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Full text of "Studies on Arthropoda"

STUDIES 
ON ARTHROPODA 



I. 



BY 



Dr. H. J. HANSEN 



WITH FOUR COPPER-PLATES 



AT THE EXPENSE OF THE RASK-0RSTED FUND 




A L 1'' R E D A, K N P F, I n c. 
220 West 42nd Street 
Nev/ York, N,Y« 



1 13.00 



COPENHAGEN 
PRINTED BY BIANCO LUNO 



Ready U'oin the Press January the ^i. 1921. 




CONTENTS 

Pag. 
The Pedipalpi, Ricinulei, and Opiliones (exc. O. Laniatores) 

collected by Mr. Leonardo Fea in tropical West Africa 

and adjacent Islands (With four Plates) 5 

On Stridulation in Crustacea Decapoda 56 

On the postembryonic Occurrence of the median "Dorsal Or- 

2;an" in Crustacea Malacostraca 66 



( 1 v- /^ ^■' 






THE PEDIPALPI, RICINULEI, AND OPILIONES 

(exc. Op. Laniatores) 

coUected by Mr. LEONARDO FEA 

in tropical West Africa and adjacent Islands. 

(With four Plates.) 



SEVERAL years ago Dr. Raffaello Gestro, Director of the 
Museo Civico di Storia Naturale di Genova, asked me 
to work out the animals of the above-named orders gathered 
by that most excellent zoological collector Mr. lyconardo Fea, 
who during 1900- — 1902 spent his last years in exploring partly 
unhealthy countries in tropical West Africa, viz. Portuguese 
Guinea, Kameroon, and French Congo, and besides the islands 
of Fernando Poo, St. Thome, Principe, and Annobon in the 
Gulf of Guinea; from this voyage Fea returned quite broken 
down to Italy and died shortly afterwards in April 1903. With 
pleasure I accepted Dr. Gestro's kind offer, as I expected to 
tind most interesting animals among the material, but the 
main portion of the collection, viz. the Opiliones Laniatores, 
I found it necessary to hand over to a colleague, and be- 
fore now I could not find the time necessary for the task. 

The material to be dealt with in the present paper is in 
reality most valuable. It consists of eleven species, but with 
two exceptions all are new to science. Furthermore I found 
it necessary to establish three new genera, and two among 
these are even important. The collection comprises three spe- 
cies of Pedipalpi Amblypygi, and one af the forms, living in 
nests of white ants, is completely blind ; furthermore two species 
of the tribe Tartarides, two species of Cryptostemma and four 
species of Opiliones, but two of these belong to the small 



studies ou Arthropoda. I. 



sub-order Op. C>'phophthalmi. As Tartarides, Ricinulei {Cryp- 
tostemma) and Op. C^'phophthalmi are scarce even in the largest 
Museums of the world, it is easily seen that the collection is 
important out of all proportion to the somewhat low number 
of species; the presence of an unknown kind of larval stage 
of the Ricinulei highly increases its value. Before concluding 
these remarks, I may beg Dr. R. Gestro to accept my sincere 
thanks for offering me the material and for his patience as to 
the long delay. 



ORDER PEDIPALPI 



A. Sub-order Amblypygi. 

In his monograph (Revision der Tarentuliden Fabr., 
1895) K. Kraepelin accepted three sub-famihes, and in 1902 
R. I. Pocock elevated them to the rank of families. Only one 
of these families, \'iz. the Phrynida?, was hitherto known from 
Africa, and a single well-known form is represented in the col- 
lection. But this contains besides two species of the famih' 
Charontidae, which hitherto was known only from the Indo- 
Australian region and the Galapagos Islands. 

It ma}^ be inserted that I am inclined to consider the second 
of the two long joints in the mandibular palps as corresponding 
to patella plus tibia in Aranese and Pedipalpi Uropygi; in the 
descriptions I can therefore follow Kraepelin who named it 
tibia. 

Titanodamon medius Herbst. 
Kraepelin mentioned three varieties, viz. typicns Kraep., 
tibialis Simon, and Johnstoni Pocock; the two last-named va- 
rieties had been established as separate species by the authors 



Pedipalpi, Ricinulei, and Opiliones. 



named, but Kraepelin was certainly right in withdrawing 
them. Var. typicus is said to occur from Senegal to the estuary 
of the Niger, var. tibialis especially in the Congo region, and 
\-ar. Johnstoni in the area between those of the two other forms. 
The material comprises specimens belonging to var. typicus 
and to var. Johnstoni, but animals from the same locahty vary 
not only much according to age in the length of the proximal 
one of the three long distal spiniform processes on the upper 
margin of the tibia, but between the numerous specimens from 
the Isl. of St. Thome, all according to the shape of the frontal 
process belonging to var. typicus, a few specimens show a trans- 
ition between typicus and Johnstoni as to the setse on the inner 
side of the antennae. 

Occurrence. — Specimens of var. typicus are to hand 
from the following places: Bolama, Portuguese Guinea (near 
lat. 12° N.), June — Dec. 1899, 6 specimens; and Isl. of St. 
Thome, at Ribeira Palma, o — 300 m above the level of the sea, 
Juli — August 1900, 1/ specimens, and 2 small specimens in 
June 1900 at the capital of the island. 

Specimens of var. Johnstoni were taken at two localities, 
\'iz. Basile on Fernando Poo, 400 — 600 m above the level of 
the sea, August — September 1901; i specimen, and N'Kogo, 
French Congo, December 1902, i specimen. 

Gharinus africanus n. sp. 

PI. I, fi.gs. I a — I c. 

This species agrees completely with the characters of the 
genus Charinus in Kraepelin's analytical key excepting as to 
one feature, y\z. that the first tarsal joint is conspicuously 
longer than the four others combined. 

Cephalothorax in adult specimens considerably broader 
than long, with six thin spines at the rounded front margin. 
The distance between each ocular tubercle and the lateral 



8 studies on Arthropod a. I. 

margin nearly or scarcely twice as long as the transverse dia- 
meter of the tubercle. 

Interior prehensile margin of the basal joint of the antennae 
with four teeth, and the most distal tooth bifid, while the ex- 
terior margin has no tooth. 

Palps about as long as the body. Trochanter with two small 
processes on the anterior margin (in fig. la the proximal spine 
is overlapped, not visible) ; the robust process on the lower 
side with a number of strong setae on the inner and lower sur- 
face. Femur in the adults at least a little shorter than the 
cephalothorax ; its upper front margin proximally with two 
small, conical, setiferous tubercles, more distally three or four 
processes, the first moderately long, the others decreasing in 
length, and the fourth, if present, quite short; the lower front 
margin (fig. i a) with at least three processes decreasing in 
length from the first to the third, and the first which originates 
near the base, is long; sometimes a small or very small fourth 
jjrocess is found. Tibia on the upper margin with four processes 
(fig. lb), the second and especially the third very long, strong, 
with a setiferous protuberance or rather an offset on the prox- 
imal side somewhat from the base; the first process is only 
about half as long as the next, but longer than the fourth; 
frequently " a setiferous conical protuberance is found before 
the first real process; the lower margin (fig. la) on the di.stal 
half with two moderately long processes, the proximal shorter 
than the distal, and this margin has besides three or four quite 
small setiferous tubercles. First tarsal joint has on the upper 
margin (fig. i c, u) near tiie middle two processes, the 
second considerably to very mucli longer than the first, while 
the lower margin has a single moderately long, jjorrected pro- 
cess a little before the end. Second tarsal joint (fig. i c) with 
two slender processes on the upper margin, the distal from con- 



Pedipalpi, Ricinulei, and Opiliones. 



siderably longer than to twice as long as the first; the lower 
margin without any process. 

The three posterior pairs of legs have the metatarsus di- 
stinctly longer than the tarsus; first tarsal joint about one- 
fourth as long again as the four other joints combined. 

The dorsal surface of the cephalothorax in the adult some- 
what dark reddish-brown, with the impressions blackish, while 
the abdomen is brownish-grey with the muscular impressions 
dark and very conspicuous. Antennae, palps, and femora of 
the legs somewhat lighter than cephalothorax; legs without 
yellowish or Ught rings. 

Length of the body of a large specimen 8.4 mm, cephalo- 
thorax 3 mm, palp stretched out 8.8 mm (claw included), se- 
cond leg (coxa not included) 14.5 mm. 

Remarks. — C. ajricamis is allied to C. australianus L. 
Koch, but is instantly distinguished in having the first tarsal 
joint in the three posterior pairs of legs very conspicuously 
longer than the four other joints combined. Furthermore adult 
specimens of C. australianus have, according to Kraepelin, a 
higher number of processes on both margins of the tibia of the 
palps than is found in C. africaniis. 

Occurrence. — Taken on three of the Islands in the Gulf 
of Guinea: Annobon, St. Thome and Principe. On Annobon 

6 specimens were taken in May 1902 in an altitude of 400 
— 500 m, and i specimen in April in o — 500 m. On St. Thome 
3 specimens were gathered in December 1900 at Agua Ize, 
altitude 400 — 700 m, and 11 specimens in September — Octo- 
ber 1900 at Visto Alegre, altitude 200—300 m. On Principe 

7 specimens were secured in Jan. — March 1901 at Roca In- 
fante Don Henrique, altitude 100 — 300 m. — Adult specimens 
from St. Thome are a little smaller than those from the two 
other islands. 



10 studies on Arthropocla. I. 



Paracharon n. gen. 

Ceplialotliorax not much broader than long, anteriorly 
somewhat produced, with a rather short, transverse, anterior 
margin. Eyes and ocular tubercles completely- want- 
ing (fig. 2 a). First sternum is a light transverse strip more 
firmly chitinized than the membrane at its front margin, but 
instead of a sternal labium only a tin}- granule with a small 
vertical seta is seen at the middle of the sternum. Each of the 
two following sterna have a rather slender, subcylindrical 
process projecting nearly vertically downward, Ingher than 
thick, and with some short, brown setse on the rounded distal 
part. — Second abdominal sternite simple, as its posterior 
part is not marked off or produced. — Palps considerably 
shorter than the bod}-, built as in the Charontidoi; first tarsal 
joint (fig. 2 d) with two spiniform processes on each margin 
and the distal processes much longer than the proximal; second 
tarsal joint with three processes on the upper margin, and the 
claw marked off but not movable. Walking legs as in the Cha- 
rontidse, excepting that the tibia of fourth pair consists only 
of two subjoints. 

Remarks. — This most interesting genus differs from 
all other genera of the suborder in having no eyes and 
no sternal labium. It shows, howe\-er, close relationship 
to the Charontidae in all other features, excepting that 
the fourth pair of tibia? ha\-e only a .single secondary arti- 
culation. 

It ma}- be noted that I have looked in vain for sensors- 
hairs on the places of the eyes. But the normal long tactile 
hairs on the walking legs and the peculiar small, clu])-shaped 
sensory hairs on the distal tarsal joints of the first pair of legs 
exist, though less numerous, as in the large form Tarcntula 
reniformis L. {Phrynichus nigrnnanus C. L. Koch) (see H. J. 
Hansen: Organs and Characters in different Orders of Arach- 



Pedipalpi, Ricinulei, and Opilioues. II 

nids, in Entom. Meddel. Kjobenhavn, Vol. IV, 1893 — 94, 
P- 154—156). 

Only one species is known. 

Paracharon caecus n. sp. 

PI. I, figs. '2 a — 2e. 

Cephalotliorax (fig. 2 a) only about one-eighth as broad 
again as long, considerably vaulted, with the radiating im- 
pressions partly deep ; the antero-lateral margin is a little con- 
cave, the front margin transverse, rather short, with about 
six minute, very short spines. 

Palps scarcely twice as long as the cephalothorax. Tro- 
chanter with two small processes on the anterior margin (fig. 2 a) , 
while the well developed process on the lower side (fig. 2 b) 
has five or six triangular, setiferous teeth on the inner margin. 
Femur not fully half as long as the cephalothorax (fig. 2 a) ; 
its upper front margin has near the middle two somewhat 
small processes; the lower margin has three to five processes, 
the first somewhat long (fig. 2 b), the second distinctly or con- 
siderably longer than the first, the others small. Tibia has on 
the upper margin (fig. 2 c) three well developed processes, the 
second process considerabh' longer than the first or the third, 
and besides generalh' two small teeth placed respectively before 
the first process and between first and second process; the 
lower margin (fig. 2 b) has three processes, the second verv' 
long and the others rather short, and besides frequently a 
minute tooth before the first and another before the second 
process. First tarsal joint large and only somewhat shorter 
than the tibia ; each margin (figs. 2 b and 2 d) has two pro- 
cesses, the distal long or very long and much longer than the 
proximal, and besides a distinct or rudimentary' tooth is fre- 
quently seen on the lower margin just before the proximal 
process, and sometimes a rudimentarv tooth before the distal. 









12 Studies on Arthropoda. 1. 



process. Second tarsal joint (figs. 2 b and 2 d) has along the 
upper margin (u) three processes increasing in length from the 
first to the third which is moderately long; the lower margin 
has only a single moderately short process. 

In second and third pairs of legs the metatarsus is about 
as long as the tarsus, and first tarsal joint is somewhat longer 
than the four distal joints combined. Fourth pair of legs (fig. 2 e) 
have the tibia divided by a single secondary articulation 
into two sub joints, the first more than three times as long 
as the second; metatarsus somewhat longer than in the 
preceding pairs of legs and conspicuously longer than the 
tarsus. 

Upper surface of cephalothorax in the adult dark yellowish 
brown, with the deep portions of the impressions dark brown; 
abdomen above light greyish with a somewhat yellowish tint. 
Antennae, palps, and femora of the legs nearly as, or a little 
lighter than, the cephalothorax; the legs without light rings. 

Length of the body of a large specimen y.^ mm, cephalo- 
thorax 2.8 mm, palp stretched out (claw included) 5 mm, se- 
cond leg (coxa not included) 8.6 mm. 

Remarks. — The specimens have been found in nests 
of Tennes (genus and species not noted) ; only a single speci- 
men has no indication on this topic on the label. 

Occurrence. — Portuguese Guinea (near lat. 12° N.) at 
two places: Bolama, June — December 1899, i spec; Rio Cas- 
sine, Jan. — Febr. 1900, i spec, and April 1900, 15 spec. 



B. Sub-order Uropygi. 

This sub-order comprises two tribes, Oxopoei and Tarta- 
rides, which differ in several important features. No specimen 
of the Oxopoei was found among the material, but it contains 
a number of specimens of the Tartarides. 



Pedipalpi, Ricinulei, and Opilioiies. I3 

Trihe Tartarides. 
In H. J. Hansen and William Sorensen: The Tartarides. 
A Tribe of the Order Pedipalpi (Arkiv for Zoologi utgif. af 
Svenska Vet. Akad. Stockholm, Band 2, Nr. 8, 1905) I described 
15 species of this most interesting group, mentioned a Califor- 
nian species established by Cook but unknown to me, and pointed 
that Artacarus liberiensis Cook must be considered a nomen 
nudum. Since 1905 Randall Jackson has described (1907) an 
interesting form taken in the Botanic Gardens, Kew; in three 
small papers (1911) F. H. Gravely established no less than 
2 species from Ceylon and 4 from India; K. Kraepelin (191 1) 
established a new species from Formosa; Hirst (1913) a species 
from the Seychelles; finally H. J. Hansen (1910) a new species 
from Kihmandjaro. Thus in all 26 species have been really 
described, but of these only 2, viz. Schizomus montanus H. J. H. 
and Trithyreus africanus H. J. H., are of African origin. And 
it may be added that both these species have been established 
on female specimens which certainly or scarcely had not arrived 
at sexual maturity, and consequently the knowledge must be 
somewhat imperfect. The Fea collection contains adult spec- 
imens of both sexes of two new species, and this material is 
therefore ver>' valuable, though the forms do not exhibit peculiar 
features expanding the general conception of the type. 

Trithyreus parvus n. sp. 

PI. I, figs. 3 a — 3 k. 

Male. 

Head with an oblong, rounded, badly limited paler eye- 
spot on each side, but without an\- cornea. Cephalic sternum 
a little longer than broad. Second thoracic tergite with a nar- 
row and whitish median strip of thinner skin. 

Palps (figs. 3 a and 3 b) somewhat slender, about two-thirds 
as long as the body. Trochanter has the lower front part some- 



lA Studies on Arthropoda. I. 

what produced, with the terminal margin straight, the comer 
angular, and this angle measures 90° — 95°. Femur uncommonly 
slender, more than two and a half times as long as deep, with 
the lower margin nearly straight though slightly sinuate, but 
without any protruding angle towards the base. Patella a little 
less than three times as long as deep. Tibia slightly more than 
three times as long as deep. Tarsus unusually long, as its upper 
margin is as long as the lower margin of the tibia; claw only 
one-fourth as long as the upper margin of the tarsus. 

First pair of legs (fig. 3 a) slender, somewhat more than 
half as long again as the body. Coxae terminate scarcely before 
the outer angle of the mandibles (maxillae auct.). Femur a little 
longer than tibia. Foot a little shorter than tibia, about nine 
and a half times as long as deep (fig. 3 d), deepest near the end 
of metatarsus; second joint of the metatarsus almost as long 
as the sum of the four proximal joints plus half of the fifth 
joint of the tarsus; terminal joint a little longer tlian fourth 
and fifth tarsal joints combined, and a little more than half 
as long as the whole metatarsus. 

Fourth legs somewhat longer than tlie body; femur scarcely 
two and one-third times as long as deep (fig. 3f). 

Three posterior abdominal segments telescoped and very 
considerably thicker (fig. 3 i) than in the other sex ; last seg- 
ment without any protuberant part above the insertion of the 
flagellum. 

Flagellum (figs. 3 h and 3 i) of very moderate size, somewhat 
less than twice as long as broad, but almost two and a half 
times as long as deep, consequently its thickened part consider- 
ably depressed. The stalk rather compressed, seen from above 
a little more than one-third as long as the main piece. Seen 
from above, the main piece is only one-fourth as long again 
as broad, broadest at the middle, with the proximal half of 
the lateral margins rather convex, and the distal half to a little 



Pedipalpi, Ricinulei, and Opiliones. 15, 

from the end conspicuously concave; the end is angular. Seen 
from the side (fig. 3 i) the lower margin of the whole flagellum 
is shaped as a line somewhat curved or convex at the middle, 
while the upper margin is incurved considerably before the end. 
The flagellum has several pairs of long or very long spiniform 
setae distributed on the sides, at the end and on the lower sur- 
face, while the upper side has only two erect shorter setse far 
from one another in the median line. 

The walking legs, the upper surface of abdomen, and the 
posterior part of the dorsal side of cephalothorax are more or 
less olive-green with a reddish or light reddish tint ; front part 
of the head, antennae and palps reddish; the lower surface of 
the body lighter. 

Body without flagellum 2.64 mm., palp 1.75 mm., first leg; 
4.1 mm., fourth leg 2.9 mm. 

Female. 

It agrees with the male in many particulars, but differs in 
several features. 

Palps (fig. 3 c) a little more robust and proportionately shorter 
than in the male, being only about half as long as the body. 
Trochanter has not the lower distal part produced, and the cor- 
ner is a rather obtuse angle measuring about 110°. Femur con- 
siderably deeper than in the male, scarcely twice as long as 
deep, and the lower margin is angularly bent a little from the 
base: the tip of this angle is a Uttle nearer to the proximal 
than to the distal end of the upper margin of the joint. Patella 
and tibia shghtly thicker than in the male, while the tarsus is 
conspicuously shorter than in that sex, and the claw is some- 
what more than one-third as long as the upper margin of the 
tarsus. 

First pair of legs slender, slightly longer than the body. 
Coxae terminate a little before the outer angle of the mandibles.. 



l6 studies on Arthropod a. I. 



Femur distinctly longer than the tibia. Foot a little shorter 
than tibia, almost ten times as long as deep (fig. 3 e) ; second 
joint of metatarsus slightly longer than the four following 
joints combined; terminal joint conspicuously more than half 
as long as the whole metatarsus. 

Fourth legs considerably shorter than the body; fenmr 
about twice as long as deep (fig. 3 g) . 

Three posterior abdominal segments (fig. 3 k) much more 
slender than in the male. 

Flagellum (fig. 3 k) a little or considerably shorter than 
in the male, nearly four times as long as deep, three-jointed, 
but the first articulation is more or less obsolete; the terminal 
joint a little more than two-thirds of the whole length. 

The largest female which has been taken as type, has the 
body 3.3 mm. long, the palp is 1.63 mm., first leg 3.4 mm., 
fourth leg 2.8 mm. Other females, which seem to be 
adult, are only about as long as the male, but the relative 
length between body, first leg, etc. about as in the large 
female. 

Colour and other characters not mentioned in this descrip- 
tion nearly as in the male. 

Remarks. — As usual the palps and the flagellum in botli 
sexes afford the best characters. T. parvus is easily distingui- 
shed from T. africanus H. J. H. (1905) in having the last tar- 
sal joint of first legs considerably shorter in proportion to more 
proximal joints, in the shape of the fenmr of the palps, etc. 
The differences between T. parvus and the next species are 
pointed out below. 

Occurrence. — Ten .specimens, among them two adult 
males, were collected on the Island of St. Thome at Vista 
Alegre, altitude 200 — 300 m., October 9, 1900. An innnature 
female was secured at Punta Frailes, Isl. of Fernando Poo, 
Oct. 1901. 



Pedipalpi, Ricinulei, ami Opiliones. I7 



Trithyreus brevicauda n. sp. 

PI. I. figs. 4 a — 4fl; PI. TI, fig.s. I a — I g. 

Male. 

Head without any distinct eye-spot on the sides. Cephahc 
sternum much longer than broad. Second thoracic tergite with 
a sharp median suture. 

Palps (figs. I a and i b) moderately robust, somewhat less 
than half as long as the body. Trochanter with the lower 
distal part sUghtly produced, the comer acute but the angle 
measuring about iio°. Fennir a little less than twice as long 
as deep, subtriangular, but the median part of the lower margin 
convex without any angle, and its deepest point a little longer 
from the proximal than from the distal end of the upper margin. 
Patella a Httle less than two and a half times as long as deep ; 
tibia slighth^ more than three times as long as deep. Tarsus 
not elongate; its upper margin a little more than half as long 
as that of the tibia. Claw distinctly less than half as long as 
the upper margin of the tarsus. 

First pair of legs (fig. i a) moderately robust, somewhat 
longer than the body. Coxae terminate somewhat before the 
outer angle of the mandibles. Femur somewhat longer than 
the tibia. Foot as long as the tibia, between eight and a half 
times and nine times as long as deep (fig. 1 e), deepest towards 
the end of the metatarsus ; second joint of metatarsus nearly as 
long as the sum of the four proximal joints and half of the 
fifth joint of the tarsus; terminal joint slightly longer than 
fourth and fifth tarsal joints combined, and distinctly more 
than half as long as the whole metatarsus. 

Fourth legs a little shorter than the body. Fenmr two and 
one-third times as long as deep (fig. 4 b). 

Three posterior abdominal segments telescoped (fig. 1 g) 
and somewhat thicker than in the female; last segment without 
any protruding part above the insertion of the flagellum. 



1 8 studies on Arthropodn. I. 



Flagellum (figs, i f and i g) uncommonly small, a little more 
than twice as long as broad, but almost three times as long as 
deep, consequently its thickened part considerably depressed. 
The stalk only a little deeper than broad, seen from above (fig. 
I f) scarcely one-third as long as the main piece, which is oblong- 
ovate, not quite half as long again as broad, with the lateral 
margins moderately convex and the end very obtuse, rounded. 
Seen from the side (fig. i g) the lower margin of the flagellum 
has its shorter proximal part feebly conca\'e and the remain- 
der somewhat convex; the upper margin has the ])roximal 
half somewhat sinuate, the distal somewhat conca\'e and nearly 
angularly bent at the middle. Several of the spiniform setae 
lost in the single specimen; the upper surface has beyond the 
middle a pair of setae and more proximally a seta in the median 
line. 

Colour nearly as in T. parvus, though a little more 
brownish. 

Length of the body 3.13 mm., palps 1.4 nun., first leg. ^.^ 
nun., fourth leg 3.0 mm. 

Female. 

Similar to the male in most respects; the sexual differences 
are less numerous than in most species. 

Palps (figs. I d) differ feebly from those in the male. Tro- 
chanter has the lower distal corner rounded and not angular, 
but the angle measures as in the male about iio^. 

First pair of legs (fig. i c) as long as the body. The coxae 
terminate distinctly longer from the outer angle of the mandibles 
than in the male. Foot slightly longer than tibia, differing 
from that in the male in having the terminal joint proportio- 
nately a little longer (fig. 4 a), somewhat longer than fifth and 
fourth joints combined, and considerably more than half as 
long as the whole metatarsus. 



Pedipalpi, Ricinulei, and Opiliones. IQ 

Fourth legs considerably shorter than the body; femur 
(fig. 4 c) a little shorter than in the male, a little more than 
twice as long as deep. 

Three posterior abdominal segments somewhat less deep 
than in the male. 

Flagellum (fig. 4 d) even a little longer than in the male, 
scarcely five times as long as deep, three-jointed ; third joint 
scarcely twice as long as the two proximal joints combined. 

Length of the body 3.15 mm., palps 1.40 mm., first leg 
5.15 mm., fourth leg 2.77 mm. 

Remarks. — T. hrevicauda is closely allied to T. afri- 
caiuts H. J. H., which was established (in 1905) on a probably 
immature female from Freetown. But the specimen of T. ajri- 
canus, which was 2.9 mm. long, thus only a httle smaller than 
the adults of T. hrevicauda, differs from these in having the 
claw of the palps conspicuously shorter, and besides especially 
in the foot which has the terminal joint proportionately much 
longer than T. hrevicauda, and though this joint is proportio- 
nately longer in immature than in adult specimens, the diffe- 
rence between the relative length of this joint in T. africanus 
and T. hrevicauda is so large that my adult specimens cannot 
belong to T. africanus. It may be added that the foot of T. afri- 
canus is nine time as long as deep, thus nearly more slender 
than those in T. hrevicauda, but in immature specimens the 
foot is less slender than in adults of the same species. — The 
small flagellum of the male differs \'ery considerably from that 
organ in males of other species hitherto known. 

Occurrence. — Bolania in Portuguese Guinea (Kerry 
coast), December 11, 1899; 3 adult specimens (i c?, 2 ?). 



20 Studies on Arthropoda. I. 



ORDER RICINULEI 

The main paper on this small but most interesting order is: 
H. J. Hansen and W . Sorcnscn: On Two Orders uf Arachnida. 
Opiliones, especially the suborder Cyphophthalmi, and Rici- 
nulei, namely the famil>- Crj^tostemmatoida^. Cambridge. 
At the l^niversity Press. 1904 (4to). — Since its publication 
nothing, as far as I know, has been added to our knowledge 
of the recent forms. But a curious attempt by Prof. Fr. Dahl 
(Berlin) may be briefly mentioned. In 191 1 he published a 
small treatise: Die Horhaare (Trichobothrien) und das System 
der Spinnentiere (Zool. Anzeiger \'ol. XXXVII, p. 522 — 532). 
He ascribes a high systematic value to the exi.stence and distri- 
bution or non-existence in the various orders of Arachnida of 
that kind of sensor}' hairs which he names "auditory hairs" ; 
besides, without an}' real discussion and without pointing out 
any feature hitherto overlooked in the orders Pedipalpi, Pal- 
pigradi, Ricinulei, Opiliones, and Acari, he refers the Palpi- 
gradi to the Pedipalpi, the Ricinulei to the Opiliones, and the 
suborder Cyphophthalmi from the Opiliones to the Acari. That 
such classification is only of the retrograde kind is rather evi- 
dent; it is scarcely necessary to prove its futility Ijy detailed 
enumerations of structural features and a lengthy discussion 
of their relative value in the orders and suborders in question. 
In a paper published in igi/'^). I have dealt with the so-called 
"auditory hairs" in Arachnida and in the two other classes of 
terrestrial Arthropoda. — On the palaeozoic forms of the order 
Ricinulei important contributions have been published by 

') H. J. Hansen. On the Trichobothria ("auditory hairs") in Arachnida, 
•Myriopoda, and Insecta, with a summary of the external sen.sory organs 
in Arachnida (Entomologisk Tidskrift utg. av I-.ntoinol iMirenineen i 
Stockhohii. Arg. 38, xgiy, p. 240 — 259. 



Pedipalpi, Ricinulei. and Opiliones. 21 



R. I. Pocock (1911) and especially by A. Petninkevitch (1913). 
It is remarkable that as to the palaeozoic genus Pnlyochera 
Scudder, which seems to be rather allied to recent forms, the 
last-named author mentions only one generic character, viz. 
that "tarsus of second leg fused with metatarsus, forming one 
joint", but this character is important, as in the recent forms 
the second pair of legs has in the adult a separate, well devel- 
oped metatarsus, and the tarsus live- jointed. 

The material collected by Fea comprises two species, one 
among them new, while the other species is (ryptostemma 
crassipalpe Hans. & Sor, which was established on immature 
specimens, but Fea has secured adult specimens of both sexes. 
Of the new form a large material is to hand, which enables me 
to add a little on a couple of points to the description in Hansen 
& Sorensen's work, and besides to show that a kind of metamor- 
phosis exists in this order, as I have two specimens of a larval 
stage with only three pairs of walking legs. 

In the paper mentioned the spiracles and the tracheae in 
the immature Cryptostemma was described (p. 131 — 132). 
I have dissected an adult male of C. Fece n. sp. and found a 
similar structure, so that only one single particular may be men- 
tioned. On the crescent-shaped spiracles we wrote: "The walls, 
particularly near the opening, are furnished with short, extre- 
mely thick, slighth' branching hairs . . . which almost have 
the character of processes, and at least in part are free of each 
other". In the adult I have found that both the convex and the 
concave margin of the crescent-shaped spiracle have a very 
close layer of processes which are very oblong, and each with 
a number of moderately short, acute spines in ever}' direction. 
Each layer has a certain thickness, as it is formed by a few rows 
of processes. The processes are on both margins shorter towards 
both ends of the spiracle, and when the processes of the two layers 
reach each other with their ends, they constitute with their 



22 Studies on Arthropoda. I. 

lateral thorns a kind af sieve for the air. The processes are 
light brownish. 

In 1904 we showed that in the female of the large form Ciyp- 
tostemma Afzelii Thor. the sternites of first and second abdo- 
minal segments are freely movable, crescent-shaped, with their 
concave margins turning against each other, and they surround 
together the genital aperture; both sternites are only visible, 
when the abdomen is bent much upwards. In C. Fcce n. sp. 
I find the same structure, but the sternites are a little less robust. 
In this species — and probably in all other forms of the order 
— the genital aperture is in the male much smaller than in the 
female; the sternite of second segment is very thin and semi- 
circular or in reality curved so nmch, that it occupies about 
three-fifths of a circle; the sternite of first segment is produced 
downwards as a freely protruding, triangular plate about as 
long as broad and with the end subacute. The same plate is 
also found in the adult male of C . crassipalpc. An internal co- 
pulator}^ organ is, as might be expected, completely wanting, 
l)ut according to my dissection of a single male, the inner sexual 
organs and the alimentar}- canal can scarcely be made out 
with certainty without fresh material or at least the sacrifice 
of several well preserved specimens of both sexes. 

The Larva. — The body is about 2.3 mm. long, second 
leg 3.3 mm. In general aspect the body of the larva is in the 
main similar to half-grown specimens, differing in being compa- 
ratively a little broader, with the grooves on cephalothorax 
partly feebly developed, partly wanting, and without grooves 
on the cucullus, while the tergites of the abdomen and the gra- 
nulation are nearly as in postlarval younger stages. But the 
larva has only three pairs of walking legs, as the 
fourth pair is completely wanting (pi. Ill, fig. i g). 
That it is the fourth pair which is wanting is seen from 
the fact, that the coxtc of first, second, and third pairs 



Pedipalpi, Ricinulei, and Opiliones. 23 

are immovabl}- united as in the adults, while in the 
adults the coxae of fourth pair are movable. Furthermore 
the larva differs from postlarv'al stages in the tarsi of second 
and third pairs of legs. The tarsi of third pair (pi. Ill, fig. i i) 
have only two joints, the second not fully twice as long as the 
first, but otherwise not showing any peculiarity worth mentio- 
ning. But the tarsi of second pair (pi. Ill, fig. i h) are interesting. 
Each of them has only two joints, both very large, somewhat 
compressed and very deep, much deeper than the metatarsus; 
the proximal joint is more than one-tliird as long as the metatar- 
sus, only a little longer and deeper than the second, which has 
the claws and even the normal subdistal sensory hair (s.) nor- 
mally developed. — The interesting fact that the Ricinulei 
thus possess a larval stage with only three pairs of walking legs 
removes the order further from all other orders of Arachnida 
except the Acari, but in nearly all other structural features 
hitherto known the Ricinulei have no affinity to the Acari. 

Cryptostemma crassipalpe Hans. & Sor. 

PI. II, figs. 2 a — 2 h. 

This species was established on two immature specimens, 
one among them not far from full-grown, with the full number 
of joints in the tarsi, but the animal was light-coloured, and the 
tergites of the three large abdominal segments were rather 
removed from each other. The material collected by Fea com- 
prises two immature specimens and five adult animals of both 
sexes. It may be convenient to begin with the young specimens. 

The largest of the immature specimens is somewhat smaller 
than the type, as its body is only 4.2 mm. long, while the type 
was 4.8 mm. It agrees well with the description and figures 
quoted, excepting that the femoral part of the palps is only 
half as thick as long, while in the type it was distinctly more 
than half as thick as long, but this difference is in other forms 



2d. Studies on Arthropoda. I. 

due to age. When viewed obliqueh- there ma>- 1)e observed a 
faint vestige of the sublateral impression on the cnculhis, wliile 
in the type this feature was not perceived. 

Adult specimens. — Body and legs brown or dark l^rown 
and on some places reddish ; the granules which are extremely 
numerous, especially on the body, are in immature specimens 
purple and therefore very conspicuous in contradistinction to 
the yellowish skin, but in the adults they are not distinctly 
marked off by colour from the skin ; v. hen the adult animal is 
flried it changes its aspect, as the exceedingly numerous scales 
are \'ery light, being somewhat slate-whitish with a yellowish 
tint, and consequently the animal looks rather clayey, darkest 
on the median ])ortion of tlie ventral surface where the clotliing 
is less dense. 

The cephalothorax has the same outline and dorsal grooves 
as the subadult type, but it differs in a curious feature. A me- 
dian area, nearly triangular with the sides convex, is raised 
l)etween the end of the median groove and the posterior margin 
as a moderately large, rounded j^rotuberance or rather low 
tubercle badly marked off, which in the male is directed mainh' 
upwards, in the female upwards and considerably backwards, 
so that in the latter sex the cephalothorax seen from above 
(fig. 2 a) has the median part of its posterior margin very con- 
spicuously ccmvex. 

Tlie cucullus (fig. 2 b) seems to be slightly broader in pro- 
])ortion to length than in young specimens, and a sublateral 
oblong impression is distinct, but real grooves are not developed. 

The palps (fig. 2 c) have the femoral part a little or conspi- 
cuously more than half as thick as long. The tibial i)art is ador- 
ned with numerous granules on more than its distal two-thirds, 
and these granules are oblong and rather coarse on the most 
distal third. - (As usual in adults the thick connecting skin bet- 
ween tlie large abdominal tergites has nearly disappeared). 



Pedipalpi, Ricinulei, and Opiliones. 25 



Sexual differences are found in the above-named protu- 
berance on the cephalothorax and in the three anterior pairs 
of legs. First legs are scarcely or distinctly thicker in the male 
(fig. 2 d), than in the female (fig. 2 e), but at least the shape of 
the metatarsus is rather different in the two sexes ; in the female 
the metatarsus is moderately slender, with the same thickness 
in its whole length, and its lower, inner margin is feebly convex; 
in the male the metatarsus is thicker, increases a little in 
thickness from the base to a little from the end, and the distal 
two-thirds of its inner margin is much more convex than in 
the female. Second legs are somewhat varying, as the}- may 
be similar in both sexes or a little stouter in the male. 

The copulatory organs (fig. 2 g) on third pair of legs on the 
whole similar to those in C. W ester mannii Guer. and C. Sjo- 
stedtii Hans. & Sor. ; of course the movable process of first tar- 
sal joint has a shape of its own and affords as usual excellent 
specific characters. It may be mention(^d that the metatarsus 
from in front (fig. 2 g, 6) is somewhat more than twice as long- 
as thick; its movable process has the most distal slender part 
curved ver\' considerably backwards. First tarsal joint {d) 
is uncommonly long, being, seen from below, only somewhat 
shorter than second and very much longer than the third, 
which is somewhat shorter than the fourth. The second tarsal 
joint {e) has the distal triangular process {e^) of the front wall 
rather small, while the lamina cyathiformis {e"), is high, and 
consequently the upper end of its wall is more than twice 
as far from the distal lower end of the joint than is the case 
with the tip of the process from the front wall. The movable 
process {d^ — d~) of first joint has the distal segment about twice 
as long as the proximal; the distal segment seen from the 
outer side (fig. 2 h) is rather narrow, the proximal process from 
its front margin is situated considerably before the middle^ 
and is somewhat short, oblong-triangular, and very acute; 



26 Studies on Arthropoda. I. 

the distal part of second segment is produced into three pro- 
cesses, but the two drawings (figs. 2 g and 2 h) may convey a 
sufficient idea of the curious shape of the whole part. 

In the male the body is 4.4 mm. long, second leg 7 mm.; 
in the female the body is 4.7 mm. long, second leg 6.9 mm. 

Remarks. — C. crassipalpe is easily separated from the 
other species hitherto known of the genus in having the bod\- 
and most of the legs covered with very broad scales, which are 
not much longer than Inroad or almost as broad as long. 

Occurrence. — Island of Fernando Poo, at two places: 
Musola, altitude 400—500 m., Januar}- 1902, 2 males, 3 females, 
I immature specimen; Basile, altitude 400 — 600 m., August — 
September 1901, i young specimen. 

Distribution. The specimens described by Hansen and 
Sorensen were from Kameroon, the part of the continent oppo- 
site to the Isl. of Fernando Poo. 

Cryptostemma Feae n. sp. 

ri. IT, figs. 3 a — 3c; PI. Ill, figs. I a — i i. 

A. Adult Female. 

Closely allied to C. Ajzclii Thorell as described by Hansen 
and Sorensen, so that it ma\ be advisable to lay special stress 
on the differences. 

Cephalothorax slightly longer than broad. Its surface is 
adorned with numerous moderately large granules conspi- 
cuously lower than high and of a special structure, as their 
outline is circular, the upper surface ver\' moderately arched, 
marked off from the sides by a sharp outline, very finely stria- 
ted, and besides ver\^ dark coloured, brownish l>lack, while 
the; sides of the granules are greyish; between the granules a 
number of feebly clavate hairs somewhat or considerably lon- 
ger than the height of the granules; the whole adornment of 
the surface differs ^rom that in C. Afzelii in having the granules 



Pedipalpi, Ricinulei, and Opilioiies. 7,y 

conspicuously less closely set and the hairs longer. The lateral 
margin of the cephalothorax is considerably less concave be- 
fore the submedian angle than in C. Afzelii, and the angle 
mentioned is less protruding. The median groove is well devel- 
oped, reaches to or nearly to the front margin, is posteriorly deep, 
and at its end is seen a short transverse groove. The marginal 
groove — along the anterior half of each lateral margin — is well 
developed; as in C. Afzelii two grooves forming a right angle 
with one another are seen on the area between the median 
groove and the submedian angle of the lateral margin. 

Cucullus (fig. 3 a) as long as broad at the base, and beyond 
the middle half as broad again as at the base ; the submarginal 
grooves moderately developed. The surface is adorned with 
numerous granules, which are smaller but proportionately 
bigher than on the cephalothorax and otherwise of the same struc- 
ture; betw^een them are found a number of minute granules 
and besides hairs; these hairs are considerably longer than the 
height of the larger granules, subclavate, and towards the front 
(lower) margin of the cucullus the hairs are longer, cylindrical, 
at the margin still longer and acute. — The cucullus differs 
from that in C. Afzelii in having the granules less closely set. 

Abdomen above with granules and hairs quite as cephalo- 
thorax ; from the middle to the posterior end the granules are 
gradually a little smaller and more numerous. On the ven- 
tral side the granules are very numerous and quite small and 
low, but somewhat larger towards the posterior end. 

The antennae (fig. 3 b) as in C. Afzelii with five teeth on the 
distal posterior margin of first joint, but they differ in being 
longer than broad; the inner margin of second joint is more ar- 
med than in C. Afzelii, as it possesses a row of about eight 
rounded teeth. 

The palps (fig. i a) ha\-e the trochanterical part on the front 
side set with numerous small granules, wdiich are much larger 



ZS Stvidies on Arthropoda. I. 

near the lower margin. The femoral part a little more than 
twice as long as thick, especially on the anterior side with a 
good number of small granules and numerous hairs, most of 
them moderately short, but no impressed points are found. 
The tibial part is slightly or a little thicker somewhat from the 
base than at the middle, but towards the end it is feebly or very 
feebly thickened, conspicuously less so than in (\ Afzclii; its 
anterior side, excepting a proximal part, is granulated, and 
the granules are nmch larger on the distal third, largest and 
l)esides oblong towards the end; on the proximal major part 
the hairs are as on the preceding joints, while several of the di- 
stal hairs are long and simple. 

The legs have their joints, excepting the tarsi, adorned 
with numerous granules of the same quality as those on 
the surface of cephalothorax, but they are on the whole pro- 
portionatel}' a little higher, less broad, with the terminal dark, 
striated area nmch smaller (fig. ; c) ; the hairs as on cephalo- 
thorax ; the adornment differs from that in C. Afzclii in having the 
granules less dense and the hairs somewhat longer. All femora 
with a conspicuous longitudinal furrow above ). Tibia of second 
pair with a distinct, that of fourth pair with a less distinct 
longitudinal dorsal furrow, which is wanting on first and third 
tibia; first tibia twice as long as thick. Fourtli joint of second 
tarsus half as long again as the fifth, which is distinctly longer 
than third joint. 

Colour dark reddish brown, with the limbs a little lighter, 
more reddish. Some ten specimens probably captured a short 
time after the last mould, have the body lighter reddish brown 
and the limbs more or less ferrugineous. 

Length of tlie body 6.4 mm., of second leg 8.5 mm. 



^) Here 1 inaj- correct a inis.soript in Hanson and Soren.sen'.s paper. In the 
description itself of C. Afzclii the femora are correctly .stated to have 
;i longitudinal dorsal furrow, but in the diagnosis is said: "femora supra 
non sulcata" and the word "non" must be cancelled. 



Pedii)al[)i, Ricinulei. ami (^piliones. 2Q 

Male. 

Similar to the females in most features, but differs strongh* 
in thickness and shape of the two anterior pairs of legs. First 
legs (fig. I b) are on the whole stouter than in the female (fig. 
I c) ; tibia and metatarsus are nmch thicker and differently 
shaped. Tibia only half as long again as thick (in the female 
twice as long as thick), thickened below but without any pro- 
cess; metatarsus widened below, with the inferior margin con- 
spicuously more convex than in the female. — Second legs 
(fig. I b) extremely altered ; femur always enormously thickened 
upwards and downwards, but yet showing considerable indi- 
vidual variation; in the specimen figured the femur is a little 
less than twice as long as deep at the middle (in the female 
(fig. I c) about four times as long as deep) , but in one specimen 
it is scarcely two-thirds as long again as deep; the femur has 
therefore both the upper and the lower margin very convex 
but it is much compressed; besides the granules on the major 
part of the anterior and the posterior side are feeblj^ developed, 
and in some specimens the dorsal longitudinal furrow has 
disappeared, while it is distinct in other specimens. Tibia on 
the lower side near the base produced into a long and thick, 
obtuse, a little curved process directed downwards and forwards, 
and granulated as the remainder of the joint. Metatarsus has 
frequently a short portion near the middle of the lower margin 
distinctly produced as an oblong, low protuberance, but in 
other specimens this expansion is very feeble. 

The copulatory organ, excepting its movable process on 
first tarsal joint, agrees so closely with that of C. Karschii 
Hans. & Sor. described and figured in 1904, that it may be 
omitted here. As usual the movable i)rocess mentioned has a 
shape of its own, affording specific characters. Its second seg- 
ment is more than twice as long as the first (fig. i e) ; seen from 
the outer side (fig. i f) its major distal part is lamellar and rather 



^O studies on Arthropoda. !. 

I 

broad ; the proximal process at its front margin protrudes before 
the middle of the segment, is long, broad at the base and tapers 
towards the broadly obtuse end. The most distal part of the 
segment has a moderately long, cur\^ed, distally slender process 
and besides a pair of short, triangular processes; the compli- 
cated shape may be judged from the two figures. 

Length of the l)ody 6.6 mm., of second leg lo mm. 

Immature Specimens. 

Specimens not far from full-grown differ as usual from the 
adult female in the colour ; the cephalothorax is yellowish ferru- 
gineous, the abdomen greyish and the legs browni.sh yellow; 
furthermore, in the palps the femoral part is more slender, the 
tibial part has the subbasal portion conspicuously thicker 
than the middle, and it increases scarcely in thickness from 
the middle to the end. 

A younger specimen, which has only four joints in the tarsi 
of fourth pair but the full number in the other pairs, is 4.1 mm. 
long, with second leg 5.6 mm.; it has the palps more slender 
than in older specimens, and the sublateral impressions on the 
cucullus are scarcely discernible. 

One of the smallest specimens (the real larvit excepted) 
is 3.1 mm. long with second leg 4.2 mm. It has only four joints 
in second tarsus, three joints in third, and two joints in fourth 
tarsus. Cephalothorax has the median groove well developed 
excepting towards the anterior margin, but some of the other 
grooves are partly or wholly indi-stinct, and the cucullus has no 
sublateral impressions. The animal is more yellowish than older 
specimens. 

The lar\a \\\\\\ three i)airs of legs has been described abov-e 
(p. 22). 

Remarks. — As already mentioned C. Fca is uUied to 
C. Afzclii, but it is much smaller, the granules on body and 



Pedipalpi, Ricinulei, and Opiliones. 3^ 

limbs are more scattered, and the distal portion of the tibial 
part of the palp is less thickened. As the male C. Afzelii is 
unknown, it is at present impossible to point the secondary 
sexual differences between the males of the two species. 

Occurrence. — Portuguese Guinea (near lat. 12° N.), at 
two places: Bolama, June — December 1899, many specimens; 
Rio Cassine, January — April iqoo, numerous specimens. 



ORDER OPILIONES 

This order comprises three sub-orders, viz. Palpatores, 
Laniatores, and Cyphophthalmi. The collection contains a 
rich material of the Laniatores, but with the consent of Dr. 
Gestro it has been handed over to a colleague. The small but 
interesting material of the two other sub-orders is dealt with 
on the following pages. 

Sub-Order O. Palpatores. 

It is an interesting and curious fact that Mr. Fea, who was a 
most excellent collector, has secured only two specimens du- 
ring his voyages in West Africa; animals of this sub-order 
must consequently be very scarce in the countries investigated. 
Both animals are fortunately adult males; they belong to two 
different genera, and both belong to the family Phalangioidse. 
A few years ago Dr. C. Fr. Roewer published the two first 
l)arts of his monograph of the Palpatores in: Abhandlungen 
aus dem Gebiete der Naturwiss. heraug. vom Naturwiss. Ver- 
ein in Hamburg. The second part of this work has the title: 
Revision der Opiliones Palpatores (^ Op. Plagiostethi) , II 
Teil; Familie der Phalangiidse (1. c. Vol. XX, I, 1912) ; it deals 



J2 Studies on Artliropoda. I. 

with three of the six sub-families into which he divided the 
family. On p. lo he separates the sub-family Phalangiina* 
from the iive others in having no horizontally protruding pro- 
cess on the lower side of first joint of the antennse (he named 
these appendages mandibles), while such a process is constantly 
found in all the other sub-families. According to this definition 
my two genera belong to his Phalangiina?. But it has been 
more difficult to refer them to any of the numerous genera 
accepted or established b}' Dr. Roewer; one of them I refer to 
an old genus, but I find it necessary to establish a new genus 
for the reception of the other. 

Dacnopilio insularis n. s]). 

ri. Ill, figs. 2a -Ml. 

Male. 

Body oblong, with the posterior margin, seen from above, 
somewhat flatly rounded; the skin coriaceous and ratlier flexible. 
Surface of cephalothorax with a considerable number of con- 
spicuous, triangular, acute denticles distributed in the following 
way : a transverse group of about sixteen such denticles occupies 
the anterior half of the area in front of the ocular protuberance, 
and some among these spines are placed on the front margin ; 
obliquely outside and in front of each antero-lateral angle of 
the ocular protuberance a single denticle is found, while each 
more lateral part of the surface has eight or nine denticles. 
The supra-antennal area has below a pair of very distinct den- 
ticles, and between this pair and the upper margin rudiments 
of a second pair. The two posterior thoracic segments are well 
marked off, and each has a transverse row of similar teeth. 

The ocular protuberance is middle-sized, longer than liigli, 
without an>' median furrow, but armed with se\'en rather 
low teeth distributed in two rows (fig. 2 b) ; the protuberance 
is scarcely as long as its distance from the front margin. 



Pedipalpi, Riciimlei, and Opiliones. 33 



The five anterior abdominal tergites each with a transverse 
band of small denticles: on the fifth tergite this band is narrow, 
consisting of a row and closeh* in front of this some few dentic- 
les; on the other tergites the band is broader and the denticles 
are placed more irregularly. On the sixth tergite denticles 
are ver^' feebly developed, while the two following tergites 
are smooth. 

The antennae are considerably elongated and thickened, 
stretched out nearly as long as the body. First joint is directed 
forwards and considerably upwards, twice as long as thick, 
somewhat cur^'ed, with most of the upper margin rather con- 
vex (fig. 2 c) , while the lower margin has its long middle part 
concave and is convex towards both ends. The upper surface 
of the joint has a number of denticles a little smaller than those 
on the cephalothorax, while the major posterior part of the 
lower surface has a lower number of somewhat longer teeth; 
the sides are nearly smooth. Second joint twice as long as, 
but not thicker than, the first, and measured to the tip somewhat 
more than three times as long as third joint; seen from the side 
the upper margin of second joint is mainly directed forwards 
and conspicuously concave which is due to a pronounced trans- 
verse impression; the joint from the upper margin to the inser- 
tion of third joint is a little fusiform, and the upper part of 
its front surface has a moderate number of somewhat small 
denticles. 

Palps somewhat longer than the bod}', moderately strong, 
%vith the tarsus slender. Trochanter obliquely triangular with 
some teeth on the upper side. Femur a little longer than the 
tarsus, increasing in thickness from base to near the end, suban- 
gular, with longitudinal rows of small, acute denticles. Patella 
without apophysis, but with some quite minute denticles on 
the upper surface. Tibia without apophysis, a little curved, 
on the lower side with one denticle or two minute denticles 

3 



]4 studies on Arthropoda. I. 



somewhat from the base, and a minute denticle at the terminal 
margin. Tarsal claw simple. 

Legs of middle length. P'irst coxae with a considerably num- 
ber of granules, each with a short, black seta; the other coxae 
with the granules and setae more feebl}^ developed. First pair 
of legs with femur, patella and tibia considerably thicker than 
in third or fourth pair, and much thicker than in second pair. 
Femora of first pair only a little shorter than the body, conspi- 
cuoush' claA'ate (lig. 2 d) and somewhat cur\'ed, with longitu- 
dinal rows of well developed, triangular denticles and a number 
of scattered denticles, but without any process at the end. Se- 
cond femora considerably longer than the body; the three po- 
sterior pairs of femora with rows of denticles distinctlj' smaller 
than those on first pair. Patellae and tibiae of all legs with the 
denticles conspicuously smaller than those on the respective 
femora and partly rudimentar\- ; tibiae more or less subangular. 

Colour of the body yellowish-grey, with some dark spots 
above; palps light yellow, but their femora, excepting at the 
end, the proximal part of tibiae and the end of the tarsi dark. 
Legs in the main as the palps, with the distal part of the femora 
and patellae brownish, while the subdistal i:)art of the tibiae 
is only slightly darker than the proximal jiart. 

Length of the body 5.6 mm., first legs (without coxae) 20 
mm., second legs 34.5 mm. 

Remarks. — This new species is more related to Dacno- 
pilio Roewer than to any other genus. Roewer has described 
two species, which differ much from one another in armature 
and probably also in size of the mandibles in the male, and the 
new species seems in some features to l)e intermediate between 
the two other African forms. 

Occurrence. — The single specimen, an adult male, was 
taken in February iqo2 in 1300 — 1400 m. above the level of 
the sea at Moka on tlie Island of Fernando Poo. 



^ Pedipalpi, Riciniilei, and Opiliones. 35 

Megistobunus n. gen. 

Allied to Megahumis, but differs in several features. 
Body very small, with the dorsal skin coriaceus and rather 
hard, excepting on the three posterior abdominal segments. 
Cephalothorax highly vaulted (fig. 3 a), even somewhat coni- 
cally raised above, and last segment distinct; no frontal median 
spine: supra-antennal area without denticles. Abdomen has 
the tergites of the five anterior segments fused nearly as a 
.shield, but each segment has a spiniform process or denticle 
in the median line. 

Ocular protuberance extremely large, constricted at the base, 
and adorned with a few pairs of extremely long processes; 
e3'es proportionately small. 

Antennse somewhat small, normal and simple in the male; 
first joint without ventral process. 

Palps (fig. 3 c and 3 d) longer than the body, moderately 
slender. Femur with longitudinal rows of triangular denticles, 
but without longer processes. Patella with a half dorsal, half 
interior apophysis which is long and somewhat slender. Tibia 
on the inner side produced into a quite short but distinct apo 
physis. Tarsus nearly as long as the femur; claw simple. 

lyCgs extremeh^ long. First coxae scabrous below, as they 
have a number of irregularly scattered granules while the 
other pairs are less scabrous, and no pair possesses any regular 
row of denticles at the anterior or posterior margin. Femora 
similar in shape and thickness, without any secondary articu- 
lation, but ^^•ith rows of small denticles; no elongate spines 
at the end of femora or patellae. 

Remarks. — The genus is established on a single adult male. 
The animal is in general a.spect rather similar to a very long- 
legged Liobunum, a genus from which it is very remote in im- 
portant features. It is related to Megahunus, but differs from 
the forms of this genus in the following features: the dorsal 

3* 



36 studies on Arthropoda. I. 



cuticle of the cephalothorax and of the major part of abdomen 
is rather hard ; the front margin of the head without any strong 
median process; the five coalesced abdominal tergites each 
with a median spine; the femora of the palps without processes 
and only with normal denticles below ; the legs extremely long, 
and all denticles quite small ). 

Megistobunus longipes n. sp. 

PI. Ill, tigs. 3a — 3d. 

Male. 

Cephalothorax highly vaulted ; its penultimate segment mo- 
derat.el}^ its last segment well, marked off. The front margin 
with a few small denticles. Above the openings of the stink- 
glands is at each side a longitudinal row of three oblong-trian- 
gular processes, and a smaller denticle is seen near the base 
of the ocular protuberance; between this protuberance and the 
frontal margin several minute denticles are scattered. The 
first thoracic segment has at the front margin a transverse row 
of three pairs of conspicuous teeth and a couple of denticles 
at the lateral margin; second segment has a transverse row 
with some few denticles. 

The ocular protuberance extremely large, with three pairs 
of conical, slender and very diverging processes (fig. 3 b) ; the 
third pair arc exceedingly long, a little longer than first pair, 
and about twice as long as second pair. The protuberance, 



^) When I looked over Dr. Roewer's descriptions of the genus M e g a b u n u s 
and its three species, I found his work very far from satisfactory. In the 
diagnosis of the genus he says: "Palpen kraftig", but in the description 
of M. diadem a: "Palpen klein und schwach", and at the two other 
species respectively: "Palpen klein" and: "Palpen kiirz". In the diag- 
nosis of the genus he says: "Beine lang and diinn", in the description 
of M. diadema: "Beine nicht sehr lang; diinn", of M. rhinoceros: 
"Beine kiirz", and of M. (irouvellei: "Beine diinn, wenig lang". That 
generic characters as those quoted on Meg a bun us are .sheer nonsense 
can scarcelv be denied. 



Pedipalpi, Ricinulei, and Opiliones. 37 

excepting a lateral portion around each eye, is set with 
numerous denticles, among which those in a narrow belt 
below the eyes are somewhat large, and the denticles diminish 
nmch in size downwards and disappear near the base of 
the protuberance; the processes are armed with conspicuous 
denticles. 

The abdominal shield has a median row of five spiniform 
processes or teeth ; the process on second segment is larger than 
that on the first, and the latter is larger than the third, while 
those on fourth and fifth segment are mere denticles; first and 
second segment each besides with a transverse row of two pairs 
of conspicuous denticles, while on the three other segments 
corresponding denticles are small or wanting. The free abdo- 
minal tergites and the ventral surface of the abdomen nearly 
smooth . 

Palps not inconsiderably longer than the body. Trochanter 
distally with a few granules above and below. Femur a little 
cur\'ed and slightly thickened towards the end, with rows of 
denticles on the lower side and granules on the upper surface. 
Patella with two marginal rows of granules above ; the apophy- 
sis slightly longer than the patella itself (fig. 3 c and 3 d) and 
somewhat shorter than the tibia, moderately slender, but with 
the subdistal part slightly thickened and everywhere clothed 
with short hairs. Tibia proximally with very incomplete rows 
of granules; the terminal inner apophysis (fig. 3 d) is short, 
shorter than broad, hair>\ 

Legs extremely long, ver>' slender. Coxae of first pair with 
a moderate number of irregularis^ scattered, setiferous granules, 
while the other pairs are less scabrous. Femora subsimilar, 
excepting that second pair are distinctly more slender than 
the first, and all with rows of small denticles; patellae with very 
few denticles, while the denticles on the tibiae are very small 
and partly rudimentary. 



38 Studies on Arthropoda. I. 

Colour of the body, antennse, palps and coxae 3'ellowish; 
the upper side of abdomen, trochanter and femur of the palps 
with a brownish tint, the patellar apophysis and the apex of 
the palps brown, the distal part of the antennal fingers black; 
ocular protuberance and its processes black excepting the 
constricted base. Legs dark yellowish brown, patella; dark 
brown, trochanter and base of femora brownish black. 

Length of the body 3.8 mm., first legs 28 mm., second legs 
64.5 mm., fourth legs 44.5 mm. 

Remarks. — In general aspect rather similar to Liohunitm ; 
second pair of legs seventeen times as long as the body. 

Occurrence. The single specimen was captured in Au- 
gust — September 1901 in an altitude of 400 — 500 m. at Basile 
on the Island of Fernando Poo. 

Sub-order O. Gyphophthalmi. 

The main work on this sub-order is H. J. Hansen and W. 
Sorensen: On two Orders of Arachnida. Cambridge 1904. It 
contains descriptions of two genera from Africa, each with a 
single species, viz. Ogovia grossa H. J. H. & W. S. from Ogove 
in sequatorial France, and Purcelliu illnstrans H. J. H. & W. S. 
from Cape. The material to hand comprises a new species of 
Ogovia and another form, for the reception of which a new 
genus must be established. 

Ogovia Hans. & Sor. 
This characteristic genus hitherto comprised a single spe- 
cies, 0. grossa, established on an adult female, and the male 
was unknown. The new species, though easily separated from 
0. grossa, agrees excellently with every cliaracter found in 
the generic description excepting as to two particulars, viz. 
that the produced median frontal plate is so large that it cannot 
be called "brevior", and that the inner marginal keel of first 



Pedipalpi, Ricinulei, and Opiliones. 39 



pair of coxse is proportionately somewhat shorter, being scar- 
cely half as long again as the distance between its end and the 
tip of coxa. As both sexes of the new species are to hand, it 
may be added to the generic description that the sexes differ 
from one another not only in the two usual points, viz. the shape 
and position of the genital aperture and the male process on 
the upper side of fourth tarsus, but besides the abdomen has 
in the male behind the genital aperture a process directed much 
backwards, and the median anterior part of the ventral abdo- 
minal surface is excavated. 

Ogovia nasuta n. sp. 

PI. Ill, figs. 4a — 4c; pi. IV, figs- I a — if. 

Male and Female. 

Body thick, scarcely three-fourths as long again as broad, 
considerably broader than deep, with the median part of the 
dorsal surface of abdomen and of the hind part of cephalothorax 
rather flatly convex. 

Cephalothorax anteriorly produced into a proportionately 
large, triangular and much vaulted plate (figs, i a and i b) 
which is somewhat broader than long with the end subacute, 
the lateral margins a little convex, and the surface densely 
and finely granulated; this plate overlaps the major part of 
first joint of the antennulse. The lateral surface between the 
lamina mentioned and the "conus foetidus" — the process with 
the aperture of the odoriferous gland at the end — is somewhat 
hollowed, so that the anterior median part of the cephalothorax 
is shaped about as a ver\" broad, rounded keel; the posterior 
transverse furrow is much arcuate, posteriorly at the middle 
flatly concave and a little or considerabh' nearer to the furrow- 
limiting the abdomen than the length of the first abdominal 
tergite at the median line. The surface is ver^'- densely and 
finely granulated and besides, excepting anteriorly on the sides. 



^.0 Studies ou Arthropoda. I. 



adorned with a number of much larger, scabrous and badly 
defined granules or rather quite small, irregular tubercles. 

The "coni foetidi" are subtriangular (fig. i a), broader than 
long, directed outwards and somewhat forwards; their anterior 
margin is, seen from above, somewhat convex to a little from 
the protruding terminal part, but seen from in front this shape 
is seen to be due to the fact that each conus has its front subter- 
minal part considerably excavated at the anterior end of the 
two labial plates limiting the terminal orifice. 

Abdomen above with granules as on the major part of ce- 
phalothorax. The seven anterior tergites divided by a shar]> 
furrow in the median line. On the \'entral side (fig. i b) the 
first furrow is curved forwards inside the spiracle and reaches 
the margin of fourth coxa. The anal operculum is somewhat 
vaulted, and seen partly from behind distinctly broader than 
long. The whole ventral surface is densely and moderately 
finely granulated, excepting the hollowed median part in the 
male mentioned later on. 

Antennae (fig. 4 a) nearly as in 0. grossa. I'irst joint has the 
dorsal transverse rounded keel very conspicuous, and the upper 
and most of the outer side of the free part of the joint is dis- 
tinctly granulated. The hand of the chela is somewhat less 
than three times as long as broad, smooth on the anterior side, 
and not fully twice as long as the movable finger. 

Stomotheca conspicuously more than half as long again as 
broad at the distal end (figs, i c and i d). 

Palps (fig. 4 b) about as long as the antenna^ measured from 
the proximal margin of the transverse dorsal keel, in the main 
as in O. grossa, yet differing in having the trochanter a little more 
slender and femur more oblique with l^otli tlie upper and the 
lower margin more convex. 

T^egs ver>' short and thick, considerably shorter than in 
0. grossa. I^ower surface of first pair of coxae not lower than 



Pedipalpi, Ricinulei, and Opiliones, 41 

on second pair. Femora of first pair (fig. 4 c) conspicuously 
cur\'ed upwards with the lower margin rather concave excepting 
towards the base, of second pair feebly curv^ed, while the two 
other pairs are straight (fig. i e). Tibiae somewhat longer and 
deeper than the patellae, in the anterior pairs about half as long 
again as deep, in fourth pair proportionately a little longer. 
Tarsi of first pair (fig. 4 c) slightly less deep than the tibiae, 
and the "solea" occupies little less than half of the lower mar- 
gin of the tarsus. 

The upper side of the body with numerous extremely short, 
thick, fusiform or rather clavate hairs which increase a httle 
in number posteriorly, so that they are closer on the posterior 
abdominal tergites than on the cephalothorax ; the ventral 
surface has a number of normal hairs which are longer and thicker 
on the coxae than on the abdomen, where they are short 
and \'ery thin. The legs, excepting their tarsi, have a number 
of \-er\^ short, fusiform hairs on the upper surface and normal 
hairs on the lower side of the joints, while the tarsi have only 
normal hairs which on the lower side are longer than on the other 
joints — excepting of course the solea of first pair which is 
densely clothed with verj^ short and thin hairs. The palps 
have the three distal joints well haired, while the hairs on their 
femoral part are shorter and less numerous, and the trochanter 
has only some hairs on the most distal part; the first joint of 
the antennae has a \'er>' moderate number of hairs. 

Sexual differences well developed. In the female the tar- 
sus of fourth pair of legs normal, shaped as the preceding pair; 
in the male the fourth tarsus (fig. i e) increases nearly gradu- 
ally conspicuously in height from rather near the base to beyond 
the middle, and it is above at the middle produced into a 
moderately large, nearly regularly conical, .slighth' curved 
process which is somewhat longer than deep at the base 
and directed l)ackwards and considerably upwards; this 



^.2 Studies on Arthropoda. I. 



process is scarcely di\-ided into a proximal and a much shorter 
distal part. 

In the male the genital aperture (tig. i c) is nearly twice as 
broad as long, with the anterior margin considerably curved, 
the posterior margin feebly concave; the genital arculi [m) 
posteriorly rather narrow, anteriorly considerably widened 
inwards, at the median line about as long as the aperture, 
and with the anterior, the inner, and the posterior margin of 
their front part distinctly raised. No genital operculum is 
marked off, but a little behind the genital aperture projects 
a large, broad, oblong, subacute process (p) directed down- 
wards and mainly backwards; this process is somewhat depres- 
sed and even slightly hollowed on the proximal part of its lower 
side; at the sides and behind this process the ventral surface 
of the abdomen is considerably excavated, and the excavation, 
which is oblong, occupies more than half of the surface between 
the base of the process and the front (lower) margin of the 
anal operculum. — In the female (fig. i d) the genital aperture 
is still shorter than in the male; the genital arculi are not mar- 
ked off from the coxa? and each terminates in a subacute angle. 
The genital operculum is not marked off posteriorly, in front 
it is very narrow with the end cut off transversely, and it is 
deeply impressed just behind that end; the surface of abdomen 
has no process and no excavation as in the male. 

Colour dark. Body and first joint of the antennae from dark 
reddish-brown to blackish brown; the distal joints of the an- 
tennae and the legs more or less dark reddish-brown, with the 
tarsi frequently a little lighter; palps brownish yellow, with 
tlie distal joints lighter. 

In the male figured the body is 3.45 mm long and 2 mm 
broad, cephalothorax 1.6 mm long, first leg 2.5 mm, fourth 
leg 2.6 mm. The single female is 3.52 mm long and 2.07 mm 
broad. 



Pedipalpi, Riciuulei, and Opiliones. 43 



Remarks. — 0. nasiita is distinguished from the female 
of 0. grossa b}' a number of features, and some among them 
may be pointed out. The frontal plate overlapping the antennae 
is much longer and broader than in 0. grossa; the granulation 
on the upper side of the body is \-ery different, the legs con- 
siderably shorter in proportion to the body, and the solea of 
the first tarsus is conspicuously longer in proportion to the- 
length of the whole lower margin of the tarsus. 

A curious feature may be mentioned here. On the two figu- 
res of the legs (figs. 4 c and i e) are seen some rings (x), viz. 
femur, tibia and tarsus each with two rings (the distal ring of 
fourth femur placed on its upper margin), patella and meta- 
tarsus each with a single ring; each ring signifies a circular 
area which is well marked off, considerably vaulted and very 
smooth and shining. Similar areas are also found both above 
and below on the body; on fig. i b an area (x) is seen on each 
■coxa, and a pair on most of the abdominal sternites. Similar 
vaulted,' shining areas has also been obser\'ed in 0. grossa, but 
they have not been found in any other genus of the suborder. 
One is inclined to think that these areas, which as to outer 
shape and aspect are somewhat similar to small eyes, may 
be organs of some quite unknown significance. 

Immature Specimen. — The animal (fig. i f), measuring 
2.3 mm in length, is proportionately broader than the adults. 
The body is yellowish brown; the articulating skin between 
the five posterior abdominal sternites constitutes darker trans- 
verse bands; the legs are light yellowish. Fig. i f as compared 
with fig. I b shows several differences due to age : lobe of second 
pair of coxae feebly developed, the surroundings of the future 
genital aperture ver>- dift"erent from the structure in the adult, 
with broad membrane around the arculi (;«) (first abdominal 
sternite), finally the bands of articulating skin between the 
abdominal sternites. 



\A Studies on Arthropoda. I. 

Occurrence. — The material, consisting of 3 adult males, 
I adult female and an immature specimen, was taken at Musola, 
Island of P'ernando Poo, January 1902, in an altitude of 400 — 
500 m. 

Parogovia n. gen. 

Cephalothorax anteriorly produced into a triangular plate 
(fig. 2 a), which covers at least the inner part of the transverse 
keel on first joint of the antennae. A median keel on cephalo- 
thorax is not developed. 

Eighth and ninth abdominal sternites completeh^ fused, 
and besides fused with the ninth tergite so that not even a 
vestige of any furrow between these plates is seen (fig. 2 b) . 

No eyes. 

"Coni foetidi" moderately long, seen from above nearly 
obUquely triangular, directed outwards and feebly upwards, 
and about half as long from the lower margin of the cephalo- 
thorax as the length of the basal diameter of the cone. The 
end of each cone has a ver}' oblong and ^'ery convex, smooth 
and shining area or piece, which looks almost as a single mode- 
rately thick lobe. 

First joint of the antenna" (fig. 2 e) not reaching to the end 
of the fenmr of the jjalps, considerably compressed, on the lower 
side raised as a large, rounded tubercle, and above with the 
proximal transverse keel well developed. The teeth on the mov- 
able finger subsimilar in shape, moderately high, rounded; 
on the fixed finger two kinds of teeth alternate (fig. 2 g), half 
of them being larger than on the movable finger, moderately 
high and rounded, while each interval between two such teeth 
is occupied with an extremely low but somewhat broad tooth, 
with the free margin a httle convex. 

The palps (fig. 2 h) have the trochanter proximally slender, 
distally much thickened with some granules, somewliat shorter 



Pedipalpi, Ricinulei, and Opiliones. 45 

than the femur which is slender and scarcely compressed; the 
tarsal joint scarcely as long as the tibial. 

First pair of coxse (figs. 2 c and 2 d) anteriorly produced 
into a triangular process; the distance between the front end 
of that process and the angle terminating the marginal keel 
of the stomotheca is only somewhat shorter than the length 
of the last-named keel. Stomotheca with the middle parts of 
its limiting keels nearly parallel, and towards the end it is a 
httle widened, only somewhat narrower at the front end — 
between the angles — than the length of the marginal keels. 
The thin-skinned part of each maxillar\' lobe (fig. 2 c, h) is 
divided into two portions distant from one another; the basal 
portion is normal, while the distal portion constitutes a freeh' 
protruding, oblong, rounded lappet. 

First pair of coxae considerably broader than those of second 
or third pair, and much narrower than fourth pair (fig. 2 b). 

Metatarsi conspicuously more than half as long as the tarsi. 
Solea of first pair of tarsi (fig. 2 i) occupies considerably more 
than half of the length of the lower margin of the tarsus, and 
is rather well defined from the proximal part of the lower side. 

Tarsi of first and second pairs without any longitudinal 
dorsal furrow. 

Claws simple, without saw-teeth; those of fourth pair of 
legs (fig. 2 k) much longer than first pair (fig. 2i), 

Sexual differences only found in the genital aperture with 
the arculi, and in the tarsi of fourth pair. 

Remarks. — The genus Parogovia, which most decidedly 
belongs to the subfamily Stylocellini H. J. H. & W. S., agrees 
with Ogovia and differs from Stylocellus especially in five cha- 
racters, viz. that the cephalothorax is anteriorly produced 
into a triangular plate, that it has no eyes, that eighth and 
ninth abdominal sternites are completely fused with one an- 
other and with the ninth tergite, that first coxae are produced 



^.6 studies on Arthropoda. J. 



into a triangular process, and that the two anterior pairs of 
tarsi have no dorsal furrow. It agrees with Styloccllns and 
differs from Ogovia in having the claws on fourth pair of tarsi 
much longer than first pair, and in some features in the palps, 
as their femoral part is not expanded and conspicuously longer 
than the trochanter, their tibial part longer than the tarsus. 
It differs from both genera in having the membranous part 
of the maxillary lobe of first coxa; developed in a peculiar way, 
in the aberrant terminal part of the coni foetidi, in having 
no trace of dorsal keel on the cephalothorax, finally in the 
antennulae, as the first joint has a single large tubercle on the 
lower side, and the fixed finger has two kinds of teeth. 

The single species known is in general aspect somewhat similar 
to Siro, though it decidedl}^ belongs to the other subfamily. 

Parogovia sironoides n. sp. 

PI. IV. fi?;.s. 2 a — 2l. 

Bod}- (fig. 2 a) • moderately thick, more than three-fourths 
as long again as broad, nmch broader tlian deep. 

Ce])halothorax almost as long as the abdomen (fig. 2 a) , 
anteriorly without real lateral excavations and without an>- 
median keel; the front lamina of moderate size, triangular, 
broader than long, subacute. The posterior transverse furrow 
is much arcuate, \'ery feebly developed excepting at the middle, 
where it is more distinct and flatly curved, and here its distance 
from the furrow limiting the abdomen is three or four times 
shorter than the length of first abdominal segment. The sur- 
face of cephalothorax is rather densely granulated; the gra- 
nules are well developed, not confluent; the lower side of the 
coxae more finely granulated. 

Coni foetidi nearly as long as broad; the end is rounded 
and shows, seen from above, an oblong and oblique, smooth, 
shining area. 



Pedipalpi, Ricinulei, and Opiliones. 47" 

Abdomen with the tergites undivided in the median line 
and densel}^ granulated; the granules, which on first tergite 
are nearly as on cephalothorax, decrease gradually in size 
posteriorly, so that they are distinctly smaller and besides 
more dense on the posterior tergites. On the ventral side the 
first furrow is moderately curved (fig. 2 b) and terminates 
without being produced forwards inside the spiracle; the anal 
operculum is somewhat vaulted, and seen partly from behind 
broader than long; the whole ventral surface is densely and 
finely granulated. 

Antennaf rather long. First joint seen in situ from above 
(fig. 2 a) four times as long as broad at the base, and here scar- 
cely as broad as the conus foetidus; it is granulated above 
and on the more proximal part of the lower side. The hand 
of the chela is a little more than five times as long as broad 
(fig. 2 f ) , smooth on the anterior side, and more than three 
times as long as the movable finger. 

Palps (fig. 2 h) about as long as the externally \-isible part 
of the antennae. Femur considerably longer than the trochanter, 
about four and a half times as long as deep, with several gra- 
nules on the lower margin. The tarsal part slightly shorter 
and a little thicker than the tibia. 

Legs (figs. 2 i and 2 k) moderately robust; first and fourth 
pairs a little shorter than the body. Surface of first pair of 
coxa- not lower than that of second pair. Femora of first pair 
with the distal half distinctly curved, so that the lower margin 
is conspicuously concave (fig. 2 i) ; the other femora nearly 
straight. Tibia somewhat longer, but not deeper, than the 
patella, about twice as long as deep. Tarsi of first pair 
(fig. 2 i) a Httle deeper than the tibia, with the lower 
margin rather angular somewhat before the middle, and 
tlie solea occupies about three-fifths of the lower margin of 
the joint. 



^.8 Studies on Arthropoda I. 



The upper side of the body moderately haired; all hairs 
\-ery thin and most of them very short, cylindrical, not acute; 
the lower surface of abdomen has but few hairs, while on the 
coxx the hairs are more numerous and moderately long. The 
legs have a good number of hairs (figs, zi and 2 k), most of 
them normal, acute, but on the proximal joints and especially 
on the femora many of the hairs are shorter and not acute. 
The palps hairy excepting the major proximal ])art of their 
trochanter (fig. 2 h) . 

Sexual differences only moderateh' conspicuous. In the 
female the tarsus of fourth pair is shaped as third pair; in the 
male the fourth tarsus is near the base on the upper side (fig. 2 k) 
produced into a process as long as the depth of the tarsus, 
rather slender, somewhat tapering from the base to beyond 
the middle, with the short distal part (fig. 2 1, ^) marked off 
and oblique-triangular, acute; the process is directed some- 
what upwards and much backwards and is distinctly cur\'ed ; 
the upper surface of the tarsus below the proximal half of the 
process is, seen from the side, somewhat concave. — In the 
male the genital aperture (fig. 2 c) is nearly circular ; a genital 
operculum is not marked off posteriorly, but its front margin 
is almost semicircularly concave; the arculi are somewhat 
narrow, almost three times as long as broad at the base, and 
not nmcli decreasing in breadth from the base forwards. In 
the female (fig. 2 d) the genital aperture is a little smaller than 
in the male and differs somewhat in shape; the genital oper- 
culum {r), which is very short, is marked off by a transverse 
impression, and has its front margin straight, and tliis margin 
is much shorter than the breadth of the aperture; the arculi 
are moderately broad and at the l)ase considerably broader 
than in the male (m), with the inner margin regularly concave. 

Colour. — The body is dark reddish brown, with the 
coxa: and the anterior part of the cephalothorax still darker, 



Pedipalpi, Ricinulei, and Opiliones. ^.Q 



blackish; legs and antenme lighter reddish brown; palps yel- 
low. 

The single male is 2.46 mm long and 1.47 mm broad, ce- 
phalothorax 1.21 mm long, antennae from the base of the trans- 
verse keel 1.58 mm long, palp 1.58 mm, first leg 2.3 mm, fourth 
leg 2.3 mm long. The female is about of the same size; its body 
is 2.48 mm long and 1.41 mm broad. 

Remarks. — Rather similar to species of the genus Siro, 
but proportionateh' broader, with the legs a little shorter and 
conspicuously thicker, and besides differing in the shape of 
the front margin of the cephalothorax and in several important 
features (shape of stomotheca, and of both pairs of maxillary 
lobes, hairs on second antennal joint, etc.). 

Occurrence. — The two specimens were captured in the 
Island of Fernando Poo; the male at Punta Frailes in Oct. — 
Nov. 1901, the female at Basile in Aug. — Sept. 1901 in an 
altitude of 400 — 600 m. 



Explanation of the Plates. 
Plate I. 

Fig. I. Charinus africanus n. sp. 
Fig. I a. Left palp, from the lower, outer side; X 8. 

— lb. Left tibia, from the upper side; X 8. 

— I c. Left tarsus, from the outer side; X 8. — n. upper 

margin. 

Fig. 2. Paracharon caecns n. gen., n. sp. 
Fig. 2 a. Cephalothorax with antennae and left palp, from 
above; x 8. 

— 2 b. I,eft palp, from the lower outer side; X 13. 

4 



^O studies on Arthropoda. I. 



Fig. 2 c. Left tibia, from the upper side; x 13. 

— 2d. Left tarsus, from the outer side; X 13. — tt. upper 

margin. 

— 2 e. Distal joints of left fourth leg, from above; X ^^/g. 

Fig. 3. Trithyreus parvus n. sp. 
Fig. 3 a. Left palp and first leg of the male, from the outer 
side; x 20. 

— 3 b. Left palp of the male, from the outer side; X 35. 

— 3 c. Left palp af the female, from the outer side; X 35. 

— 3d. Left first foot of the male, from the side; x 47. 

— 3 e. Left first foot of the female, from the side; X 47. 

— 3 f . Left fourth leg of the male, from the side ; x 20. 

— 3 g. Left fourth leg of the female, from the side ; x 20. 

— 3 h. The three posterior abdominal segments with fiagel- 

lum of a male, from above; X 60. 

— 3 i. The same segments with fiagellum of a male, from 

the side; x 60. 

— 3 k. Posterior abdominal segments with flagellum of a 

female, from the side; x 60. 

Fig. 4. Trithyreus brevicauda n. sp. (continued). 
Fig. 4 a. Foot of first left leg of the female, from the side ; X 46. 

— 4 b. Left fourth leg of the male, from the side; X 20. 

— 4 c. Major part of left fourth leg of the female, from the 

side; x 20. 

— 4 d. Posterior abdominal segment witli fiagellum of a 

female, from the side; X 62. 

Plate IL 

Fig. I. Trithyreus brevicauda n. sp. (continued). 
Fig. I a. Left palp and first leg of the male, from tlie outer 
side; x 20. 



Pedipalpi, Riciiiulei, and Opiliones. 5 1 

Fig. I b. Left palp of the male, from the outer side; X 43. 

— I c. Left palp and first leg of a female, from the outer 

side; x 20. 

— id. Left palp of a female, from the outer side; 

X 43- 

— I e. Foot of first leg of the male, from the outer side; 

X 46. 

— if. Posterior abdominal segments with flagellum of a 

male, from above; X 62. 

— I g. Posterior abdominal segments with flagellum of a 

male, from the side; X 62. 

Fig. 2. Cryptostemma crassipalpe Hans. & Sor. * 

Fig. 2 a. Cephalothorax of an adult female, from above; X 12. 

— 2 b. Cucullus of an adult female, from in front; X 16. 

— 2 c. Left mandibular palp of an adult female, from in 

front; X 22. 

— 2d. The two anterior right legs of the adult male, from 

above; x 9. — Hairs omitted. 

— 2 e. The two anterior right legs of the adult female, from 

above; x 9. — Hairs omitted. 

— 2 f. Major distal part of first right leg of the female, from 

above; X 21. 

— 2 g. Distal part of third left leg of an adult male, from 

in front; X 25. — a. terminal portion of the tibia; 
b. metatarsus; c. its movable process; d. first tarsal 
joint; d^. first joint of its movable process; d'^. se- 
cond joint of the same process; e. second tarsal joint; 
e^. front margin of its distal lower wall of its 
'lamina cyathiformis' ; e-. posterior high wall of 
the 'lamina cyatluformis' ; /. third tarsal joint. 

— 2 h. Tarsal movable process of the organ shown in fig. 2g, 

essentially from below; X 58. 



52 studies on Arthropod a. I. 

Fig. 3. Cryptostemma FecB n. sp. 
Fig. 3 a. Cucullus of a male, from in front; X 15. 

— 3 b. Left antenna, from behind (from below) ; x 23. 

— 3 c. Portion of the front side of the right first femur of 

a female; X 65. 

Plate III. 

Fig. I. Cryptostemma Fece n. sp. (continued). 
Fig. I a. Left mandibular palp of an adult male, from in front ; 
X 20. 

— lb. First and second right leg of an adult male, from 

above; X ^^/g. 
— =- I c. First and second right leg of an adult female, from 
above; x ^^/2. — Granules and hairs omitted. 

— id. Major part of first right leg of the same adult female, 

from above; X 15. 

— I e. Tarsal movable process of the copulatory organ in 

third left leg of an adult male, from in front; X 42. 

— if. Distal half of the same mo\-able process, from the 

outer side; X 47. 

— I g. Body with left palp and the more proximal joints 

of all three pairs of legs of a larva, from below; X 15. 

— I h. Metatarsus and tarsus of second right leg of a larva, 

from above; x 21. — s. sensory seta. 

— 1 i. Metatarsus and tarsus of third leg of a larva, from 

above; x 21. 

Fig. 2. Dacnopilio insidaris n. sj). 
Fig. 2 a. Body with left antenna and palp of a male, from 
the side; X "/a- 

— 2 b. Ocular protuberance of the male, from the side; X 24. 

— 2 c. Left antenna of the male, from the outer side; 

-^ /2- 



Pedipalpi. Riciniilei, and Opiliones. 53 

Fig. 2 d. Femur of first right leg of the same male, from in 

front; X ^'/a- 

Fig. 3. Megistohumis longipes n. gen., n. sp. 
Fig. 3 a. Body with left antenna, palp, and the proximal 
parts of the legs of an adult male, from the side; 
X scarcel}^ 8. 

— 3 b. Ocular protuberance of the same male, from the 

side; x 13. 

— 3 c. Left palp of the same male, from the outer side; 

X 13. 

— 3d. Patella, tibia and proximal part of tarsus of the same 

palp, from above; X 16. 

Fig. 4. Ogovia nasuta n. sp. 
Fig. 4 a. Left antenna of a male, from the outer side; X 24. 

— 4 b. Left mandible with palp of a male, from the outer 

side; X 24. 

— 4 c. First left leg of a male, from the outer side; X 23. 

— X. vaulted, smooth and shining circular areas. 

Plate IV. 

Fig. I. Ogovia nasuta n. sp. (continued). 
Fig. I a. Cephalothorax and the two anterior abdominal seg- 
ments of a male, from above; x 16. The plate from 
the front end covers the major part of first joint of the 
antennae; of the palps only trochanter and femur, 
of the three following pairs of legs the protruding 
ends of the coxae and the proximal part of the 
trochanter are drawn; of fourth legs only the end 
of the coxae is rendered. 

— lb. Body with coxse and trochanteres of all walking legs 

of a male, from below; X 14. Antennae and man- 



<4 studies on Arthropoda. I. 



dibles with palps removed; granulation and hairs 
omitted. — x. vaulted, smooth and shining circular 
areas. 
Fig. I c. Middle portion of the anterior half of the body of 
the same male, from below; X 23. — /. angle limit- 
ing the stomotheca in front; //. soft portion of the 
maxillary lobe of lirst coxa; m. genital arculi; p. 
process originating a little behind the genital aper- 
ture. 

— id. Middle anterior portion of the body of a female ; 

from below; X 26. 

— I e. Left fourth leg of a male, from the outer side; X 23. 

— X. vaulted, smooth and shining circular areas. 

— if. Ventral side of an immature specimen; x 15. — m. 

genital arculi. 

Fig. 2. Parogovia sironoides n. gen., n. sp. 
Fig. 2 a. Body with the proximal joints of the appendages 
of a male, from above; x 10. 

— 2b. Body of the same specimen, from below; x 16. — 

Granulation and hairs omitted. 

— 2 c. Middle portion of the anterior half of the body of 

the same male, from below; x 32. — /. angle limit- 
ing the stomotheca in front; h. soft part of the 
maxillan.' lobe of first coxa divided into two portions 
somewhat removed from one another; m. genital 
arculi. 

— 2d. Middle portion of the anterior half of the body of 

a female, from below; X 32. — r. genital operculum. 

— 2 e. Left antenna of a male, from the outer side; X 32. 

— 2 f . Second and third joints of the same antenna, obliquely 

from the upper and outer side; X 36. 

— 2 g. Distal portion of the same antenna; X lOtS. 



Pedipalpi, Ricinulei, and Opiliones. 55 

Fig. 2 h. I/cft palp of a male, from the outer side; X 32. 

— 2 i. First left leg of a male, from the outer side ; x 27. 

— 2 k. Fourth left leg of a male, from the outer side; X 27. 

— 2 1. Upper part of the tarsus of the same fourth leg more 

highl^^ magnified. — s. proximal, and f. terminal 
part of the dorsal process. 



ON STRIDULATION 
IN CRUSTACEA DECAPODA 

I. Introductory Remarks. 

It is a well-known fact that stridulating organs exist in a 
number of species belonging to genera of very different families 
of Crustacea Decapoda; most of our knowledge on this topic 
down to the year 1899 has been put together by Ortmann in 
Bronn: Klassen und Ordnungen des Thierreichs, Bd. V, II. 
Abth., Crust. 2, p. 1245. Structural features interpreted as 
stridulating organs have been discovered in three species of 
PencBOpsis (de Man, 191 1) of the family Penseidae; in several 
species of some genera of the Palinuridse (Ortmann, 1. c.) ; in 
Thalassina anomala Herbst (Pearse, 191 1) of the family Thalas- 
sinida?; in Coenobita ritgosa H. M.-Edw. (Hilgendorf, 1869) of 
the family Coenobitidse ; in Clibanarius strigimanus White (Hen- 
derson, 1888) of the family Paguridse, and in a number of crabs. 
Among the tribe Oxj'stomata such organs are known only in 
the species of the genus Matida (Hilgendorf, 1869; Ortmann, 
I.e.); in the tribe Oxyrrhyncha they are unknown ; in the old 
tribe Cyclometopa they exist in Pscudozius Edwardsii Barr. 
(Barrois, 1888) of the family Xanthidae, in Ovalipes bipustulatus 
H. M.-Edw. (Wood-Mason, 1878) of the family Portunidse, and 
in a few African species of the genus Potamon (Caiman, 1908) 
of the family Potamonidse. In the tribe Catometopa organs are 
found in all species excepting one of the genus Ocypoda (various 
authors), in some species of MacrophtJmlmiis, in Helice and 
Metaplax (Hilgendorf, de Man, Ortmann), all l)elonging to the 



StricUilation in Crustacea Decapoda. 1. ^'J 

family Ocypodidae; finally in some species of Sesarma and Hete- 
rograpsus (Hilgendorf, de Man, Ortmann) of the family Grapsidae. 
In most cases it is eas}' to recognize a stridulating organ. 
Excepting in the genus Potamon and perhaps in Thalassina 
(see later on) it consists always of a very regular row of granules 
or small tubercles or a file-like series of ridges, playing the role 
of a fiddle, while the role of the bow is performed b}^ a ridge 
or a small, regular row of granules or very short ridges, or by a 
sharjD margin. In some cases, as in Ocypoda, the fiddle is moved 
to and fro against the ridge-shaped bow; in other cases, f. inst. 
in Penceopsis, the sharp margin acting as bow is moved forwards 
and backwards against a ridged area. The structure in Potamon 
is completely different: a patch of modified spines on the coxa 
of each of the two anterior pairs of thoracic legs is moved 
against ver>' modified spines on thickened lobes from the lower 
margin of the carapace (Caiman, Ann. Mag. Nat. Hist., ser. 8„ 
vol. I, 1908, p. 469); the structure in Thalassina is unknown to 
me. — The sound has been heard in living animals of at least 
two species of Palinurus (several observ^ers), some species of 
Ocypoda (Hilgendorf, Ortmann, Anderson, Alcock), Thalassina 
anomala (Pearse)^), and MattUa (Ortmann). But even if living 
animals have not been obser\'ed, the organ is easily interpreted 
when both the fiddle and the bow are well developed, or, as is 
the case in Penceopsis, the fiddle is so characteristic that no 
doubt remains. But in some cases, f. inst. in Coenobita mgosa,. 
the interpretation of the structures in question is less certain, 
as either the fiddle or the bow is not developed in such a way, 
that it is possible to decide with absolute certainty whether the 



^) Pearse's treatise scarcely exists in Copenhagen. But according to Cai- 
man's "record" the animal stridulates, and as I have been unable to dis- 
cover the organ in question it may be supposed that it is either aber- 
rant in structure or perhaps to be found in some unexpected or hidden 
place, consequently that Pearse who studied the habits of the animal, 
heard the stridulation. 



58 studies on Arthropoda. 1. 

parts are used by the living animals as a stridulating 
organ. 

Many years ago I discovered casually a highly developed 
stridulating organ in a species of Ovalipcs (family Portunidse), 
and afterwards a fine organ in the genus Acanthocarpus , belong- 
ing to the family Calappidse in the tribe Oxystomata. The aim of 
the present small treatise is not only to describe these new organs, 
but to mention the structure in forms of the genus Ocypoda, to 
consider the curious occurrence of organs both in terrestrial 
forms, in amphibious Decapoda and in real marine species, 
two among them inhabiting even moderately deep water, 
finally from comparison with other classes of animals to point 
out analogies and the need of further biological observ^ations 
of living Decapods. 

II. Organs in the genera Ovalipes and Acanthocarpus. 

I. Ovalipes {Platyofiichus) ocellatus Herbst. — On the lower 
side of each half of the carapace and considerably from the 
antero-lateral margin is seen a long, narrow, slightly raised area, 
which is parallel with the direction of that margin and con- 
sequently arcuate; it begins somewhat from third maxillipeds 
and reaches to outside the insertion of the cheliped. This area 
has a large number of rather coarse, transverse ridges completely 
similar to each other in breadth and extremely regularly arranged, 
radiating in the direction of the postero-exterior angle of the 
coxa of the cheliped. The area is broadest near the middle and 
tapers towards both ends and especially towards the anterior 
end; consequently the ridges towards this end are gradually 
shorter, and those near the end only oblong, small tubercles. 
The part acting as bow is found on the merus of the chelipeds, 
the upper side of which has proximalh* in front an oblong area 
with ver>' regular oblique ridges — in a large female to hand 
■8 ridges — and these terminate as slight protul)erances on the 



Stridulation in Crustacea Decapoda. I. 59 



front margin. When the chehpeds are moved forwards and 
backwards a moderatel}' strong sound is produced. 

(More than fort^^ years ago Wood-Mason pointed out a 
special feature in another species of the genus Ovalipes {Platyo- 
nichus), viz. 0. bipustulatus M.-'Edw., and interpreted it as a 
stridulating organ. The lower surface of both chelse have a good 
number of transverse, serrated ridges, but these are not fully 
as regular as usual in a stridulating organ, and I have been un- 
able to point out with certainty any part which can act as bow ; 
in this case observation of living animals are necessary for 
settling the question.) 

2. Acanthocarpus Alexandri Stimps. — Both chelae are 
similar; each is large, ver}' high, with the inner side somewhat 
flat ; on this smooth surface is seen a somewhat curved, vertical, 
distinctly raised area somewhat before the insertion of the 
movable finger. The area reaches about to the upper margin of 
the chela and occupies scarcely three-fourths of its height; its 
upper half is somewhat broad, but it tapers rapidly above and 
slowly from the middle downwards; the area is adorned with a 
high number of very fine and very closely set ridges which are 
ver\' obhque on the long median line of the area, and the ridges 
are equally close in its whole length. On the lower side of the 
orbit and further outwards somewhat below and a little behind 
the front margin of the carapace runs an oblique, thick keel, 
which is broadest at its middle, narrower towards both ends 
and especially outwards ; each keel has 8 or 9 transverse, rounded, 
rather thick ridges arranged v^ry regularly, and each interval 
is about twice as broad as one of the ridges. When the chela is 
moved forwards-upwards and in the opposite direction below 
the front part of the carapace, it is rubbed against the ridged 
keel described, and produces a rather high sound. — This 
species has been taken in West Indies and somewhat more 
northwards in depths from 71 to 150 fath. ; one of the 



*<fcr«^-V- 



/:st, 






6o Stiuli-s oil Arthropoda. I. 



specimens in the Copenhagen Museum is from near 200 
fathoms. 

Another species of the same genus is A . bispi)iosus A. M.-lvdw.^ 
which is described and figured by A. Milne-Edwards and Bouvier 
in Memoirs Mus. Comp. Zool. vol. XXVII, no. i, 1902, p. 127; 
the specimen was taken in 140 fathoms. Fig. 6 on pi. XXV in 
that work represents the inner side of a chela and shows a striated 
vertical area not mentioned in the text; this area agrees much 
with that found in A. Alexandfi, and accordingly A. hisfinosiis 
has a stridulating organ completely as developed as in the other 
species. 

III. Organs and their use in the genus Ocypoda. 

Though the organs found in various species are well known 
and even afford excellent specific characters, it is necessar\' for 
the following considerations to mention two main types. At 
least 14 species of this genus are accepted as \'alid, and a 
stridulating organ is found in all excepting a single form. In 
this genus the part acting as fiddle is always found in both 
sexes on the inner side of the large chela, and that area is rubbed 
against a ridge on the lower side of the ischium of the same leg. 
In the American O. quadrata F. a robust, subvertical keel is 
found somewhat before the fingers, and this keel has a rather 
low number of coarse and regularly but not closel}' set granules; 
0. platytarsis H. M.-Kdw. from India shows a rather similar 
structure. In the Asiatic O. macroccra H. M.-Edw. the keel is 
somewhat low, flattened above, with the lower half of its surface 
moderately broad, the upper half tapering to the end. Near 
the upper end this area has rather coarse ridges moderately 
distant from each other; from tlic upper end downwards to 
beyond the middle the ridges become gradually finer and more 
closely set, and on the lower third they are very fine, and the 
impressions between them extremely narrow. In the connnon 



Stridulation iu Crustacea Decapoda. I. 6l 



Indo-Australian 0. ceratophthalma Pall, the area differs some- 
what from that in 0. macroccra ; the area tapers upwards from 
a little below the middle, and nearly these three-fifths have 
somewhat coarse ridges moderatel}- remote from each other, 
while further below the ridges are suddenly ver\' fine, all 
equally fine and close together. The area can therefore 
be compared with a fiddle bearing two chords; 
when its major upper part is moved against the 
ridge on ischium the tone produced is deep; when 
only its lower part is used the tone is very much 
higher. — 0. macroccra can even produce three tones in using 
respectively the upper, the middle or the lower part of the area. 
On these musical possibilities see later on. 

The species of the genus Ocypoda are not marine, but un- 
fortunately our knowledge of their biology, especially of their 
musical faculties, is imperfect. The best account of a species 
is written by Alcock in "Administration Report of the marine 
Survey of India for the official year 1891 — 92"; he observed 
0. macrocera, and the part of his account (p. 16) necessary for 
my purpose shall be reprinted here. "They are gregarious, and 
though at times they ma}- be seen marching (migrating?) in 
battalions across the sand, the}' usually live in "warrens" at and 
above high- water mark, where they excavate tortuous burrows 
.... and when alarmed flying each one to its burrow". "The 
sounds can be heard, and their effect seen, by forcing one crab, 
which we will call the intruder, into the burrow of another, 
which we will call the rightful owner. The intruder shows the 
strongest reluctance to enter, and will take all the risks of open 
flight, rather than do so, and, when forced in, he keeps as near 
the mouth of the borrow as possible. When the rightful owner 
discovers the intruder he utters a few broken tones of remon- 
strance, on hearing which the intruder, if permitted, will at 
once leave the burrow. If the intruder be prevented from making 



62 Studies on Arthropoda. I. 

his escape, the low and broken tones of the rightful owner 
gradually rise in loudness and shrillness and frequency until 
they become a continuous low-pitched whirr, or high-pitched 
growl, the burrow acting as a resonator". In a paper published 
in 1900 the same author says: "One almost certain use of the 
stridulating-organ is to give warning to intending trespassers, 
of its own species". 

On 0. ceratophthalma Anderson wrote (Journ. Asiatic Soc. 
Beng. vol. LXIII, 1894, p. 138) : "A loud croaking noise .... At 
first I imagined it must be caused by frogs, so perfectly did it 
resemble the croaking of these animals .... it proceeded from 
the burrows of the Ocypoda crab which here fringed the beach 
at high-water mark. The burrows are frequently, in coral sand, 
very wide at their mouth (6 to 8 ins.) and then taper gradually 
downwards, so that they act as excellent resonators. The cause 
of the stridulation of the crabs was by no means apparent, the 
animals were all lying hidden in their burrows, and several 
were croaking at the same time, as if in concert". — Ortmann 
visited a place inhabited b}' the same species; he says (1. c.) 
that all specimens were in their burrows, and the sound produced 
was "a deep bass-tone". 

It is easily seen that the curious structure of the fiddle in 
0. macrocera and O. ceratophthalma is not explained by the 
obser\'ations quoted. 

IV. On Stridulation. 

Among the above-named stridulating Decapoda the species 
belonging to the family Palinurida? and the genera Pcnceopsis 
and Acanthocarpus are purely marine animals. Ocypoda is ter- 
restrial. Ovalipes ocellatus is amphibious, "not unfrequently taken 
at the surface of the sea", and "perfectly at home among the 
loose sands at low-water mark, and also abundant on sandy 
bottoms off shore" ; "when living at low- water mark on the sand 



Stridulation in Crustacea Decapoda. 63 

beaches it generally buries itself up to its eyes" (Stebbing, 1893, 
from Verrill and Smith) ; Matuta seems to live under rather 
similar conditions. The other forms, possessing either decidedly 
stridulating organs or features interpreted as such are either 
terrestrial [Coenobita rugosa) or amphibious or inhabiting 
shallow water at the shore. 

Stridulating organs of types built according to the same 
principles as in terrestrial or amphibious Crustacea are known 
in many Insects and in some Aranese, but it is interesting that 
the organs in really marine Decapoda are in principle similar 
to those in terrestrial forms. {Potamon and Thalassina are left 
out of consideration). The first thing to be noted is that the 
organs in Decapoda exist in both sexes; scarcely any species 
has been obser\'ed, in which only the male possesses an organ 
absent in the female; only in Penceopsis there seems to be some 
insignificant sexual difference in the size of the stridulating 
area (De Man, 1911). 

With good reason Caiman writes (1909) : "In the case of 
purely aquatic species, the function of these organs is less easy 
to understand". But analogies can be pointed out. A good 
number of fishes of very different families are known to produce 
sounds ; in many forms the sound is produced by the air-bladder 
specially adapted for that purpose, in others by the first strong 
ray in the pectoral fins, or the anterior rays in the first dorsal 
fin, or the first ray in the ventral fins, or by tlie praeoperculum. 
( Readers taking interest in this matter may be referred to the 
excellent work by Dr. William Sorensen: Om Lydorganer hos 
Fiske. 1884. Sorensen describes and explains the instruments, 
but says nothing on their biological use) . A more close analogy 
is Corixa, a genus of Hemiptera, of which several species 
inhabit fresh water in Europe, and they stridulate ver>^ 
well; Pelohiiis Hennanni of the family Dytiscidai may also 
be noted. 



64 Studies on Arthropod a. I. 



Stridulating organs built according to the same principle 
as in Penceopsis, Acanthocarpus, Palinuridcp, etc. are known in 
males of European Aranese as Asagam and Steatoda and in nu- 
merous Insects, thus in both sexes of Necrophoridae, in most 
Cerambycidae, in the genus Crioccris (fam. Chrysomelidse) , in 
several genera of lyamellicornia and in the larvae of at least most 
types of that big family (the Lucanidae included), in some other 
Coleoptera, in the Reduviidae (Hemiptera), etc. (The organs in 
Cicadidae, lyocustidae etc. are omitted here) . In all the terrestrial 
Arthropoda enumerated we find an .area, or two areas, with 
regularly arranged and generally fine to extremely fine, trans- 
verse ridges, and a sharp margin, a row of minute tubercles, 
or the end of rostrum acts as bow. We generally know little 
and frequently nothing on the biological role of the organ in 
most of these Insects, but it must be emphasized that in every 
form known to me the ridges on a stridulating are sim- 
ilar as to thickness and distance; consequently one 
might suppose that only a single tone could be produced, and 
that modulation must be rather limited. But that is in reality 
not always the case; for the best proof of the opposite, and the 
most interesting obser\^ation on the use of stridulating organs 
of the normal structure we are indebted to J. C. Scliiodte (Naturh. 
Tidsskrift, 3. Raekke, B. VII, 1870, p. 188). Schiodte experi- 
mented with specimens of Nccrophorus, especially A^ vespillo, 
and writes: "When listening to a flower-pot in which several 
specimens of Nccrophorus are about letting down a carcass [of 
a small mammal or bird], one hears distinctly that they during 
the work underground communicate mutually by the language 
of stridulation. The sound is now higher and hasty, now deeper 
and smoothered, and on the whole modulated in tlie most 
manifold way. Sometimes a pause is suddenly interrupted by 
a single sharp tone instantly answered by a similar one from 
another specimen or simultaneously from se\'eral specimens. 



Stridiilatioii iu Crustacea Decapoda. 65 



and one gets the impression of the different moods agitating 
the animals during the pains of the work: eagerness, impatience, 
fretfuhiess, weariness, exultation; and sometimes the whole 
company makes a kind of chorus-cry for the measured regula- 
tion of the work". 

And let us now turn to the stridulating organs in such forms 
as the terrestrial crabs Ocypoda ceratophthalma and 0. macrocera 
described above. It is instantly seen that they are more 
highly developed than, for instance, those inNecropho- 
rus, because they in the first-named species can produce two 
\-er\- different tones, in the latter form even three tones. Whether 
the animals really use this faculty is unknown (see p. 62), but 
it may be supposed that they occasionally do so. And the topic 
is, in my opinion, very interesting. But well planned — and, 
for the rest, probably somewhat difficult — observations of 
stridulating Decapoda and above all of species of Ocypoda with 
highly developed organs are needed ; I even hope that this little 
treatise may create interest in some Zoologist, who has an oppor- 
tunity for biological obser\'ations in the field or perhaps in a 
good "terrarium". 



ON THE POSTEMBRYONIC OCCURRENCE 

OF THE MEDIAN "DORSAL ORGAN" IN 

CRUSTACEA MALACOSTRACA 

I. Introductory Remarks. 

In a paper on Sergcstcs (Proc. Zool. Soc. I^ondon 1896) the 
present author wrote in the description of the youngest Masti- 
io'o/)ws-stage of S. arcticus Kr. : "just is front of the gastro-hepatic 
groove is observed a short protuberance in the median Hue". 
— In his valuable paper: Zur Kenntnis der Metamorphose 
von Sergcstes arcticus Kr. (Zool. Anz. Bd. XXXIII, 1908) 
E. Wasserloos writes (p. Ji8) in the description of the second 
Protozoea-stage : "In der Mittellinie des Cephalothorax bemerkt 
man iiV)er dem Gehirn und dem Naupliusauge eine linsenartige 
Her\'orwolbung des Chitins .... die bisher bei keiner Sefgcstes- 
Larve ausser bei einigen Mastigopen \-on Sergcstes arcticus von 

Hansen beschrieben worden ist An Schnitten habe icli 

ausser der Chitinbucht und der darunter liegenden, allerdings 

undeuthche Matrix nichts wahrgenommen Bine genaue Be- 

schreibung und genaue Angaben iiber die erwahnte Hervorstiil- 
jjung kann ich nicht geben, doch mochte ich eine Vermutung 
aussprechen: Die Lage des Organs iiber dem Naupliusauge und 
der Um.stand, dass es mit der Zuriickbildung des Naupliusauges 
ebenfalls verschwindet, lassen es als wahrscheinlich erkennen, 
dass die Protuberanz als Sammellinse fitr das Naupliusauge 
dient". His suggestion on the function of the i)r()tul)erance is 
certainly erroneous. 

While working out the rich collection of Sergestidie collected 
1)y tlie Prince of Monaco, I obser\ed the dorsal ])rotuberance in 



The "dorsal or^an" in Crustacea Malacostraca. 67 



Acanthosoma-stages of five species and in young Mastigopus- 
specimens of several species, furthermore a rudiment of the same 
organ in adult specimens. The idea struck me that it must be 
the so-called "dorsal organ" known in embryos of Crustacea 
of most orders, but unknown in almost all adult Malacostraca 
and in larvae of the same sub-class. In examining adults of a 
good number of species belonging to all orders of Malacostraca, 
I was surprised that the dorsal organ, though frequently looking 
only as a rudiment, exists in some or several representatives of 
most orders. It is found on the surface most frequently nearly 
vertically above the mandibles or their base. 

II. On our earlier Knowledge. 

Nearh- all original observations on the dorsal organ are 
scattered in papers very different in contents, though the 
majority deal with embryolog}^ or development. In 1904 W. T. 
Caiman's excellent hand-book on Crustacea was published; in 
1913 W. Giesbrecht's part on Crustacea in Lang's Handbuch 
der Morphologic was edited. These careful authors had written 
"records" on the Carcinology during a number of years and 
were uncommonly well versed in the vast literature; the state- 
ments of both authors together in the books mentioned on 
the occurrence of the dorsal organ may therefore be considered 
nearly sufficient, and an extract is given here. 

On Anaspides (the order Anaspidacea) Caiman writes (p. 
164) : "On the dorsal surface [of the head], in front of the cervical 
groove, is a pigmented area with a circular central spot sur- 
rounded by four minute pits. The significance of this structure 
is quite unknown, but it may be comparable to an obscure 
"dorsal organ" apparently glandular in nature, occupying a 
similar position in certain other Malacostraca". It may be point- 
ed out, that Caiman in his treatment of the other orders of 
Malacostraca does not mention the occurrence of a "dorsal 

5* 



68 Studies on Arthropoda. I. 

organ" in the adult of any other form, but possibly he refers 
in the lines quoted to the organ found in embryos of other 
orders. — On the Tanaidacea he says (p. 194) : "A paired "dorsal 
organ" is present" in the embryo; on p. 213: "A "dorsal organ" 
is present in many Isopod embrj'os, and assumes very different 
forms", which are mentioned. On the Amphipoda (p. 237) : 
"A "dorsal organ" is early developed as a median thickening 
of the ectoderm . . . . " — On the embryo in Mysidse (p. 179 — 80) : 
"A pair of lateral thickenings of the epiblast appear \-ery earl)' 
and, approaching each other on the dorsal side, fuse to form an 
invaginated "dorsal organ"." But I may remark that Nusbaum 
und W. Schreiber in 1898 (Biolog. Centralbl. Bd. 18, p. 742) 
figured a transverse section of an embryo of My sis Lamorna' 
showing the unpaired dorsal organ and a pair of dorso-lateral 
organs, and all three organs have already begun to degenerate. 
— At the Nebaliacea, Cumacea, Euphausiacea, Decapoda, and 
Stomatopoda Caiman does not mention any dorsal organ. 

Giesbrecht writes (p. 156 — 57) that not only in Cladocera 
but in "vielen anderen Crustacea ein anderes Organ von meist 
driisiger Struktur, ebenfalls in der Dorsalwand des Kopfes, das 
aber nur transitorisch (bei lunbryonen, seltener Larven) auftritt 
und sich meistens schon friili zuriickbildet ; das ist die Nacken- 
driise (Nackenorgan, Nackenschild, Dorsalorgan . . . .)". Among 
the Malacostraca it is found early in the embryo in Arthro- 
straca (consequently Tanaidacea, Isopoda, and Amphipoda), 
in Cumacea, Mysidacea, perhaps also some Decapoda; "bei Lep- 
tostraken scheint es eine unpaarige Leiste zu bilden"; finally 
"als Nackendriise aufgefasst wird auch .... eine dorsale 
Driisenscheibe in the Maxillargegend von jungen und adulten 
ICuphausien etc.". I have been unable to find in the special 
literature on the Euphausiacea any statement on the organ. 

In a small paper published in Jouni. Linn. Soc. Lond. vol. 
XXIX (1903) the ]jre.sent writer ])ointed out an organ on the 



The "dorsal organ" in Crustacea Malacostraca. 69 



surface of the head near the posterior margin in the Isopod 
Bathynomus giganteus; the organ — which shall be mentioned later 
on — was briefly described, figured and compared with the organ 
discovered by Caiman (1896) in Anaspides tasmanice; it was 
added, that I had found a vestige of the organ in the allied form 
Cirolana borealis. — The statements in the literature on the 
organ in larv'se of Sergestes arcticus are quoted above. 

Let us sum up. The median dorsal organ has been found 
in embryos of representatives of the following orders of Mala- 
costraca: Nebaliacea, Mysidacea, Cumacea, Tanaidacea, Isopoda, 
.\mphipoda and perhaps some Decapoda. The Embryologists 
tell us that it, at least generally, is of glandular nature and dis- 
appears early; on its function nothing is known, besides it is 
unknown in the embryo of Stomatopoda. — In adult forms it 
has been found in Anaspides of the order Anaspidacea; in Ba- 
thvnomiis and a species of Cirolana of the order Isopoda; finally, 
according to the quotation from Giesbrecht, in young and adult 
forms of the order Euphausiacea. In larval stages it is only 
known in Sergestes arcticus. 

III. The "dorsal organ" in adult Malacostraca 
and some larvae. 

It has not been my intention to inspect every genus avail- 
able, but types of families or genera have been selected and 
examined with a good pocket-lens ; it was found that within the 
same family, sometimes even within the same genus, an organ 
could not infrequently be discovered in a large species, but not 
with any certainty in small forms. The following enumeration 
shows that the organ has a much wider occurrence than might 
have been supposed. 

I. Nebaliacea. 

Nehalia bipes O. Vabi. — In turning the animal a little to 
and fro so that the light changes on the smooth, shining dorsal 



JO .studies on Arthropoda. I. 

surface of the head it is generally possible to perceive a faint 
vestige of a median protuberance situated not inconsiderabh- 
behind the base of the movable rostral plate. 

2. Anaspidacea. 

Anaspides tasmanice G. Thonis. — Caiman described and 
figured a very conspicuous dorsal organ (Trans. Roy. Soc. 
Edinburgh, XXXVIII, 1896). Nothing is to be added. 

Paranaspides lacustris Geoff. Smith. — The organ is more- 
reduced than in Anaspides, as there is no pigmentation; the ver\ 
small pits are irregularly arranged, and their number in three 
specimens varies from 3 to 5. 

3. Mysidacea. 

Gnathophausia. — In a gigantic specimen of G. ingcus Uohrn 
a part of the dorsal keel a little in front of its base and about 
15 mm. long is flattened above and behind the middle distinctly 
thickened, this narrow area has a pair of very conspicuous, 
very oblique, convex spots marked off all around b>- an de- 
pression, and the spots look almost as scars. In G. zuea Will.- 
Suhni about one mm. of the dorsal keel above the mandibles 
is, seen from above, rather considerably thickened but not flat ; 
seen from the side with the light transmitted the dorsal chitinc 
of this part is thin and the contents opaque, indicating that the 
tissue is of another quality than in the surroundings. 

Eucopia. — In one specimen of E. sculpticauda h'axon a 
nearly circular, somewhat shallow depression situated above 
the mandibles somewhat in front of the anterior transverse 
furrow; the depression has a minute, dark point as if a scar 
after a ])rick with a needle. In another specimen the depression 
is wanting, but the dark point exists; in a third specimen noth- 
ing could be seen with certaintw In specimens of E. ungui- 
cnlata Will.-Sulnn nothing was found. 



The "(lorsal oryan" in Crustacea Malacostraca. 71 

1_ 

Lophogaster. — In L. typiais M. Sars no vestige is discover- 
able. 

Borcomysis. — B. scyphops G. O. S. has somewhat in front 
of the gastric groove a moderately large and rather shallow 
depression, at the middle of which is seen a somewhat low, a 
little oval, rounded and \-er\' distinct protuberance with one 
to three minute pits. — B. nob His G. O. S. has very near the 
gastric groo\'e a small but rather high, a little oval protuberance 
without visible pits. — B. microps G. O. S. has a little in front 
of the gastric groove a conspicuous, nearly semiglobular, smooth 
protuberance. 

Of the subfamily Mj'sinas two moderately large forms, viz. 
Mysideis grandis Goes and Mvsis mixta Lilljeb., have been 
inspected, but no vestige of any organ was found. 

4. Cumacea. 
Some few larger forms ha^'e been examined, but nothing 
could be found. 

3. Tanaidacea. 
Some large specimens of Apseudes and Alaotanais, thus 
representatives for both families of the order, have been in- 
spected with negative result. 

6. Isopoda. 
Idothea. — In /. balthica Pall, is frequently found near the 
posterior margin of the head a very small area a little broader 
than long, and with a few tiny pits or, in one specimen, with 
impressed transverse lines; the area is generally a little or 
somewhat depressed, but in one specimen it is larger than usual, 
raised and shining. In a well-sized specimen from Iceland the 
area is uncommonly conspicuous, moderately large, very slightly 
depressed, shining, with a large, oblong pit in the middle, and 



72 studies on Arthropoda. I. 



around this 8 or 9 extremely minute, oblong pits. — In /. cmar- 
ginata F. and /. ungulata Pallas organs somewhat similar to 
those in /. halthica are found. 

Among other animals belonging to the sub-order Valvifera 
Mesidothca sibirica Bir., M. Sahini Kr., and Arcturus Baffin i 
Sab. have been inspected, but with negative result. 

Scrolls. — S. paradoxa Fabr. has a very distinct though 
somewhat small, a little oblong, depressed area near the posterior 
margin of the head; the area has several tin}' pits, especially 
near the masgin. — .S. Schythei Ltk. has a similar nearly circular 
area. 

Bathynomiis. — In B. giganteus A. M.-Edw. an organ was 
discovered, briefly described and figured by me in 1903 (see 
p. 68). I have examined a large immature female. Near the 
posterior margin of the head it has a most conspicuous, whitish 
spot, which does not show the numerous, irregularly impressed 
points on the surrounding brownish chitine. The spot is as a 
whole a little arched but depressed along the middle; each half 
has a small group of 6 or 7 minute pits; the median part has a 
x'ery oblong area well marked off by an depressed line and with 
a linear depression in the median line; somewhat behind each 
group of pits is found a shallow excavation which has a number 
of oblique, parallel, impressed striae; at the front end of the area 
is seen a group of some nearh' rudimentary pits. 

Cirolana. — In C. borealis Lilljeb. a \ery small group of three 
to six tiny pits is found near the posterior margin of the head; 
they are sometimes situated in a very shallow or very distinct 
depression. — A specimen of C. Rossii Miers has some irregularly 
scattered minute pits. 

^ga psora L. and specimens of Rocinda, Anilocra and Cy- 
mothoa have been inspected with negative result. 

Among the sulvorder Asellota Asellus aquaticus L., Mun- 
nopsis iypica M. Sars and Muniuypsurus giganteus G. O. Sars, 



The "dorsal organ" in Crustacea Malacostraca. 73 



among the Oniscoidea Ligia oceanica L. and the gigantic Tylos 
granulatiis Krauss have been examined without finding any 
vestige of the dorsal organ. 

7. Amphipoda. 
Large specimens of genera comprising forms of very consider- 
able size, as Aiionyx, Stegocephalus, Gammarus, Maera, have 
been inspected, but the result was absolute negative. 

8. Euphausiacea. 

Thysanopoda. — In T. egregia H. J. H. the gastric keel — 
between the front margin and the gastric groove — has somewhat 
behind the middle a short part raised rather considerably^; this 
part is cut off horizontally so that its upper surface is flattened, 
and this surface shows an aspect different from that of the 
surrounding integument. (The raised part is shown in fig. 21 
in my paper on Schizopoda in Bull. Mus. Ocean. Monaco, no. 
30, 1905, p. 23) — T. cornuta Illig shows a similar structure. — 
In T.cpgualis H. J. H. nothing could be perceived. — T. microph- 
thalma G. O. S. has a feeble vestige on the gastric keel which 
on its highest part is a little flattened above. 

In Bentheuphausia amhlyops G. O. S. a vestige is scarcely 
perceptible. 

In Meganyctiphanes norvegica M. Sars the gastric keel has 
a somewhat high and rather short part thickened upwards with 
the dorsal surface distinctly flattened; seen from the side with 
transmitted light the contents of the raised part is generally 
lighter than the surrounding tissue. 

Euphausia. — In E. superha Dana the median keel is at some 
distance in front of the gastric groove more or less distinctly 
raised and thickened, and at each side of this part the surface 
of the carapace is distinctly excavated, but the keel itself is 
rounded above without vestige of any special structure. 



74 Studies on Arthropoda. I. 



' In E. KroJinii Brandt no \'estige. 

In N ematohrachion boopis Calm, and Ncmatoscelis mcgalops 
G. O. Sars the gastric crest is at its highest point slightly angular 
and shows a vestige of the organ ; seen from the side with trans- 
mitted light the contents of that part of the keel is lighter than 
the surrounding tissue. 

lyarvse. — Some larva? belonging to the genus Euphausia 
have been examined, but it was not possible to discover ain- 
rudiment of a dorsal organ. At a future occasion I will have to 
examine the vast material of larval forms secured together with 
adults in the North Atlantic by the Prince of Monaco, and then 
there will be a good opportunity to look for the dorsal organ in 
representatives of almost all genera. 

q. Decapoda. 
The dorsal organ is found in the adults of most species of the 
Penaeida? inspected by me, and in all genera and species of the 
Sergestidse excepting Lucifer. Furthermore it was detected in 
several genera of the Caridea. Besides it was disco\ered in cer- 
tain larv^al stages of the genus Scrgcstcs and in a few larvae of 
Penseida?. In the following an account shall be given, but as to 
the genus Sergestes a brief abstract may be sufficient, because 
in a monograph of the North-Atlantic species secured by the 
Prince of Monaco and now in composition a more detailed report 
will be embodied. — In vain I have looked for the organ in a 
species of the genus Cambanis, and it is scarcely to be found in 
Astacidse, Palinuridse, Galatheidse, etc. or in the crabs. 

A. Pena'idte. 

Aristeus. — A. Edwardsianus Johns, has a little behind the 

first dorsal spine the median keel thickened with its ui)per 

surface expanded ; ni the middle of this part is found a somewhat 

oblong area well marked off by a depression, this area has 



The "dorsal organ" in Crustacea Malacostraoa. 75 



slightly before the middle an oblong, dark-brown spot sharph' 
defined and with at least one minute pit. — In .4 . scmidentatus 
Bate is found a moderately small but uncommonly deep de- 
pression a little behind the first dorsal spine; the depression has 
a small, circular, somewhat convex area behind the middle. 

Solenocera Agassizii Fax. — A little behind the first dorsal 
spine an oblong, rather depressed area with a small knot at the 
middle. 

Pencens. — P. setijerus L. has considerably behind the first 
dorsal spine a proportionately somewhat long part of the median 
keel thickened and flattened above or even longitudinally a 
little excavated ; the broadest part of that area has in one spec- 
imen an oblong pit, in another specimen nothing. It must be 
remarked that the area in question is situated more backwards 
than in any other genus of the Malacostraca. — In P. caramotc 
Risso nothing could be discovered. 

Larvae. — In a gigantic larva in the Myszs-stage, measuring 
20.5 mm. from the tip of rostrum to the end of telson and cap- 
tured in the Pacific, a small, oblong and rather low protuberance 
is easily seen a little behind the origin of the rostral keel. — 
In a larva in an older Mys/s-stage, belonging to a quite different, 
somewhat small type, an oblong and somewhat high protuber- 
ance is found a little in front of the gastro-hepatic groove; seen 
from the side the protuberance is directed upwards and forwards ; 
the larva is from lat. 4^/3° N., long. 1071/,° E. 

B. Sergestidse. 
Sergestes. — The organ has been found in adult specimens of 
everyone of the 15 species hitherto captured north of equator, 
but it could not be seen in every specimen of some of the smaller 
species, as S. mollis Smith, S. arcticus Kr. , S. atlanticus M. Edw., 
S. vigilax Stimps., and S. Edwardsii Kr. The organ is situated 
a Httle or somewhat in front of the gastro-hepatic groove. It 



76 studies on Arthropoda. I. 



is a small or very small area, in most cases distinctly or consider- 
ably raised as a rounded, nearly circular or transYerse-o\'al 
protuberance, sometimes, but far from always, marked off b}' 
an depressed line; in a few cases the whole area is depressed. 
Frequently the area has a ver>'' distinct granule or raised point 
in front of the middle, in other cases a few tiny pits or no pit 
could be made out. In a few cases the area is brown, but gener- 
ally of the same colour as its surroundings. There is also some 
individual variation in the area as to its height or the depth of 
its surrounding impression. 

Petalidium. — In one specimen of P. obesuni Kr. is found a 
moderately small, a little oblong and feebl}' elevated area with 
a tiny and more raised point in front. In some specimens an 
area was not discoverable with certainty, but the quality of the 
integument makes the investigation difficult and uncertain. 

Sicyonella. — In one specimen of 5. maldivensis Borr. a 
distinct, small group of fi\'e minute pits close behind the dorsal 
crest; in another specimen the group has nearly vanished. 

Acetes. — In one specimen of A. vulgaris H. J. H. a group of 
six minute pits close together and somewhat in front of the 
rudimentary^ gastro-hepatic groove; in another specimen a 
rudimentary^ protuberance with vestiges of a few pits. 

Lucijer. — No trace of any organ. 

Larvae of Sergestes. — As mentioned above (p. 66) Wasser- 
loos discovered the dorsal organ — but did not inter])ret it as 
such — in the second Protozoea-stage and the Acanthosoma- 
stages of 5. arcticus Kr. ; the present writer had found the 
protuberance in question in the youngest Mastigopus of the same 
species. The dorsal organ is an erect or distinctly ])rocurved, 
rounded protuberance in the Atlantic Acanthosoma-stag'^s known 
to me and belonging to .S". arcticus Kr., S". tcniiiremis Kr., .S". ro- 
bustus Smith, .S. c<)r)iicnlitni Kr., and .S". vigilax Stimps. ; the 
protuberance is generally easily seen from llie side. It is also 



The "dorsal organ" in Crustacea Malacostraca. 77 

found in the younger Mastigopus-stages of several and probably 
all species, but it differs sometimes considerably in aspect, as 
in S. pectinatiis Sund it is shaped as a thick, short spine directed 
obliquel}' forwards. In the older lars'al stages it has apparently 
disappeared, but 3'et it exists in all probability, because it is 
found in adult specimens, though its aspect is quite dif- 
ferent; I have not undertaken the certainl}^ difficult inve- 
stigation of the dorsal integument in front of the gastro- 
hepatic groove in older lar\'?e or very young specimens with 
black eyes. 

Larvae of Lucifer. — In the youngest Mys?'s-stage — 
without pleopods — of L. Faxonii Borr. a rather large and 
considerably vaulted dorsal protuberance is found above the 
base of the mandibles ; it is not marked off in any way from the 
surrounding integument, and I am unable to decide whether it 
is homologous with the very characteristic protuberance in the 
Acanthosoma — or Mys/s-stages — of Sergestes. In older stages, 
with pleopods, the protuberance is proportionately^ smaller. 

C. Caridea. 

Acanthephyra muUispina Cout. — A little behind the first 
dorsal spine the median line has a small, circular depression sur- 
rounding a more or less distinct protuberance. 

In Nematocarcinus exilis Bate a very smaU organ close at 
the base of the first dorsal spine. 

In Ephyrina sp. from the North Atlantic the anterior crest 
of the carapace is somewhat from its base flattened above with 
a small but ven,^ distinct, circular, rounded protuberance, the 
surface of which has two or three pits so tiny that they could 
not be counted with certainty. 

Pasiphae. — In P. principalis Sund the dorsal keel is distinctly 
thickened considerably behind the rostrum, but a protuberance 
is wanting and no pit could be discovered. — A young P. sivado 



yS studies on Arthropoda. I. 

Risso about 13 mm. long lias a just perceptible, rudimentary 
protuberance considerably behind the rostrum. 

In Spirontocaris microceros Kr. an area marked off by 
a circular depression very near the base of the first dorsal 
spine. 

In Bythocaris Icucopis G. O. S. a distinct rudiment in the 
keel a little behind the first spine. 

In Alpkeus avams F. a small, but well developed depression 
a little or slightly behind the median keel. 

PalcBmon brasilicnsis Hell., Pandalus Bonnicri Caull. and 
Pontonia sp. have been inspected with negative result. 

10. Stomaiopoda. 

To begin with it may be remarked that moderately large to 
\'ery large specimens of the present order have frequently the 
surface of the median part of the carapace more or less rubbed 
and are consequently badly fit for the investigation, as in such 
cases the dorsal organ is frequently difficult or impossible to 
point out with certainty. 

The dorsal organ has been mentioned in descriptions — and 
besides shown in figures — of some species of the genus Squilla 
by at least two able Zoologists, viz. Stanley Kemp in his excellent 
memoir on the Indo-Pacific forms of the order (1913) and by 
Caiman (1917). They name it the "dorsal pit", which is only 
a descriptive term, as they did not recognize what this pit 
really is. — It may also be pointed out that Giesbrecht in his 
useful, extremely elaborate, long-winded and as to literature 
not always \'ery fair monograph of the Mediterranean Stomato- 
poda (1910) has not observ^ed the "dorsal pit", though it is con- 
spicuous in the common Squilla mantis h., at least sometimes 
not difficult to see in .S". Desniarestii Risso, and ought at least 
to have been indicated on his large figures of the carapace of 
these two species. 



The "dorsal organ" in Crustacea Malacostraoa. 7Q 



Squilla. — In .S. mantis L. the median keel is considerably 
widened and flattened above somewhat behind the bifurcation 
and far in front of the mandible; the widened, rather short part 
has a very oblong depression, the bottom of which is a little 
convex and looks as pricked feebly with a needle. — In 5. 
raphidea F. the organ is conspicuous and nearly as in 5. mantis. 

— In 5. Desmarcstii Risso was found in two of four specimens 
examined a small, circular, low depression with a rudimentary 
median pit; the depression is situated considerably in front of 
the mandibles. — In 5. gibba Nobili the median carina is flattened 
and much widened a little behind the middle and has there a 
small, a little oblong, convex area surrounded by a rather deeply 
depressed ring, and placed a little in front of the mandibles. 

Lysiosquilla. — In a rather large specimen of L. cusebia 
Risso a very small and feeble depression a little in front of the 
middle of the carapace above the mandibles; in another rather 
large specimen and in small specimens nothing could be observed. 

— In L. maculata F. no vestige was found. 

Pseudosquilla ciliata F. — In two specimens a nearly circular, 
\-ery conspicuous and somewhat deep depression with two or 
three tiny pits on the flat or a little convex bottom, situated a 
little in front of the middle of the carapace and somewhat in 
front of the mandibles. In a third specimen the depression is 
small and shallow, in a fourth quite rudimentary'. 

Odontodactylns scyllarus L. — Slightly behind the middle of 
the carapace a rather large, moderately shallow depres- 
sion, in the middle of which a small, circular, somewhat convex 
area. 

Gonodactylus Oerstedii H. J. H. — A deeper or more shallow, 
small, circular, depressed area a little behind the middle of the 
carapace and above the mandibles. In young specimens the 
depression is either considerably more feebly developed or not 
discernible. 



So studies on Arthropoda. I. 

Larvae. — Specimens of Alima, Pseuderichthus and Lvsic- 
richthus, also the first stage, have been inspected, and the result 
was always negative. 

IV. Summary. 

The preceding chapter shows that the "dorsal organ" can 
be observed on the posterior part of the dorsal surface of the 
head or on the carapace in adults of some or several genera of 
all orders of Malacostraca, excepting in Cumacea, Tanaidacea 
and Amphipoda. Whether it sometimes or frequently or even 
generally has a special function must be made out by a student, 
who has a good material either of living animals or of specimens 
especially preserved for histological research, for cutting b>- 
microtome. As the organ has such wide occurrence, it is no very 
probable that it has no function in the adult. And this sup- 
position is strenghtened by the fact that it exists in many and 
probably in most adult Stomatopoda, but seems to be wanting 
in their larvae; that it should exist in the embrj^os of this order 
— on which nothing is known — then be absent in the larvae 
but reappear in the adults, is highly improbable. We are in 
reality completely ignorant on the significance of the dorsal 
organ both in embryos, in larvic and in adults, so ignorant 
that it seems scarcely possible to produce even a hypothesis. 
Perhaps experiments on living animals similar to those carried 
out by some authors in order to investigate the excretor\' 
system might yield some result. 



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studies on Arihropo4iay I. 



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