (J). 
S:te: intestine. 


Allomegasolena spinosa Siddiqi & Cable, 
1960 

Hosts: Chaetodipterus faber (J); *Luti- 
anus apodus (J). 

Swe. ceca. 

Five specimens agree with the description 
given by Siddiqi and Cable except that suck- 
er ratio may reach 1:1.68 and the eggs at- 
tain a length of 75 up. 


Megasolena archosargi Sogandares-Bernal - 
& Hutton, 1959 

Host: *Archosargus unimaculatus (J). 

Site: intestine. 

Eight specimens from 5 fish are in agree- 
ment with the description given by Sogan- 
dares-Bernal and Hutton (1959a) except in 
the size of the eggs which in our material 
measure 60-75 by 39-47 p» as compared with 
73.5-88.2 by 42-48.3. 


FAMILY HAPLOSPLANCHNIDAE 
Roche, 1925 
Haplosplanchnus mugilis n.sp. 
Figure 10 

Host: Mugil curema (C). 

Site: intestine. 

Holotype: U.S.N.M. 60259. 

Description based on 13 specimens. Body 
elongated, tapering posteriorly, 0.780-1.15 
long, 0.220-0.467 wide. Cuticle aspinose; 
eye-spot pigment present. Oral sucker 0.075- 
0.120 long, 0.083-0.135 wide; ventral suck- 
er 0.138-0.180 long, 0.096-0.168 wide, on 
a short peduncle; sucker ratio 1:1.27-1.55. 
Prepharynx short; pharynx 0.037-0.063 in 
diameter; esophagus as long as pharynx, 
cecum extending to about anterior level of 
ovary. Testis entire, 0.150-0.165 long, 0.083- 
0.120 wide, about midway between acetabu- 


180 


lum and posterior end of body; seminal 
vesicle tubular, sometimes reaching level of 
Ovary; pars prostatica large, spherical to 
ovoid; prostate glands diffuse, inconspicu- 
ous; ejaculatory duct short. Ovary entire, 
anterior to testis, 0.090-0.120 long, 0.053- 
0.105 wide; seminal receptacle dorsal, near 
Ovary; uterus extending posterior to testis. 
Genital pore midway between pharynx and 
acetabulum. Eggs 48-63 by 30-36 p, con- 
taining oculate miracidia. Vitellaria of 10-12 
inconspicuous follicles scattered between 
posterior edge of seminal vesicle and anterior 
third of testis. Excretory bladder not ob- 
served; its pore terminal. 

This is the first species of Haplosplanchnus 
to be reported from the Gulf-Caribbean 
region. Of the 3 previously described spe- 
cies, H. mugilis is most like H. pachysomus 
(Eysenhardt, 1829), but differs in having 
a shorter seminal vesicle, shorter peduncle 
of the acetabulum, a spherical or ovoid pars 
prostatica and the uterus extending posterior 
to the testis. It differs from H. caudatus 
(Srivastava, 1939) in the same respects and 
also in body shape. In H. puri Srivastava, 
1939, the testis is in the extreme posterior 
end of the body, the cecum and seminal 
vesicle are long, the vitellaria are compact, 
and the pars prostatica tubular. 


Schikhobalotrema acutum (Linton, 1910) 
Skrjabin & Guschanskaja, 1955 
Synonyms: Deradena acuta Linton, 1910; 
Haplosplanchnus acutus (Linton) Manter 
£957. 
Hosts: *Strongylura ardeola (C); S. ra- 
phidoma (J); S. timucu (C). 


Site: intestine. 


> 


Schikhobalotrema adacutum (Manter, 1937) 
Skrjabin & Guschanskaja, 1955 

Synonym: Haplosplanchnus adacutus Man- 
ter, 1937. 

Hosts: Abudefduf saxatilis (J); *Hali- 
choeres pictus (J); *Hemiramphus brasi- 
liensis (C, J); *Hepsetia stipes (J). 

Site; intestine. 


Schitkhobalotrema obtusum (Linton, 1910) 
Skrjabin & Guschanskaja, 1955 
Synonyms: Deradena obtusa Linton, 1910; 
Haplosplanchnus obtusus (Linton) Manter, 
LOST 
Host: Acanthurus hepatus (C). 
Site; intestine. 


Tulane Studies in Zoology 


Vol. 11 


Schikhobalotrema adbrachyurum Siddiqi & 
Cable, 1960 
Hosts: *Sparisoma flavescens (C); *S. 
viride (C); *Pseudoscarus guacamaia (C, 
J); *P. plumbaeus (J). 


Site: intestine. 


Schtkhobalotrema pomacentri (Manter, 
1937) Skrajabin & Guschanskaja, 1955 


Synonym: Haplosplanchnus pomacentri 
Manter, 1947. 
Hosts: *Pomacentrus analis (C); *P. 


fuscus (C, J); *P. leucosticus (C); *Micro- 
spathodon chrysurus (C, J). 
Site: intestine. 


Schikhobalotrema sparisomae (Manter, 1937) 
Skrjabin & Guschanskaja, 1955 


Synonym:  Haplosplanchnus 
Manter, 1937. 

Hosts: *Pseudoscarus guacamaia (C, J); 
*Scarus crotcensts (C); *Scaras spo ())-: 
*Sparisoma abildgaardi (C, J); *S. brachiale 
(J); *S. flavescens (J); *S. radians (J). 


Site; intestine. 


Sparisomae 


Schikhobalotrema bivesiculum n.sp. 
Figure 11 


Host: Abudefduf saxatilis (J). 
Site: intestine. 


Holotype: U.S.N.M. 60260. 


Description based on 6 specimens. Body 
elongated, more tapered posteriorly than 
anteriorly, 0.965-1.40 long, 0.347-0.467 
wide. Cuticle aspinose; eye-spot pigment 
present. Oral sucker 0.098-0.180 long, 0.153- 
0.200 wide; ventral sucker preequatorial, 
0.195-0.300 long, 0.180-0.233 wide, with 
elongated aperture but no _ posterolateral 
lobes; sucker ratio 1:1.35-1.48. Prepharynx 
short; pharynx 0.060-0.090 in diameter; 
esophagus short; cecum extending to mid- 
level of ovary. Gonads postequatorial. Testis 
entire, at about midlevel of hindbody, 0.200- 
0.267 long, 0.167-0.200 wide. Seminal vesi- 
cle bipartite; long tubular posterior portion 
extending to about midlevel of ovary; thick- 
walled anterior division with conspicuous 
circular muscles and 2 large nuclei on inner 
surface, protruding into lumen; pars pro- 
statica difficult to interpret; possibly part 
of a narrow tube leaving muscular portion 
of seminal vesicle or within conspicuous 
mass filling genital atrium and probably cor- 
responding to what has been called genital 


No. 4 Trematodes of Marine Fishes 181 


Q ws 
CF 


Nae 

Ga) a} 

eee 
FiO) 


Figure 7. Infundibulostomum anisotremi, holotype, dorsal view. Figure 8. Pseudobac- 
ciger manteri, holotype, ventral view. Figure 9. Bacciger opisthonemae, holotype, ventral 
view. Figure 10. Haplosplanchnus mugilis, holotype, ventrolateral view. Figure 11. 
Schikhobalotrema bivesicula, holotype, ventral view. 


182 


bulb; prostate cells inconspicuous. Ovary 
entire or slightly irregular, 0.120-0.140 long, 
0.078-0.105 wide; anterior to testis; seminal 
receptacle dorsolateral to ovary; uterus pre- 
ovarian. Genital pore median, at pharyngeal 
level. Eggs few, 60-78 by 40-53 p. Vitellaria 
of large follicles extending from anterior 
margin of ventral sucker to near posterior 
end of body. Excretory vesicle not observed; 
excretory pore terminal. 


Schtkhobalotrema bivesiculum differs from 
all other species of the genus in its conspicu- 
ous bipartite seminal vesicle. It resembles 
S. acutum and S. adacutum in general topog- 
raphy and although it has a longitudinal 
aperture in the acetabulum as in those spe- 
cies, the sucker lacks posterolateral lobes. 
The bipartite seminal vesicle with a thick 
anterior division in this species may be of 
generic value but the species is placed in 
Schikhobalotrema because others showing 
gradations in that respect may exist. 


Schikhobalotrema elongatum n.sp. 
Figure 12 
Hosts: Mugil cephalus (C); M. curema 
(p: 
Site; intestine. 
Holotype: U.S.N.M. 60261. 


Description based on 20 specimens. Body 
elongated, 0.734-1.49 long, 0.200-0.367 
wide, with prominant cuticular rings, espe- 
cially, in hindbody. Eye-spot pigment pres- 
ent. Oral sucker 0.090-0.150 long, 0.105- 
0.180 wide; ventral sucker without lobes, in 
anterior third or fourth of body, 0.120-0.195 
in diameter, aperture circular; sucker ratio 
1:1-1.32. Prepharynx about half length of 
pharynx; pharynx 0.060-0.075 in diameter; 
esophagus about as long as pharynx; cecum 
extending to midlevel of testis. Gonads in 
posterior half of body. Testis usually elon- 
gated, median, 0.167-0.366 long, 0.098-0.174 
wide; seminal vesicle tubular, reaching al- 
most midway between acetabulum and ovary; 
prostate cells granular, conspicuous, their 
ducts forming a bulbous mass just posterior 
to muscular genital atrium. Genital pore 
median, near posterior edge of pharynx. 
Ovary entire, anterior to testis, 0.090-0.133 
long, 0.060-0.090 wide, well removed from 
ventral sucker except in contracted speci- 
mens; seminal receptacle dorsal, near ovarian 
level; uterus preovarian. Eggs 60-84 by 
42-54 p. Vitelline follicles large, extending 


Tulane Studies in Zoology 


Vol. :11 


from pharyngeal level to posterior end of 
body, tending to fuse in hindbody. Excretory 
vesicle not observed; pore terminal. 


The most distinctive features of Schik- 
hobalotrema elongatum are its long hind- 
body with prominent cuticular rings, and 
its well-developed bulb of prostatic ducts. In 
body shape, S. elongatum is similar to sey- 
eral species. S. acutum and S. adacutum have 
a ventral sucker with a longitudinal aperture 
and lateral lobes. S. obtusum lacks a pro- 
static vesicle; the genital atrium is non- 
muscular and the ovary close to the acetabulu- 
lum. S. pomacentri has an equatorial ventral 
sucker and S. kyphost (Manter, 1947) a 
lobed ovary. S. girellae (Manter & Van 
Cleave, 1951) is most like S. elongatum but 
differs from that species in the extent of the 
seminal vesicle, in having a long tubular 
genital atrium and a more anterior ovary. 
S. manteri Siddigi & Cable, 1960, differs in 
the distribution of the vitellaria, position of 
the testis and posterior extent of the uterus. 
S. robustum Pritchard & Manter, 1961, is 
much larger, has a thin-walled genital atrium 
and the genital pore on a finger-like pro- 
jection. 


Schikhobalotrema heterocotylum n.sp. 
Figure 13 


Host: Pseudoscarus guacamaia (C). 

Site: intestine. 

Holotype: U.S.N.M. 60262. 

Description based on 5 specimens. Body 
subspherical to pyriform, 1.29-1.50 long, 
0.869-0.965 wide. Cuticle thick, aspinose; 
eye-spot pigment present. Oral sucker 0.128- 
0.186 long, 0.233-0.266 wide, ventral lip 
with papilla bearing openings of salivary 
gland in forebody. Ventral sucker at mid- 
body, 0.466-0.533 long, 0.533-0.613 wide, 
its interior with 2 anterior and 2 posterior 
tuberculated projections. Sucker ratio 1:2.6- 
2.8. Prepharynx short; pharynx 0.130-0.150 
long, 0.105-0.135 wide; esophagus short; 
cecum extending to midlevel of acetabulum. 
Gonads in posterior third of body. Testis 
0.386-0.486 long, 0.213-0.333 wide, to right 
of midline; seminal vesicle long, extending 
to posterior level of ventral sucker or slightly 
beyond; pars prostatica small, inconspicuous. 
Ovary entire to slightly lobed, 0.133-0.180 
long, 0.098-0.135 wide, median, posterior to 
acetabulum; seminal receptacle postovarian 
and much larger than ovary; uterus pre- 


No. 4 


ovarian. Genital pore ventral, near midlevel 
of pharynx. Eggs 84-99 by 53-75 p. Vitel- 
line foliicles numerous, extending from mid- 
level of pharynx to posterior end of body. 
Excretory vesicle not observed. 

No haplosplanchnid has been described as 
having an acetabulum with the tuberculated 
lobes characteristic of this species. Most like 
it in other respects is S. brachyurum (Man- 
ter, 1937) a paratype of which we have ex- 
amined and found to lack such lobes. It 
further differs from S. heterocotylum in havy- 
ing a different body shape, a smaller ventral 
sucker and a different sucker ratio. Although 
similar in body shape, S. glomerosum Pritch- 
ard and Manter, 1961, has a smaller ventral 
sucker, a preacetabular ovary, more anterior 
testis and a well-developed metraterm. 

The next species has haplosplanchnid char- 
acteristics but differs from known genera in 
having the ventral sucker so near the pos- 
terior end of the body that the gonads neces- 
sarily are preacetabular. In life, that sucker 
was nearer the posterior end of the body in 
our specimen than it appears in the whole 
mount (Fig. 14). Thus the Haplosplanch- 
nidae is another example of families in 
which certain species evidently are second- 
arily amphistomatous. To receive the spe- 
cies, a new genus is erected and character- 
ized as follows: 


Haplosplanchnoides n.g. 


Diagnosis: Family Haplosplanchnidae. 
Body with thick unarmed cuticle. Oral suck- 
er terminal; ventral sucker near posterior end 
of body. Prepharynx and pharynx present; 
cecum single, extending to posterior end of 
body. Testis single, anterior to acetabulum; 
Cirrus sac absent; seminal vesicle long and 
tubular. Ovary anterior to testis; seminal 
receptacle present; uterus pretesticular. Geni- 
tal pore in anterior half of body. Vitellaria 
extensive, from prepharyngeal level to pos- 
terior end of body. Eggs large. Parasites in 
intestine of marine fishes. Type and only 
species: 

Haplosplanchnoides hemiramphi 

n.g., 0.sp. 
Figure 14 

Host: Hemiramphus brasiliensis (J). 

Site; intestine. 

Holotype: U.S.N.M. 60263. 


Description based on a single specimen. 
Body elongated, rounded at both ends, .1.73 


Trematodes of Marine Fishes 183 


long by 0.720 wide. Oral sucker 0.233 long, 
0.267 wide; ventral sucker near posterior end 
of body, 0.420 long, 0.313 deep; ratio of 
sucker lengths 1:1.7. Prepharynx short; 
pharynx 0.133 long, 0.160 wide; cecum long, 
terminating near posterior end of body. 
Testis entire, 0.188 by 0.210, near anterior 
margin of ventral sucker; seminal vesicle 
long, sinuous, extending from genital pore 
tO testis; pars prostatica and prostate cells 
not seen, probably obscured by vitelline fol- 
licles; ejaculatory duct short. Ovary entire, 
0.153 by 0.158, a short distance anterior to 
testis; seminal receptacle anterodorsal to 
ovary; uterus short. Genital pore probably 
median, about midway between ovary and 
pharynx. Vitelline follicles large, filling most 
of body between prepharynx and posterior 
end. Eggs few, 75-84 by 60-68 pw. Excretory 
vesicle not observed; pore dorsal, a short 
distance from posterior end of body. 

Although certain other haplosplanchnids 
have short hindbodies, their gonads are never 
anterior to the acetabulum except perhaps in 
Schikhobalotrema glomerosum which Pritch- 
ard and Manter (1961) described from two 
specimens in poor condition. 


FAMILY MEGAPERIDAE Manter, 
1934 
Thysanopharynx elongatus Manter, 1933 
Host: Lactophrys bicaudalis (C). 
Site; intestine. 


Megapera gyrina (Linton, 1907 ) 
Manter, 1934 

Synonyms: Distomum gyrinus Linton, 
1907; Eurypera gyrina (Linton) Manter, 
1935: 

Hosts: Lactophrys bicaudalis (C); L. tri- 
cornis (J); *L. triqueter (C). 

Site; intestine. 


Megapera pseudogyrina n.sp. 
Figure 15 


Host: Cantherines pullus (J). 

Site: intestine. © 

Holotype: US.N.M. 60264. 

Description based on 17 specimens. Body 
0.667-1.60 long, greatest width 0.246-0.506, 
in forebody, gradually tapering towards pos- 
terior end; sides sometimes incurved pos- 
terior to testes; anterior end broadly rounded: 
Cuticular spines extremely fine, not extend- 
ing posterior’ to testes; eye-spot. pigment 
present. Oral‘sucker 0.150-0.333 Jong, 0.1654 


184 


0.400 wide, with histology peculiar to the 
family; ventral sucker 0.060-0.100 in diame- 
ter, near junction of first and second thirds 
of body length; sucker ratio 1:0.3-0.45. 
Pharynx 0.045-0.090 long, 0.130-0.150 wide 
with lobed anterior border; esophagus ab- 
sent; ceca long and wide, opening separately 
through ani at posterior end of body. Testes 
2, elongated, 0.120-0.333 long, 0.060-0.166 
wide, symmetrical, extending posterolaterally 
from pharynx to about posterior level of 
ventral sucker, overlapping cecal arch. Semi- 
nal vesicle free in parenchyma, spherical, 
sometimes larger than ventral sucker, usually 
extending posterior to it; pars prostatica 
dorsal to genital atrium, small, spherical, 
surrounded by prostate cells. Ovary indented 
to trilobed, 0.053-0.150 long, 0.045-0.080 
wide; seminal receptacle anterodorsal to 
ovary; uterus short, mostly preovarian. Geni- 
tal pore median, immediately preacetabular. 
Vitelline follicles elongated, in two ventro- 
lateral groups extending from near testes to 
posterior end of body, and 2 dorsal groups 
usually enmeshed in midline, extending from 
posterior margin of ovary to near posterior 
end of body. Eggs thin-shelled 48-53 by 
31-33 p. Excretory vesicle tubular as far as 
ovarian level. 


This species is most like Megapera gyrina 
(Linton, 1907) but differs from it chiefly 
in body shape and the form and distribution 
of the vitellaria. Posterior to the testes the 
body tapers gradually or is moderately in- 
curved instead of narrowing abruptly and 
becoming tail-like as in M, gyrina. Although 
the vitelline follicles of M. pseudogyrina are 
elongated, they are by no means as narrow, 
uniformly shaped, or as numerous as in M. 
gyrina. 


Superorder Epitheliocystidia La Rue, 1957 
Order Plagiorchtida La Rue, 1957 
Suborder Plagiorchiata La Rue, 1957 
Superfamily Plagiorchioidea Dollfus, 1930 
FAMILY MICROPHALLIDAE 
Travassos, 1921 


Carneophallus lactophrysi Siddiqi & Cable, 
1960 


Hosts: *Myrichthys acuminatus (C); *M. 
oculatus (C). 

Site: intestine. 

Although this species was a common 
parasite of eels of the genus Myrichthys in 


Tulane Studies in Zoology 


Voli 


Curagao, it was not found there in the trunk 
fish which is the type host in Puerto Rico. 


Microphallus excellens (Nicoll, 1907 ) 
Baer, 1943 
Synonym: Spelotrema excellens Nicoll, 
1907. 

Host: *Myrichthys oculatus (C). 

Site: intestine. 

Fifteen worms from a single host are 
identified as M. excellens using Baer’s (1943) 
key and that of Biguet, Deblock and Capron 
(1958). They differ, however, from Nicoll’s 
specimens in being smaller (0.547-0.781 
compared with 0.71-1.39) and also in hav- 
ing relatively larger suckers and pharynx, 
Belopolskaja in Skrjabin (1952) considers 
Spelotrema feriatum Nicoll, 1907, a syno- 
nym of S. excellens. The present material 
falls within the combined range of measure- 
ments given under those names. 

A species of Microphallus has not hitherto 
been reported from a marine fish. 


Superfamily Lepocreadioidea Cable, 1956 
FAMILY LEPOCREADIIDAE Nicoll, 
1934 
Homalometron elongatum Manter, 1947 
Host: Gerres cinereus (J). 

Site; intestine. 


Homalometron foliatum Siddigi & Cable 
1960 

Hosts: Haemulon album (C); H. flavo- 
lineatum (C, J); *H. sciurus (J); *Luti- 
anus mahogoni (C); *Brachygenys chry- 
Sargyreus (C). 

Site; intestine. 

Thirty individuals of this species agree 
with the original description except that the 
body length is up to 3.956, the sucker ratio 
ranges from 1:0.70-1.03, the vitellaria may 
reach the anterior margin of the ovary, and 
egg size is 66-90 by 45-60 p. 


Crassicutis marina Manter, 1947 
Host: Gerres cinereus (J). 
Site: intestine. 


Crassicutis gerridis n.sp. 
Figure 16 
Host: Gerres cinereus (C, J). 
Site: intestine. 
Holotype: U.S.N.M. 60265. 
Description based on 15 specimens. Body 
elongated, 1.04-2.6 long, 0.400-0.710 wide. 


No. 4 Trematodes of Marine Fishes 185 


Figure 12. Schikhobalotrema elongata, holotype, ventral view. Figure 13. Schikhobal- 
otrema heterocotyla, holotype, ventral view. Figure 14. Haplosplanchnoides hemiramphi, 
holotype, ventrolateral view. Figure 15. Megapera pseudogyrina, holotype, ventral view. 
Figure 16. Crassicutis gerridis, holotype, ventral view. 


186 


Long, very narrow, widely spaced spines im- 
bedded in thick cuticle of larger specimens 
but not of immature ones. Eye-spot pig- 
ment present. Oral sucker 0.173-0.280 in 
diameter; ventral sucker equatorial, 0.206- 
0.400 long, 0.267-0.333 wide; sucker ratio 
1:1.13-1.43. Prepharynx about 3/4 length of 
pharynx; pharynx 0.055-0.105 long, 0.075- 
0.114 wide; esophagus shorter than pharynx, 
intestinal bifurcation well anterior to ventral 
sucker; ceca long, extending almost to pos- 
terior end of body. Testes tandem, close to- 
gether, 0.100-0.300 long, 0.180-0.333 wide; 
posttesticular space 1/6-1/8 body length. 
Cirrus sac absent; ejaculatory duct relatively 
long, opening into indistinct genital atrium; 
pars prostatica small, with indistinct pros- 
tate cells; seminal vesicle saccular, extend- 
ing to or slightly beyond posterior edge of 
ventral sucker. Genital pore small, median, 
immediately anterior to ventral sucker. 
Ovary smooth, 0.100-0.233 in diameter, 
separated from anterior testis by Mehlis’ 
gland; seminal receptacle small, inconspicu- 
ous, antero- or laterodorsal to ovary; Laurer’s 
canal opens dorsally, median to ovary; uterus 
pretesticular; metraterm not observed. Vitel- 
laria of large follicles extending from pos- 
terior edge of oral sucker or pharynx to 
posterior end of body, occupying all avail- 
able space between ventral sucker and phar- 
ynx. Eggs 83-113 by 46-60 p. Excretory 
vesicle short, sac-shaped, not quite reaching 
posterior testis. 


Three species of Crassicutis have been de- 
scribed. C. cichlasomae Manter, 1936, from 
a fresh-water host, C. marima Manter, 1947, 
and C. archosargi Sparks & Thatcher, 1960. 
C. gerridis differs from all 3 in having a 
long forebody filled with vitelline follicles. 
In addition it differs from C. cichlasomae in 
habitat and in having tandem rather than 
diagonal testes. It differs from C. archosargi 
in the position of the ventral sucker and 
position of the gonads. C. marina, described 
from the same host in Florida and also found 
in this study, differs from C. gerridis chiefly 
in having a more anterior ventral sucker and 
much less extensive preacetabular vitellaria, 
differences observed in living specimens as 
well as whole mounts; other possible differ- 
ences are the less conspicuous seminal vesicle 
and seminal receptacle in C. gerridis. 


Tulane Studies in Zoology 


Vol. 11 


Lepidapedon trachinott Hanson, 1950 

Hosts: *Epinephelus morio (C); E. Stri- 
atus (C). 

Site: ceca. 

The most distinctive feature of this spe- 
cies is the lateral depression at the left mar- 
gin of the body opposite the genital pore. 
Our specimens agree with Hanson’s descrip- 
tion except for a slightly larger size (up to 
3.76 long by 0.594 wide) and an entire 


rather than irregular ovary. 


Lepidapedon truncatum Sogandares-Bernal, 
O59 
Synonym: **Lepidapedon holocentri Sid- 
digi & Cable, 1960. 
Host: Holocentrus ascensionts (C, J). 
Site: ceca and upper intestine. 


Lepidapedon holocentri is obviously a 
synonym of L, truncatum. The descriptions 
of both were in press at the same time. 


Neolepidapedon mycteropercae Siddigi & 
Cable, 1960 
Hosts: *Mycteroperca bonaci (C); *M. 
venonosa (C). 
Site: intestine. 


Fifty-five specimens are in general agree- 
ment with the original description of the 
species, based on a single specimen, but show 
certain variations. The vitellaria may or 
may not be confluent between the gonads 
and may reach the midlevel of the external 
seminal vesicle. The metraterm is muscular, 
the genital pore is usually posterolateral to 
acetabulum but may be near its midlevel. 
The sucker ratio is 1:1.38-1.48 and eggs 
measure up to 62 » 1n length. 


Neolepidapedon hypoplectri n.sp. 
Figure 17 

Host: Hypoplectrus unicolor (J). 

Site: intestine. 

Holotype: U.S.N.M. 60266, 

Description based on 13 specimens. Body 
elongated, 1.35-2.02 long, 0.280-0.380 wide. 
Entire cuticle spinose; eye-spot pigment pres- 
ent. Oral sucker 0.090-0.112 in diameter; 
ventral sucker 0.063-0.083 in diameter; 
sucker ratio 1:0.70-0.76. Prepharynx very 
short; pharynx 0.033-0.054 in diameter; 
esophagus 2-3 times length of pharynx; in- 
testinal bifurcation about midway between 
suckers; ceca extending’ to near posterior 
end of body. Testes 2, entire, 0.068-0.135 


No. 4 


long, 0.090-0.150 wide, tandem, not close 
together. Cirrus sac extending well posterior 
to ventral sucker, containing small spherical 
internal seminal vesicle, large conspicuous 
pars prostatica and cirrus; external seminal 
vesicle tubular, sinuous, surrounded by pros- 
tatic cells along most of its length and ex- 
tending about halfway from ventral sucker 
to ovary. Ovary entire, smooth, pretesticular, 
0.063-0.098 long, 0.080-0.105 wide, seminal 
receptacle postovarian; uterus preovarian; 
metraterm about same length as cirrus sac. 
Genital atrium small; genital pore sinistral, 
at about midlevel of ventral sucker. Eggs 
45-60 by 28-33 p. Vitelline follicles in lat- 
eral fields, from about end of cirrus sac to 
posterior end of body; confluent in post- 
testicular space, usually between testes, rarely 
between ovary and anterior testis. Excretory 
vesicle tubular, extending to intestinal bi- 
furcation; sphincter well developed, pore 
terminal. 

The excretory vesicle is known to termi- 
nate between the anterior margin of the 
acetabulum and the intestinal bifurcation in 
3 species of Neolepidapedon besides N. 
hypoplectri: N. medialunae Montgomery, 
1957, N. epinmepheli Siddiqi & Cable, 1960, 
and N. mycteropercae Siddiqi & Cable, 
1960. The anterior extent of the vesicle is 
not described for N. cablez Manter, 1954, 
and N. retrusum (Linton, 1940). In all other 
species reported to date, it reaches only to 
the anterior testis or ovary. N. hypoplectrus 
differs from N. cablez chiefly in lacking the 
preacetabular, glandular hump-like protuber- 
ance, in sucker ratio and position of the ace- 
tabulum; from N. retrusum and N. myc- 
teropercae in sucker ratio; from N. epine- 
phelt in the anterior extent of the vitellaria, 
position of the genital pore and in having 
a smaller pharynx and narrow eggs; and 
from N. medialunae in position of the ven- 
tral sucker, extent of spination, shape of 
oral sucker and in having a shorter pre- 
pharynx and esophagus. 


Multitestis inconstans (Linton, 1905 ) 
Manter, 1931 
Synonym: Distomum inconstans Linton, 
1905. 
Host: Chaetodipterus faber (J). 
Site: intestine. 


Multitestis blenni Manter, 1931 
Host: Chaetodipterus faber (J). 


Site: intestine. 


Trematodes of Marine Fishes 


187 


Multitestis chaetodont Manter, 1947 
Synonym: Distomum sp. of Linton, 1907, 
parlis: 
Hosts: Chaetodon capistratus (J); C. stri- 
atus (J). 
Site: ceca and intestine. 


It is of interest to note that none of the 
Chaetodon species examined in Curacao 
harbored M. chaetodonz. It seeems to be re- 
placed there by the next species, at least in 
C. capistratus. 


Multitestis rotundus Sparks, 1954 
Hosts: *Archosargus unimaculatus (J); 
*Chaetodon capistratus (C, J). 
Site; intestine. 


Our specimens actually agree more with 
those of Sogandares-Bernal and Hutton 
(1959b) than with the species as originally 
described. The vitellaria extend anteriorly 
as far as the pharyngeal level and sometimes 
to the oral sucker; the follicles may or may 
not be confluent posteriorly. 


Opechona chloroscombri n.sp. 
Figure 18 
Synonym: Opechona sp. Siddigi & Cable, 
1960. 
Host: Chloroscombrus chrysurus (J). 
Site: intestine. 


Holotype: US.N.M. 60267. 


Description based on 12 mature specimens. 
Body elongated, rounded at both ends, 1.06- 
1.50 long, 0.220-0.300 wide. Entire cuticle 
spinose; eye-spot pigment present. Oral 
sucker 0.039-0.053 in diameter; ventral 
sucker in anterior region of mid-third of 
body, 0.055-0.068 long, 0.063-0.084 wide; 
sucker ratio 1:1.4-1.5. Prepharynx 0.030- 
0.098 long; pharynx 0.048-0.060 long, 0.033- 
0.042 wide; esophagus 0.180-0.255 long, 
including a posterior glandular portion 
(pseudesophagus) measuring 0.105-0.150 in 
length; ceca extend to near posterior end of 
body. Testes 0.068-0.100 in diameter, tan- 
dem, close together, in posterior third of 
body; posttesticular space 1/5-1/4 body 
length. Cirrus sac long, extending to about 
midway between acetabulum and ovary, 
containing small spherical internal seminal 
vesicle, larger pars prostatica and long cir- 
rus; external seminal vesicle tubular, about 
half length of cirrus sac. Ovary entire, 0.045- 
0.054 long, 0.053-0.078 wide; seminal re- 
ceptacle between ovary and anterior testis; 


188 


uterus preovarian; metraterm about same 
length as cirrus; genital pore slightly sinis- 
tral, about midway between acetabulum and 
intestinal bifurcation. Vitelline follicles ex- 
tending from level of ventral sucker to pos- 
terior end of body, not confluent posterior 
to testes. Opaque eggs 45-53 by 30-36 yp. 
Excretory vesicle, thick-walled (epithelial), 
varying in anterior extent from anterior 
margin of acetabulum to intestinal bifurac- 
tion. 


Opechona chloroscombri is most similar 
to O. gracilis (Linton, 1910) in the extent 
of the vitellaria and length of the excretory 
vesicle but differs from that species in size 
and ratio of suckers and the ratio of pseud- 
esophagus to esophagus. 


Opechona sardinellae n.sp. 
Figure 19 


Host: Sardinella macrophthalmus (J). 

Site: intestine. 

Holotype: US.N.M. 60268. 

Description based on 9 specimens. Body 
1.03-1.158 long, 0.286-0.536 in maximum 
width near testicular level, tapering toward 
anterior end; posterior extremity rounded, 
usually with indentation at excretory pore. 
Cuticle spinose, spines extending only to 
about intestinal bifurcation; eye-spot pig- 
ment present. Oral sucker 0.075-0.135 long, 
0.105-0.150 wide; ventral sucker somewhat 
preequatorial, 0.066-0.113 in diameter; suck- 
er ratio 1:0.7-0.8. Prepharynx short, phar- 
ynx massive, 0.090-0.150 in diameter; 
esophagus about same length as pharynx, 
its epithelial region (pseudesophagus) al- 
most as long as simple anterior portion; ceca 
wide, reaching midway between testes and 
posterior end of body. Testes tandem, 0.075- 
0.153 long, 0.098-0.160 wide, in posterior 
third of body; cirrus sac extending to about 
midway between acetabulum and ovary, con- 
taining small internal seminal vesicle, pars 
prostatica which is sometimes indistinctly 
bipartite, and cirrus. External seminal vesicle 
tubular, about 2/3 length of cirrus sac. Ovary 
entire, 0.055-0.100 long, 0.080-0.114 wide, 
separated from anterior testis by seminal 
receptacle and vitelline reservoir; uterus pre- 
testicular, terminating in short, muscular 
metraterm. Genital pore sinistral, midway 
between ventral sucker and intestinal bi- 
furcation. Vitelline follicles extending from 
level of acetabulum to posterior end of body, 


Tulane Studies in Zoology 


Vol. 11 


rarely confluent behind testes. Eggs 58-68 
by 35-45 p. Excretory vesicle tubular, sig- 
moid in living specimens, crossing left ce- 
cum ventrally and extending to midlevel of 
pharynx; excretory pore terminal, with 
sphincter. 


Species of Opechona with the extent of 
vitellaria and excretory vesicle more or less 
as in O. sardinellae are: O. orientalis (Lay- 
man, 1930) and O. pharyngodactyla Manter, 
1940. O. sardinellae differs from O. orten- 
talis chiefly in shape of ovary and oral 
sucker and in having a shorter prepharynx 
and esophagus, and from O. pharyngodactyla 
in lacking the finger-like projections on the 
pharynx and in sucker ratio. Opechona gra- 
cilis reported by Linton (1910) from Claw- 
panodon pseudohispanicus and by Manter 
(1947) from Harengula (Sardinella) ma- 
crophthalmus is distinguished from O. sar- 
dinellae by extent of body spination, sucker 
ratio, proportion of pseudesophagus to the 
esophagus, size of pharynx and length of 
the excretory vesicle which was reported by 
Manter to extend only to the acetabulum. 
Opechona sardinellae differs from O. chlo- 
roscombri in body shape and spination, size 
and ratio of suckers, and length of excretory 
vesicle and esophagus. 


Lepocreadium bimarinum Manter, 1940 
Host: *Bodianus rufa (J). 
Site: intestine. 


Lepocreadium trulla (Linton, 1907 ) 
Linton, 1910 
Synonym: Distomum trulla Linton, 1907. 
Host: Ocyurus chrysurus (C, J). 
Site; intestine. 


Lepocreadtum pyriforme (Linton, 1900) 
Linton, 1940 

Synonym: Duistomum pyriforme Linton, 
1900. 

Host: Peprilus paru (J). 

Site: intestine. 

Linton (1940) reported as this species 
trematodes from 9 hosts including Peprilus 
paru. We doubt that all of them are the 
same species. Our single specimen lacks 
eggs but otherwise agrees with Linton’s de- 
scription and is referred to L. pyriforme on 
the basis of its similarity to his Figure 47 
of a specimen from Palinurichthys perct- 
formis, the type host of L. pyriforme. 


No. 4 


Lepocreadinm opsanust Sogandares-Bernal 
& Hutton, 1960 

Hosts: *Calamus arctifrons (J); *C. ba- 
janado (J). 

Site: ceca and intestine. 

In the extent and distribution of vitel- 
laria, position of gonads, extent of the ex- 
cretory vesicle and general topography, our 
numerous specimens are most like ones 
which Sogandares-Bernal and Hutton (1960) 
named provisionally as Lepocreadium opsa- 
nust. In the Jamaican specimens, the body 
is somewhat smaller (0.346-0.586 long, 
0.185-0.253 wide) as are the eggs (54-62 by 
33-38 «) and the sucker ratio is 1:0.83-1.00 
whereas Sogandares-Bernal and Hutton gave 
a ratio of 1:1-1.85. However, their drawing 
of the holotype shows an acetabulum that 
is smaller than the oral sucker as is true of 
most of our specimens. 


Lepocreadium hemiramphi n.sp. 
Figure 20 
Host: Hemiramphus brasiliensis (C). 


Site; intestine. 
Holotype: US.N.M. 60269. 


Description based on 25 specimens. Body 
from pyriform to more elongated with blunt- 
ly pointed posterior end, 0.286-0.513 long, 
0.160-0.200 wide. Entire cuticle spinose; 
eye-spot pigment present. Oral sucker 0.039- 
0.055 in diameter; ventral sucker in middle 
third of body, 0.037-0.060 in diameter; suck- 
er ratio 1:0.91-1.00. Pharynx 0.036-0.045 
long, 0.030-0.033 wide; prepharynx and 
esophagus about as long as pharynx; in- 
testinal bifurcation close to acetabulum; ceca 
extend almost to posterior end of body. 
Testes 2, entire, tandem, near posterior end 
of body, 0.030-0.060 in diameter; cirrus sac 
about 1/5 body length, reaching ovarian 
zone, usually ovoid, containing spherical in- 
ternal seminal vesicle, prominent pars pro- 
statica and long, winding cirrus; external 
seminal vesicle large, spherical. Genital 
atrium small; genital pore anterosinistral to 
acetabulum. Ovary entire, median, 0.027- 
0.039 long, 0.030-0.048 wide; seminal re- 
ceptacle postovarian; uterus short, pre- 
Ovarian, terminating in conspicuous, thick- 
walled metraterm. Eggs few (no more than 
2 observed in uterus of any one worm), 
45-57 by 30-37 p. Vitelline follicles extend- 
ing from esophageal level almost to pos- 
terior end of body, slightly if at all over- 


Trematodes of Marine Fishes 


189 


reaching posterior testis. Excretory vesicle 
tubular, its anterior extent not determined; 
excretory pore terminal. 

Lepocreadium hemiramphi is most similar 
to L. floridanum Sogandares-Bernal and Hut- 
ton, 1959 and L. pyriforme (Linton, 1900). 
It differs from both in being much smaller 
and more compact with the cirrus sac reach- 
ing the ovary, and in having the testes nearer 
the posterior end and the vitellaria extend 
farther anteriorly but not appreciably pos- 
terior to the testes. 


Lepocreadium truncatum n.sp. 
Figure 21 

Synonym: Lepocreadium sp. Siddiqi & 
Cable, 1960. 

Host: Ocyurus chrysurus (C). 

Site: intestine. 

Holotype: US.N.M. 60270. 

Description based on 4 specimens (1 from 
Curacao and 3 from Puerto Rico). Body 
0.467-0.714 long, 0.293-0.393 wide, pyri- 
form, tapering anteriorly, truncated poste- 
riorly. Entire cuticle spinose, spines becom- 
ing sparse posteriorly; eye-spot pigment 
present. Oral sucker 0.053-0.075 long, 0.060- 
0.090 wide; ventral sucker 0.099-0.120 in 
diameter; sucker ratio 1:1.3-1.55. Prephar- 
ynx present; pharynx 0.045-0.060 in diame- 
ter; esophagus short; intestinal bifurcation 
about midway between suckers; ceca extend- 
ing to posterior end of body. Testes 2, en- 
tire, tandem, 0.042-0.083 long, 0.105-0.150 
wide; cirrus sac extending midway from ace- 
tabulum to ovary, containing spherical in- 
ternal seminal vesicle, well developed pro- 
static complex and thick cirrus; external 
seminal vesicle saccate. Ovary 0.068-0.083 
long, 0.030-0.068 wide, trilobed, to right of 
median line; seminal receptacle present; 
uterus pretesticular; metraterm distinct. 
Genital pore about midway between ventral 
sucker and intestinal bifurcation. Eggs few, 
53-60 by 28-33 pw. Vitelline follicles large, 
extending from anterior level of ventral 
sucker to posterior end of body. Excretory 
vesicle tubular, anterior extent not deter- 
mined; pore terminal. 


This species is most like L. trulla (Linton, 
1907) and L. maris (Caballero, 1957) but 
differs from them chiefly in sucker ratio 
and in having tandem rather than diagonal 
testes. 


190 Tulane Studies in Zoology 


Apocreadium balistis Manter, 1947 
Host: Balistes vetula (J). 
Site; intestine. 


Of 3 specimens, one of which was im- 
mature, none shows ridges on the testes. 


Apocreadium mexicanum Manter, 1937 
Host: *Monacanthus hispidus (J). 
Site: intestine. 


Our many specimens are more like those 
of Siddiqi and Cable (1960) from Puerto 
Rico than the species as originally described. 
The posttesticular space usually is less than 
half as long as the body but sometimes the 
2 regions are about equal in length. Opaque 
eggs measure 63-71 by 42-45 yp, collapsed 
ones are 30-40 » wide. The anterior limit 
of the vitellaria varies between the posterior 
and anterior margins of the ventral sucker. 


Neoapocreadium coili (Sogandares-Bernal, 
1959 ) Siddiqi & Cable, 1960 
Synonym: Apocreadium coili Sogandares- 
Bernal, 1959. 
Host: Balistes vetula (J). 
Site: intestine. 


Neoapocreadium angustum (Sogandares- 
Bernal, 1959) Siddiqi & Cable, 1960 
Synonym: Apocreadium angustum Sogan- 
dares-Bernal, 1959. 
Host: Lactophrys trigonus (C). 
Site: intestine. 


Postporus epinepheli (Manter, 1947 ) 
Manter, 1949 
Synonyms: Optisthoporus epinepheli Man- 
1947; Postporus mycteropercae (Man- 
ter) Manter, 1949. 
Hosts: Epinephelus adscensionis (C, J); 
E. guttatus (C); E. morio (C); E. striatus 
(CG, J). 


Site: intestine. 


The variations observed in this species by 
Siddiqi and Cable (1960) are confirmed. 
The sucker ratio is 1:0.61-0.97 except for 
one of 1:0.53 in a single, apparently ex- 
cessively flattened specimen from Epine- 
phelus striatus. 


Myzoxenus lachnolaimi Manter, 1947 
Host: Lachnolaimus maximus (J). 
Site: intestine. 


Rhagorchis odhneri Manter, 1931 
Synonym: Gargorchis varians Linton, 
1940. 


Vol. 11 


Host: Alutera schoepfi (J). 

Site; intestine. 

Manter (1931) described the excretory 
vesicle as being tubular but in our living 
specimens it was distinctly Y-shaped, bi- 
furcating dorsal to the ventral sucker to form 
voluminous arms extending to the sides of 
the pharynx. The main excretory tubules 
evidently leave the stem of the vesicle and 
divide into anterior and posterior tubules 
before reaching the acetabular level. 


Cadenatella kyphosi n.sp. 
Figures 22 and 23 


Host: Kyphosus sectatrix (C). 
Site; intestine. 
Holotype: U.S.N.M. 60271. 


Description based on 13 specimens; meas- 
urements on 8 mature ones. Body elongated, 
2.3-4.15 long, 0.267-0.366 wide. Cuticle 
spinose from anterior end almost to ventral 
sucker dorsally, to level of testis ventrally. 
Eye-spot pigment present; brownish yellow 
pigment scattered through parenchyma. 
Forebody with 14-18 midventral accessory 
suckers; one, 2 or rarely 3 near anterior 
edge of pharynx, distinctly separated from 
others extending from near posterior edge 
of pharynx to acetabulum. Oral sucker with 
8 muscular preoral lobes, rather uniform in 
length, in a dorsal and a ventral row of 4 
each; lobes not subdivided or prominently 
extended in either living or fixed and stained 
specimens; sucker 0.133-0.166 long includ- 
ing lobes, 0.107-0.140 wide. Ventral sucker 
in anterior fourth of body, 0.146-0.200 in 
diameter; sucker ratio 1:1.14-1.40. Pre- 
pharynx wide, about same length as pharynx; 
pharynx massive, pyriform, 0.150-0.220 long, 
0.107-0.155 wide; esophagus very short, 
ceca extending to posterior end of body, 
joining excretory vesicle to form uroproct 
with terminal pore. Single testis elongated, 

0.333-0.446 long, 0.127-0.167 wide, about 
1/3 body length from posterior end. Cirrus 
sac absent; seminal vesicle long, sinuous, ex- 
tending about halfway from ventral sucker 
to Ovary; pars prostatica ovoid, near anterior 
edge of acetabulum; ejaculatory duct short. 
Genital atrium inconspicuous, genital pore 
midventral, immediately anterior to ventral 
sucker. Ovary entire, 0.080-0.133 long, 
0.106-0.140 wide, anterior to testis, separated 
from it by vitelline reservoir; seminal re- 
ceptacle of uterine type, anterior to ovary; 


No. 4 Trematodes of Marine Fishes 191 


“4 


11m 
Ce 


UT i. 
HFN E 


Oo 
£ AY ay = 
aor’ ey Eps ne RY: rox} 
> e . => HER ay pe wage l 


Sots a 
ey ee SCE 
Syste 
eek 
) > ae) 
weet Heed 
| 21 
Gem bast 
te Fee 
ry hr 
19 > Xf A 


ew 


Figure 17. Neolepidapedon hypoplectri, holotype, ventral view. Figure 18. Opechona 
chloroscombri, holotype, ventral view. Figure 19. Opechona sardinellae, holotype, dorsal 
view. Figure 20. Lepocreadium hemiramphi, holotype, ventral view. Figure 21. Lepo- 
creadium truncatum, holotype, dorsal view. Figure 22. Cadenatella kyphosi, holotype, 


ventral view. Figure 23. Same, oral sucker from dorsal aspect, drawn free-hand from 
living specimen. 


192 


Mehlis’ gland near receptacle; Laurer’s canal 
present, opening dorsal to ovary; uterus pre- 
ovarian, intercecal; metraterm absent. Eggs 
broadly lunate, 48-68 by 22-30, usually 52- 
63 by 24-28 p. Vitelline follicles numerous, 
extending from posterior third of testis to 
end of body. Excretory vesicle Y-shaped, its 
stem bifurcating at ovarian level to form 
wide arms extending to about midway be- 
tween ovary and ventral sucker; excretory 
canals leave tips of arms to extend to sides 
of oral sucker and turn posteriorly, receiving 
first branch at prepharyngeal level; flame 
cells numerous. Lymphatic channels not evi- 
dent but a number of ventral glands on each 
side of forebody have ducts in a bundle 
accompanying excretory canals with some at 
least opening at anterior end of body. 

This species represents a peculiar group 
of trematodes known only from chubs of 
the genus Kyphosws. Uncertainty as to its 
affinities was mentioned earlier in this 
paper in connection with the Superfamily 
Haploporoidea. Generic concepts within the 
group remain to be clarified. 

Dollfus (1946) described 3 species from 
Kyphosus sectatrix in Senegal and allocated 
each to a new subgenus in the genus Exen- 
terum, naming them E. (Enenterum) pseu- 
daureum, E. (Cadenatella) cadenati and E. 
(Jeancadenatia) brumpti. The subgenus 
Enenterum had 10 oral lobes, no accessory 
suckers, and 2 testes; Cadenatella had 8 oral 
lobes, one accessory sucker and one testis; 
and Jeancadenatia had 10 oral lobes, nu- 
merous accessory suckers, and probably one 
testis which Dollfus misinterpreted as 2 in 
his macerated specimens. Nagaty (1948) 
followed Manter’s (1947) suggestion and 
raised the subgenera to generic rank. 

Winter (1957) described Jeancadenatia 
dohenyt from Kyphosus elegans in the Mexi- 
can Pacific; it has 10 oral lobes, one testis, 
and only 2 accessory suckers. Sogandares- 
Bernal (1959) identified as J. brumpti 2 
specimens from K. sectatrix at Bimini, each 
with but one testis. Manter (1949) de- 
scribed Cadenatella americana as having a 
cirrus sac but reexamination of the holotype 
reveals that the pars prostatica probably was 
misinterpreted as a cirrus sac. The presence 
of accessory suckers and absence of a cirrus 
sac seems to be correlated in these trema- 
todes, as in the opecoelids. 

We believe that the presence of accessory 
suckers is a generic character whereas their 


Tulane Studies in Zoology 


Vol. 11 


number distinguishes species; the same is 
concluded for the oral lobes some of which 
may be more or less distinctly subdivided in 
some species and not in others. Jeancadenatia 
thus is considered a synonym of Cadenatella 
which has page priority. It includes the fol- 
lowing species: 
C. cadenati Dollfus, 1946, type species 
C. americana Manter, 1949 
C. brumpti (Dollfus, 1946) n.comb. 
Syn. Enenterum (Jeancadenatia) 
brumpti Dollfus 
Jeancadenatia brumpti (Doll- 
fus) Nagaty, 1948 
C. dohenyi (Winter, 1957) n.comb. 
Syn. Jeancadenatia dohenyt Winter 
C. Kkyphosi n.sp. 


Cadenatella kyphost is most like C. brump- 
tz but differs from that species in the ar- 
rangement of the accessory suckers; number 
size and shape of the oral lobes; shorter pre- 
pharynx; and smaller size. 


Diploproctodaeum plicitum (Linton, 1928) 
Sogandares-Bernal & Hutton, 1958 


Synonyms: Distomum sp. of Linton, 1898 
and 1905; Psilostomum plicitum Linton, 
1928; Bianium concavum Stunkard, 1930; 
Bianium adplicitum Manter, 1940; Bianinm 
plicitum (Linton) Stunkard, 1931. 

Hosts: Spheroides spengleri (J); *S. tes- 
tudineus (J). 

Site: intestine. 


Diploproctodaeum baustrum (MacCallum, 
1919) La Rue, 1926 

Synonyms: Hemistomum haustrum Mac- 
Callum, 1919; Bianium lecanocephalum Pé- 
rez Vigueras, 1955. 

Host: *Cantherines pullus (C, J). 

Site; intestine. 

The ovary in this species is variable. Mac- 
Callum (1919) and Pérez Vigueras (1955b) 
show an entire or subtriangular ovary. The 
majority of the specimens in our collection 
show the ovary in various degrees of lobula- 
tion with a maximum of 5 lobes. Dr. 
Sogandares (personal communication) con- 
firmed this variation in his material from 
Bimini. In many specimens, a few vitelline 
follicles extend to the mid- or anterior level 
of acetabulum. 


No. 4 


Diploproctodaeum diodontis n.sp. 
Figure 24 
Host: Diodon hystrix (J). 
Site: intestine. 
Holotype: US.N.M. 60272. 


Description based on 10 mature specimens. 
Body discoid, ventral surface concave, in- 
rolled at sides, 0.965-1.74 long, 0.887-1.39 
wide. Entire cuticle spinose; eye-spot pig- 
ment present. Oral sucker well removed 
from anterior end of body, 0.135-0.200 long, 
0.180-0.270 wide; ventral sucker equatorial 
to somewhat postequatorial, 0.180-0.266 in 
diameter; sucker ratio 1:1-1.2. Prepharynx 
absent; pharynx massive, 0.120-0.150 long, 
0.140-0.195 wide, usually with 8 anterior 
lobes; esophagus absent; ceca wide, arching 
from pharynx and extending posteriorly to 
converge at sides of posterior testis where 
each opens dorsally at an anus far from pos- 
terior end of body. Testes usually diagonal, 
rarely tandem, 0.120-0.233 long, 0.080-0.150 
wide; cirrus sac extending to near midlevel 
of ventral sucker, usually to left of that 
sucker but may be displaced to right, con- 
taining spherical internal seminal vesicle, 
pars prostatica of about same size, and cir- 
rus; external seminal vesicle prominent. 
Ovary with 10-15 distinct lobes, to left of 
midline, usually opposite anterior testis but 
may be intertesticular; seminal receptacle 
antero- or laterodorsal to ovary; uterus short, 
pretesticular, terminating in muscular me- 
traterm. Genital atrium wide; genital pore 
ventral, to left of midline, immediately pos- 
terior to intestinal bifurcation. Vitellaria of 
numerous follicles extending in a wide, more 
or less circular zone from oral sucker to near 
posterior end of body; confluent or not at 
level of intestinal bifurcation. Eggs 53-62 
by 30-45 py. Excretory vesicle somewhat sig- 
moid, slightly overreaching ventral sucker; 
excretory pore mid-dorsal, between anal 
openings. 

The discoid body shape and dorsal anal 
openings well removed from the posterior 
end of the body distinguish this species from 
all others in the genus Diploproctodaeum. 
Except for anal openings, it is very similar 
to species of Pseudocreadium with which it 
could easily be confused. Because both gen- 
era have all the characteristics of the Lepo- 
creadiidae, there is no justification for main- 
taining the Diploproctodaeidae Ozaki, 1928, 


Trematodes of Marine Fishes 


193 


or Dermadenidae Yamaguti, 1958, as fami- 
lies distinct from the Lepocreadiidae. 

Although Yamaguti (1958) gives in his 
key a number of features to distinguish the 
genera Pseudocreadium Layman, 1930, and 
Hypocreadium Ozaki, 1936, descriptions of 
their species show that differences, even at 
the generic level, are relative rather than 
absolute. Thus Yamaguti has referred P. 
scaphosomum Manter, 1940, to Hypocread- 
ium but retained P. lamelliforme (Linton, 
1907 ) in the genus Psewdocreadiuwm when the 
only difference between the 2 species seems 
to be the shape of the external seminal 
vesicle. Bravo-Hollis and Manter (1957) 
accepted the 2 genera “on the basis of an 
intertesticular ovary and the uterus extend- 
ing posterior to the ovary.” In the new spe- 
cies described below, the ovary is between 
the testes but the uterus does not extend pos- 
terior to it. Because Sogandares-Bernal 
(1959) observed variations in the position 
of the ovary and posterior extent of the 
uterus in his specimens of P, scaphosomum, 
he considered Hypocreadium a synonym of 
Pseudocreadium. We agree with that opin- 
ion. 


Pseudocreadium \actophrysi n.sp. 
Figures 25 and 26 

Synonym: Pseudocreadium sp. Siddiqi & 
Cable, 1960. 

Hosts: Lactophrys tricornis (C); L. tri- 
gonus (C); L. triqueter (C, J). 

Site: intestine. 

Holotype: US.N.M. 60273. 


Description based on 25 mature specimens. 
Body broadly pyriform to almost circular, 
0.333-0.667 long, 0.280-0.710 wide. Entire 
cuticle spinose; spines partially lost in some 
specimens; eye-spot pigment present. Oral 
sucker subterminal, 0.033-0.078 long, 0.053- 
0.083 wide; ventral sucker subequatorial, 
0.056-0.102 in diameter; sucker ratio 1:1.0- 
1.3. Prepharynx absent; pharynx 0.033- 
0.068 in diameter; esophagus about same 
length as pharynx; ceca arching to enclose 
reproductive system, ending about midway 
between testes and posterior end of body. 
Testes symmetrical, irregular, 0.083-0.165 in 
diameter; cirrus sac to right, not extending 
posterior to midlevel of ventral sucker, con- 
taining large internal seminal vesicle, bipar- 
tite pars prostatica and relatively long cirrus. 
External seminal vesicle an elongated sac 


194 Tulane Studies in Zoology 


overlapping Cirrus sac posterodorsally. Ovary 
irregular to trilobed, 0.045-0.090 long, 0.060- 
0.120 wide, immediately posterior to ventral 
sucker, median or rarely submedian; seminal 
receptacle large, ovoid, to left of ventral 
sucker; uterus short, not extending posterior 
to Ovary; metraterm about half length of cir- 
rus sac. Vitelline follicles numerous, extend- 
ing from level of oral sucker to posterior end 
of body, confluent dorsally at intestinal bi- 
furcation and posterior to ovary. Eggs few, 
58-68 by 33-45 p. Genital pore to left of 
midline, at level of, or immediately pos- 
terior to intestinal bifurcation. Excretory 
vesicle tubular, extending to posterior edge 
of ovary; excretory pore dorsal, far removed 
from posterior end of body. Siddiqi and 
Cable (1960) reported the flame cell form- 
ula for their immature specimen to be 
eee) ye ra 

Manter (1945) pointed out that Linton 
(1907) confused 2 species as Distomum 
lamelliforme. Linton’s Figure 75 probably 
is P. lactophryst; thus 3 rather than 2 spe- 
cies may have been misinterpreted as a 
single one by Linton. 

The broadly pyriform body in combination 
with a uterus that does not extend posterior 
to the ovary distinguish Pseudocreadium 
lactophryst from all species of Psewdocread- 
mm except P. spinosum Manter, 1940. A 
comparison of the present material with 
Manter’s description and 3 specimens of P. 
spinosum reveals the following: P. lacto- 
phrysi is a smaller species but its measure- 
ments overlap those of P. spimoswm; the 
anterior end is somewhat pointed rather than 
truncated; a prepharynx is absent; the testes 
are more anterior and the seminal receptacle 
is ovoid rather than tubular. 


Pseudocreadium anandrum Manter, 1947 


Hosts: *Calamus arctifrons (J); *C. ba- 
janado (J). 
Site; intestine. 


Pseudocreadinm lamelliforme (Linton, 
1907 ) Manter, 1946 
Synonym: Distomum lamelliforme Linton, 
1907. 
Host: Balistes vetula (J). 
Site; intestine. 


Pseudocreadium galapagoensis Manter, 
1946 
Synonym: Pseudocreadinm scaphosomum 
Manter, 1940 in part. 


Vol. 11 


Host: *Balistes ringens (C). 

Site; intestine. 

Sogandares-Bernal (1959) described Psew- 
docreadium biminensis from 2 specimens 
and indicated a close resemblance to P. 
galapagoensis. For his species, he gave 5 
distinguishing features which, except for the 
anterior extent of the vitellaria, are variable 
in our 17 specimens from Curagao. Sucker 
ratios are intermediate, being 1:0.70-1.00 
compared with 1:0.51-0.89 for P. biminen- 
sis and 1:1-1.13 for P. galapagoensis; the 
external seminal vesicle is usually median 
and transverse but lateral and diagonal in 
a few of our specimens. In the posterior 
extent of the cirrus sac, the Curacao material 
is like P. galapagoensis, but in the position 
of the ventral sucker, it is like P. biminenstis. 
In all 17 specimens, however, the vitellaria 
extend to the mid- or anterior level of the 
oral sucker and thus provide the only dis- 
tinguishing feature between the 2 species. 
Dr. Manter examined one of our specimens 
and verified its identification. Minute spines 
which were observed on the cirrus in living 
material were not reported by Manter but 
were seen in a paratype provided by him. 
The excretory vesicle is pyriform, with its 
narrower anterior portion receiving 2 canals 
which extend slightly anterior to the ventral 
sucker and divide into anterior and posterior 
secondary tubules, each joined by 2 large 
groups of flame cells. 


Dermadena lactophrysi Manter, 1946 
Synonym: Distomum lamelliforme Linton, 
1907 in part. 
Hosts: Lactophrys tricornis (C); 
gonus (C); L. triqueter (C). 
Site: intestine. 


L. trt- 


Superfamily Opecoeloidea Cable, 1956 
FAMILY OPECOELIDAE Ozaki 1925 
Hamacreadium oscitans Linton, 1910 

Synonyms: Podocotyle breviformis Man- 
ter, 1940; **Pseudoplagioporus brevivitellus 
Siddiqi & Cable, 1960. 

Hosts: Antsotremus Opie 
*Archosargus unimaculatus (J); *Bathy- 
stoma aurolineatum (J); *Calamus cala- 
mus (J); *Haemulon album (C); *H. 
bonariense (J); *H. melanurum (C); H. 
scmurus (J). 

Site: intestine. 


vir ginicus 


No. 4 


Our specimens from the various hosts 
show that the anterior extent of the excretory 
vesicle depends on the degree of maturity 
of the trematodes; it reaches the posterior 
margin of the acetabulum in the immature 
specimens and only to the ovarian level in 
mature ones. Reexamination of a paratype 
of Pseudoplagioporus brevivitellus indicates 
that species to be a synonym of H. oscitans. 


Hamacread‘um mutabile Linton, 1910 

Hosts: Lutianus apodus (J); L. griseus 
Ops Lojocw (J). 

Site: intestine. 


Hamacreadium consuetum Linton, 1910 
Host: Haemulon scturus (J). 
Site; intestine. 


Helicometrina nimua Linton, 1910 
Hosts: Haemulon sciurus (J); *Hypo- 
plectrus indigo (J); *Lachnolaimus maxt- 
mus (J); *Lutianus jocu (J); *Platophrys 
lunatus (J); *Spheroides spengleri (J). 
Site: intestine. 


Helicometrina trachinoti Siddiqi & Cable, 
1960 

Host: *Trachinotus glaucus (J). 

Site; intestine. 

Six specimens from one fish agree with 
the original description of the species ex- 
cept that in all of them, the cirrus sax ex- 
tends to the posterior margin of the ventral 
sucker. 


Helicometra equilata (Manter, 1933) 
Siddiqi & Cable, 1960 
Synonym: Stenopera equilata Manter, 
1933: 
Host: Holocentrus ascenstonis (J). 
Site: intestine. 


Helicometra execta Linton, 1910 
Host: *Halichoeres pictus (J). 
Site: intestine. 


Horatrema crassum Manter, 1947 
Synonym: Manteriella crassa (Manter) 
Yamaguti, 1958. 
Hosts: Eques acuminatus (J); *E. punc- 
tatus (C). 
Site: intestine. 


Pinguitrema lobatum Siddiqi & Cable, 
1960 
Host: Gerres cinereus (J). 
Site: intestine. 


Trematodes of Marine Fishes 


195 


Pseudopecoeloides carangi (Yamaguti, 
1938) Yamaguti, 1940 
Synonym: Cymbephallus carangt Yama- 
guti, 1938. 
Hosts: 
coe 


Site: intestine. 


(C); C. ruber 


*Caranx crysos 


Pseudopecoeloides equesi Manter, 1947 

Hosts: Eques acuminatus (C); *E. punc- 
tats (C). 

Site; intestine. 


Pseudopecoeloides gracilis Manter, 1947 
Host: Selar crumenophthalmus (J). 
Site: intestine. 


Neopecoelus scorpaenae Manter, 1947 
Host: Scorpaena plumieri (C). 
Site: intestine. 


Our material is identified as this species 
even though the anal openings could not be 
confirmed by careful examination of living 
specimens or whole mounts. In some stained 
specimens, strands were seen extending pos- 
teriorly from the end of each cecum. In 
other respects, there is close agreement with 
the original description of the species ex- 
cept that interruption of the vitellaria at 
the level of each testis occurred in a minor- 
ity of our specimens. 


Opecoeloides brachyteleus Manter, 1947 

Hosts: Upeneus maculatus (C, J); U. 
martinicus (C, J). 

Site: intestine. 


Opecoeloides elongatus Manter, 1947 
Hosts: Upeneus maculatus (C, J); U. mar- 
timicus (C, J). 


Opecoeloides vitellosus (Linton, 1900 ) 
Von Wicklen, 1946 

Synonyms: Distomum vitellosum Linton, 
1900; Anisoporus mantert Hunninen & 
Cable, 1940; Opecoeloides manteri (Hunni- 
nen & Cable) Hunninen & Cable, 1941; 
*Cymbephallus vitellosus (Linton) Linton, 
1934. 

Host: *Epinephelus adscenstonis (J). 

Site: intestine. 

Von Wicklen (1946) pointed out that 
Linton had identified as Distomum vitel- 
losum more than one species and that Ope- 
coeloides vitellosus should be restricted to 
trematodes that agree with the descriptions 
given originally by Linton (1900) and later 
by Hunninen and Cable (1941). 


196 


Pseudopecoelus barkeri Hanson, 1950 
Hosts: Holocentrus ascensionis (C, J); 
*H. vexillarius (C). 
Site: intestine. 


According to the key to the genus Psez- 
dopecoelus given by Manter (1954), our 
specimens could be either P. barkeri Han- 
son, 1950, or P. tortugae (Manter, 1934). 
In general body shape and indented testes, 
they are more like P. tortugae but egg meas- 
urements (45-52 by 27-32 ») are more like 
those of P. barkeri. The sucker ratio is in- 
termediate (1:1.7-2.3). Thus the 2 species 
seemingly differ only in egg size. 


Pseudopecoelus holocentri n.sp. 
Figure 27 
Synonym: **Pseudopecoelus elongatus of 
Hanson, 1950, nec (Yamaguti, 1938) 
Host: Holocentrus ascensionis (C). 
Site: intestine. 


Holotype: US.N.M. 60274. 


Description based on 2 specimens. Body 
slender, 2.45-3.28 long, 0.600-0.667 wide. 
Oral sucker 0.120-0.130 long, 0.128-0.135 
wide; ventral sucker in anterior fourth of 
body, 0.240-0.280 in diameter, without papil- 
lae; sucker ratio 1:2-2.15. Prepharynx short; 
pharynx 0.105 long, 0.090-0.114 wide; 
esophagus 0.098-0.112 long; ceca end blind- 
ly near posterior end of body. Testes 2, tan- 
dem, lobed, not contiguous, 0.293-0.346 long, 
0.213-0.273 wide; seminal vesicle tubular, 
extending about 1/3 distance from ventral 
sucker to ovary; ejaculatory duct short; pars 
prostatica indistinct. Ovary pretesticular, 
slightly irregular, 0.133-0.166 in diameter; 
uterine seminal receptacle, Mehlis’ gland and 
uterus preovarian; metraterm well-developed. 
Genital pore sinistral, near anterior margin 
of pharynx. Eggs 52-54 by 27-30 p. Vitel- 
line follicles extending from near posterior 
level of ventral sucker to the posterior end 
of body, interrupted opposite gonads. Excre- 
tory vesicle extending to ovarian level; pore 
terminal. 

The only other species of Psewdopecoelus 
with vitellaria interrupted opposite the gon- 
ads is P, elongatus (Yamaguti, 1938). That 
species differs from P. holocentri in having 
a smaller pharynx, a different sucker ratio 
(1:1.56 compared with 1:2-2.15) and a 
longer posttesticular space. 

Some of the specimens which Hanson 
(1950) reported as P. elongatus have been 


Tulane Studies in Zoology 


Voli 


examined and found to agree with the pres- 
ent species except in having somewhat less 
irregular gonads. 


Pseudopecoelus gymnothoracis n.sp. 
Figure 28 

Host: Gymnothorax moringa (C). 

Site: intestine. 

Holotype: U.S.N.M. 60275. 

Description based on 8 specimens. Body 
pyriform to linguiform, 1.40-1.85 long, 
0.637-0.830 wide. Cuticle smooth. Oral 
sucker 0.113-0.133 long, 0.120-0.153 wide. 
Ventral sucker 0.200-0.233 long, 0.233-0.273 
wide; sucker ratio 1:1.7-2.1. Prepharynx 
short; pharynx 0.038-0.060 in diameter; 
esophagus 3-5 times length of pharynx; in- 
testinal bifurcation about midway between 
suckers; ceca converge posteriorly, ending 
blindly at about midlevel of posttesticular 
space. Testes 2, extremely lobed, constricted 
medially, 0.113-0.266 long, 0.333-0.440 wide. 
Cirrus sac absent; seminal vesicle tubular, 
extending well posterior to acetabulum but 
not reaching ovary; pars prostatica weakly 
developed, prostate cells few; ejaculatory duct 
short. Ovary lobed, pretesticular, submedian, 
0.100-0.120 long, 0.173-0.200 wide; uterine 
seminal receptacle, Mehlis’ gland and uterus 
preovarian; Laurer’s canal present; metra- 
term well-developed. Genital atrium small, 
genital pore sinistral, at about midesophageal 
level. Eggs 54-69 by 30-45 p, usually 63-67 
by 37-42. Vitelline follicles numerous, ex- 
tending along entire length of ceca, con- 
fluent at intestinal bifurcation. Excretory 
vesicle tubular, extending to ovarian level. 

The combination of highly lobed and me- 
dially constricted testes with vitelline fol- 
licles broadly confluent at the intestinal bi- 
furcation distinguishes this species from all 
others in the genus Psewdopecoelus. 


Pseudopecoelus minutus n.sp. 
Figure 29 
Host: Doratonotus megalepis (C). 
Site: intestine. 


Holotype: U.S.N.M. 60276. 


Description based on 6 specimens, meas- 
urements on 4 mature ones. Body elongated, 
rounded at both ends, 0.606-0.720 long, 
0.180-0.233 wide. Oral sucker 0.063-0.075 
in diameter; ventral sucker at junction of 
anterior and middle third of body, 0.105- 
0.135 in diameter; sucker ratio 1:1.7-2.0. 


No. 4 Trematodes of Marine Fishes 197 


Figure 24. Diploproctodaeum diodontis, holotype, dorsal view. Figure 25. Pseudo- 
creadium lactophrysi, holotype, ventral view. Figure 26. Same, cirrus sac of another 
specimen drawn free-hand. Figure 27. Pseudopecoelus holocentri, holotype, dorsal view. 
Figure 28. Pseudopecoelus gymnothoracis, holotype, ventral view. Figure 29. Pseudo- 
pecoelus minutus, holotype, ventral view. 


198 


Prepharynx absent; pharynx 0.030-0.040 
long, 0.042-0.048 wide; esophagus 1-2 times 
length of pharynx; ceca end blindly a short 
distance from posterior end of body. Testes 
2, tandem, entire, contiguous, 0.045-0.120 
in diameter; seminal vesicle long, reaching 
about midway between ventral sucker and 
Ovary; pars prostatica not evident. Ovary 
pretesticular, subtriangular, smooth, 0.030- 
0.075 in diameter; Mehlis’ gland, uterine 
seminal receptacle and uterus preovarian; 
metraterm well-developed. Genital atrium 
muscular; genital pore sinistral, near pos- 
terior margin of pharynx. Eggs few, 45-54 
by 22-30 p». Vitelline follicles extending an- 
terior to ventral sucker, usually interrupted 
lateral to acetabulum. Excretory vesicle tubu- 
lar, extending to level of ovary; excretory 
formula 2{(2+2) + (2+2)} = 16 flame 
cells. 

Only Pseudopecoelus gibbonsiae Manter 
and Van Cleave, 1951, shares with this spe- 
cies the combined features of an entire ovary 
and vitellaria that extend anterior to the ace- 
tabulum. However, P. gibbonsiae is much 
larger (2.26-2.55 by 0.643-0.780) and its 
eggs almost twice the size of those of P. 
minutus, 

The pygmy wrasse which harbors this spe- 
cies lives in clumps of rockweed in close 
association with the snail, Columbella mer- 
catoria, and an amphipod which probably 
serve as intermediate hosts. A minute ope- 
coelid cercaria, to be described elsewhere, 
develops in that snail and was observed to 
penetrate and encyst in the amphipod. 


Coitocaecum sp. 
Figure 30 
Host: Labrisomus bucciferus (J). 


Site; intestine. 
Deposited specimen: U.S.N.M. 60277. 


Description based on a single specimen. 
Body elongated, rounded at both ends, 0.750 
by 0.267. Cuticle smooth. Oral sucker 0.067 
by 0.084; ventral sucker subequatorial, 0.126 
by 0.150, with transverse aperture; sucker 
ratio 1:1.84. Prepharynx short; pharynx 
0.045 by 0.065; esophagus 0.115 long; in- 
testinal bifurcation about midway between 
pharynx and acetabulum; cyclocoel gut ex- 
tending to near posterior end of body. Testes 
tandem, contiguous, 0.045-0.050 long, 0.065- 
0.072 wide; seminal vesicle pyriform, pre- 
acetabular; followed by a narrow duct lead- 


Tulane Studies in Zoology 


Vol. 11 


ing to inconspicuous Cirrus sac anterior to 
arch of left cecum; content of cirrus sac not 
evident; genital pore sinistral, at esophageal 
level. Ovary pretesticular, slightly dextral, 
0.045 by 0.065. Mehlis’ gland preovarian; 
seminal receptacle absent; uterus not extend- 
ing tO posterior testis; metraterm simple. 
Eggs collapsed, 48-56 by 27-33 p. Vitellaria 
extending from esophageal level to posterior 
end of body, confluent in posttesticular space 
but not at intestinal bifurcation. 

This species is described and figured but 
not named because only one specimen in 
poor condition was found. 


FAMILY OPISTHOLEBETIDAE 
Fukui, 1929 
O pistholebes diodontis Cable, 1956 
Host: Diodon hystrix (C, J). 
Site: intestine. 


Pachycreadinm crassigulum (Linton, 1910) 
Manter, 1954 

Synonyms: Lebourta crassigula Linton, 
1910; Plagioporus crassigulus (Linton) Price, 
1934. 

Hosts: *Calamus arctifrons (J); C. baja- 
nado (J); *Archosargus unimaculatus (J). 

Site: intestine. 


Superfamily Allocreadioidea Nicoll, 1934 
FAMILY GORGODERIDAE Looss, 1901 


Xystretrum solidum Linton, 1910 

Synonyms: Catoptroides aluterae MacCal- 
lum, 1917; Catoptroides magnum MacCal- 
lum, 1917; Macia pulchra Travassos, 1921; 
Xystretrum pulchrum (Travassos) Manter, 
1947; Xystretrum papillosum Linton, 1910. 

Hosts: *Balistes vetula (J); *Cantherines 
pullus (J); *Canthigaster rostratus (C); 
*Lactophrys tricornis (J); Spheroides tes- 
tudineus (J). 

Site: urinary bladder and kidney ducts. 


The effect of crowding on the size of this 
species, mentioned by Manter (1947), was 
shown by the more than 100 specimens with 
which the kidney ducts and bladder of one 
Canthigaster rostratus were literally stuffed. 


Phyllodistomum pomacanthi n.sp. 
Figure 31 
Host: Pomacanthus arcuatus (J). 
Site: posterior intestine. 
Holotype: U.S.N.M. 60278. 
Description based on a single specimen. 
Body foliate, sides inrolled ventrally, 3.28 


No. 4 


long, 1.49 in maximum width, at level of 
posterior testis. Oral sucker 0.334 by 0.374; 
ventral sucker slightly preequatorial, 0.267 
by 0.240; sucker ratio 1:0.71. Pharynx ab- 
sent; esophagus 0.188 long; ceca wide, ex- 
tending almost to excretory pore. Testes 2, 
diagonal, intercecal, lobed, separated by coils 
of the uterus; anterior testis 0.266 by 0.334, 
to left of midline; posterior testis 0.280 by 
0.306, 0.70 from posterior end of body; 
seminal vesicle anterior to genital pore, 
globular, wall poorly defined; prostate cells 
free in parenchyma; ejaculatory duct short, 
curves from seminal vesicle posteriorly to 
enter thick-walled genital atrium; genital 
pore about midway between intestinal bifur- 
cation and acetabulum. Ovary pretesticular, 
indented or irregular, to right of midline, 
0.240 by 0.200; seminal receptacle absent; 
Mehlis’ gland between vitellaria; uterus ex- 
tending to near ends of ceca; metraterm 
well-developed. Eggs few, 24-36 by 14-23 yp. 
Vitellaria 2 lobed masses, close together but 
separated by uterus, symmetrically placed, a 
short distance posterior to ventral sucker. 
Excretory vesicle not evident; pore dorsal, 
some distance from posterior end of body. 

Phyllodistomum pomacanthi is most like 
P. carangis Manter, 1947, but differs from 
that species chiefly in being much smaller, 
and in having lobed testes, a more posterior 
ventral sucker and ceca extending farther 
posteriorly. 


FAMILY ZOOGONIDAE Odhner, 1911 

The present study includes species belong- 
ing to the genera Diplangus, Deretrema, 
Steganoderma and Diphtherostomum. Skrja- 
bin (1957) has placed all but Diphtherosto- 
mum in the family Steganodermatidae Doll- 
fus, 1952, in which the vitellaria are more 
extensive and the eggs thicker-shelled than 
in the Zoogonidae. He also erected the 
superfamily Zoogonoidea to include those 
families. Yamaguti (1958) assigned Dji- 
plangus to the family Callodistomidae Poche, 
1926, but left the others in the Zoogonidae. 
Until those arrangements can be evaluated 
on the basis of life history studies, we prefer 
to leave all 4 genera in the Zoogonidae and, 
tentatively, in the superfamily Allocreadi- 
oidea. 

Diplangus paxillus Linton, 1910 

Hosts: Anisotremus virgintcus (J); Hae- 
mulon scturus (J); *Gerres cinereus (J). 

Site; intestine. 


Trematodes of Marine Fishes 


199 


Diplangus parvus Manter, 1947 
Host: *Haemulon sciurus (J). 
Site: intestine. 


Deretrema fusillum Linton, 1910 
Host: *Mycteroperca bonact (C). 
Site: intestine. 


Steganoderma nitens (Linton, 1898) 
Manter, 1947 

Synonyms:  Distomum nitens Linton, 
1898; Lecithostaphylus nitens (Linton) Lin- 
ton, 1940; **Steganoderma elongatum Man- 
ter, 1947. 

Host: *Strongylura ardeola (C). 

Site; intestine. 


5 


Variations in our 3 specimens overlap 
features used by Manter (1947) to separate 
Steganoderma elongatum from S. nitens. 
Reexamination of type specimens shows 
that the genital pore in S. nitens is more 
lateral than Linton (1898) figured and 
that its relatively far anterior position in 
S. elongatum is due to contraction of the 
forebody. 

A striking feature of this species, not 
mentioned by either Linton or Manter, is 
the presence of conspicuous glands occupy- 
ing the full width of the body from near the 
vitellaria to a short distance anterior to the 
acetabulum. They were seen in Manter’s 
type specimen but could not be recognized 
with certainty in Linton’s because of its con- 
dition. On each side, anterior to those 
glands, is a group of several less conspicuous 
ones with ducts extending anteriorly in a 
distinct bundle to separate and open at 
clusters of pores on the anterior margin of 
the oral sucker. 


Steganoderma hemiramphi Manter, 1947 
Figure 32 

Hosts: Hemiramphus brasiltensis (C, J); 
Gerres cinereus (J). 

Site: intestine. 

That the larva of this species is an oph- 
thalmoxiphidiocercaria is evident from the 
presence of eye-spot pigment in the forebody 
and a stylet (Fig. 32) in the oral sucker of 
one of 2 specimens from Curagao. In known 
zoogonid life histories, the larva has a stylet 
but no tail or eye-spots. In the Monorchiidae, 
cercariae with eye-spots have a well de- 
veloped tail whereas the absence of eye-spots 
is usually accompanied by more or less re- 
duction of the tail. Should that situation 


200 


apply to the Zoogonidae, the life history of 
Steganoderma hemiramphi could be decisive 
in determining the affinity of its family to 
others, a matter that is still obscure. 


Steganoderma atherinae (Price, 1934) 
Manter, 1947 
Synonym: Lecithostaphylus atherinae Price, 
1934. 
Hosts: Hepsetia stipes (J); *Strongylura 
timucu (C). 
Site: intestine. 


The single specimen from Curacao, al- 
though from a new host, is in close agree- 
ment with the species as described else- 
where in the Caribbean region. However, 
none of many individuals of Hepsetia stipes 
examined in Curacao harbored this species 
whereas it was found in almost all H. stipes 
in Jamaica. 


Diphtherostomum anisotremi n.sp. 
Figure 33 
Host: Antsotremus virginicus (J). 


Site; intestine. 
Holotype: U.S.N.M. 60279. 


Description based on 8 specimens. Body 
plump, tapering toward both ends, 0.440- 
0.767 long, 0.173-0.267 wide. Cuticle of 
forebody with large spines; hindbody smooth. 
Oral sucker 0.060-0.090 in diameter; ventral 
sucker equatorial to slightly postequatorial, 
somewhat quadrangular, 0.120-0.210 in di- 
ameter; sucker ratio 1:2-2.35. Prepharynx 
absent; pharynx 0.026-0.037 in diameter; 
esophagus 3-4 times length of pharynx; ceca 
not quite reaching midlevel of acetabulum. 
Testes 2, symmetrical to diagonal, immedi- 
ately posterior to or overlapping posterior 
margin of ventral sucker; cirrus sac arcuate, 
elongated, 0.195-0.233 long, 0.039-0.060 
wide, extending to and sometimes over- 
lapping anterior margin of acetabulum; con- 
taining bipartite seminal vesicle, ovoid pars 
prostatica, prostate cells and spiny cirrus. 
Genital pore sinistral, just posterior to level 
of intestinal bifurcation. Ovary smooth, dor- 
sal to acetabulum, 0.064-0.105 long, 0.050- 
0.083 wide; seminal receptacle postovarian; 
uterus occupying most of hindbody; metra- 
term muscular, spiny, about same length as 
cirrus sac. Eggs very thin-shelled, 27-36 by 
9-14 w. Vitellaria in 2 masses, 0.030-0.078 
in diameter, near posterior margin of ace- 
tabulum. Excretory vesicle short, sac-shaped; 
pore terminal. 


Tulane Studies in Zoology 


Vol. 11 


This species hesitantly is reported as new 
because of its similarity to Dzphtherosto- 
mum americanum. However, Manter (1947) 
described that species as having a very short 
esophagus and a cirrus sac with a width of 
1/2-3/4 its length whereas in D. anisotremi, 
the esophagus is at least 3 times as long as 
the pharynx and the cirrus sac is about 4 
times as long as wide. The figure of a 
trematode identified by Sogandares-Bernal 
and Hutton (1959b) as D. americanum 
shows an elongated cirrus sac but a short 
esophagus. 


FAMILY MONORCHIIDAE Odhner, 
1Ovs 

The status of Genolopa Linton, 1910, 
Proctotrema Odhner, 1911, Lasiotocus Looss, 
1907, and other related genera has been re- 
viewed by many authors including Yama- 
gutt (1934), Hopkins (1941), Manter 
(1942), Nagaty (1948) and more recently 
by Thomas (1959) and Manter and Pritch- 
ard (1961). We accept the genus Genolopa 
for species with atrial spines as suggested by 
Manter and Pritchard. These authors also 
suggested that Laszotocus be separated from 
Proctotrema on the basis of an entire ovary 
versus a 3- or 4-lobed one. In some trema- 
todes this character is variable. In the hemi- 
urid, Dichadena acuta, for instance, the 
ovary may be entire or distinctly 4-lobed. 
Moreover, lobation may be a matter of 
degree which can vary with handling of 
specimens or with their age. However, our 
material can be allocated between Lasiotocus 
and Proctotrema as distinguished by Manter 
and Pritchard and, for that reason, the va- 
lidity of both genera is accepted at this 
time. Actually Lasiotocus was never pub- 
lished by Looss as a formal name; instead it 
was mentioned in a subjunctive sense in 
criticizing the Rules of Nomenclature. How- 
ever facetious the intent of Looss may have 
been, the Law of Priority establishes the 
validity of such names. Thus Lasiotocus 
would take priority over Proctotrema if 
those genera are considered to be synony- 
mous. 


Genolopa ampullacea Linton, 1910 

Synonym: **Genolopa longicaudata Sid- 
digi & Cable, 1960. 

Hosts: Bathystoma striatum (J); Hae- 
mulon album (J); *H. bonariense (J); H. 
flavolineatum (C, J); *H. melanurum (C); 
H. sciurus (J). 


No. 4 


Site: ceca and intestine. 

Siddiqi and Cable (1960) described Geno- 
lopa longicaudata from Odontoscion dentex 
and, in a key, distinguished it from G. am- 
pullacea on the basis of having a post- 
testicular space “3 or 4 times length of tes- 
tis” and a metraterm sac “reaching well pos- 
terior to ventral sucker.” Our more abundant 
material shows that these features are highly 
variable; G. longicaudata accordingly is re- 
duced to synonymy with G. ampullacea. 


Genolopa brevicaecum (Manter, 1942 ) 
Manter and Pritchard, 1961 

Synonym: Paraproctotrema brevicaecum 
Manter, 1942. 

Host: Caranx bartholomaei (J). 

Site: intestine. 

Seventeen specimens from 2 fish are in 
close agreement with Manter’s description 
and measurements. The majority of the 
worms are elongate, spindle-shaped but a 
few are pyriform. They also confirm the 
presence of the atrial spines reported by 
Manter and Pritchard (1961). In most 
specimens, the spines were difficult to dis- 
tinguish from those on the cirrus but 2 
worms with that organ retracted, show a 
ring of spines around the genital atrium. 
The metraterm is unspined. 


Lastotocus longicaecum (Manter, 1940) 
Yamagut, 1953 
Synonym: Proctotrema longtcaecum Man- 
ter, 1940. 
Host: Antsotremus virginicus (J). 
Site: ceca and intestine. 


Lastotocus truncatus (Linton, 1910) 
Thomas, 1959 
Synonyms: Genolopa truncatum Linton, 


1910; Proctotrema truncatum (Linton) 
Manter, 1940. 
Hosts: *Bathystoma aurolineatum (J); 


*Brachygenys chrysargyreus (C); *Cala- 

mus calamus (J); Haemulon album (C); 

*H. bonariense (J); H. flavolineatum (C, 

J); A. scturus (J); *Lutianus mahogoni (C). 
Site: ceca and intestine. 


Lasiotocus longovatus ( Hopkins, 1941) 
Thomas, 1959 
Synonyms: Genolopa longovatum Hop- 
kins, 1941; Proctotrema longovatum (Hop- 
kins) Manter, 1942. 
Hosts: *Bathystoma aurolineatum (J); 


Trematodes of Marine Fishes 


201 


*Haemulon bonariense (J); *H. flavoline- 
atum (C); *H. scwrus (J). 
Site: ceca and intestine. 


Seventy trematodes are referred to this 
species on the basis of egg size and other 
measurements, length of ceca and general 
topography of organs. We did not observe 
the urn-shape described by Hopkins (1941) 
in either living or mounted specimens. How- 
ever, Hopkins states (p. 401) “This is al- 
most certainly the same species as the speci- 
men shown in Figure 223 of Linton (1910) 
under the name ‘Monostomum sp..” Ob- 
viously, he was referring to Linton (1905) 
since Figure 223 of Linton’s 1910 paper 
represents a bucephalid. Our material is 
very similar to Linton’s in the shape of the 
body and oral sucker. 


Proctotrema pritchardae n.sp. 
Figure 34 
Host: Haemulon album (C). 
Site; intestine. 


Holotype: US.N.M. 60280. 


Description based on 2 specimens. Body 
elongated, rounded at both ends, 1.158 to 
1.22 long, 0.40 wide. Cuticular spines ex- 
tend along entire length of body; eye-spot 
pigment absent. Oral sucker somewhat 
funnel-shaped to spherical, 0.142-0.160 long, 
0.172-0.180 wide; ventral sucker in middle 
third of body, 0.105-0.113 long, 0.090-0.098 
wide; sucker ratio 1:0.60-0.64. Prepharynx 
short; pharynx 0.072-0.075 long, 0.075-0 083 
wide; esophagus about same length as phar- 
ynx; intestinal bifurcation about midway 
between pharynx and ventral sucker; ceca 
long, extending to near posterior end of 
body. Gonads in middle third of body. Tes- 
tis median, entire, 0.150-0.246 in diameter; 
cirrus sac 0.233-0.255 long, 0.105 wide, to 
right of midline, extending to posterior 
margin of ovary; containing spherical semi- 
nal vesicle, tubular pars prostatica and cirrus 
with small, inconspicuous spines. Ovary 
with 3 or 4 distinct lobes, to right of mid- 
line, immediately anterior to, or overlapped 
by testis. Metraterm sac 0.210-0.213 long, 
0.105-0.109 wide, consisting of large pos- 
terior vesicle without spines and smaller 
spinose anterior portion, separated by 
sphincter; metraterm spines distinct, about 
12 p» long, larger than those of cirrus. 
Uterus voluminous, mostly  posttesticular; 
distal end entering spiny portion of metra- 


202 


term sac just anterior to sphincter. Genital 
atrium unarmed, genital pore median, about 
midway between ventral sucker and intes- 
tinal bifurcation. Eggs numerous, 18-21 by 
10-12 w. Vitellaria in lateral groups of 9-10 
follicles each extending from anterior mar- 
gin of ovary to midlevel of testis. Excretory 
vesicle tubular, anterior extent not deter- 
mined; pore terminal. 

Manter and Pritchard (1961) recognize 
the following species of Proctotrema: P. 
bacilliovatum Odhner, 1911, P. macrorchis, 
Yamaguti, 1934, P. plectorhynchi Yama- 
guti, 1934, P. chaetodipteri (Thomas, 1959), 
P. himezi (Yamaguti, 1951), P. parvum 
Manter, 1942, and P. latwm (Manter, 1942). 
Proctotrema pritchardae differs from P. ba- 
cilliovatum, P. plectorhynchi, and P. macror- 
chis chiefly in having smaller eggs, longer 
ceca, and in the shape of the oral sucker; 
from P. chaetodipteri in body shape and 
length of posttesticular space; from P. himezi 
in having longer ceca; from P. parvum in 
having smaller eggs and much longer ceca, 
and from P. latum in body shape and ac- 
companying topography of internal struc- 
tures. 

The species is named in honor of Mrs. 
Mary Hanson Pritchard in recognition of her 
work in the field of trematodology. 


Proctotrema anisotremi n.sp. 
Figure 35 
Host: Anisotremus virginicus (J). 


Site: ceca and intestine. 
Holotype: U.S.N.M. 60281. 


Description based on 10 specimens. Body 
oval, 0.500-0.714 long, 0.267-0.400 in maxi- 
mum width, at level of vitellaria. Entire 
cuticle spinose, eye-spot pigment absent. 
Oral sucker 0.090-0.113 long, 0.105-0.135 
wide; ventral sucker somewhat preequa- 
torial, 0.055-0.067 long, 0.060-0.075 wide; 
sucker ratio 1:0.52-0.70. Prepharynx very 
short; pharynx subspherical, 0.038-0.051 in 
diameter; esophagus very short; ceca extend- 
ing just posterior to testis. Gonads in mid- 
dle third of body. Testis entire, to right of 
midline, 0.120-0.180 long, 0.099-0.155 wide; 
cirrus sac crescent-shaped, to right of mid- 
line, 0.140-0.195 long, 0.060-0.090 wide, ex- 
tending to posterior margin of acetabulum 
or slightly beyond, containing large spherical 
seminal vesicle, short tubular pars prostatica 
and cirrus with spines about 8 , long. 


Tulane Studies in Zoology 


Vol. 19 


Ovary immediately pretesticular, at level of 
ventral sucker, with 3 almost separate lobes; 
uterus filling posttesticular space and left 
side of hindbody, joining metraterm sac 
immediately posterior to sphincter. Metra- 
term sac smaller than cirrus sac, 0.113-0.160 
long, 0.045-0.075 wide, rarely extending 
posterior to ventral sucker; consisting of 
large, posterior vesicle without spines sepa- 
rated from a smaller anterior spinose division 
by a sphincter; metraterm spines slightly 
larger than those of cirrus. Genital atrium 
unarmed, genital pore median, about mid- 
way between acetabulum and intestinal bi- 
furcation. Eggs thin-shelled, 17-20 by 9- 
11 pw. Vitellaria in 2 lateral groups of 8 or 
9 follicles each, extending from about mid- 
level of acetabulum to that of testis. Ex- 
cretory vesicle not observed; pore terminal. 

Proctotrema anisotremi is to be compared 
with P. parvum and P. latum which have a 
more or less spherical oral sucker. It dif- 
fers from P. parvum in having smaller eggs, 
longer ceca and large cirrus sac, and in the 
position of the testis. Although described 
from the same host as P. anisotremi, P. la- 
tum differs in having a characteristically 
broad shape emphasized by Manter (1942), 
a proportionally larger cirrus sac extending 
posterior to ventral sucker, a more anterior 
genital pore, and longer ceca. Similarities 
include shape of oral sucker, position of tes- 
tis, and lobation and position of ovary. P. 
anisotremt differs from P. pritchardae in 
shape of ovary, in position of testis and in 
having shorter ceca. 


Chrisomom decapteri n.sp. 
Figures 36 and 37 
Host: Decapterus macarellus (C). 


Site: intestine. 
Holotype: US.N.M. 60282. 


Description based on 6 specimens. Body 
elongated, rounded at both ends, 0.900- 
1.50 long, 0.300-0.400 wide (one additional 
specimen without eggs measured 0.772 by 
0.220). Cuticular spines numerous to pos- 
terior edge of ventral sucker, then become 
sparse and shortly disappear. Eye-spot pig- 
ment present. Oral sucker transversely elon- 
gated, 0.045-0.063 long, 0.060-0.090 wide; 
ventral sucker about 1/3 body length from 
anterior end, 0.084-0.090 long, 0.054-0.084 
wide; sucker ratio 1:1-1.15. Prepharynx 
about half length of pharynx; pharynx 0.053- 


No. 4 Trematodes of Marine Fishes 


i) 
>) 
1S) 


Ta dbok 
9G.08: 
OU So 4 


i 
Baro 


pares 


Figure 30. Coitocaecum sp., dorsal view. Figure 31. Phyllodistomum pomacanthi, 
holotype, dorsal view. Figure 32. Stylet of Steganoderma hemiramphi, drawn free-hand 
from living specimen. Figure 33. Diphtherostomum anisotremi, holotype, ventrolateral 
view. Figure 34. Proctotrema pritchardae, holotype, ventral view. Figure 35. Procto- 
trema anisotremi, holotype, ventral view. Figure 36. Chrisomon decapteri, holotype, dor- 
sal view. Figure 37. Same, eggs enlarged. 


204 


0.083 in diameter; esophagus 2-2.5 times 
length of pharynx; ceca long, extending to 
near posterior extremity. Testis elongated, 
almost half body length; cirrus sac to right 
of midline, 0.150-0.374 long, 0.050-0.064 
wide, containing spherical seminal vesicle, 
tubular pars prostatica and cirrus with small 
spines 9-12 » long. Ovary irregularly lobed, 
to left of and usually overlapping testis an- 
teriorly. Metraterm sac almost as large as 
cirrus sac, consisting of posterior vesicle 
with a few scattered spines and anterior 
division with numerous spines similar to 
those of cirrus. Uterus extending posterior 
to testis, overlapping it ventrally and en- 
tering metraterm sac near its anterior spiny 
portion. Genital atrium small, without 
spines; its pore median, preacetabular. Vitel- 
Jaria in lateral fields, extending from an- 
terior edge of ovary or posterior margin of 
Cirrus sac to tips of ceca; follicles elongated, 
tending to fuse. Eggs 20-24 by 12-17 un, 
rounded at one end, tapering at other (Fig. 
37). Excretory vesicle sac-shaped, very 
short; pore terminal. 

This species is so similar to C. tropicum 
(Manter, 1940) that further collections of 
C. tropicus from the Pacific might prove 
the 2 to be identical. The main differences, 
which may be due to development in differ- 
ent host species, are the larger and more 
elongated testis, slightly more extensive vitel- 
Jaria and a somewhat more anterior ovary 
in C, decaptert, Similarities concern such de- 
tails as shape of the eggs, the presence of 
3 or 4 large spines in the posterior portion 
of the metraterm sac, extent of spination 
and measurements. Manter (1940a) de- 
scribed a very short, Y-shaped excretory 
vesicle in C. tropicum whereas it was ob- 
served to be short but sac-shaped in living 
specimens of C. decapteri; Manter may have 
interpreted the expanded main excretory 
ducts as part of the vesicle. In living C. 
decaptert, the uterus was seen to join the 
metraterm sac at its anterior, more spinose 
portion whereas Manter described that junc- 
tion as being at the posterior end of the sac. 
We studied the type specimen of C, tropicus 
and concluded that the uterus enters the 
metraterm sac as in C. decapteri. Dr. Man- 
ter reexamined the specimen and agreed 
with that interpretation. Thus the diagnosis 
of the genus must be emended as follows: 


Tulane Studies in Zoology 


Vol. 11 


Genus Chrisomom emended 


Monorchiidae: Body elongated; esophagus 
more than twice length of pharynx; testis 
single, elongated, near posterior end of body; 
Cirrus sac and metraterm sac spinose; ovary 
irregularly lobed; uterus extending posterior 
to testis Or not, joining more spinose an- 
terior portion of metraterm sac; vitelline 
follicles numerous, extending most of length 
of hindbody but not reaching acetabulum; 
excretory vesicle sac-shaped, short; para- 
sites in intestine of marine fishes. Type 
species: C. tropicus (Manter, 1940) Manter 
& Pritchard, 1961 (Synonym: Telolecithus 
tropicus Manter); other species: C. decap- 
tert .sp. 

A single specimen from Selar crumenoph- 
thalmus from Jamaica is in agreement with 
the measurements and general topography 
of both species of Chrisomon except that 
the testis, due to distortion and poor fixation, 
is more anterior, and the cirrus sac overlaps 
the metraterm covering it more or less com- 
pletely. Its identification, therefore, remains 
undetermined. 


Postmonorchis orthopristis Hopkins, 1941 

Synonym: Pristisomum orthopristis (Hop- 
kins) Yamaguti, 1958. 

Hosts: *Gerres cinereus (J); *Haemulon 
album (J); *H. flavolineatum (C, J); *H. 
scirus (J). 

Site: intestine. 


Pseudohurleytrema eucinostomt (Manter, 
1942) Yamaguti, 1954 
Synonym: Hurleytrema eucinostomi Man- 
ter, 1942. 
Hosts: *Eucinostomus pseudogula (J); 
Gerres cinereus (C). 
Site: intestine. 


Our specimens differ from Manter’s only 
in having somewhat smaller eggs (22-26 
by 12-16 » compared with 26-28 by 20). 
Siddiqi and Cable (1960) reported the same 
species from Puerto Rico; in their speci- 
mens, eggs measured 27-30 by 11-15 p 
(unpublished data). 


Hurleytrematoides chaetodont (Manter, 
1942) Yamaguti, 1954 
Synonym: Hurleytrema chaetodoni Man- 
ter, 1942. 
Hosts: Chaetodon capistratus (C, J); C. 
striatus (J). 
Site; intestine. 


No. 4 


Hurleytrematoides curacaensis n.sp. 
Figures 38 and 39 
Hosts: Chaetodon capistratus (C); Chae- 
todon ocellatus (C). 
Site: intestine. 
Holotype: US.N.M. 60283. 


Description based on 20 specimens. Body 
elongated, more rounded posteriorly, 0.880- 
1.26 long, 0.175-0.267 wide. Entire cuticle 
densely spinose; eye-spot pigment present. 
Oral sucker 0.053-0.075 in diameter; ventral 
sucker in anterior fourth or third of body, 
0.045-0.060 in diameter; sucker ratio 1:0.8- 
0.9. Prepharynx very short; pharynx 0.030- 
0.039 long, 0.039-0.054 wide; esophagus 
2-3 times length of pharynx; intestinal bi- 
furcation well anterior to ventral sucker; 
ceca long, extending to midlevel of post- 
testicular space or slightly beyond. Gonads 
median, in about middle third of hindbody. 
Testis entire, 0.143-0.246 long, 0.113-0.173 
wide. Posttesticular space 0.240-0.426 long. 
Cirrus sac well-developed, to right of mid- 
line, 0.158-0.195 long, 0.040-0.50 wide, con- 
taining a bipartite seminal vesicle, a short 
pars prostatica and a cirrus with needle-like 
spines, 8-11 m long. Ovary entire, pretes- 
ticular, 0.067-0.100 in diameter; seminal 
receptacle not evident; uterus mainly post- 
testicular, coils mostly transverse; metra- 
term sac absent; metraterm well-developed, 
0.083-0.105 long, 0.030-0.033 wide; entire 
length with spines 7-10 » long, similar in 
shape to those of cirrus. Genital atrium 
without spines; genital pore median, im- 
mediately preacetabular. Eggs 27-33 by 
16-23 yw, exclusive of single unipolar fila- 
ment, 1-1.5 times length of egg. Vitellaria 
with 10-15 follicles on each side, well pos- 
terior to acetabulum, mostly pretesticular, 
a few sometimes extending to midlevel of 
testis. Excretory vesicle tubular; its pore 
terminal. 


This species is very similar to H. chaeto- 
dont, Both are from Chaetodon capistratus 
and one individual harbored both trema- 
todes. H. curacaensis differs from H. chae- 
todomi mainly in having a smaller ventral 
sucker and wider eggs with much shorter 
filaments. The uterine coils tend to be more 
transverse than longitudinal and they lack 
the strand-like appearance characteristic of 
H. chaetodoni, The eggs of H. chaetodoni 
are highly variable in length. Exclusive of 
the filament, they measure 37-42 by 15-17 » 


Trematodes of Marine Fishes 


205 


in our specimens. A single specimen re- 
ported from Puerto Rico by Siddiqi and 
Cable (1960) has eggs measuring 52-54 by 
15-16 (unpublished data). Sogandares- 
Bernal and Sogandares (1961) gave a length 
of 30-32 and suggested that it may vary 
with populations. In H. coronatum Manter 
and Pritchard, 1961, the filament is 10-15 
times the length of the egg. H. malaboensis 
Velasquez, 1961, has a longer egg filament 
(7-8 times length of egg), apparently a 
non-spiny cirrus and longer ceca. 

The next species could be assigned to the 
genus Hurleytrematoides if it had one in- 
stead of 2 testes. Hence a new genus is 
erected for it, and diagnosed as follows: 


Diplohurleytrema n.g. 


Monorchiidae: subfamily Hurleytremati- 
nae: Body spinose, elongated; eye-spot pig- 
ment absent. Acetabulum  preequatorial; 


ceca short; esophagus long. Testes 2, diag- 
onal; cirrus sac long, containing bipartite 
seminal vesicle and spiny cirrus. Ovary en- 
tire, pretesticular, in anterior half of body; 
seminal receptacle present; metraterm sac 
absent; uterus occupying most of hindbody. 
Vitelline follicles lateral, mostly in anterior 
half of body. Genital pore preacetabular. 
Eggs with single unipolar filaments. Excre- 
tory vesicle tubular. Parasitic in intestine of 
marine fishes. Type and only species: 


Diplohurleytrema brevicaecum 


n.g., 1.sp. 
Figure 40 

Host: Echidna catenata (C). 

Site; intestine. 

Holotype: US.N.M. 60284. 

Description based on 25 specimens. Body 
elongated, rounded anteriorly, tapering pos- 
teriorly, 0.566-1.25 long, 0.213-0.407 wide, 
Entire cuticle spinose, with spines becom- 
ing smaller posteriorly; eye-spot pigment 
absent. Oral sucker 0.107-0.180 long, 0.135- 
0.200 wide; ventral sucker preequatorial, 
0.083-0.146 long, 0.090-0.160 wide; sucker 
ratio 1:0.70-0.80. Prepharynx absent; phar- 
ynx 0.039-0.070 in diameter; esophagus 
thick-walled, 3-4 times length of pharynx, 
usually sinuous, surrounded by gland cells; 
ceca short, terminating in zone of anterior 
testis. Testes 2, entire, usually diagonal, 
rarely almost symmetrical or nearly tandem, 
0.083-0.200 in diameter; anterior testis to 


206 


right of midline, posterior testis slightly to 
left. Cirrus sac 0.233-0.467 long (1/3-1/2 
body length), 0.045-0.080 wide; slightly to 
left of midline; containing bipartite seminal 
vesicle, very small and indistinct pars pro- 
statica and long cirrus armed with minute 
spines, difficult to see in stained specimens 
but evident in living material. Ovary en- 
tire, to left of midline, anterior to testes; 
seminal receptacle large, posterodorsal to 
ovary, overlapping tip of left cecum; Mehlis’ 
gland posteromedian to ovary; Laurer’s canal 
opens dorsal to posterior end of cirrus sac; 
uterus strand-like in appearance, filling a 
large portion of the posttesticular space and 
terminating in muscular, thick-walled metra- 
term with finely stippled lining (spines ?). 
Genital atrium spacious; its pore median, 
immediately posterior to intestinal bifurca- 
tion. Eggs 30-37 long by 13-17 p wide, 
exclusive of single unipolar filament 2-3 
times length of egg. Vitellaria in lateral 
groups of 25-30 follicles each, extending 
from posterior level of pharynx to midlevel 
of anterior testis. Excretory vesicle tubular, 
extending to anterior testis; pore terminal. 


Diplomonorchis myrophitis n.sp. 
Figures 41 and 42 


Host: Myrophis punctatus (J). 
Site: intestine. 
Holotype: US.N.M. 60285. 


Description based on 3 specimens. Body 
oval, 0.887-1.062 long, 0.347-0.513 wide. 
Cuticle with spines close together anteriorly 
becoming sparse posteriorly. Eye-spot pig- 
ment present. Oral sucker 0.097-0.108 long, 
0.105-0.113 wide; ventral sucker in middle 
third of body length, 0.072-0.075 long, 
0.054-0.072 wide; sucker ratio 1:0.63-0.70. 
Prepharynx absent; pharynx 0.045-0.053 in 
diameter; esophagus about as long as phar- 
ynx; ceca extending short distance posterior 
to testes. Gonads in middle third of body. 
Testes 2, 0.113-0.140 long, 0.108-0.167 
wide, entire, symmetrical, lateral portions 
extracecal. Cirrus sac to right of acetabulum, 
0.200-0.253 long, 0.090-0.100 wide, extend- 
ing posteriorly to mid- or posterior level of 
Ovary, enclosing seminal vesicle, small in- 
conspicuous pars prostatica, and spiny cir- 
rus. Metraterm sac 0.167-0.213 long, 0.067- 
0.090 wide, posterior 3/5 non-spiny, ante- 
rior part spinose; spines of metraterm and 
cirrus wedge-shaped 15-17 p» long. Ovary 


Tulane Studies in Zoology 


Vola 


distinctly trilobed, to right of midline, 0.113- 
0.133 long, 0.063-0.098 wide; seminal re- 
ceptacle absent; Mehlis’ gland posteromedian 
to cirrus sac; uterus voluminous, mainly post- 
testicular, entering median side of spinose 
anterior portion of metraterm sac. Genital 
atrium unarmed, but appears to be spinose 
when occupied by partly everted cirrus; 
genital pore midway between acetabulum 
and intestinal bifurcation. Eggs numerous, 
20-24 by 15-17 pw. Vitellaria in lateral 
groups of 10-12 follicles, extending from 
about midacetabular level to ends of ceca. 
Excretory vesicle tubular; pore terminal. 

In both Drplomonorchis leiostomi Hop- 
kins, 1941, and D. bivitellosus (Manter, 
1940) the testes are extracecal and the ceca 
extend almost to the posterior end of the 
body whereas in D. myrophitis the ceca are 
overlapped by the testes and terminate a 
short distance posterior to them. Other dif- 
ferences are the more anterior position of 
the testes and distribution of the vitellaria 
in D. myrophitis. 


Diplomonorchis micropogoni n.sp. 
Figure 43 
Hosts: Muicropogon furniert (J); Archo- 
sargus unimaculatus (J). 
Site: intestine. 
Holotype: U.S.N.M. 60286. 


Description based on 17 specimens. Body 
oval to pyriform, 0.233-0.620 long, 0.186- 
0.420 wide. Entire cuticle spinose; eye-spot 
pigment present. Oral sucker 0.046-0.083 
long, 0.066-0.098 wide; ventral sucker in 
middle third of body, 0.037-0.067 in diame- 
ter; sucker ratio 1:0.61-0.84. Prepharynx 
absent; pharynx 0.022-0.037 long, 0.027- 
0.053 wide; ceca terminating near posterior 
margin of testes. Testes 2, symmetrical, 
extracecal, immediately postequatorial, 0.054- 
0.166 long, 0.038-0.080 wide; cirrus sac 
0.090-0.180 long, 0.045-0.090 wide, to right 
of midline, extending short distance poste- 
rior to ventral sucker; containing spherical 
seminal vesicle, short tubular pars pro- 
statica and cirrus with spines. Metraterm 
sac 0.083-0.105 long, 0.035-0.042 wide, with 
large, unarmed posterior vesicle and anterior 
portion with a few spines 6-8 » long, similar 
to those of cirrus. Ovary 4-lobed, 0.060- 
0.165 long, 0.053-0.068 wide, partly anterior 
to, and partly overlapping level of right 
testis; uterus voluminous, occupying almost 


No. 4 


all available space posterior to intestinal bi- 
furcation; entering metraterm sac near an- 
terior spinose end. Vitelline follicles rela- 
tively large, in 2 lateral groups, coinciding 
with or slightly exceeding zone occupied by 
gonads. Eggs numerous, thick-shelled, 22-30 
by 14-18 pw, usually 24-27 by 15-17. Ex- 
cretory vesicle tubular; pore terminal. 

Worms from some hosts could be sepa- 
rated into 2 size groups but otherwise were 
identical. The presence of such groups may 
be expected occasionally because monorchiid 
cercaria may emerge from, and reenter the 
same clam to encyst in large numbers. Thus 
various age groups of adult worms would re- 
sult from the host’s feeding on infected 
clams at different times. 

This species is distinguished from D. 
leiostomt and D. bivitellosus by its short 
ceca and the extent of the uterus. It further 
differs from D. bivitellosus in the distribu- 
tion of the vitellaria, sucker ratio and egg 
size, and from myrophitis in body shape 
and size, extracecal position of testes, more 
extensive uterus and in having fewer spines 
in the metraterm. 


Diplomonorchis hopkinsi n.sp. 
Figure 44 
Host: Micropogon furniert (J). 
Site: intestine. 
Holotype: US.NM. 60287. 


Description based on 16 specimens. Body 
oval, 0.247-0.380 long, 0.180-0.233 wide. 
Entire cuticle spinose; eye-spot pigment ab- 
sent. Oral sucker 0.040-0.053 long, 0.052- 
0.070 wide; ventral sucker just within mid- 
dle third of body length, 0.033-0.039 in di- 
ameter; sucker ratio 1:0.65-0.80. Prephar- 
ynx absent; pharynx 0.025-0.035 in diame- 
ter, esophagus short; ceca extending just 
posterior to testes. Gonads mainly post- 
equatorial. Testes 2, entire, 0.037-0.068 in 
diameter, symmetrical to somewhat oblique, 
mainly extracecal; cirrus sac to right of mid- 
line, 0.084-0.130 long, 0.038-0.045 wide, 
extending at least to midlevel of ovary, con- 
taining spherical seminal vesicle, pars pro- 
statica and relatively long spiny cirrus. 
Metraterm sac indistinct; its spines and 
those of cirrus minute, difficult to see. 
Ovary indistinctly 4-lobed. Immediately an- 
terior to right testis which it may overlap; 
uterus voluminous, filling most available 
space posterior to intestinal bifurcation, en- 


Trematodes of Marine Fishes 


207 


tering metraterm sac near its spinose an- 
terior region. Genital atrium wide, unarmed; 
genital pore median, approximately midway 
between acetabulum and intestinal bifurca- 
tion. Vitellaria 4-6 follicles on each side, 
mainly dorsomedian to testes, rarely extend- 
ing anteriorly to midlevel of ovary. Eggs 
small, thin-shelled, 13-15 by 9-11 p. Excre- 
tory vesicle tubular; its pore terminal. 

This species is named in honor of Prof. 
S. H. Hopkins. Because it and D. micro- 
pogont were found together in the same host 
individual, they were not immediately recog- 
nized as being distinct species. Later we 
found that D. hopkinsi lacked eye-spot pig- 
ment, had an indistinct metraterm sac, and 
contained much smaller eggs. Hopkins did 
not mention eye-spot pigment in describing 
D. leiostomi but our examination of the type 
revealed its presence. Cercaria of the Mo- 
norchiidae, unlike most other families, may 
or may not have eye-spots whose pigment 
can be readily found in the adults. Even so, 
it is unexpected to find in the same genus, 
species with or without such pigment, but 
there may be other such instances as many 
descriptions are not explicit in that matter. 


Diplomonorchis sphaerovarium n.sp. 
Figures 45 and 46 


Host: Spheroides testudineus (J). 
Site: intestine. 
Holotype: US.N.M. 60288. 


Description based on 15 specimens. Body 
oval to elongated, rounded at both ends, 
0.984-1.41 long, 0.386-0.579 wide. Entire 
cuticle spinose, with spines becoming sparse 
posteriorly; eye-spot pigment present. Large 
glands in forebody, characteristic of many 
monorchiids, especially prominent. Oral 
sucker 0.105-0.120 long, 0.135-0.153 wide; 
ventral sucker about one-third body length 
from anterior end, 0.080-0.108 long, 0.099- 
0.120 wide; sucker ratio 1:0.7-0.82. Pre- 
pharynx absent; pharynx 0.054-0.067 in di- 
ameter; esophagus shorter than pharynx; 
ceca relatively long, extending to about mid- 
dle of posttesticular space. Gonads equa- 
torial. Testes 2, entire, symmetrical, 0.100- 
0.150 long, 0.068-0.105 wide; cirrus sac 
elongated, on right, 0.200-0.330 long, 0.066- 
0.090 wide, extending to ovarian zone, con- 
taining large, spherical seminal vesicle, small 
pars prostatica and cirrus with spines 8-10 p 
long. Metraterm sac 0.133-0.226 long, 0.070- 


208 Tulane Studies in Zoology Vol. 11 


Nut) 6) 
Ai 
O 


v= 


Figure 38. Hurleytrematoides curacaensis, holotype, ventral view. Figure 39. Same, 
terminal reproductive organs enlarged. Figure 40. Diplohurleytrematoides brevicaecum, 
holotype, ventral view. Figure 41. Diplomonorchis myrophitis, holotype, ventral view. 
Figure 42. Same, terminal reproductive organs enlarged. Figure 43. Diplomonorchis 
micropogoni, holotype, ventral view. Figure 44. Diplomonorchis hopkinsi, holotype, ven- 
tral view. Figure 45. Diplomonorchis sphaerovarium, holotype, ventral view. Figure 
46. Same, terminal reproductive organs enlarged. 


No. 4 


0.120 wide, with unarmed posterior vesicle 
and anterior region with spines similar to 
those of cirrus. Ovary smooth, immediately 
anterior to right testis, 0.075-0.100 long, 
0.067-0.090 wide; uterus mostly posttesticu- 
lar, entering metraterm sac near middle of 
spinose portion. Genital atrium unarmed; 
genital pore median, preacetabular. Vitel- 
laria consisting of 2 lateral groups of nu- 
merous follicles, mostly posttesticular. Eggs 
24-30 by 16-20 yw. Excretory vesicle long, 
tubular, extending to ventral sucker: pore 
terminal. 


The present species is referred to Dzplo- 
monorchis even though the ovary is spherical 
and not lobed as in other members of the 
genus. That feature, the more extensive 
vitellaria, and perhaps much longer excre- 
tory vesicle distinguish D. sphaerovarium 
from all other. species of Diplomonorchis. 
Together these characteristics may be of 
generic value but until other species having 
those features are found, we prefer to 
broaden the concept of the genus Diplo- 
monorchis to include those characters. The 
genus Diplomonorcheides Thomas (1959) 
would then differ from Dzplomonorchis 
only in having a bipartite seminal vesicle. 


Diplomonorchis emended 

Monorchiidae; subfamily Monorchiinae; 
body oval to elongated. Cuticle spinose. 
Ventral sucker in anterior half of body; 
ceca extending to posterior margin of testes 
or beyond. Testes 2, usually symmetrical, 
inter- or extracecal. Cirrus sac with uni- 
partite seminal vesicle, pars prostatica and 
spiny cirrus. Ovary entire or lobed, pre- 
testicular, to right of midline; metraterm 
sac present, anterior region spinose; uterus 
extensive. Genital pore midventral, pre- 
acetabular. Vitelline follicles postacetabular, 
lateral, in gonadal zone. Excretory vesicle 
tubular. Parasites in intestines of marine 
fishes. Type species: D. leiostomi Hop- 
kins, 1941; other species: D. brevivitellosus 
(Manter, 1940) Hopkins, 1941 (Synonym: 
Paramonorcheides brevivitellosus); D. my- 
rophitis nsp.; D. micropogoni ssp.; D. 
hopkinsi n.sp.; D. sphaerovarium nssp. 


Order Opisthorchiida La Rue, 1957 
Suborder Opisthorchtata La Rue, 1957 
Superfamily Opisthorchioidea ( Witenberg, 
1929) Vogel, 1930 


Trematodes of Marine Fishes 


209 


FAMILY CRYPTOGONIMIDAE Ciurea, 
1933 


Metadena adglobosa Manter, 1947 
Hosts: Lutianus apodus (C, J); *L. aya 
CC). enseus. (C, J) * i yocn (Ce aie 
synagris (J). 
Site: ceca and intestine. 
Metadena crassulata Linton, 1910 
Hosts: Lutianus analis (J); *L. aya (C). 
Site: intestine. 


Metadena globosa (Linton, 1910) 
Manter, 1947 
Synonym: Stegopa globosa Linton, 1910. 
Hosts: *Lutianus apodus (J); L. aya (C); 
Ocyurus chrysurus (J). 
Site: intestine. 


Paracryptogonimus neoamericanus Siddiqi & 
Cable, 1960 

Hosts: Lutianus aya (C); Ocyurus chry- 
surus (C). 

Site: intestine. 

Certain dimensions given by Siddiqi and 
Cable (1960) are extended by a single speci- 
men found in L. aya. It measures 1.293 by 
0.772 and has oral spines up to 26 pu long. 


Siphodera vinaledwardsu (Linton, 1899) 
Linton, 1910 
Synonym: Monostomum vinaledwardsu 
Linton, 1899. 
Hosts: Lutianus analis (C, J); *L. aya 
(C); *L. buccanella (C); L. synagris (J); 
Ocyurus chrysurus (C, J). 


Site: intestine. 


FAMILY HETEROPHYIDAE Odhner, 
1914 
Scaphanocephalus sp. 

Host: Epinephelus striatus (J). 

Site: intestine. 

The single specimen was large and well 
developed but lacked eggs. It probably was 
a recently ingested metacercaria that had 
excysted in the intestine of the fish but could 
not have persisted and matured there. The 
genus Scaphanocephalus is closely related to 
Galactosomum and species of both genera 
are parasites of piscivorous birds. 

Suborder Acanthocolpiata n.subo. 
Superfamily Acanthocolpoidea n.superf. 
FAMILY ACANTHOCOLPIDAE Lihe, 
1909 

After LaRue (1957) placed the family 
Acanthocolpidae in the superfamily Allo- 


210 


creadioidea of the suborder Plagiorchiata, 
Peters (1961) described the embryology of 
the cercarial excretory system and, on that 
basis, suggested a closer affinity of that 
family with the Echinostomatoidea. The 
morphology of their adults is indeed very 
similar as is also the development of the 
excretory system in their cercariae, except 
that the bladder is large and epithelial in the 
Acanthocolpidae whereas the echinostomes 
have a small bladder that lacks an epithelium. 
In that respect and in the location of the 
primary excretory pores, the embryology of 
the excretory system in acanthocolpids agrees 
with that of cercariae of the order Opisthor- 
chiida as characterized by La Rue. However, 
differences in both larval and adult mor- 
phology exceed those occurring between 
families, superfamilies or even suborders of 
La Rue’s scheme. For that reason, a new 
suborder is proposed and characterized as 
follows: 
Suborder Acanthocolptata 

Spinose, distomatous trematodes with bio- 
cellate cercaria developing in rediae in ma- 
rine prosobranch gastropods. Excretory sys- 
tem stenostomate; primary pores of cercarial 
embryo in tail, well removed from body-tail 
furrow; excretory vesicle sac- to Y-shaped 
with short arms, wall with conspicuous gran- 
ular cells. Cercaria with well-developed 
suckers; oral sucker not protrusible, with or 
without stylet. Tail well-developed, with or 
without longitudinal fins, possibly zygocer- 
cous or otherwise modified in species whose 
life histories are unknown. Metacercariae 
in fishes. 


The new superfamily has the characters 
of the suborder but is not further character- 
ized at this time because of the possibility 
that it will eventually include the Campu- 
lidae. No life histories in that family have 
yet been determined. However, its affinity 
with the acanthocolpids is strongly suggested 
by unpublished studies made in this labora- 
tory concerning the adult morpohlogy of 
Orthosplanchnus fraterculus, a  campulid 
from the gall bladder of the sea otter, En- 
hydra lutris. 


Stephanostomum casum (Linton, 1910) 
McFarlane, 1936 
Synonyms: Stephanochasmus casus Linton, 
1910; Lechradena edentula Linton, 1910 


Tulane Studies in Zoology 


Vol. 11 


Hosts: *Lutianus aya (C); *L. buccanella 
(C), *L. synagris (J). 
Site: intestine. 


Stephanostomum coryphaenae Manter, 
1947 
Host: Coryphaena hippurus (C). 
Site: intestine. 


Stephanostomum dentatum (Linton, 1900) 
Manter, 1931 
Synonym: Distomum dentatum Linton, 
1900. 
Hosts: Epinephelus striatus (J); *Myc- 
teroperca bonacit (C). 
Site: intestine. 


Stephanostomum sentum (Linton, 1910) 
Manter, 1947 

Synonym: Stephanochasmus sentus Linton, 
1910. 

Hosts: *Antisotremus virginicus (J); 
*Caranx latus (J); Gerres cinereus (C); 
*Haemulon album (C); H. scwmrus (J); 
*Lutianus sp. (C). 

Site: intestine. 


Stephanostomum ditrematis (Yamaguti, 
1939) Manter, 1947 

Synonyms:  Echinostephanus  ditrematis 
Yamaguti, 1939; Stephanostomum longisom- 
um Manter, 1940; Stephanostomum fili- 
forme Linton, 1940. 

Hosts: *Caranx bartholomaet (J); *C. 
crysos (J); C. hippos (J); C. latus (J); 
"“Caran% sp. (@)e 

Site; intestine. 


Stephanostomum pseudocarangis Sogandares- 
Bernal, 1959 
Host: Holocentrus ascensionis (J). 
Site; intestine. 


Stephanostomum megacephalum Manter, 
1940 

Host: Caranx latus (J). 

Site: intestine. 


Stephanostomum aulostomi n.sp. 
Figures 47 and 48 


Host: Aulostomus maculatus (C). 
Site: junction of stomach and intestine. 
Holotype: U.S.N.M. 60289. 


Description based on one complete and 2 
incomplete specimens. Body elongated, 6.37 
long, 0.547-0.667 in maximum width at level 
of acetabulum. Entire cuticle spinose, spines 
becoming sparse posteriorly; eye-spot pig- 


No. 4 


ment present. Oral sucker 0.193 by 0.273, 
with 36 perioral spines 21-37 by 11-15 yp, 
alternating in 2 rows of 18 each; ventral 
sucker 0.333-0.387 in diameter; sucker ratio 
1:1.6.  Prepharynx 0.700 long; pharynx 
0.300-0.334 long, 0.167-0.180 wide; esopha- 
gus short; intestinal bifurcation close to ace- 
tabulum; ceca extending to near posterior 
end of body, joining excretory vesicle to 
form uroproct; feces seen discharged from 
terminal pore. Gonads in posterior third of 
body. Testes 2, 0.360-0.587 long, 0.187- 
0.234 wide, tandem, separated by vitelline 
follicles. Cirrus sac long, not quite reach- 
ing midway between acetabulum and ovary, 
containing saccate seminal vesicle, pars pro- 
statica and long spiny cirrus. Ovary 0.300- 
0.327 long, 0.253-0.267 wide, anterior to, 
and separated from testes by vitelline fol- 
licles; uterine seminal receptacle, Mehlis’ 
gland and uterus preovarian; metraterm well- 
developed, without spines, joining male duct 
near posterior margin of acetabulum. Geni- 
tal atrium tubular; genital pore immediately 
preacetabular. Eggs 60-75 by 45-51 up. 
Vitelline follicles extending from posterior 
end of body to near posterior margin of 
ventral sucker. Excretory vesicle obscured by 
vitellaria. 


The combination of 2 uninterrupted rows 
of 18 oral spines each and vitellaria that ex- 
tend to near the posterior margin of the 
ventral sucker distinguishes S. aulostomi 
from most species of Stephanostomum. It 
differs from S. caswm in having a more elon- 
gated body, more anterior testes, less pos- 
terior extent of the cirrus sac and gonads 
separated by vitelline follicles; and from S. 
coryphaenae chiefly in the shape of the cir- 
rus sac, more anterior extent of the vitellaria, 
and in having a much longer genital atrium 
and somewhat wider eggs. 

In habitat, §. awlostomi is unusual, with 
the oral sucker anchored just above the 
pyloric sphincter of the host and most of 
the body in the duodenum. Thus it was not 
until the digestive tract was opened without 
separating the stomach and intestine that a 
complete specimen was obtained. 


_ Stephanostomum metacercaria 
Host: Cypselurus bahiensis (C). 
Site: cyst on gill arch. 
The single. acanthocolpid metacercaria 
found in this study had an oral sucker that 


Trematodes of Marine Fishes 


Za 


was smaller than the acetabulum and 48 
perioral spines in 2 uninterrupted rows. 


Tormopsolus orientalis Yamaguti, 1934 
Host: *Seritola dumerili (C). 
Site; intestine. 


Manteria brachydera (Manter, 1940) 
Caballero, 1950 
Synonyms: Dthemistephanus bachyderus 
Manter, 1940; Stephanostomum sp. Linton, 
1940. 
Host: Oligoplitis saurus (J). 
Site; intestine. 


Suborder Hemiurata Skrjabin & 
Guschanskaja, 1954 
Superfamily Hemiuroidea Faust, 1929 
FAMILY HEMIURIDAE Lihe, 1901 


Parahemiurus merus (Linton, 1910) 
W oolcock, 1935 

Synonyms: Hemiurus merus Linton, 1910; 
Parahemturus parahemiurus Vaz & Pereira, 
1930; P. platichthyi Lloyd, 1938; P. atheri- 
nae Yamagutt, 1938; P. harengulae Yama- 
guti, 1938. 

Hosts: Abudefduf saxatilis (J); *Caranx 
erysos (J); *C. hippos (J); *C. latus (J); 
*Echenets naucrates (C); Opisthonema og- 
linum (J); *Sardinella anchovia (C, J); S. 
macrophthalmus (J); *Seriola dumerili (J). 

Site: stomach. 


Sterrhurus fusiformis (Luhe, 1901) 
Looss, 1907 
Synonym: Lecithochirium fusiformis Lahe, 
1901. 
Hosts: Gymnothorax moringa (C, J); *G. 
vicinus (C). 
Site: stomach. 


Sterrhurus musculus Looss, 1907 

Synonyms: Sterrhurus floridensis Manter, 
1934, in part; Sterrhurus laeve (Linton) of 
Manter, 1931. 

Hosts: *Achirus lineatus (J); *Ept- 
nephelus adscenstonis (C); *E. morio (C); 
*Haemulon album (C); *H. scturus (J); 
Holocentrus. ascenstonis (C, J); *H. vexil- 
larius (J); *Leptocephalus conger (J); 
*Tutianus apodus (C); *L. aya (C); *L. 
griseus (C); Malacanthus plumieri (J); 
*Platophrys lunatus (C, J); *Prionotus 
punctatus (J);.*Rypticus saponaceus (C); 
Scorpaena plumiert (C, J); *Selar crume- 


212 


nophthalmus (J); Synodus intermedius 
WO: *Trachinotus glaucus (J)% 


(C, 


Site: stomach. 

Lecithochirium microstomum Chandler, 
L935 

Synonym: Lecithochirium  sinaloense 


Bravo-Hollis, 1956. 

Hosts: Synodus intermedius (J); *Selar 
crumenophthalmus (J); *Sertola dumerili 
(Pp). 

Site: stomach. 

Lecithochirium parvum Manter, 1947 

Synonym: Sterrhurus floridensis Manter, 
1934, in part. 

Hosts: * Abudefduf saxatilis (J); *Bathy- 
stoma striatum (J); *Caranx bartholomaet 
CDE tGaiggs: “Cs *G. Gappos' 4G; J); 
*Dules dispilurus (J); *Epinephelus ad- 
scensionis (J); *Holocentrus ascenstonts 
(J); *Lutianus apodus (C); *L. aya (C); 
*I. griseus (J); *Sardinella macrophthal- 
mus (J); *Scorpaena plumiert (J); *Selar 
crumenophthalmus (J); *Seriola dumerili 
(C, J); *Upeneus martinicus (J); Synodus 
intermedius (J). 

Site: stomach. 


Ectenurus americanus (Manter, 1947 ) 
Manter & Pritchard, 1960 

Synonyms: Parectenurus americanus Man- 
ter, 1947; Magnacetabulum americanum 
(Manter) Yamaguti, 1954. 

Hosts: Caranx bartholomaet (J); *C. ery- 
sos (J); *C. hippos (J); *Epinephelus stri- 
atus (J); *Selar crumenophthalmus (J); 
*Seriola dumerili (J); *Synodus inter- 
medius (J). 


Site: stomach. 


Ectenurus virgulus Linton, 1910 
Hosts: *Caranx bartholomae (J); *C. 
hippos (J); *Priacanthus cruentatus (C); 
Sardinella macrophthalmus (J); Selar cru- 
menophthalmus (J); *Trachinotus glaucus 
C]): 
Site: intestine. 
Dinurus barbatus (Cohn, 1902) 
Looss, 1907 
Synonym: Lecithocladium barbatum Cohn, 
1902. 
Host: Coryphaena hippurus (C). 
Site: stomach. 


Dinurus breviductus Looss, 1907 
Host: Coryphaena hippurus (C). 
Site: stomach. 


Tulane Studies in Zoology 


Vol. 11 


Dinurus tornatus (Rudolphi, 1819) 
Looss, 1907 
Synonyms: Distomum tornatum Rudolphi, 
1819; Lecithocladium tornatum (Rud.) 
Luhe, 1901. 
Host: Coryphaena hippurus (C). 


Site: stomach. 


Stomachicola rubea (Linton, 1910) 
Manter, 1947 

Synonyms: Dinurus rubeus Linton, 1910; 
Pseudostomachtcola rubea (Linton) Skrjabin 
& Guschanskaja, 1954. 

Hosts: Gymnothorax moringa (J); *G. 
vicinus (J). 

Site: stomach. 


Neogenolinea opisthonemae Siddigi & 
Cable, 1960 

Hosts: Opisthonema oglinum (J); *Sar- 
dinella anchovia (J); *S. macrophthalmus 
CIid:- 

Site: stomach. 

Brachadena pyrtformis Linton, 1910 

Synonyms: “Distomum  bothryophoron 
Olsson” of Linton, 1905; Lecithaster aniso- 
tremit MacCallum, 1921; L. gibbosus (Rud.) 
of Linton, 1940 in part; **Aponurus sym- 
metrorchis Siddiqi & Cable, 1960. 

Hosts: Antsotremus virginicus (J); 
*Archosargus unimaculatus (J); Bathy- 
stoma striatum (J); Calamus bajanado (J); 
*Eucinostomus pseudogula (J); *Haemulon 
bonariense (J); *H. flavolineatum (C, J); 
HI. sciurus (J). 

Site; stomach. 

A reexamination of the type and para- 
types of Aponurus symmeterorchis Siddiqi 
and Cable, 1960, reveals that the vitellaria 
unite centrally, a characteristic of the genus 
Brachadena, and the measurements given 
overlap those of Brachadena pyriformis. 


Genolinea noblei n.sp. 
Figure 49 


Host: Abudefduf saxatilis (C). 

Site: stomach. 

Holotype: US.N.M. 60290. 

Description based on a single specimen, 
Body thick, rounded at both ends, 1.25 long, 
0.367 wide. Oral sucker subterminal, 0.087 
long, 0.082 wide, surmounted by conspicu- 
ous fleshy lobe; ventral sucker 0.227 long, 
0.213 wide, with longitudinal aperture; 
sucker ratio 1:2.59. Prepharynx absent; 
pharynx 0.045 long, 0.063 wide; esophagus 


No. 4 


absent; ceca wide, extending to near pos- 
terior end of body. Testes 2, entire, slightly 
diagonal, 0.075 long, 0.083-0.090 wide, sepa- 
rated by uterine coils; seminal vesicle long, 
sinuous, not reaching midlevel of acetabu- 
lum, with 4 conspicuous swellings connected 
by narrow ducts; prostate vesicle ovoid, 0.045 
long, 0.030 wide, surrounded by prostate 
cells; duct very short. Ovary entire, 0.060 
long, 0.105 wide, posterior to, and separated 
from testes by uterine coils; Mehlis’ gland 
dorsal to vitellaria; seminal receptacle not 
evident, possibly concealed by uterus and 
ceca; uterus extending to near posterior end 
of body; metraterm well-developed, spiny, 
joining prostatic duct at base of sinus sac. 
Hermaphroditic duct with swollen posterior 
region and elongated anterior portion. Sinus 
sac apparently of open or incomplete type. 
Genital pore ventral, opposite intestinal bi- 
furcation. Eggs 28-33 by 10-12 pw. Vitellaria 
2 compact, tandem masses immediately post- 
ovarian. Excretory vesicle short, lined with 
epithelial cells; excretory ducts uniting dor- 
sal to pharynx. 

The spiny metraterm and _ longitudinal 
aperture of the acetabulum distinguish this 
species from all others in the genus Geno- 
linea. G. tanyopa Montgomery, 1957, has 
a longitudinal aperture but differs from G. 
noble: in sucker ratio, posterior extent of 
the seminal vesicle, nature of the sinus sac 
and also in having a much longer prostatic 
duct. 


The species is named in honor of the late 
Alden E. Noble of the University of the 
Pacific in recognition of his contributions 
to trematodology. 


Aponurus elongatus Siddiqi & Cable, 1960 
Synonym: Aponurus sp. Linton, 1940. 
Host: Chaetodipterus faber (J). 

Site: stomach. 


Leurodera decora Linton, 1910 
Hosts: Anisotremus virginicus (J); Hae- 
mulon flavolineatum (J); H. sciurus (J). 
Site: stomach. 


Dichadena acuta Linton, 1910 
Synonym: Lecithaster acutus (Linton) 
Manter, 1947. 
Hosts: Acanthurus bahianus (C); A. coe- 
ruleus (J); A. hepatus (C, J). 
Site: stomach. 


Trematodes of Marine Fishes 


213 


Macradena perfecta Linton, 1910 
Host: * Acanthurus hepatus (C, J). 
Site: stomach. 


Hysterolecitha rosea Linton, 1910 
Host: Acanthurus hepatus (J). 
Site: stomach. 


Hysterolecitha sogandaresi n.sp. 
Figure 50 

Host: Acanthurus coeruleus (J). 
Site: stomach. 
Holotype: US.N.M. 60291. 
Description based on 3 specimens. Body 
non-appendiculate, tapering posteriorly, 1.54- 
2.22 long, 0.467-0.533 in maximum width at 
level of acetabulum. Oral sucker subterminal, 
0.147-0.173 long, 0.167-0.220 wide; ventral 
sucker near midbody, 0.367-0.433 long, 
0.387-0.400 wide; sucker ratio 1:2.14-2.33. 
Prepharynx absent; pharynx 0.060-0.070 
long, 0.070-0.090 wide; esophagus very 
short; ceca swollen near intestinal bifurca- 
tion, extending to near posterior end of 
body. Testes 2, entire, diagonal (nearly tan- 
dem in one specimen), 0.082-0.123 in di- 
ameter; anterior testis somewhat dorsal to 
acetabulum; seminal vesicle long, coiled tube, 
mostly anterior to ventral sucker; pars pro- 
statica short, surrounded by prostate cells. 
Ovary entire, submedian,  posttesticular, 
0.068-0.105 in diameter, usually overlapping 
posterior level of testes; uterus extending to 
near tips of ceca; metraterm simple, joining 
short prostatic duct at base of small, spheri- 
cal sinus sac. Hermaphroditic duct short. 
Genital pore median, ventral, some distance 
posterior to intestinal bifurcation. Eggs nu- 
merous, 26-31 by 15-19 pw. Vitellaria of 7 
subglobular follicles, immediately _ post- 
ovarian. Excretory arms uniting dorsal to 
pharynx; excretory pore terminal. 

Of the 11 species described in Hystero- 
lecitha, H. sogandarest is most similar to H. 
acanthuri Annereaux, 1947, from a related 
host in the Phillippines but differs from 
that species in having testes closer to the 
acetabulum, a shorter prostatic duct and 
more compact vitellaria. 

This species is named in honor of Dr, 
Franklin Sogandares in recognition of his 
contributions to knowledge of the Trema- 
toda. 


214 


Theletrum pomacentri n.sp. 
Figure 51 
Host: Pomacentrus leucosticus (J). 


Site: stomach. 
Holotype: U.S.N.M. 60292. 


Description based on a single specimen. 
Body non-appendiculate, tapering posterior- 
ly, 1.54 long, 0.500 in maximum width at 
level of acetabulum; forebody 0.413 long, 
hindbody 0.865. Oral sucker 0.099 long, 
0.105 wide; ventral sucker 0.262 long, 0.240 
wide, with longitudinal aperture; sucker 
ratio 1:2.43. Prepharynx absent; pharynx 
spherical, 0.054 in diameter; esophagus 
short; ceca extending to near posterior end 
of body. Testes smooth, diagonal, 0.075- 
0.090 in diameter; seminal vesicle coiled, 
tubular, mostly preacetabular; prostate vesi- 
cle reniform, surrounded by poorly-developed 
prostate cells. Ovary bilobed, 0.060 long, 
0.135 wide, posterior to, and separated from 
testes by coils of uterus; Mehlis’ gland not 
evident; uterus voluminous, not reaching 
ends of ceca; metraterm simple, ventral to 
seminal vesicle, joining very short prostatic 
duct at base of sinus sac. Hermaphroditic 
duct widest anteriorly; sinus sac subglobular, 
0.060 by 0.075. Genital pore midventral, 
posterior to intestinal bifurcation. Eggs nu- 
merous, 27-30 by 10-12 pw. Vitellaria im- 
mediately postovarian; in 3 Compact masses 
with 2 anterior ones possibly connected by 
an isthmus; posterior mass slightly indented. 
Excretory system not observed. 

Theletrum pomacentri differs from all 
the other species in the genus in having a 
bilobed ovary. It further differs from T. 
fustiforme Linton, 1910, in lacking the post- 
acetabular fold and in having 3 rather than 
2 vitelline masses; from T. gravidum Man- 
ter, 1940, in sucker ratio and shape and 
extent of seminal vesicle; from T. /issoso- 
mum Manter, 1940, in sucker ratio; and 
from T. magnasaccum Sogandares-Bernal 
and Sogandares, 1961, in shape and extent 
of the seminal vesicle, shape of the sinus sac 
and in the position of the genital pore. 


The type specimen was damaged after it 
was studied and drawn. 


In Jamaica and especially Curacao, sur- 
geon fish were commonly infected with a 
monorchid hemiurid which otherwise re- 
sembled species of Macradena. To receive 
that species, a new genus is proposed and 
characterized as follows: 


Tulane Studies in Zoology 


Vol. 11 


Monorchimacradena n.g. 


Hemiuridae. Medium size distomes with- 
out ecsoma. Cuticle smooth. Oral sucker 
subterminal; ventral sucker preequatorial. 
Ceca extending to near posterior end of 
body. Testis single; seminal vesicle post- 
acetabular; pars prostatica long, tubular, 
mostly posterior to ventral sucker, sur- 
rounded by prostate cells. Sinus sac present. 
Ovary entire, posttesticular; seminal recep- 
tacle present. Vitellaria of elongated lobes, 
postovarian. Eggs small and numerous. Ex- 
cretory commissure present. Parasitic in in- 
testine of marine fish. Type and only species: 


Monorchimacradena acanthuri 
n.g., N.sp. 
Figure 52 
Host: Acanthurus hepatus (C, J). 
Site: intestine. 


Holotype: US.N.M. 60293. 


Description based on 38 specimens; meas- 
urements on 10. Body usually elongated, 
1.1-2.57. long, 0.240-0.374 in maximum 
width at level of acetabulum. Oral sucker 
subterminal, 0.082-0.145 long, 0.105-0.180 
wide; preoral lobe fleshy, often expanded, 
resembling head of a planarian, especially 
noticeable in living specimens. Ventral 
sucker in anterior third or fourth of body, 
0.150-0.266 in diameter, aperture transverse; 
sucker ratio 1:1.3-1.8. Prepharynx absent; 
pharynx 0.060-0.097 in diameter; esopha- 
gus about same length as pharynx; ceca 
without epithelium for a short distance from 
intestinal bifurcation, ending blindly near 
posterior end of body. Testis about equa- 
torial, 0.105-0.200 long, 0.068-0.130 wide; 
seminal vesicle sac-like, immediately pre- 
testicular; pars prostatica long, tubular, usu- 
ally in hindbody, sometimes partly dorsal to 
ventral sucker, surrounded by conspicuous 
prostate cells along entire length; ejaculatory 
duct as long as pars prostatica when not 
contracted. Ovary smooth, 0.045-0.160 in 
diameter, immediately posttesticular; seminal 
receptacle as large or larger than ovary; 
uterus extending to near posterior extrem- 
ity; metraterm simple, joining pars pro- 
statica at base of sinus sac. Hermaphroditic 
duct wide. Sinus sac spherical to pyriform, 
0.060-0.112 in diameter. Genital pore mid- 
ventral, posterior to intestinal bifurcation. 
Eggs numerous, 20-28 by 9-15 p. Vitellaria 
immediately postovarian, of 7 digitiform or 


No. 4 Trematodes of Marine Fishes MS: 


BOO 
Yes 


Sea! 


Sf 


eure 


Way 
SS 


AIP 


UY 


S 


5| 


Figure 47. Stephanostomum aulostomi, holotype, dorsal view. Figure 48. Same, an- 
terior end enlarged. Figure 49. Genolinea noblei, holotype, ventrolateral view. Figure 
50. Hysterolecitha sogandaresi, holotype, ventral view. Figure 51. Theletrum poma- 
tA holotype, ventral view. Figure 52. Monorchimacradena acanthuri, holotype, ven- 
tral view. 


216 


slightly branched lobes, united centrally. Ex- 
cretory system with commissure dorsal to 
pharynx; pore terminal. 


Dictysarca virens Linton, 1910 
Hosts: Gymnothorax funebris (J); G. 
moringa (J); *G. vicinus (J). 
Site: swim bladder. 


FAMILY ACCACOELIIDAE Looss, 1912 


Tetrochetus coryphaenae Yamaguti, 1934 

Hosts: Coryphaena hippurus (C); *Dto- 
don hystrix (J). 

Site: intestine. 


FAMILY HIRUDINELLIDAE Dollfus, 
1952 
Hirudmella sp. 
Host: Scorpaena plumieri (C). 
Site: stomach. 
A single immature specimen was taken 
from the stomach of a scorpion fish. 


FAMILY SCLERODISTOMIDAE 
Dollfus, 1932 


Sclerodistomum sphoeroidis Manter, 1947 
Host: *Diodon hystrix (C, J). 
Site: stomach. 


FAMILY PROSOGONOTREMATIDAE 
Pérez Vigueras, 1940 


Prosogonotrema b:labiatum Pérez 
Vigueras, 1940 
Host: Ocyurus chrysurus (J). 
Site: stomach. 


V. GEOGRAPHICAL DISTRIBUTION 


The geographical distribution of the di- 
genetic trematodes of marine fishes in the 
Gulf-Caribbean region has been discussed 
by a number of investigators. Manter 
(1940b, 1947, 1955) included them in re- 
viewing the zoogeography of the group on 
a world-wide basis. A more limited ap- 
proach was that of Sparks (1960) who 
concluded that the “hydrographic, climato- 
logical, physiographic and geological condi- 
tions existing both now and in the past in 
the northern gulf” are responsible for the 
differences seen between the trematode 
faunas of the Dry Tortugas and Grand Isle, 
Louisiana. Siddiqi and Cable (1960) dis- 
cussed geographical distribution and the re- 
lated factors of isolation, speciation and host 
specificity. They compared the trematode 
fauna of Puerto Rico with that of Tortugas, 


Tulane Studies in Zoology 


Vol. 11 


Bermuda, Galapagos Islands, Woods Hole, 
Beaufort and Hawaii and found that the 
per cent of Puerto Rican species common 
also to each of those localities decreased in 
the order listed, from a maximum of 76% 
at Tortugas. 

The present study provides data from 2 
additional localities in the Caribbean region. 
Table 1 compares by families the number 
of new and previously described species 
from those localities. With respect to the 
total found there, the differences in their 
trematode faunas are striking in certain 
families. Curagao is notably poor in fel- 
lodistomatid species where but 3, all pre- 
viously known and widely distributed, were 
found in 4 of 124 species of fishes. In con- 
trast, 12 species of fellodistomatids were 
found in Jamaica where 127 species of 
fishes were examined. The difference may 
be due to the limited variety of habitats in 
Curacao for lamellibranch molluscs which 
serve as intermediate hosts of the fello- 
distomatids. On the other hand the richness 
of such habitats in Jamaica and the life 
history pattern in that family would seem 
to favor speciation in the group as indi- 
cated by the discovery there of 4 new spe- 
cies. In known life histories, fellodistomatid 
metacercariae occur in molluscs or possibly 
amphipods and would be ingested only by 
birds feeding in shallow water or by bottom- 
feeding fishes where the food source could 
serve to isolate populations of shallow water 
host species. 

The same may be observed concerning the 
family Lepocreadiidae except that in it, 
gastropods serve as the mulluscan hosts. 
Their relative abundance and variety in 
Curacao, as compared with lamellibranchs, 
is reflected in the discovery there of 5 new 
lepocreadiids, whereas 6 were found in 
Jamaica. A comparable situation with re- 
spect to new species of monorchiids, which 
have lamellibranch molluscan hosts, seems 
to contradict what was said above concern- 
ing the fellodistomatids but may be ex- 
plained by a lower degree of specificity of 
the monorchiids for those hosts. For ex- 
ample, Cercaria caribbea XXXVI, a monor- 
chid larva, occurs in at least 2 distantly re- 
lated species of bivalves in different local- 
ities. 

Curacao is rather removed from other 
Gulf-Caribbean areas that have been in- 
vestigated. The number of known trema- 


No. 4 


Trematodes of Marine Fishes 


TABLE 1. 
Distribution of Digenetic Trematode Species. 


Number of Species 


common to 


Curacao Jamaica both localities 

Family total old new old new old new 
Aspidogastridae 2 0 0 2 0 0 
Acanthocolpidae 10 6 il 7 0 4 
Accacoeliidae il 1 0 1 0 1 
Bivesiculidae 1 0 1 0 1 0 1 
Bucephalidae 15 eZ 1 10 0 8 
Cryptogonimidae 5 5 0 4 0 4 
Fellodistomatidae 12 3 ) 8 4 3 
Gorgoderidae 2 il 0 1 il 1 
Hemiuridae 23 11 2 16 3 8 1 
Haploporidae 4 1 0 4 0 1 
Haplosplanchnidae 11 6 3 5 3 5 1 
Lepocreadiidae 36 11 5 22 6 6 2 
Megaperidae 3 2 0 i 1 1 
Microphallidae 2 2 0 0 0 0 
Monorchiidae 7 6 4 8 5 6 
Opecoelidae Pal 8 3 16 0 6 
Opistholebetidae % 1 0 2 0 1 
Paramphistomatidae it 0 0 il 0 0 
Pronocephalidae 1 0 0 1 0 0 
Prosogonotrematidae 1 0 0 1 0 0 
Sclerodistomatidae iL iL 0 1 0 1 
Zoogonidae tl 4 0 4 il 2 

178 80 20 115 25 57 5 


Not included in this table are Stephanostomum metacercaria, Hirudinella sp. (immature), 


Scaphanocephalus sp., and Alcicornis siddiqii. 


tode species found there, however, reflects 
its nearness to the mainland, along the 
coast of which many such species probably 
have a continuous distribution toward both 
Central America and eastward along the 
South American coast. Other species are 
cosmopolitan parasites of far-ranging fishes 
throughout the region. However, a degree 
of isolation is suggested by the number of 
new species in certain families including 
those discussed above, and by the absence of 
certain others. Thus 3 of 11 opecoelids 
found in Curacao are new whereas all of 
16 Jamaican species are known ones. On 
the other hand, the opecoelids, Hamacrea- 
dium mutabile and Helicometrina nimia, 
were not found in Curacao although num- 
bers of their host species were examined. 
The same was observed for several other 
species indicated by a dagger in the Host- 
Parasite List. 

Table 2 includes, by family, the number 
of species common to each of several lo- 
calities and Tortugas where 146 species are 
known excluding those reported from deep- 
water fishes. Cuba is not included in the 


table because Pérez Vigueras described as 
new several species which were not ade- 
quately compared with known ones and 
probably are not distinct from them. Most 
similar to the trematode fauna of Tortugas 
is that of the Bahamas (Bimini, Nassau and 
Eleuthera) and Bermuda, as is to be ex- 
pected from their relationship to the Gulf 
Stream. When Jamaica and Curacao are 
compared with Tortugas, the percent of 
their trematode species in common with 
that locality is the same even though Cu- 
racao is much farther from Tortugas. The 
somewhat lower percentage for Puerto Rico 
reflects its isolated position at the eastern 
end of the Greater Antilles chain. Differ- 
ences in the trematode faunas of the north- 
ern Gulf of Mexico and Tortugas have been 
discussed by Sparks (1960) and are indi- 
cated by the percentages given in Table 2. 
The physiographic factors which he stressed 
probably determine the distributional limits 
not only of definitive host species but per- 
haps more significantly those of molluscan 
hosts as well. Several molluscs that harbor 
a variety of larval trematodes throughout 


218 Tulane Studies in Zoology Vol. 11 


TABLE 2. 
Resemblance of the trematodes of marine fishes from the Gulf-Caribbean areas to those of 
Tortugas, Florida, as indicated by the number of species common to 
both localities. 


os 
o 
n 3 S 
PP aoe ° oo 
is % = % Pe % % 3 = bos 
Se ge nt. 85 Ge so Soe 
B= Su FS ‘gst Ppa SS =o =x 20 
So ee ee 
is cS AS ahi b=! ae Mud Hi oa 
Family = ZA —Q 5 Ay S) —Q ) Qe 
Aspidogastridae 1 1 1 1 1 
Acanthocolpidae i 2 4 5 4 5 2 2 2 
Accacoeliidae 1 1 1 il 1 
Bivesiculidae 1 1 1 
Bucephalidae 13 1 3 5 3 a 1 1 
Cryptogonimidae 4 il 3 4 2 4 2 1 1 
Fellodistomatidae 9 1 4 7 4 3 if 1 
Gorgoderidae Pe 1 1 1 il 
Hemiuridae 32 3 9 14 14 ial 4 14 4 
Hirudinellidae 1 1 1 
Haploporidae 4 2 2 1 tL 1 
Haplosplanchnidae 7 il 4 3 4 2 
Lepocreadiidae 16 7 11 W 3 5 1 1 
Megaperidae 4 2 1 3 2 1 
Microphallidae 
Monorchiidae 9 1 2 a 4 5 2 
Opecoelidae 21 f{ i 10 9 6 1 
Opistholebetidae Be 1 il 1 
Paramphistomatidae 1 1 1 
Pronocephalidae 2 il 1 2 1 
Sclerodistomatidae 1 1 i 1 
Zoogonidae 8 1 3 3 7) ul 
Total 146 19 48 79 60 57 28 All 12 


Percent of species 
in common 100 89 80 57 50 57 65 48 40 


-* Number in parenthesis following locality indicates total number of trematode species 
known from that area. 


the Caribbean region disappear before reach- VI. Host SPECIFICITY 

ing the latitude of Boca Ciega Bay or Grand Because the host is the immediate en- 
Isle in the Gulf of Mexico. Hence adults vironment of the parasite, host specificity 
of trematodes that are specific for such as well as distribution is an isolating factor 
molluscs would not be found in non-migra- of major importance in the zoogeography 
tory hosts north of that latitude. A notable of parasites in general. Manter (1957) sum- 
exception to that situation allows the cryp- marized the extent to which digenetic trema- 
togonimid, Siphodera vinaledwardsu, to oc- todes have been reported from one or more 
cur from Cape Cod to Curacao at least. species of marine fishes in Japan, Tortugas, 
That species has as its molluscan host snails the Mediterranean, and the British Isles. 
that are very similar if not the same species According to him, the trematodes found in 
throughout that range. However, north of but one host species ranged from 48% of 
Tortugas, whether along the Atlantic or Gulf the known species in the British Isles to 
coasts, the definitive host is a toadfish which 76.4% in Japan; the average for all local- 
is replaced by very different fishes, the snap- ities, calculated from Manter'’s data, is 60.9% 
pers, from Tortugas southward. and that for Curagao and Jamaica combined — 


219 


Trematodes of Marine Fishes 


No. 4 


VI 


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220 


is 599%. Trematodes recovered from 2 host 
species varied between 12.1% in Japan to 
22.6% in Tortugas with an average of 
17.7% compared with 18% in the present 
study. For 3 host species, the number 
ranged from 6.76% in Japan to 9.3% in 
the British Isles, and averaged 7.74% as 
compared with 9.5% in Curagao and Ja- 
maica. For 4 hosts, the range was from 
1.68% in Japan to 6.6% in the British Isles, 
averaging 3.7% which is very close to 
3.4% calculated from the present study. 
From 3% of the trematode species in Japan 
to 18.6% in the British Isles have been 
reported from 5 or more host species, with 
an average of 9.77%; present data show 
10.6% in Curacao and Jamaica. 

The above data indicate a rather high de- 
gree of host specificity for trematodes of 
marine fishes but do not take into account 
the differences between trematode families 
or the degree to which more than one host 
species of a trematode may be related. Those 
aspects are included in Table 3 from which 
the above calculations for Curacao and Ja- 
maica were made. In it, each trematode 
family is analyzed according to the number 
of its species found in various host species, 
genera and families. Although certain trem- 
atode families are represented by only one 
or a few species, the table shows in general 
that when host species are grouped into 
higher taxa, viz., genera and families, the 
number of trematode species reported from 
hosts belonging to more than one such taxon 
progressively decreases. On that basis, he- 
miurid species are least host specific whereas 
the lepocreadiids show the opposite extreme. 
The striking difference between those groups 
probably is correlated with the localization 
of the majority of hemiurids in the stomachs 
of their hosts whereas all of the lepocreadiids 
reported in this study are intestinal parasites. 
Another factor is how much the parasite 
must grow and develop to mature after 
reaching the host; in this case it is less in the 
hemiurids than the lepocreadiids. A group 
with even more advanced metacercariae than 
the hemiurids is the microphallids, of which 
one species, previously known only from 
birds, is here reported from snake-eels. Until 
trematode life histories and the food habits 
of potential hosts are better known, it will 
be impossible to distinguish trematodes that 
are truly host-specific from those that do 
not have the opportunity to infect more than 


Tulane Studies in Zoology 


Vol. 11 


a few of the potential available hosts. 
Enough is known, however, to indicate rather 
clearly that larval trematodes are usually 
more specific in their molluscan hosts than 
are adults with respect to vertebrates. 


VII. ALPHABETICAL HOstT-PARASITE LIST 


Following each host species is the number 
of individuals examined from Curagao (C) 
and/or Jamaica (J). For each parasite spe- 
cies, the number of infected fish is given for 
one or both localities. A dagger (+) indi- 
cates a trematode species found in one lo- 
cality but not the other where the number 
of potential hosts examined indicates rarity 
if not absence of the parasite. 


(6) J 
Abudefduf saratilis (Linnaeus), sergeant 
major 12 22 
+Genolinea noblei 1 
Lecithochirium parvum 1 
Parahemiurus merus 2 
Schikhobalotrema adacuta 1 
tSchikhobalotrema bivesiculum 1 
Acanthurus bahianus Castelnau, ocean tang 
Dichadena acuta 
Schikhobalotrema obtusa 
Acanthurus coeruleus Bloch & Schneider, 
blue tang 5 
Dichadena acuta 4 
Hapladena varia 5 
Hysterolecitha sogandaresi 1 
Mesolecitha linearis i 
Acanthurus hepatus (Linnaeus), doctor fish 24 22 
Dichadena acuta 13 
Hapladena varia 4 
Huysterolecitha rosea 
Macradena perfecta 1 
Monorchimacradena acanthuri 8 
Schikhobalotrema obtusa 2 
Achirus lineatus (Linnaeus), striped sole 
Sterrhurus musculus 
Angelichthys ciliaris (Linnaeus), queen 
angelfish 1 
Glyphicephalus candidulus 
Anisotremus virginicus (Linnaeus), porkfish 1 
Brachadena pyriformis 
Diphtherostomum anisotremi 
Diplangus paxillus 
Hamacreadium oscitans 
Infundibulostomum anisotremi 
Lasiotocus longicaecum 1 
Leurodera decorum 
Proctotrema anisotremi 
Stephanostomum sentum 
Archosargus unimaculatus (Bloch), brim 
Brachadena pyriformis 
Diplomonorchis micropogoni 
Hamacreadium oscitans 
Megasolena archosargi 
Multitestis rotundus 
Pachycreadium crassigulum 
Aulostomus maculatus Valenciennes. 
trumpet fish 9 
Stephanostomum aulostomi 3 
Balistes ringens Linnaeus, cocuyo a 
Pscudocreadium galapagoensis 
Balistes vetula Linnaeus, queen triggerfish 2 
Apocreadium balistis 
Neoapocreadium coili 
Pseudocreadium lamelliforme 
Xustretrum solidum 
Bathystoma aurolineatum (Cuy. & Val.), 
yellow tomtate 
Hamacreadium ositans 
Lasiotocus longovatus 
Lasiotocus truncatus 
Bathystoma striatum (Linnaeus), common 
tomtate 
Brachadena pyriformis 
Genolopa ampullacea 
Lecithochirium parvum 


ol ell) 


NO NEO RH OR CRORE WH he ly 


= 
HOH BEEN HNeHT 


No. 4 


Bodianus rufa (Linnaeus), Spanish hogfish 


Lepocreadium bimarinum 
Brachygenys chrysargyreus (Giinther), 
Bronze grunt 
Homalometron foliatum 
Lasiotocus truncatus 
Calamus arctifrons Goode & Bean, grass 
porgy 
Lepocreadium opsanusi 
Pachycreadium crassigulum 
Proctoeces lintoni 
Pseudocreadium anandrum 
Calamus bajanado (Bloch & Schneider), 
jolt-head porgy 
Brachadena pyriformis 
Cotyloyaster basiri 
Lepocreadium opsanusi 
Pachycreadium crassigulum 
Proctoeces lintoni 
Pscudocreadium anandrum 


Calamus calamus (Cuy. & Val.), saucer-eye 
’ 


porgy 
ITamacreadium oscitans 
Lasiotocus truncatus 


Cantherines pullus (Ranzani), gray filefish 


Apocreadium (immature) 
Diploproctodaeum haustrum 
Megapera pseudogyrina 
Xuystretrum solidum 
Canthigaster rostratus (Bloch), sharp- 
nosed puffer 
Xystretrum-solidum 
Caranez bartholomuaei (Cuy. & Val.), 
yellow jack 
Alcicornis carangis 
Bucephalus varicus 
Ectenurus americanus 
Hetenurus virgulus 
Genolopa brevicaecum 
Lecithochirium parvum 
Stephanestomum ditrematis 
Tergestia acuta 
Tergestia pectinata 


Carane chrysos (Mitchill), hard-tailed jack 


Bucephalus varicus 
Betenurus americanus 
Parahemiurus merus 
Pseudopecoeloides carangi 
Stephanostomum ditrematis 
Tergestia acuta 
Caranz hippos (Linnaeus), common jack 
Bucephalus varicus 
Betenurus americanus 
Betenurus virgulus 
Lecithochirium parvum 
Parahemiurus merus 
Stephanostomum ditrematis 
Tergestia pectinata 
Carangz latus Agassiz, horse-eye jack 
Bucephalus varicus 
Lecithochirium parvum 
Parahemiurus merus 
Stephanostomum ditrematis 
Stephanostomum megacephalum 
Stephanostomum sentum 
Tergestia pectinata 
Carang ruber (Bloch), skip-jack 
Alcicornis carangis 
Bucephalus varicus 
Pseudopecoeloides carangi 
Caranz-sp.. jack 
Bucephalus varicus 
Stephanostomum ditrematis 
Centropomus undecimalis (Bloch), snook 
Bucephalus sp. 
Ceratacanthus schoepfi (Walbaum), 
orange filefish 
Rhagorehis odhneri 
Chaetodipterus faber (Broussonet), 
spadefish 
Allomegasolena spinosa 
Aponurus clongatus 
Gymnotergestia chaetodipteri 
Multitestis blenni 
Multitestis inconstans 


Chaetodon capistratus Linnaeus, four-eyed 


butterfly fish 
Hiurleytrematoides chaetodoni 
tHurleytrematoides curacaensis 
+Multitestis chaectodoni 
Muttitestis rotundus 


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Trematodes of Marine Fishes 


Chaetodon ocellatus Bloch, common 
butterfly fish 
Hurleytrematoides chaetodoni 
Hurleyltrematoides curacaensis 
Chaetodon striatus Linnaeus, banded 
butterfly fish 
Hurleytrematoides chaetodoni 
Multitestis chaetodoni 
Chloroscombrus chrysurus (Linnaeus), 
bumper 
Bucephalus varicus 
Opechona chloroscombri 
Tergestia pectinata 
Clepticus parrae (Bloch & Schneider) 
creole 
Tergestia laticollis 
Coryphaena hippurus Linnaeus, dolphin 
Dinurus barbatus 
Dinurus breviductus 
Dinurus tornatus 
Hirudinella-sp. 
Ntephanostomum coryphaenae 
Tetrochetus coryphaenae 
Decapterus macarellus (Cuy. & Val.), 
mackerel scad 
Chrisomon decapteri 
Diodon hystrix Linnaeus, porcupine fish 
{Diploproctodaeum diodontis 
Opistholebes diodontis 
Sclerodistomum diodontis 
Tetrochetus coryphaenae 
Doratonotus megalepis Giinther, mottled 
sea basslet 
Pseudopecoelus minutus 
Dules dispilurus Giinther, sandfish 
Lecithochirium parvum 


Eeheneis naucrates Linnaeus, shark remora 


Parahemiurus merus 
Echidna catentata Bloch, chained moray 
Diplohurleytrema brevicaecum 
Epinephelus adscensionis (Osbeck), 
rock hind 
Lecithochirium parvum 
Opecoeloides vitellosus 
Postporus epinepheli 
Sterrhurus musculus 
Epinephelus guttatus (Linnaeus), 
red hind 
Postporus epinepheli 
Epinephelus morio (Cuy. & Val.), red 
grouper 
Sterrhurus musculus 
Lepidapedon trachinoti 
Postporus epinepheli 
Epinephelus striatus (Bloch), Nassau 
grouper 
Ectenurus americanus 
Lepidapedon trachinoti 
Postporus epinepheli 
Scaphanocephalus sp. 
Stephanostomum dentatum 
EBques acuminatus (Bloch & Schneider), 
ecubbyu 
Horatrema crassum 
Pseudopecoeloides equesi 
Eques punctatus Bloch & Schneider, 
ribbonfish 
Horatrema crassum 
Pseudopecoeloides equesi 
Bucinostomus pseudogula (Cuy. & Val.), 
mojarra 
Brachadena pyriformis 
Pseudohurleytrema eucinostomi 
Gerres cinereus (Walbaum), gray mojarra 
Crassicutis gerridis 
Crassiculis marina 
Diplangus pavillus 
+Homalometron elongatum 
Pinguitrema lobata 
Postmonorchis orthopristis 
Pseudohurleytrema eucinostomi 
Steganoderma hemirhamphi 
Stephanostomum sentum 
Gymnothoraxe funebris Ranzani, green 
moray 
Dictysarca virens 
Dollfustrema muraenae 
Gyumnothorax moringa (Cuvier), spotted 
moray 
{Dictysarea virens 
Dollfustrema macracanthum 
Dollfustrema muraende 


we oo 


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mre bo bok 


> He Ot 


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14 


Re eo RA 


13 
9 


aw oo 


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Pscudopecoelus guymnothoracis 
Sterrhurus fusiformis 
Stomachicola rubea 
Gymnothorar vicinus (Castelnau), brown 
moray 
Diectysarca virens 
Dollfustrema gumnothoracis 
Dollfustrema macracanthum 
Dollfustrema muraenae 
Sterrhurus fusiformis 
Stomachicola rubea 
Haemulon album (Cuv. & Val.), 
margate fish 
Genolopa ampullacea 
Hamacreadium oscitans 
Homalometron foliatum 
Lasiotocus truncatus 
Postmonorchis orthopristis 
Proctotrema pritchardi 
Stephanostomum sentum 
Sterrhurus musculus 
Haemulon bonariense (Cuy. & Val.), 
black grunt 
Brachadena pyriformis 
Genolopa ampullacea 
Hamacreadium oscitans 
Lasiotocus longovatus 
Lasiotocus truncatus 
Haemulon flavolineatum (Desmarest), 
yellow grunt 
Brachadena pyriformis 
Diplangus pacillus 
Genolopa ampullacea 
Homalometron foliatum 
Lasiotocus longovatus 
Lasiotocus truncatus 
jLeurodera decora 
Postmonorchis orthopristis 
Haemulon melanurum (Linnaeus), 
black-tailedgrunt 
Genolopa ampullacea 
Hamacreadium oscitans 
Hlaemulon sciurus Shaw, 
blue-striped grunt 
Brachadena pyriformis 
Diplangus parvus 
Diplangus paxillus 
Genolopa ampullacea 
Hamacreadium consuetum 
Hamacreadium oscitans 
Helicometrina nimia 
Homalometron foliatum 
Lasiotocus longovatus 
Lasiotocus truncatus 
Leurodera decora 
Postmonorchis orthopristis 
Stephanostomum sentum 
Sterrhurus musculus 
Halichoeres pictus (VPoey), 
painted doncella 
Helicometra execta 
Schikhobalotrema adacuta 
SLED ars brasiliense (Linnaeus), 
balao 
Haplosplanchnoides hemirhamphi 
+Lepocreadium hemirhamphi 
Schikhobalotrema adacuta 
Steganoderma hemirhamphi 
Hepsetia stipes (Miiller & Troschel), 
hard-head silverside 
Schikhobalotrema adacuta 
tSteganoderma atherinae 
Holacanthus tricolor (Bloch), 
rock beauty 
yAntorchis holacanthi 
Cleptodiscus havanensis 
Holocentrus ascensionis (Osbeck), 
squirrel fish 
Telicometra equilata 
Lecithochirium parvum 
Lepidapedon truncatum 
Pseudopecoelus barkeri 
{Pseudopecoelus holocentri 
Stephanostomum pseudocarangis 
Sterrhurus musculus 
ITolocentrus verillarius (Poey), 
squirrel fish 
Pseudopecoelus barkeri 
Sterrhurus musculus 
Hypoplectrus unicolor indigo (Poey), 
butter hamlet 
Helicometrina nimia 


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Tulane Studies in Zoology 


Hypoplectrus unicolor (Walbaum), 
butter hamlet 
Neolepidapedon hypoplectri 
Kyphosus sectatriz (Linnaeus), 
white chub 
Cadenatella kyphosi 
Labrisomus bucciferus Poey, blenny 
Coitocaeccum sp. 


Vol. 11 


c 


Lachnolaimus marimus (Walbaum), hogfish 


Helicometrina nimia 
Myzoxrenous lachnolaimi 
Lactophrys bicaudalis (Linnaeus), 
trunkfish 
Megapera gurina 
Thysanopharyne elongatus 
Lactophrys tricornis (Linnaeus), 
common trunkfish 
7Dermadena lactophrysi 
Megapera gyrina 
Proctoeces maculatus 
Pseudocreadium lactophrysi 
Xystietum solidum 
Lactophrys trigonus (Linnaeus), 
trunkfish 
Dermadena lactophrysi 
Neoapocreadium angustum 
Pseudocreadium lactophrysi 
Lactophrys triqueter (Linnaeus), 
trunkfish 
Dermadena lactophrysi 
Megapera gyrina 
Pseudocreadium lactophrysi 
Leptocephalus conger Linnaeus, conger eel 
Sterrhurus musculus 
Lutianus analis (Cuy. & Val.), 
muttonfish 
Metadena crassulata 
Siphodera vinaledwardsii 
Lutianus apodus (Walbaum), 
schoolmaster 
Allomegasolena spinosa 
jHamacreadium mutabile 
Lecithochirium parvum 
Metadena adglobosa 
Metadena globosa 
Sterrhurus musculus 
Lutianus aya (Bloch), West Indian 
red snapper 
Lecithochirium parvum 
Metadena adglobosa 
Metadena crassulata 
Metadena globosa 
Paracryptogonimus neoamericanus 
Siphodera vinaledwardsii 
Ntephanostomum casum 
Sterrhurus musculus 
Lutianus buccanella (Cuy. & Val.), 
black-finned snapper 
Siphodera vinaledwardsit 
Stephanostomum casum 


Lutianus griseus (Linnaeus), gray snapper 


Hamacreadium mutabile 
Lecithorchirium parvum 
Metadena adglobosa 
Sterrhurus musculus 
Lutianus jocu (Bloch & Schneider), 
dog snapper 
TTamacreadium mutabile 
Helicometrina nimia 
Metadena adglobosa 
Lutianus mahogani (Cuy. & Val.), 
mahogany snapper 
Homalometron foliatum 
Lasiotocus truncatus 
Lutianus synagris (Linnaeus), 
lane snapper 
Metadena adglobosa 
Siphodera vinaledwardsii 
Stephanostomum casum 
Lutianus sp. 
Stephanostomum sentum 
Matacanthus plumieri (Bloch), sandfish 
Sterrhurus musculus 
Micropogon furnieri (Desmarest), croaker 
Diplomonorchis hopkinsi 
Diplomonorchis micropogoni 
Lobatestoma ringens 
Microspathedon chrysurus (Cuy. & Val.), 
yellow-tailed demoiselle 
Schikhobalotrema pomacentri 
Monacanthus hispidus (Linnaeus), 
efish 


ee 


_ 
a 


DE a fk ek ek pk 


me Col 


if 


= RPh 


me bo 


Re 


ie’ 2) 


~ eS Re 


Ge 


Hee 


ll ell eh od 


CIR HD 


Cl ett OCOD 


No. 4 


Apocreadium mexicanum 
Mugil cephalus Linnaeus, striped mullet 2 
Haplosplanchnus mugilis 1 
Schikhobalotrema elongatum Za 
Mugil curema (Cuy. & Val.), white mullet 10 
jHaplosplanchnus mugilis 9 
Schikhobalotrema elongatum 
Mycteroperca bonaci (Poey), black grouper 
Deretrema fusillus 
Neolepidapedon muycteropercae 
Prosorhunchus atlanticum 
Prosorhynchus ozakii 
Ntephanostomum dentatum 
Stephanostomum ditrematis ; 
Mycteroperca falcata (Poey), scamp 
Prosorhynchus atlanticum 
Mycteroperca venonosa (Linnaeus), 
yellow-fin grouper 
Neolepidapedon mycteropercae 
Prosorhynchus atlanticum 
Myrichthys acuminatus (Gronow), 
sharp-tailed eel 
Carneophallus lactophrysi 
Myrichthys oculatus (Kaup), black-spotted 
snake eel 
Carneophallus lactophrysi 
Microphallus excellens 
Myripristis jacobus (Cuy. & Val.), big-eyed 
squirrel fish 
Bivesicula caribbensis 
Myrophis punctatus Lutken, speckled 
worm eel 
Diplomonorchis myrophitis 
Ocyurus Chrysurus (Bloch), yellowtail ial 
Lepocreadium trullae 7 
Lepocreadium truncatum ail 
Metadena globosa 
Paracryptogonimus neoamericanus 4 
Prosogonotrema bilabiatum 
Siphodera vinaledivardsii 2 
Oligoplitis saurus (Bloch & Schneider), 
leather-jacket 
Manteria brachydera 
Tergestia pectinata 
Opisthonema oglinum (Le Sueur), 
thread-fin herring 
Bacciger opisthonemae 
Neogenolinea opisthonemae 
Parahemiurus merus 
Tergestia pectinata 
Peprilus paru (Linnaeus), harvest fish 
Lepocreadium pyriforme 
Platophrys lunatus (Linnaeus), peacock 
flounder 
Helicometrina nimia 
Sterrhurus musculus 
Pomacanthus arcuatus (Linnaeus), black 
angelfish 
Antorchis urna 
Glyphicephalus candidulus 
Phyllodistomum pomacanthi 
Pomacanthus paru (Bloch), French 
angelfish 
Antorchis urna 
Pomacentrus analis Poey, blue-spotted 
demoiselle 
Schikhobalotrema pomacentri 
Pomacentrus fuscus Cuy. & Val., brown 
demoiselle 
Schikhobalotrema pomacentri 
Pomacentrus leucostictus Miill. & Trosch., 
beau-gregory 
Schikhobalotrema pomacentri 
Theletrum pomacentri 
Priacanthus cruentatus (La Cépede), big-eye 
Betenurus virgulus 
Prionotus punctatus (Bloch), spotted 
sea-robin 
Sterrhurus musculus 
Promicrops itaiara (Lichtenstein), jewfish 
Prosorhynchus promicropsi 
Pseudoscarus guacamaia (Cuvier), rainbow 
parrotfish 
Hapladena ovalis 
Schikhobalotrema adbrachyura 
Schikhobalotrema heterocotylum 
Schikhobalotrema sparisomae 
Pseudoscarus plumbaecus Bean, purple 
parrotfish 
Schikhobalotrema adbrachyura 
Rypticus saponaceus (Bloch & Schn.), 
soapfish 


EE EEO REE on 


= 


Ore rhb cho 


Rho 


Be wor NR eB Ee 


bo 


BPO PR WO 


Como 


co 


mC Ot 


= 


ow) 


So EEOC et 


Eh © PHO parents 


iw) 


Trematodes of Marine Fishes 


Prosorhynchus aquayoi 1 
Sterrhurus musculus 1 
Sardinella anchovia (Cuy. & Val.), Spanish 
sardine 6 
tN eoyenolinea opisthonemae 
Parahemiurus merus 1 
Sardinella macrophthalmus (Ranzani), 
big-eyed sardine 6 
Ectenurus virgulus 
Lecithochirium parvum 
Neogenolinea opisthonemae 
Opechona sardinellae 
Parahemiurus merus 
Pseudobacciger manteri 
Scarus croicensis (Bloch), Bahama 
parrotfish 6 
Schikhobalotrema sparisomae 3 
Scarus sp. 
Schikhobalotrema sparisomae 
Scomberomorus cavalla (Cuy. & Val.), king 
mackerel 2 
Rhipidocotyle baculum 1 
_ Bucephatloides arcuatus 1 
Scorpaena plumieri Bloch, West Indian 
scorpion fish 1 
Bucephalus scorpaenae 1 
Hirudinella sp. 
Lecithochirium parvum 
tNeopecoelus scorpaenae 10 
Sterrhurus musculus 
Selar crumenophthalmus (Bloeh), 
goggle-eyed scad 
Chrisomon sp. 
Hetenurus americanus 
Betenurus virgulus 
Lecithochirium microstomum 
Lecithochirium parvum 
Pseudopecoeloides gracilis 
Sterrhurus musculus 
Tergestia pectinata 
Seriola dumerili (Risso). great amberjack 1 
Bucephalus varicus 
Hetenurus americanus 
Lecithochirium microstomum 


fon) 


Lecithochivium parvum if 
Parahemiurus merus 
Tormopsolus orientalis 1 
Sparisoma abildgaardi (Bloch), red 
parrotfish 2 
Schikhobalotrema sparisomae 1 


Sparisoma brachiale (Poey), parrotfish 
Hapladena ovalis 
Schikhobalotrema sparisomae 
Sparisoma flavescens (Bloch & Schn.), 


mud parrottish 25 
Hapladena ovalis 
Schikhobalotrema adbrachyura 3 


Schikhobalotrema sparisomae 
Sparisoma radians (Cuy. & Val.), radiant 
parrotfish 
Schikhobalotrema sparisomae 
Sparisoma viride (Bonnaterre), green 
parrottish 1 
Schikhobalotrema adbrachyura 
Spheroides spengleri (Bloch), southern 
swellfish 
Diploproctodaeum plicitum 
Helicometrina nimia 
Spheroides testudineus (Linnaeus), 

West Indian puffer 4 
Diplomonorchis sphaerovarium 
Diploproctodaeum plicitum 
Xystretrum solidum 


Sphyraena barracuda (Shaw), barracuda 6 
Bucephaloides longicirrus ul 
Bucephaloides longoviferus 2 


Strongylura ardeola (Cuyv. & Val.), needlefish 1 
Schikhobalotrema acuta 
Steganoderma nitens 

Strongylura raphidoma (Ranzani), 


houndfish 
Schikhobalotrema acuta 
Strongylura timucu (Walbaum), timucu 16 
Schikhobalotrema acuta 1 
Steganodcrma atherinae 1 
Synodus intermedius (Agassiz), lizardfish 3 


Betenurus americanus 

Lecithochirium microstomum 

Lecithochirium parvum 

Sterrhurus musculus 2 
Trachinotus glaucus (Bloch), palometa 

Betenurus virgulus 


J 


=) 


Wake TO= 


2 


hobo bot 


en 


eee 


i 
OVS 


> 


w 


Soe 


Lal | 


224 


Helicometrina trachinoti 
Sterrhurus musculus 


Upeneus maculatus Bloch, red goatfish 6 4 
Opecoeloides brachyteleus 1 2 
Opecocloides elongatus 5 3 

Upeneus martinicus Cuy. & Val., yellow 

goatfish 13 9 
Lecithochirium parvum 1 
Opecoeloides brachyteleus 4 
Opecoeloides elongatus 9 5 


VIII. List OF NEGATIVE FISHES 


Numbers preceding the letters C and J 
are of individuals examined in Curacao and 
Jamaica, respectively. 


Albula vulpes (Linnaeus), bonefish, 3C 

Anchoviella epsetus (B onnaterre), striped 
fee Alay) 

Antennarius ocellatus (Bloch & Sehn.), 
frogfish, 1C 

Apogon binotatus (Poey), cardinal fish, 2C 

Apogon conn (Silvester), Conklin’s cardinal 
fish, 

Apogon ae ulatus (Poey), 

Apogon sp., 2C 

Apogonichthys stellatus Cope, conehfish, 1C 

Bathygobius soporator (Cuyv. & Val.). mapo, 
19C, 63 

Cephalacanthus volitans (Linnaeus), flying 
gurnard, 4C 

Cephalopholis fulvus (Linnaeus), 

Ceratacanthus scripta (Osbeck), 
filefish, 2J 

Cetengraulis edentulus (Cuvier), 
anchovy, 17J 

Chilomycterus atinga (Linnaeus), 
boxfish, 2J 

Chromis marginatus (Castelnau), reef-fish, 19C 

Cypselurus bahiensis (Ranzani), flyingfish, 8C 

Decapterus punctatus Agassiz round sead, 2C 

Diapterus rhombeus (Cuy. & Val.), rhomboild 
mojarra, 8J 

Bleotris perniger (Cope), goby, TJ 

Elops saurus Linnaeus, ten-pounder, 5C 

Protelis smaragdus (Cuv. & Val.). emerald goby, 3C 

Etropus crossotus Jordan & Gilbert, fringed 
flounder, 2J 

Fistularia tabacaria Linnaeus, cornet fish, 1C 

Gramma hemichrysos, royal gramma, 1C 

Gymnothorae ocellatus Agassiz, ocellated moray, 
3C 

Haemulon macrostomum Giinther. 

Halichoeres bivittata (Bloch), 

Halichoeres kirschii (Jord. 
wrasse, 38C 

Hippocampus punctulatus Guichenot, spotted 
seahorse, 1C, 1J 

Hirundichthys affinis (Giinther), 
flyingfish, 1C 

Hypoplectrus unicolor nigricans (Poey), vaca, 10 

Labrisomus nuchipennis (Quoy & Gaim.), hairy 
blenny, 9C, 6J 

Lophogobius cyprinoides (Pallas). crested goby, 67 

Lophogobius glaucofraenum (Gill), bridled goby, 5C 

Microgobius sp., goby, 4C 

Monacanthus tuckeri Bean, Tucker's filefish, 5C 

Mollinesia vandepolli (Van Lindth de Jeude), 
killifish, 4C 

Odontoscion dentex (Cuy. & Val. ), corvina, 1J 

Ogeocephalus vespertilio (Linnaeus), batfish, 1C 

Ophichthus sp., snake eel, 1C 

Opisthognathus aurifrons (Jord. 
jawfish, 1C 


anchovy, 


spotted 


cardinal fish, 9C 


coney, 5C, 1J 
scrawled 


whalebone 


spotted 


grunt. 1J 
dick, 7C 
Kirsche’s 


gray 
slippery 
& Ever.), 


four-winged 


& Thomp.), 


Pempheris muclleri Poe y. glassy pempherid, 2C 
Priacanthus arenatus Cuy. & Val., common big-eye, 
4C 


Prionodes tigrinus (Bloe h), 

Rupiscar te s atlanticus (Cuy. 
| OR ar’. 

Ryptic us bistrispinus (Mitechill), 
soapfish, 2J 

Scarus caeruleus (Bloch). 

Stegastes niveatus (Poey), 
demoiselle, 1C 

Strongylura notatus (Voey), needlefish, 1J 

Syngnathus elucens Poey, Poey’s pipefish, 11J 

Thalassoma bifasciatum (Bloch), blue-head, 


harlequin 
& Val.), 


serranid, 1C 
rock-skipper, 


two-spined 


blue parrotfish, 1J 
turquoise-spotted 


12C 


Tulane Studies in Zoology 


Vol. 11 


IX. LITERATURE CITED 


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BIGUET, J., DEBLOCK, S. and CAPRON, A. 
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CABLE, R. M. 1954a. Studies on the marine 
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: 1954b. Studies on the 
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Pee eee eee 5 . 1956a. Opistholebes dio- 
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the allocreadioid problem. Parasitology 
46: 1-13. 


a 1956b. Marine cercariae 
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CABLE, R. M. 1962. A cercaria of the trema- 
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1963. Marine cercariae 

of Curacao and Jamaica. Zeitschr. f. Par- 

asitenk. 23: 429-469. 


ie . and NAHHAS, F. M. 1962. 
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Digenea) and its life history. J. Parasi- 
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Linton, 1910 sue ce Hemiuridae). 
Proc. Helm. Soc. Washington 30: 206-210. 


Douirus, R. PH. 1946. Sur trois espéces de 
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ee _. 1955. Some digenetic 
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No. 4 


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415-428. 

LA Rug, G. R. 1957. The classification of 
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LINTON, E. 1898. Notes on trematode para- 
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507-548. 

=n 1900. Fish parasites col- 
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nn 1901. Parasites of fishes 
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ee 1905. Parasites of fishes 

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me 1907. Notes on parasites 
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33 785-126. 


—. 1910. Helminth fauna 
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~~ fishes mainly from the Woods Hole re- 
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are _... 1918. Notes on the ge- 
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MAnNTER, H. W. 1931. Some digenetic tre- 
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1 


1940a. Digenetic trema- 
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. 1940b. The geographical 
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marine fishes of the tropical American 


Pacific. Rep. Allan Hancock Pac. Exp. 
Zee Sole pAle 

. 1940c.  Gasterostomes 
~ (Trematoda) of Tortugas, Florida. Car- 
negie Inst. Washington Pub. No. 524: 


1-19. 

en 1942. Monorchidae (Tre- 
‘matoda) ‘from fishes of Tortugas, Flori- 
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360. 

eee IAD ID Enmaoena LaCto= 
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tol; oleAlle4ai7. 

4 . 1947. The digenetic tre- 

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Trematodes of Marine Fishes 225 


1949. An additional tre- 
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Sept Ret oe es 1954. Some digenetic tre- 
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: . 1955. The zoogeography 
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= = 1957.) lost sspeciucihy 
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fe SRE ee 1963b. Studies on dige- 
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fb SR Na ae ONS and PRITCHARD, MARY 
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MeEuRA, H. R. 1961. New Superfamily Hap- 
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NICHOLL, W. 1914. The trematode parasites 
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PEREZ VIGUERAS, I. 1940a. Macrorchitrema 
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kde ANGEL EE te Ee 1940b. Prosogonotremi- 
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226 Tulane Studies in Zoology 


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=i ean ae 
1959c. Studies on helminth “parasites 
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(18%: 


Vol. 11 


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vertebrates, in 2 parts, New York. 1575 
pp. 


X. ABSTRACT 


Examination of 1527 fishes repre- 
senting 185 species yielded 178 species 
of trematodes including 39 new spe- 


cies, for 5 of which new genera are 
erected. New species in previously 
known genera, their hosts, localities 


(C = Curagao, J = Jamaica), and fam- 
ilies are: Dollfustrema gymnothoracis 
from Gymnothorax vicinus, C, (Buce- 
phalidae); Infundibulostomum aniso- 
tremi from Anisotremus virginicus, J; 
and Bacciger opisthonemae from Opis- 
thonema oglinum, J, (Fellodistomati- 
dae); Haplosplanchnus mugilis from 
Mugil curema, C; Schikhobalotrema 
bivesiculum from Abudefduf saxatilis, 
J; S. elongatum from Mugil cephalus 
and M. curema, C, J; and S. hetero- 
cotylum from Pseudoscarus guacamaia, 
C, (Haplosplanchnidae); Megapera 
pseudogyrina from Cantherines pullus, 
We (Megaperidae) ; Crassicutis gerri- 


dis from Gerres cinereus, C, J; Neo- 
lepidapedon hypoplectri from ” Hypo- 
plectrus unicolor, J; Opechona chloro- 


scombri from Chloroscombrus  chry- 
surus, J; O. sardinellae from Sardi- 
nella macrophthalmus, J; Lepocreadi- 
um truncatum from Ocyurus chrysurus, 
C; L. hemiramphi trom Hemiramphus 
brasiliensis, C; Cadenatella kyphost 
from Kyphosus sectatrix, C; Diploproc- 
todaeum diodontis from Diodon hys- 
trix, J; and Pseudocreadium lactophry- 
si from Lactrophrys tricornis, C, J, 
(Lepocreadiidae) ; Pseudopecoelus holo- 
centri from Holocentrus ascensionis, C; 
P. gymnothoracis from Gymnothorax 
moringa, C; and P. minutus from Dora- 
tonotus megalepis, C, (Opecoelidae) ; 
Phyllodistomum pomacanthi from Po- 


No. 4 


Trematodes 


macanthus arcuatus, J, (Gorgoderidae) ; 
Diphtherostomum anisotreni from Ani- 
sotremus virginicus, J, (Zoogonidae) ; 
Proctotrema pritchardae from Haemu- 
lon album, C; P. anisotremi from Ani- 
sotremus virginicus, J; Chrisomon de- 
capteri from Decapterus macarellus, C; 
Hurleytrematoides curacaensis from 
Chaetodon capistratus, C; Diplomonor- 
chis myrophitis from Myrophis puncta- 
tus, J; D. micropogoni from Micropo- 
gon furnieri and Archosargus unimacu- 
latus, J; D. hopkinsi from Micropogon 
furniert, J; and D. sphaerovarium from 
Spheroides testudineus, J, (Monorchii- 
dae); Stephanostomum aulostomi from 
Aulostomus maculatus, C, (Acanthocol- 
pidae) ; Genolinea noblei from Abudef- 


of Marine Fishes 


macradena acanthuri from Acanthurus 
hepatus, C, J (Hemiuridae). 

New names for previously misidenti- 
fied species are: Pseudobacciger man- 
teri, Pseudopecoelus holocentri (listed 
above), and Alicornis siddiqii (Puerto 
Rico). New combinations are: Clepto- 
discus havanensis for Macrorchitrema 
h., Cadenatella brumpti for Jeancade- 
natia b., and Cadenatella dohenyi for 
Jeancadenatia d. 

The following species are reduced to 
synonomy as indicated: Proctoeces neo- 
magnoris = Mesolecitha linearis; Lepi- 
dapedon holocentri = L. truncatum; 
Pseudoplagioporus brevivitellus = Ha- 
macreadium oscitans; Steganoderma 
elongatum = S. nitens; Genolopa lon- 


227 


duf saxatilis, C; Hysterolecitha sogan- 
daresi from Acanthurus coeruleus, J; 
and Theletrum pomacentri from Poma- 
centrus leucosticus, J, (Hemiuridae). 


New genera and species, their hosts, 
localities and families are: Gymnoter- 
gestia chaetodipteri from Chaetodip- 
terus faber, J; Pseudobacciger man- 
teri from Sardinella macrophthalmus, J 
(Fellodistomatidae) ; Haplosplanchnoi- 
des hemiramphi from Hemiramphus bra- 
siliensis, J (Haplosplanchnidae) ; Diplo- 
hurleytrema brevicaecum from Hchidna 
catenata, C (Monorchiidae) ; Monorchi- 


gicaudata = G. ampullacea; and Apo- 
nurus symmetrorchis = Brachadena py- 


riformis. 


Possible affinity of the Enenterum 
group with the Haploporoidea is dis- 
cussed. The Suborder Acanthocolpiata 
and Superfamily Acanthocolpoidea are 
proposed for the Family Acanthocol- 
pidae, and placed in the Order Opis- 


thorchiida, Superorder 
dia of La Rue’s system. 


Epitheliocysti- 


Data concerning geographical distri- 
bution and host-parasite specificity are 


presented in tables and discussed. 


XI. INDEX OF TREMATODE SPECIES 
(New species and genera in boldface) 


Alcicornis carangis, 171 
siddiqu, 172 

Allomegasolena spinosa, 179 

Antorchis holacanthi, 174 
urna, 174 

Apocreadium balistis, 190 
mexicanum, 190 

Aponurus elongatus, 213 


Bacciger opisthonemae, 177 
Bivesicula caribbensis, 178 
Brachadena pyriformis, 212 
Bucephaloides arcuatus, 173 
longicirrus, 173 
longoviferus, 173 
Bucephalus scorpaenae, 171 
varicus, 170 
Spee 


Cadenatella kyphosi, 190 
Carneophallus lactophrysi, 184 
Chrisomon decapteri, 202 
Cleptodiscus havanensis, 178 
Coitocaecum sp., 198 
Cotylogaster basiri, 170 
Crassicutis gerridis, 184 
marina, 184 


Deretrema fusillum, 199 
Dermadena lactophrysi, 194 
Dichadena acuta, 213 
Dictysarca virens, 216 


Dinurus barbatus, 212 
breviductus, 212 
tornatus, 212 
Diplangus parvus, 199 
paxillus, 199 
Diphtherostomum anisotremi, 200 
Diplohurleytrema brevicaecum, 205 
Diplomonorchis hopkinsi, 206 
micropogoni, 2(6 
myrophitis, 206 
sphaerovarium, 206 
Diploproctodaeum diodontis, 193 
haustrum, 192 
plicitum, 192 
Doilfustrema gymnothoracis, 172 
macracanthum, 172 
muraenae, 172 


Ectenurus americanus, 212 
virgulus, 212 


Genolinea noblei, 212 

Genolopa ampullacea, 200 
brevicaecum, 201 

Glyphicephalus candidulus, 179 

Gymnotergestia chaetodipteri, 176 


Hamacreadium consuetum, 194 
mutabile, 194 
oscitans, 194 

Hapladena ovalis, 179 
varia, 179 


228 Tulane Studies in Zoology 


Haplosplanchnoides hemiramphi, 183 


Haplosplanchnus mugilis, 179 

Helicometra equilata, 195 
execta, 195 

Helicometrina nimia, 195 
trachinoti, 195 

Hirudinella sp., 216 

Homalometron elongatum, 184 
foliatum, 184 

Horatrema crassum, 195 

Hurleytrematoides chaetodoni, 204 
curacaensis, 205 

Hysterolecitha rosea, 213 
sogandaresi, 213 ' 


Infundibulostomum anisotremi, 176 


Lasiotocus longicaecum, 201 
longovatus, 201 
truncatus, 201 
Lecithochirium microstomum, 212 
parvum, 212 
Lepidapedon trachinoti, 186 
truncatum, 186 
Lepocreadium bimarinum, 188 
hemiramphi, 189 
opsanusti, 189 
pyriforme, 188 
trulla, 188 
truncatum, 189 
Leuwrodera decora, 213 
Lobatostoma ringens, 170 


Macradena perfecta, 213 
Manteria brachydera, 211 
Megapera gyrina, 183 
pseudogyrina, 183 
Megasolena archosargi, 179 
Mesolecitha linearis, 174 
Metadena adglobosa, 209 
crassulata, 209 
globosa, 209 
Microphallus excellens, 184 
Monorchimacradena acanthuri, 214 
Multitestis blenni, 187 
chaetodoni, 187 
inconstans, 187 
rotundus, 187 
Myzoxenous lachnolaimi, 190 


Neoapocreadium angustum, 190 
coili, 190 

Neogenolinea opisthonemae, 212 

Neolepidapedon hypoplectri, 186 
mycteropercae, 186 

Neopecoelus scorpaenae, 195 


Opechona chloroscombri, 187 
sardinellae, 188 

Opecoeloides brachyteleus, 195 
elongatus, 195 
vitellosus, 195 

Opistholebes diodontis, 198 


Pachycreadium crassigulum, 198 


Paracryptogonimus neoamericanus, 209 


Parahemiurus merus, 211 
Phyllodistomum pomacanthi, 198 
Pinguitrema lobatum, 195 
Postmonorchis orthopristis, 204 
Postporus epinepheli, 190 
Proctoeces lintoni, 174 
maculatus, 174 
Proctotrema anisotremi, 202 
pritchardae, 201 
Prosogonotrema bilabiatum, 216 
Prosorhynchus aguaot, 174 
atlanticum, 174 
ozakii, 174 
promicropsi, 174 
Pseudobacciger manteri, 177 
Pseudocreadium anandrum, 194 
galapagoensis, 194 
lactophrysi, 193 
lamelliformae, 194 
Pseudopecoeloides equesi, 195 
carangi, 195 
gracilis, 195 
Pseudopecoelus barkeri, 196 
gymnothoracis, 196 
holocentri, 196 
minutus, 196 
Pseudohurleytrema eucinostomi, 204 


Rhagorchis odhneri, 190 
Rhipidocotyle baculum, 173 


Scaphanocephalus sp., 209 
Schikhobalotrema acutum, 180 
adacutum, 180 
adbrachyurum, 180 
bivesiculum, 180 
elongatum, 182 
heterocotylum, 182 
obtusum, 180 
pomacentri, 180 
sparisomae, 189 
Sclerodistomum sphoeroidis, 216 
Siphodera vinaledwardsii, 209 
Steganoderma atherinae, 200 
hemiramphi, 199 
nitens, 199 
Stephanostomum aulostomi, 210 
casum, 210 
coryphaenae, 210 
dentatum, 210 
ditrematis, 210 
megacephalum, 210 
pseudocarangis, 210 
sentum, 210 
Sterrhurus fusiformis, 211 
musculus, 211 
Stomachicola rubea, 212 


Tergestia acuta, 174 

laticollis, 174 

pectinata, 174 
Tetrochetus coryphaenae, 216 
Theletrum pomacentri, 214 
Thysanopharyn« elongatus, 183 
Tormopsolus orientalis, 211 


Xystetrum solidum, 198 


TULANE STUDIES IN ZOOLOGY 
VOLUME 11 


(New genera, species, and subspecies in boldface) 


A blabesmyia 

aequifasciata, 113, 150, 158 

basalis, 114 

illinoensis, 113 

janta, 114 

johannseni, 114 

peleensis, 113,157, 159, 161 

ramphe, 114, 116, 150, 157, 159, 161 
Adocia neens, 48, 67 
Agkistrodon piscivorus, 19 
Akiefferiella, 121 
Alternanthera philoxeroides, 121, 152 
Amblyomma 

maculatum, 12 

americanum, 12 
Anthosigmella varians, 55, 69 
Apodemus sylvaticus, 20 
Aristida stricta, 5 
Aspiculuris, 19, 21 

americanum, 18, 20 
Aulena columbia, 34 
Austroiceles cruciata, 91 
Axinella polycapella, 51, 67 


Bombyx mori, 91 

Bubaris, 50 

Buckley, Burton H., article, 151-166 
Bullis, Harvey R., Jr., article, 99-107 


Callyspongia, 42 
procumbens, 42, 53 
repens, 42, 43, 67, 71 
vaginalis, 41, 42 
Calopsectra, 146 
Cambarus propinquus, 93 
Campeloma lewisi, 156 
Camptokief feriella, 121 
Capillaria, 21 
hepatica, 18, 19 
Carcinus, 92 
maenas, 79, 80 
Carmia macilenta, 50, 53, 68 
Carya tomentosa, 5 
Chaoborus, 154, 155, 160, 165, 166 
punctipennis, 156 
Chironomus, 125 
abbreviatus, 131 
attenuatus, 123, 157, 159, 161 
brunneipennis, 124 
carinatus, 133, 157, 159, 161 
chaetoala, 133, 150 
chelonia, 124 
digitatus, 129, 131 
adiectus, las, £50, 157, 159, 162) es 
edwardsi, 134, 136, 157, 159, 161, 163, 
165 
emorsus, 138, 150, 157, 159, 161 
fulvipilus, 123, 157, 158, 161, 162 


fulvus, 129, 131, 132, 157, 159, 161 
fumidus, 127 
galeator, 134, 150, 157, 159, 161, 163 
incurvus, 126, 128, 150 
insolita, 124 
malochi, 131 
modestus, 126, 127, 157, 159, 161 
monochromus, 134, 157, 159 
natchitocheae, 124, 125, 150, 157, 159, 
161 
neomeodestus, 127 
nervosus, 126, 127, 157, 159, 161, 162, 
165 
nigricans, 128, 157,159, 161 
nigrovittatus, 138, 157, 159, 161 
parvilamellatus, 131 
ponderosus, 129, 130, 150, 157, 159, 161 
sorex, 130 
stigmaterus, 123 
xenolabis, 128, 157, 159, 161, 163 
Chondrilla nucula, 63, 67 
Cinachyra 
alloclada, 59, 60, 69 
cavernosa, 60 
Cladotaenia, 18 
Cliona 
caribboea, 57, 68 
celata, 57, 68 
lampa, 57, 69 
trwitti, 57, 68 
vastifica, 57, 58, 68 
viridis, 58, 61, 68 
Coelosphaera 
actinoides, 44 
fistula, 44, 45, 67, 71 
tunicata, 44 
Coelotanypus 
concinnus, 112 
scapularis, 112 
tricolor, 111 
Crangon armillatus, 79 
Cranilla 
cinachyra, 60, 62, 63 
crania, 60, 63, 69 
laminaris, 60, 69 
Crassostrea virginica, 57 
Cricotopus 
bicmmetus, 1115, 157, 159, 1695 163 
lebetis, 118, 119, 120, 150 
remus, 115, 120, 150, 157, 161, 162, 165 
sylvestris, 118 
tricinctus, 117, 119 
trifasciatus, 117, 118 
Cryptochironomus, 131, 132 
Ctenophthalmus pseudagyrtes, 13, 14, 15 
Cuterebra 17 
angustifrons, 15 
Cyamon vickersi, 48, 69 


** Volume 11, number 5 which consists of a single paper, DIGENETIC AND ASPIDO- 
GASTRID TREMATODES FROM MARINE FISHES OF CURACAO AND JAMAI- 
CA, is self indexed. The index for the number appears on pages 227-228. 


INDEX TO AUTHORS AND SCIENTIFIC NAMES—Continued 


Darwinella 

australiensis, 37 

joyeuxi, 37, 66 

mullert, 37 
Dermacentor variabilis, 12, 20 
Diataraxia oleracea, 91 
Diospyros virginiana, 5 
Dysidea 

crawshayi, 36, 37 

etheria, 36, 67 
Fimeria, 17, 20 
Endamoeba muris, 17, 20 
Esperiopsis obliqua, 49 
Eukiefferiella, 121 
Eulaelaps stabularis, 6, 7, 10 
Eurypon claratella, 49, 53, 68, 71 
Euryspongia rosea, 36, 66 
Euschongastia peromysci, 6, 8, 20 
Eutrombicula batatas, 8 


Faxonella, 84, 85 
clypeata, 80, 83, 86, 88, 96 
Fingerman, Milton, article, 75-78 


Gahrlepia americana, 6, 8 
Gecarcinus lateralis, 79, 90, 92 
Geodia gibberosa, 43, 44, 48, 51, 59 
Giardia muris, 17, 20 
Gly ptotendipes 
lobiferus, 135, 159, 161 
meridionalis, 135, 157, 159, 161, 164, 165 


Haemogamasus liponyssoides, 6, 7, 10 
Haemolaelaps glasgow, 6, 7, 10, 11, 20 
Halichondria 

bowerbanki, 52, 53 

malanadocia, 54, 67 

panicea, 51, 52, 53, 67 
Haliclona, 41, 42, 67 

erina, 40 

permollis, 40, 51, 67 

rubens, 39, 67 

viridis, 39, 67 
Halicometes stellata, 56, 68 
Halisarca 

dujardini, 39 

magellinica, 39 

purpora, 37, 38, 66, 71 
Heliosoma trivolvis, 156 
Hippiospongia 

gossypina, 34, 66 

lachne, 34, 66 
Holoplocamia 

delaubenfelsi, 45, 47, 68, 71 

penneyt, 48 
Homaxinella waltonsmithi, 51, 69 
Hoplovleura hirsuta, 13, 20 
Hoplopsyllus affinis, 18, 15, 20 
Hymeniaciden heliophila, 55, 68 
Hymenolepis, 21 

nana, 18, 20 
Hymenosoma 

orbiculare, 91 


Tanthella 
ardis, 36, 66 
basta, 36 


Idotea 

balthica, 75 

metallica, 75 
lTotrochota birotulata, 386 
Ircinia 

campand, 35, 66 

lasciculata, 34, 35, 66 

ramosa, 34,66 

strobilina, 35, 66 
Ixodes 

bishoppi, 12 

minor, 12, 13 


Krenosmittia, 121 


Labrundima, 113 

longipalpi, 113 
Larsia, 112 
Lauterborniella 

varipennis, 136, 157, 159, 161 
Layne, James E., article, 3-27 
Leucosolenia 

canariensis, 48, 65, 67 
Ligia 

exotica, 75, 77 

olfersi, 75, 77, 78 
Lissodendoryx isodactyalis, 48, 68 
Little, Frank J., Jr., article, 31-71 
Lysmata seticaudata, 80, 93 


Maia squinada, 80 
Mellita quinquiesperforata, 44 
Merriamium tortugasensis, 48, 52, 53, 68 
Metertetrancyhus ulmi, 91 
Microciona prolifera, 49, 58, 68 
Microcricotopus, 121 
Mobberly, William C., Jr., article 79-96 
Murex, 100 

atlantis, 106-107 

beawi, 99, 104,105 

brevifrons, 99, 105 

cabritu, 101 

cailleti, 108, 104 

carnicolor, 106 

consuelae, 99, 103 

donmoorei, 99, 101, 102, 107 

elenensis, 101 

kugleri, 104 

margaritensis, 105 

messorius, 99, 103 

nuceus, 99, 107 

nuttingi, 106 

oregonia, 99, 102, 106, 107 

pazi, 196 

pomum, 99, 105 

pulcher, 103 

recurvirostris, 103 

ribidus, 1038 

sallasi, 103 

springeri, 99, 102, 104, 105, 107 

thompsoni, 99, 102, 108, 104, 105, 107 

tryoni, 99, 101 
Mycale, 50 


Natrix fasciata, 19 

Nanocladius 
alternantherae, 121, 157, 159, 165 
brevinervis, 121 
sordens, 121 


INDEX TO AUTHORS AND SCIENTIFIC NAMES—Continued 


Nematospiroideas dubius, 24 


Oguro, Chitaru, article, 75-78 
Ondatra zebethicus, 19 
Orconectes 

immunis, 79, 87, 89, 92, 93 

limosa, 80 

rusticus, 79 

virilis, 79, 87, 90, 93, 94 
Ornithonyssus bacoti, 6, 7, 10 
Ostrea permollis, 59 


Pachygrapus crassipes, 79 
Palaemon serratus, 80, 93 
Palaemonetes vulgaris, 77 
Panulivus argus, 91, 93 
Parakiefjeriella, 121 
Paraleuterborniella elachista, 136, 157, 
IL) aleal 
Paratya compressa, 77 
Parazoanthus parasiticus, 42 
Paruterina, 18 
Patuloscula plicifera, 42 
Pedionomus, 137 
beckae, 137, 139, 150, 158, 160, 161 
Pellina, 40 
Pentaneura 
americana, 115 
moiilis, 115 
peleensis, 115 
joulKorsoliliar., Wil, slabs}, itssee WSF alSjal 
planensis, 112, 157, 159, 161 
Peromyscus 
[NOTOCMARMISS Bo Dy WO, Was, Ws, 1G BO; BAe 
el Ol 
gossypinus, 15 
leucopus, 16-19, 21 
Phliseodictyon 
nodosum, 45 
oleracea, 45 
Physa integra, 156 
Pinus 
australis, 5 
clausa, 4 
elliottii, 5 
laevis, 5 
taeda, 5 
Placospongia 
carinata, 56, 57, 60, 61, 69 
melobesoides, 57 
Plocamilla, 47 
plena, 48 
Podomys, 3 
Polygenis 
floridanus, 13-15, 20, 22, 28, 25 
guy, 13-15 
Polypedilum, 137, 138-140 
digitifer, 140, 158, 160, 161, 163-166 
illinoense, 140, 158, 160, 161 
trigonum, 140, 158, 160, 161 
Porocephalus, 20, 21, 24 
crotali, 19 
Procambarus clarki, 79, 87, 89, 90, 92 
Procladius 
bellus, 111, 157, 159, 161, 168, 165 
procladius, 111 


Psectrocladius 
elatus, 119 
stratiotis, 119 
vernalis, 119, 122 

Pseudochironomus 
ave, 121, 122, 184, 138, 157, 159, 161, 

162, 164, 165 
fulviventris, 121 
pseudoviridis, 121 

Quercus 
chapmanni, 4 
falcata, 5 
laurifolia, 5 
myrtifolia, 4 
virginiana, 4 


Rictularia, 19, 21 
coloradensis 18 
Rhaphisia 
laxa, 54 
menzeli, 54, 68, 71 
myxa, 54 
Rhizochalina oleracea, 44-46, 67 


Salix nigra, 152 
Sargassum, 48-50, 65 
Scypha acanthoxea, 63, Orem 
Sesarma, 91 
Siderastrea siderea, 57 
Sigmodon hispidus, 13 
Spheciospongia 

coccinea, 55 

coccinopsis, 55 

vesparia, 55,69 
Spongia, 66 

barbara, 33, 66 

cheris, 33 

gramined, 33, 66 

zimmocea, 33 
Stelletta grubii, 58, 59, 69 
Stenochironomus, 137, 140 

macateei, 137, 158, 159, 161, 163 
Suberites undulatus, 58 
Sublette, James E., articles, 109-150, 

151-166 

Syphacia, 19 

peromysci, 18 


Tadania ignis, 48, 68 
Taenia lyncis, 18 
Tanypus, 116 
punctipennis, 111 
stellatus, 110,111, 157, 159, 16l, 163 
Tanytarsus, 109, 121, 140 
allicis, 146, 147, 150 
buckleyi, 143, 144-146, 150, 158, 160, 
161 
conjusus, 141, 158, 160, 161 
dendyi, 141, 142, 143, 145, 150, 158, 
160, 161 
disimilis, 144 
glabrescens, 144 
limneticus, 146, 147, 150 
neoflavellus, 142, 143, 146, 147, 158, 
160, 161, 168 
pusio, 144 


INDEX TO AUTHORS AND SCIENTIFIC NAMES—Continued 


quadratus, 148, 144, 147, 150, 158, 160, 
1 


61 
recens, 143, 144, 145, 150, 158, 160, 
161 

recurvatus, 146 

varela, 143 

virdiventris, 141, 158, 160, 161 

xanthus, 141, 142, 148, 145, 150, 158, 

160, 161 

Tenaciella obliqua, 49 
Tethya 

aurantia, 58, 68 

extensa, 58 
Tetilla laminaris, 60 
Thalassia, 382, 49, 50 

testudinum, 31, 37 
Thalyseurypon vasiformis, 50, 68 
Toxemma tubulata, 51, 68 
Trachygellius cinachyra, 61, 63 
Tribelos, 137 
Trichomonas muris, 17, 20 
Trichostrongylus, 19 

ransomi, 18 
Trombicula crossleyi, 6, 8, 20, 22 


Trophon, 99 
actionophorus, 107 
Typha, 119 
Typhis, 99 
alatus, 107 


Uca 
pugilator, 79, 93 
pugnax, 91 
rapax, 91 

Ulva, 50 

Unimia trisphaera, 58, 69 


Verongia, 47 
aurea, 35 
fistularis, 35 
fulva, 35 
longissima, 35; 36, 66 


Walchia, 6 


Xenopsylla cheopis, 15 
Xytopsena sigmatum, 40, 67 


Zizaniopsis miliacea, 157 


TULANE STUDIES IN ZOOLOGY 
VOLUME 10 
INDEX TO AUTHORS AND SCIENTIFIC NAMES 
(New genera, species, and subspecies in boldface) 


Abastor, 142, 145, 169, 177, 209, 210 Cambarus, 60 
erythrogrammus, 147, 149, 167, 168, bartoni, 49, 54, 55 
208, 212 clypeatus, 57, 61 
Acteonia, 34 Carphophis, 167, 171, 172 
Agkistredon, 157, 175, 203 amoenus, 138-144, 166, 208, 210, 212 
bilineatus, 200 Castor, 210 
contortrix, 200, 202, 209, 212 Causus, 200 
halys, 200 Cemophora, 184, 185, 187, 188, 207, 212 
piscivorus, 140, 142, 143, 144, 145, 147, coccinea, 139 
200-202, 205, 208, 212 Cenio, 34 
Alligator, 138, 142 Cercaria, 85 
Alsophis, 176 ftimbriata, 86 
Ambystoma tigrinum, 125 glareosa, 86 
Anabaena, 15 munita, 86 
Anadera, 145 opaca, 86 
Aneides, 124 paradoxa, 86 
Anilioides, 163, 164 protensa, 86 
minuatus, 134, 156, 164, 165, 207, 208, zusilla, 86 
22, stupulosa, 86 
Anilius, 163 Charina, 157, 158, 162, 163, 207 
Anolis carolinensis, 145, 146 prebottae, 165 
Arizona, 180, 185 Cheilophis, 163, 207 
Artemia salina, 92,114 Chionactis, 171 
Atractus trilineatus, 166 Chirocephalus 
Astacus astacus, 54 grubei, 92,117 
Auffenberg, Walter, article, 131-216 nankinensis, 92, 106, 117 
Azemiops, 200 diaphanus, 92, 112 
stagnalis, 92 
Baqai, Izhar U., article, 91-120 Chlorophis, 176, 190 
Bitis, 175 Coluber, 154, 174, 175, 178, 179, 189, 190 
arietans, 200 constrictor, 188-146, 149, 150, 176-178, 
Boa, 157, 158 208, 212 
Bolyeria, 156 oaxaca, 177 
Boreostracon, 145 spinalis, 177 
Bothrops, 175 Congeria leucophaeata, 22, 29 
alternata, 200 Conophis, 190, 207 
jararacussu, 200 Coronella, 171 
neuweidii, 200 Constrictor, 158, 207 
schlegelii, 200 constrictor, 157, 159 
Brachidontes recurvus, 29 Crassostrea, 43 
Branchinecta virginica, 30 
paludosa, 92 Crotalus, 175, 201, 202, 209 
coloradensis, 92 adamanteus, 138-146, 149, 150, 200, 2038- 
occidentalis, 92, 93, 106, 110, 117 206, 208, 212 
Branchipus atrox, 202, 203 
stagnalis, 92, 110, 116, 118 durrissus, 200, 203 
grubet, 92,116 giganteus, 143, 144, 147, 203-206, 208, 
Buccinum, 34 PiIl, Bly 
Bufo, 141, 145 horridus, 200, 203, 212 
Bungarus, 198 mitchelli, 202, 203 
Bush, Francis M., article, 121-128 molossus, 200, 203 
ruber, 200, 203 
Caesarea, 156 tigris, 200, 203 
Calamagras, 134, 157, 158, 161, 207, 209 Cylindrophis, 163 
angulatus, 162, 163, 165 
floridanus, 134, 156, 162-165, 208, 212 Dasypus bellus, 143 
murivorus, 162, 165 Dendroaspis, 198 
primus, 162, 163 Denisonia, 198 
talpivorus, 162, 165 Derocerus, 145, 146 
truxalis, 162, 165 Desmognathus fuscus, 125 
Callinectes sapidus, 29 Diadophis, 169, 172, 209 
Callocardia, morhuana, 41 amabilis, 166, 170 
Cambarellus, 50, 51, 53, 54, 129 elinorae, 134, 156, 170, 171, 208, 212 


shufeldti, 49, 55, 56 punctatus, 142, 1438, 166, 170, 208, 212 


INDEX TO AUTHORS AND SCIENTIFIC NAMES—Continued 


Donax, 145 
Dromicus, 176 
Dryinoides, 207 
Drymarchon, 176 
corais, 138, 139, 141, 142, 144-147, 149, 
150, 178-180, 208, 212 
Drymobius, 178 
Dryophis, 176, 190 
Dundee, Harold A., article, 129-130 


Elaphe, 154, 176, 177, 179, 185, 206, 207, 
209 
climacophora, 181 
dione, 181 
guttata, 138-144, 149, 180-183, 208, 212 
kansensis, 182 
obsoleta, 138-142, 145, 149, 180-183, 
208, 212 
situla, 181 
subocularis, 180 
taeniurus, 181 
vulpina, 181 
Ensatina, 126 
Enulius, 167, 171 
flavitorquatus, 166 
Epicrates, 157, 158, 207 
inornatus, 162 
Equus, 148, 145, 149 
Erethizon, 210 
Eryx, 209 
Eubranchipus 
oregonus, 92 
serratus, 92 
Euglandina, 145 
Eumeces fasciatus, 142, 210 
Eunectes, 157, 158 
Eurycea 
aquatica, 121-128 
bislineata, 121, 123-126, 128 
longicauda, 121, 123 
lucifuga, 121, 123 
multiplicata, 121 
nana, 121 
neotenes, 121 
troglodytes, 121 
tynerensis, 121 


Fairbanks, Laurence D., article, 3-47 
Farancia, 142, 169, 177, 209, 210 
abacura, 145-147, 149, 167, 168, 208, 212 
Faxonella 
beyeri, 61, 62, 64, 69 
clypeata, 57, 59-61, 64, 69-71, 75-77, 79 
clypeatus, 61, 62, 69 
Faxonius, 57, 60 
clypeatus, 61 
Fingerman, Milton, article, 49-56 
Fitzpatrick, J. F., Jr., article, 57-79 


Gastrocopta, 145 

Gastrophryne carolinensis, 125 

Gemma, 34 

Geochelone, 137, 148, 210, 211 

Geomys, 138 

Gigantophis, 158 

Goniobasis, 148 

Gyrinophilus, 124, 126 
porphyriticus, 125 


Gyrodactylus 
shorti, 83-85 
funduli, 85 


Haemachatus, 198 
Haldea, 192, 198 
striatula, 191, 212 
valeriae, 191 
Helicops, 193, 209 
leopardinus, 191 
caricaudae, 191 
Heliocodiscus, 145 
Helisoma, 145 
Hemidactylum, 126 
Heterodon 
brevis, 134, 156, 173, 174, 208, 212 
nasicus, 172-174, 208, 212 
platyrhinos, 138, 189, 141-144, 149, 172- 
174, 208, 209, 212 
plionasicus, 174 
semus, 141, 142, 143, 172, 173, 174; 2085 
2 
Hipparion, 134 
Holliman, Rhodes B., article, 83-86 
Holmesina, 145, 168 
Hydrophis, 198 
Hypsirrhinchus, 151 


Ictalurus furcatus, 29 
Immantodes, 176 


Kinosternon, 145 


Lacuna, 34 
Lagodon rhomboides, 33 
Lampropeltis, 154, 167, 177, 180, 186, 187, 
207, 209 
calligaster, 184 
doliata, 145, 170, 171, 184, 185, 188, 208, 
PA 
getulus, 138, 140-142, 144-146, 149, 184, 
185, 208, 212 
multicincta, 184 
polyzona, 184 
zonata, 184 
Laticauda, 198 
Leimadophis, 176, 184 
Leptomicrurus, 198 
Leptophis, 176, 188-190 
Lichanura, 157, 158, 163, 165, 207 
roseofusca, 162 
Liodytes alleni, 145, 149, 191, 193, 208, 
209, 212 
Liophis, 171, 184 
Littoridina 
sphinctostoma, 22, 29 
Lonchophylla 
concava, 87 
hesperia, 87 
mordax, 87 
robusta, 87, 90 
thomasi, 87 
Loxocemus, 158, 163 
Lygophis, 176 


Machairodus, 134 

Macoma, 8 
mitchelli, 22, 29 

Macroclemys, 210 


INDEX TO AUTHORS AND SCIENTIFIC NAMES—Continued 


Manculus, 126 
Masticophis, 154, 174, 179, 189, 190 
flagellum, 138, 139, 141-144, 176-178, 
208, 212 
lateralis, 176, 177 
taeniatus, 177 
Mastodon, 149 
americanus, 148 
Megatherium, 145 
Menetus, 145 
Mentha spicata, 125 
Mercenaria, 29 
mercenaria, 30, 41, 42 
Micropogon undulatus, 29 
Micruroides, 198 
Micrurus, 198, 209 
circinnalis, 199 
corralinus, 199 
frontalis, 199 
fulvius, 138, 141-144, 199, 208, 212 
nigrocinctus, 199 
Milneria kelseyi, 34 
Mulinia, 145 
Mya arenaria, 30, 40, 41 
Mytilus, 43 


Nagabhushanan, R., article, 49-56 
Naja, 198 
Nasturtium officinale, 125 
Natrix, 142, 146, 154, 198, 206 
cyclopion, 140, 144, 149, 191, 198, 194, 
116 )5),, 7AOies, PAL 
erythrogaster, 140, 191, 195, 196, 208, 
210-213 
grahami, 191 
harteri, 191 
natrix, 191 
rhombifera, 191, 193 
rigida, 191, 212 
septemvittata, 191, 212 
sipedon, 138, 140, 141, 144, 145, 147, 
ial), Gil, Sek, IU, UNS Aes) 2a 
stolata, 191 


taxispilota, 147, 149, 191, 193-195, 208, 


212 
tesselatus, 191 
Neofiber, 138 
Neoseps, 209 
Neurodromicus 
dorsalis, 157, 158 
barbouri, 212 
Ninia, 166, 193 
atrata, 191 
sebae, 191 
Notechis, 198 
Nucula delphinodonta, 34 


Odocoileus, 146, 149 

Ogmophis, 157, 158, 163, 187, 207, 209 
arenarum, 161, 165 
compactus, 161, 162, 165 
oregonensis, 161, 165 
pauperrimus, 134, 156, 160-162, 164, 

165, 208, 212 

Opheodrys, 174, 176 
aestivus, 138-140, 142, 175, 208, 212 
vernalis, 175 

Ophiophagus, 198 


Ophisaurus 
compressus, 140 
ventralis, 145 

Orconectes, 61, 69, 79 
beyeri, 57, 60, 62 
clypeata, 61 
clypeatus, 58, 61, 62 
inermis, 60 
lancifer, 60 
leptogonopodus, 60 
pellucidus, 60 

Oxybelis, 176, 188, 190, 207, 212 


Pacifastacus trowbridgi, 49 
Palaemon serratus, 53 
Palaemonetes, 55 
vulgaris, 49, 54 
Paleoelaphe, 181, 182, 209 
kansensis, 182 
Paleofarancia, 135, 168, 209, 210 
brevispinosus, 156, 168, 169, 208, 212 
Paleopython, 158 
Paleryx, 158 
Pandalus montagui, 53 
Paracambarus, 61 
Paraepicrates, 157,158 
Paraoxybelis, 187, 188, 190, 207 
floridanus, 134, 155, 189, 190, 208, 212 
Parasterte, 34 
Pecten irridians, 31 
Pelamis, 198 
Penaeus setiferus, 29 
Peringia ulvae, 33 
Pinctada, 38 
albina, 30, 33 
Pituophis, 176, 180, 185 
catenifer, 183 
melanoleucus, 139, 142, 143, 145, 146, 
149, 183, 184, 208, 212 
Pitymys, 138 
Plethodon, 126 
Polygyra, 145 
Polymesoda, 8 
caroliniana, 22, 29 
Praunus flexosus, 53 
Procambarus, 61, 75 
Procyon lotor, 145 
Psammophis, 176 
Pseudemys, 142 
caelata, 134 
scripta, 210 
Pseudobranchus, 138 
robustus, 144, 210 
striatus, 144, 210 
Pseudocemophora, 187, 207 
antiqua, 134, 156, 187, 188, 208, 212 
Pseudoepicrates, 157, 158, 207 
barbouri, 158-160 
stanolseni, 134, 155, 157-160, 208, 212 
Pseudotriton, 212, 124, 127, 128 
ruber, 125 
Rana, 83, 145, 147 
grylio, 140 
Rangia, 5-9, 17-21, 25, 26, 29, 30, 31, 34, 
Ae 
cuneata, 4, 33, 35-39, 44, 46, 47 
Rangianella, 20, 35-38 


INDEX TO AUTHORS AND SCIENTIFIC NAMES—Continued 


Rhadinea, 184 
flavilata, 142, 144, 166, 171, 172, 208, 
212 
Rhinocheilus, 184, 185, 187 
Rhineura, 209 
Rose, Francis L., article, 212-128 


Salvadora, 174, 176, 185 
hexalepis, 175 
lineata, 175 
mexicanus, 175 
Sanzinia, 156, 157 
Seminatrix pygaea, 191, 192 
Serridentineus, 134 
Siren, 133, 138 
lacertina, 145 
Sistrurus, 157, 175 
catenatus, 158, 200-202 
miliarius, 141-1438, 150, 200-202, 208, 
212 
Sonora, 167,171 
episcopa, 166 
Spilotes, 179 
Sternotherus, 148 
Stilosoma, 185, 209 
extenuatum, 139, 186, 187, 208 
vetustum, 134, 156, 186, 208, 212 
Streptocephalus 
proboscideus, 117 
purcelli, 117 
seali, 91, 92, 106, 108, 115-118, 120 
texanus, 116,117 
torticornis, 106 
Strobilops, 145 
Storeria 
dekayi, 188, 142, 191-193, 208, 212 
occipitomaculata, 191, 192 
Sylvilagus 
floridanus, 150 
palustris, 150 
Synaptomys, 210 
Syngnathus scovelli, 83, 85 


Tagelus plebius, 29 
Tantilla 


coronata, 138, 142, 166, 188, 189, 208, 
212 


eisent, 166, 188 
melanocephalus, 188 
nigriceps, 166, 188 
Tanupolama, 146 
Tapirus, 138 
veroensis, 146 
Terrapene carolina, 211 
Texadina, 29 
Thamnophis, 145, 154, 193 
brachystoma, 191 
cyrtopsis, 191, 197 
elegans, 191 
marciana, 19% 
ordinoides, 191 
radix, 191, 197 
sauritus, 191, 197 


sirtalis, 138-145, 147, 149, 150, 191, 197, 


198, 208, 212 

Tivela stultorum, 29 
Toluca, 166, 171 

lineatus, 166 
Trachyboa, 156 
Transenella, 34 
Tridacna, 38 
Trimeresurus, 175 

viridesoma, 200 
Troglocambarus, 61 
Tropidoclonion lineatum, 191 
Tripidophis, 156 


Uromacer, 176, 188, 190 
Uropeltis, 163 
Urosalpinx, 34 


Vipera, 175, 200 
Vulpes, 210 


Walton, Dan W., article, 87-90 
Xenopeltis, 163, 166 
Zamenis, 177 


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